Inland navigation and land use interact to impact European freshwater biodiversity.

Inland navigation in Europe is proposed to increase in the coming years, being promoted as a low-carbon form of transport. However, we currently lack knowledge on how this would impact biodiversity at large scales and interact with existing stressors. Here we addressed this knowledge gap by analysing fish and macroinvertebrate community time series across large European rivers comprising 19,592 observations from 4,049 sampling sites spanning the past 32 years. We found ship traffic to be associated with biodiversity declines, that is, loss of fish and macroinvertebrate taxonomic richness, diversity and trait richness. Ship traffic was also associated with increases in taxonomic evenness, which, in concert with richness decreases, was attributed to losses in rare taxa. Ship traffic was especially harmful for benthic taxa and those preferring slow flows. These effects often depended on local land use and riparian degradation. In fish, negative impacts of shipping were highest in urban and agricultural landscapes. Regarding navigation infrastructure, the negative impact of channelization on macroinvertebrates was evident only when riparian degradation was also high. Our results demonstrate the risk of increasing inland navigation on freshwater biodiversity. Integrative waterway management accounting for riparian habitats and landscape characteristics could help to mitigate these impacts.

Multi-decadal improvements in the ecological quality of European rivers are not consistently reflected in biodiversity metrics.

Humans impact terrestrial, marine and freshwater ecosystems, yet many broad-scale studies have found no systematic, negative biodiversity changes (for example, decreasing abundance or taxon richness). Here we show that mixed biodiversity responses may arise because community metrics show variable responses to anthropogenic impacts across broad spatial scales. We first quantified temporal trends in anthropogenic impacts for 1,365 riverine invertebrate communities from 23 European countries, based on similarity to least-impacted reference communities. Reference comparisons provide necessary, but often missing, baselines for evaluating whether communities are negatively impacted or have improved (less or more similar, respectively). We then determined whether changing impacts were consistently reflected in metrics of community abundance, taxon richness, evenness and composition. Invertebrate communities improved, that is, became more similar to reference conditions, from 1992 until the 2010s, after which improvements plateaued. Improvements were generally reflected by higher taxon richness, providing evidence that certain community metrics can broadly indicate anthropogenic impacts. However, richness responses were highly variable among sites, and we found no consistent responses in community abundance, evenness or composition. These findings suggest that, without sufficient data and careful metric selection, many common community metrics cannot reliably reflect anthropogenic impacts, helping explain the prevalence of mixed biodiversity trends.

The recovery of European freshwater biodiversity has come to a halt.

Owing to a long history of anthropogenic pressures, freshwater ecosystems are among the most vulnerable to biodiversity loss1. Mitigation measures, including wastewater treatment and hydromorphological restoration, have aimed to improve environmental quality and foster the recovery of freshwater biodiversity2. Here, using 1,816 time series of freshwater invertebrate communities collected across 22 European countries between 1968 and 2020, we quantified temporal trends in taxonomic and functional diversity and their responses to environmental pressures and gradients. We observed overall increases in taxon richness (0.73% per year), functional richness (2.4% per year) and abundance (1.17% per year). However, these increases primarily occurred before the 2010s, and have since plateaued. Freshwater communities downstream of dams, urban areas and cropland were less likely to experience recovery. Communities at sites with faster rates of warming had fewer gains in taxon richness, functional richness and abundance. Although biodiversity gains in the 1990s and 2000s probably reflect the effectiveness of water-quality improvements and restoration projects, the decelerating trajectory in the 2010s suggests that the current measures offer diminishing returns. Given new and persistent pressures on freshwater ecosystems, including emerging pollutants, climate change and the spread of invasive species, we call for additional mitigation to revive the recovery of freshwater biodiversity.

Long-term forecast of thermal mortality with climate warming in riverine amphipods.

