Studying interactions among anthropogenic stressors in freshwater ecosystems: A systematic review of 2396 multiple-stressor experiments.

Understanding the interactions among anthropogenic stressors is critical for effective conservation and management of ecosystems. Freshwater scientists have invested considerable resources in conducting factorial experiments to disentangle stressor interactions by testing their individual and combined effects. However, the diversity of stressors and systems studied has hindered previous syntheses of this body of research. To overcome this challenge, we used a novel machine learning framework to identify relevant studies from over 235,000 publications. Our synthesis resulted in a new dataset of 2396 multiple-stressor experiments in freshwater systems. By summarizing the methods used in these studies, quantifying trends in the popularity of the investigated stressors, and performing co-occurrence analysis, we produce the most comprehensive overview of this diverse field of research to date. We provide both a taxonomy grouping the 909 investigated stressors into 31 classes and an open-source and interactive version of the dataset (https://jamesaorr.shinyapps.io/freshwater-multiple-stressors/). Inspired by our results, we provide a framework to help clarify whether statistical interactions detected by factorial experiments align with stressor interactions of interest, and we outline general guidelines for the design of multiple-stressor experiments relevant to any system. We conclude by highlighting the research directions required to better understand freshwater ecosystems facing multiple stressors.

Local stressors mask the effects of warming in freshwater ecosystems.

Climate warming is a ubiquitous stressor in freshwater ecosystems, yet its interactive effects with other stressors are poorly understood. We address this knowledge gap by testing the ability of three contrasting null models to predict the joint impacts of warming and a range of other aquatic stressors using a new database of 296 experimental combinations. Despite concerns that stressors will interact to cause synergisms, we found that net impacts were usually best explained by the effect of the stronger stressor alone (the dominance null model), especially if this stressor was a local disturbance associated with human land use. Prediction accuracy depended on stressor identity and how asymmetric stressors were in the magnitude of their effects. These findings suggest we can effectively predict the impacts of multiple stressors by focusing on the stronger stressor, as habitat alteration, nutrients and contamination often override the biological consequences of higher temperatures in freshwater ecosystems.

Rapid evolution generates synergism between multiple stressors: Linking theory and an evolution experiment.

Global change encompasses many co-occurring anthropogenic stressors. Understanding the interactions between these multiple stressors, whether they be additive, antagonistic or synergistic, is critical for ecosystem managers when prioritizing which stressors to mitigate in the face of global change. While such interactions between stressors appear prevalent, it remains unclear if and how these interactions change over time, as the majority of multiple-stressor studies rarely span multiple generations of study organisms. Although meta-analyses have reported some intriguing temporal trends in stressor interactions, for example that synergism may take time to emerge, the mechanistic basis for such observations is unknown. In this study, by analysing data from an evolution experiment with the rotifer Brachionus calyciflorus (~35