Forecasting long-term consequences of global warming requires knowledge on thermal mortality and how heat stress interacts with other environmental stressors on different timescales. Here, we describe a flexible analytical framework to forecast mortality risks by combining laboratory measurements on tolerance and field temperature records. Our framework incorporates physiological acclimation effects, temporal scale differences and the ecological reality of fluctuations in temperature, and other factors such as oxygen. As a proof of concept, we investigated the heat tolerance of amphipods Dikerogammarus villosus and Echinogammarus trichiatus in the river Waal, the Netherlands. These organisms were acclimated to different temperatures and oxygen levels. By integrating experimental data with high-resolution field data, we derived the daily heat mortality probabilities for each species under different oxygen levels, considering current temperatures as well as 1 and 2°C warming scenarios. By expressing heat stress as a mortality probability rather than a upper critical temperature, these can be used to calculate cumulative annual mortality, allowing the scaling up from individuals to populations. Our findings indicate a substantial increase in annual mortality over the coming decades, driven by projected increases in summer temperatures. Thermal acclimation and adequate oxygenation improved heat tolerance and their effects were magnified on longer timescales. Consequently, acclimation effects appear to be more effective than previously recognized and crucial for persistence under current temperatures. However, even in the best-case scenario, mortality of D. villosus is expected to approach 100% by 2100, while E. trichiatus appears to be less vulnerable with mortality increasing to 60%. Similarly, mortality risks vary spatially: In southern, warmer rivers, riverine animals will need to shift from the main channel toward the cooler head waters to avoid thermal mortality. Overall, this framework generates high-resolution forecasts on how rising temperatures, in combination with other environmental stressors such as hypoxia, impact ecological communities.

A 34-year survey under phosphorus decline and warming: Consequences on stoichiometry and functional trait composition of freshwater macroinvertebrate communities.

Worldwide, freshwater systems are subjected to increasing temperatures and nutrient changes. Under phosphorus and nitrogen enrichment consumer communities are often thought to shift towards fast-growing and P-rich taxa, supporting the well-known link between growth rate and body stoichiometry. While these traits are also favoured under warming, the temperature effect on stoichiometry is less clear. As recently shown, there is a general link between functional traits and body stoichiometry, which makes the integration of stoichiometric traits a promising tool to help understanding the mechanisms behind taxonomic and functional community responses to nutrient changes and/or warming. Yet, such approaches have been scarcely developed at community level and on a long-term perspective. In this study, we investigated long-term responses in stoichiometry and functional trait composition of macroinvertebrate communities to nutrient changes (decreasing water P; increasing water N:P) and warming over a 34-year period in the Middle Loire River (France), testing the potentially opposing responses to these drivers. Both drivers should cause shifts in species composition, which will alter the overall community stoichiometry and functional composition following assumptions from ecological stoichiometry theory. We found that the macroinvertebrate community shifted towards P-poor taxa, causing significant trends in overall community stoichiometry which indicates long-term changes in the nutrient pool provided by these consumers (i.e. decrease in %N and %P, increase in N:P). Further, while the former high-P conditions favoured traits associated to detritus feeding and fast development (i.e. small maximum body size, short life duration), recent conditions favoured predators and slow-developing taxa. These results suggest nutrients to be a more important driver than temperature over this period. By providing a pivotal link between environmental changes and functional trait composition of communities, approaches based on stoichiometric traits offer sound perspectives to investigate ecological relationships between multiple drivers operating at various scales and ecosystem functioning.

Direct habitat descriptors improve the understanding of the organization of fish and macroinvertebrate communities across a large catchment.

In large-scale aquatic ecological studies, direct habitat descriptors (e.g. water temperature, hydraulics in river reaches) are often approximated by coarse-grain surrogates (e.g. air temperature, discharge respectively) since they are easier to measure or model. However, as biological variability can be very strong at the habitat scale, surrogate variables may have a limited ability to capture all of this variability, which may lead to a lesser understanding of the ecological processes or patterns of interest. In this study, we aimed to compare the capacity of direct habitat descriptors vs. surrogate environmental variables to explain the organization of fish and macroinvertebrate communities across the Loire catchment in France (105 km2). For this purpose, we relied on high-resolution environmental data, extensive biological monitoring data (>1000 sampling stations) and multivariate analyses. Fish and macroinvertebrate abundance datasets were considered both separately and combined to assess the value of a cross-taxa approach. We found that fish and macroinvertebrate communities exhibited weak concordance in their organization and responded differently to the main ecological gradients. Such variations are probably due to fundamental differences in their life-history traits and mobility. Regardless of the biological group considered, direct habitat descriptors (water temperature and local hydraulic variables) consistently explained the organization of fish and macroinvertebrate communities better than surrogate descriptors (air temperature and river discharge). Furthermore, the organization of fish and macroinvertebrate communities was slightly better explained by the combination of direct or surrogate environmental variables when the two biological groups were considered together than when considered separately. Tied together, these results emphasize the importance of using a cross-taxa approach in association with high-resolution direct habitat variables to more accurately explain the organization of aquatic communities.

Long-term impacts of nutrient control, climate change, and invasive clams on phytoplankton and cyanobacteria biomass in a large temperate river.

Recent studies suggest that climate change, with warmer water temperatures and lower and longer low flows, may enhance harmful planktic cyanobacterial growth in lakes and large rivers. Concomitantly, controlling nutrient loadings has proven effective in reducing phytoplankton biomass especially in North America and Western Europe. In addition, the impact of invasive benthic filter-feeder species such as Corbicula on phytoplankton has largely been overlooked in large rivers, leading to even more uncertainty in predicting future trajectories in river water quality. To investigate how nutrient control, climate change and invasion of benthic filter-feeders may affect phytoplankton biomass and composition, we assembled a large database on the entire water course of the River Loire (France) over three decades (1991-2019). We focus on cyanobacteria to provide an in-depth analysis of the 30-year trend and insights on future possible trajectories. Since 1991, total phytoplankton and cyanobacteria biomasses have decreased 10-fold despite warmer water temperature (+0.23 °C·decade-1) and lower summer flow (-0.25 L·s-1·km-2·decade-1). In the long-term, the contribution of planktic cyanobacteria to total biomass was on average 2.8%. The main factors driving total phytoplankton and cyanobacteria biomasses were total phosphorus (4-fold decrease), the abundance of Corbicula clams (from absence before 1998 to 250-1250 individuals·m-2 after 2010), the duration of summer low flows and the intensity of summer heatwaves. The River Loire constitutes an example in Europe of how nutrient control can be an efficient mitigation strategy, counteracting already visible effects of climate change on the thermal regime and flow pattern of the river. This may hold true under future conditions, but further work is needed to account for the climate trajectory, land and water use scenarios, the risk of enhanced benthic biofilm and macrophyte proliferation, together with the spread of invasive filter-feeding bivalves.

Increasing climate-driven taxonomic homogenization but functional differentiation among river macroinvertebrate assemblages.

Global change is increasing biotic homogenization globally, which modifies the functioning of ecosystems. While tendencies towards taxonomic homogenization in biological communities have been extensively studied, functional homogenization remains an understudied facet of biodiversity. Here, we tested four hypotheses related to long-term changes (1991-2016) in the taxonomic and functional arrangement of freshwater macroinvertebrate assemblages across space and possible drivers of these changes. Using data collected annually at 64 river sites in mainland New Zealand, we related temporal changes in taxonomic and functional spatial ß-diversity, and the contribution of individual sites to ß-diversity, to a set of global, regional, catchment and reach-scale environmental descriptors. We observed long-term, mostly climate-induced, temporal trends towards taxonomic homogenization but functional differentiation among macroinvertebrate assemblages. These changes were mainly driven by replacements of species and functional traits among assemblages, rather than nested species loss. In addition, there was no difference between the mean rate of change in the taxonomic and functional facets of ß-diversity. Climatic processes governed overall population and community changes in these freshwater ecosystems, but were amplified by multiple anthropogenic, topographic and biotic drivers of environmental change, acting widely across the landscape. The functional diversification of communities could potentially provide communities with greater stability, resistance and resilience capacity to environmental change, despite ongoing taxonomic homogenization. Therefore, our study highlights a need to further understand temporal trajectories in both taxonomic and functional components of species communities, which could enable a clearer picture of how biodiversity and ecosystems will respond to future global changes.

Long-term trends in trait structure of riverine communities facing predation risk increase and trophic resource decline.

Many large European rivers have undergone multiple pressures that have strongly impaired ecosystem functioning at different spatial and temporal scales. Global warming and other environmental changes have favored the success of invasive species, deeply modifying the structure of aquatic communities in large rivers. Some exogenous species could alter trophic interactions within assemblages by increasing the predation risk for potential prey species (top-down effect) and limiting the dynamics of others via resource availability limitation (bottom-up effect). Furthermore, large transboundary rivers are complex aquatic ecosystems that have often been poorly investigated so that data for assessing long-term ecological trends are missing. In this study, we propose an original approach for investigating long-term combined effects of global warming, trophic resource decrease, predation risk, and water quality variations on the trait-based structure of macroinvertebrate and fish assemblages over 26 yr (1985-2011) and 427-km stretch of the river Meuse (France and Belgium). The study of temporal variations in biological, physiological, and ecological traits of macroinvertebrate and fish allowed identifying community trends and distinguishing impacts of environmental perturbations from those induced by biological alterations. We provide evidence, for this large European river, of an increase in water temperature (close to 1°C) and a decrease in phytoplankton biomass (-85%), as well as independent effects of these changes on both invertebrate and fish communities. The reduction of trophic resources in the water column by invasive molluscs has dramatically affected the density of omnivorous fish in favor of invertebrate feeders, while scrapers became the major feeding guild among invertebrates. Macroinvertebrate and fish communities have shifted from large-sized organisms with low fecundity to prolific, small-sized organisms, with early maturity, as a response to increased predation pressure.

At what scale and extent environmental gradients and climatic changes influence stream invertebrate communities?

In a context of increasing landscape modifications and climatic changes, scale hierarchy becomes an ever more crucial issue to integrate in the analysis of drivers and stressors of biological communities, especially in river networks. To cope with this issue, we developed (i) spatial hierarchical models of functional diversity of stream invertebrate communities to assess the relative influence of local- vs. regional-scale factors in structuring community assembly, and (ii) analysis of metacommunity elements to determine the ecological processes behind the structuring. The spatial structuring of benthic invertebrate communities was investigated over 568 sites in South-eastern France. Community structure was mainly driven by the altitudinal gradient and spring flow variation at broad scales, with functional diversity gradually decreasing with elevation and being maximized at intermediate levels of flow variability. According to the 'elements of metacommunity structure' analysis, the prevailing influence of the altitudinal gradient was also supported by a Clementsian structuration of invertebrate communities. Conversely, the influence of observed climatic changes in temperature and rainfall was weak and observed only at fine scales. As a result, natural environmental filters were stronger drivers of the functional diversity of communities than human-induced stressors (e.g. water pollution and hydromorphological alterations). More broadly, our results suggest that management needs to embrace the possibilities of gathering high spatial and taxonomical resolution data when analysing and predicting flow variation and climate change effects in order to preserve and restore functionally diverse communities. Moreover, to develop environmental flow schemes or restoration and climate change adaptation strategies for freshwater communities, local and regional processes need to be addressed simultaneously; equally responsible as drivers of community diversity.

Assessing long-term effects of multiple, potentially confounded drivers in ecosystems from species traits.

Although species traits have the potential to disentangle long-term effects of multiple, potentially confounded drivers in ecosystems, this issue has received very little attention in the literature. We aimed at filling this gap by assessing the relative effects of hydroclimatic and water quality factors on the trait composition of invertebrate assemblages over 30 years in the Middle Loire River (France). Using a priori predictions on the long-term variation of trait-based adaptations over the three decades, we evaluated the ability of invertebrate traits to indicate the effects of warming, discharge reduction and water quality improvement. Hydroclimatic and water quality factors contributed to up to 65% of the variation in trait composition. More than 70% of the initial trait response predictions made according to observed long-term hydroclimatic changes were confirmed. They supported a general climate-induced trend involving adapted resistance and resilience strategies. A partial confounding effect of water quality improvement acting on trophic processes was also highlighted, indicating that improved water quality management can significantly help to reduce some adverse effects of climate change. This trait-based approach can have wider implications for investigating long-term changes driven by multiple, potentially confounded factors, as frequently encountered in the context of global change.

Long-term changes in temperate stream invertebrate communities reveal a synchronous trophic amplification at the turn of the millennium.

The positive effects of water quality improvement on stream biodiversity in the temperate regions are expected to be at risk with the projected climatic changes. However, the processes and mechanisms behind the predicted threats remain uncertain. From long-term series of benthic invertebrate samples from temperate rivers and streams in France, we analyzed diversity and composition shifts over time in relation to geographic elements and human stressors. Mechanisms for community changes were investigated with a trait-based analysis for the entire dataset and for a selected caddisfly community module. We observed a 42% increase in the taxonomic richness of stream invertebrate communities over the last 25years. A gradual trend induced by water quality improvement was distinguished from a more abrupt climate change-induced shift in communities around the year 2000. Trophic amplification - the intensification of trophic interactions and pathways through the food web - was identified as the mechanism behind the strong community shift. Four lines of evidence for this trophic amplification are highlighted: (i) higher dissolved oxygen concentrations indicated a shift in primary production, (ii) the trait-based analysis of entire communities showed a bottom-up food web amplification, (iii) the trait-based analysis of the community module evidenced feeding strategy shifts and increased food web interactions, and (iv) the abundance analysis of the community module showed a productivity increase. These results lend credit to persistent investments in water quality for improving stream biodiversity, and contrary to expectation, climate change impacts seem so far to have reinforced these positive effects.

Global climate change in large European rivers: long-term effects on macroinvertebrate communities and potential local confounding factors.

Aquatic species living in running waters are widely acknowledged to be vulnerable to climate-induced, thermal and hydrological fluctuations. Climate changes can interact with other environmental changes to determine structural and functional attributes of communities. Although such complex interactions are most likely to occur in a multiple-stressor context as frequently encountered in large rivers, they have received little attention in such ecosystems. In this study, we aimed at specifically addressing the issue of relative long-term effects of global and local changes on benthic macroinvertebrate communities in multistressed large rivers. We assessed effects of hydroclimatic vs. water quality factors on invertebrate community structure and composition over 30 years (1979-2008) in the Middle Loire River, France. As observed in other large European rivers, water warming over the three decades (+0.9 °C between 1979-1988 and 1999-2008) and to a lesser extent discharge reduction (-80 m(3) s(-1) ) were significantly involved in the disappearance or decrease in taxa typical from fast running, cold waters (e.g. Chloroperlidae and Potamanthidae). They explained also a major part of the appearance and increase of taxa typical from slow flowing or standing waters and warmer temperatures, including invasive species (e.g. Corbicula sp. and Atyaephyra desmarestii). However, this shift towards a generalist and pollution tolerant assemblage was partially confounded by local improvement in water quality (i.e. phosphate input reduction by about two thirds and eutrophication limitation by almost one half), explaining a significant part of the settlement of new pollution-sensitive taxa (e.g. the caddisfly Brachycentridae and Philopotamidae families) during the last years of the study period. The regain in such taxa allowed maintaining a certain level of specialization in the invertebrate community despite climate change effects.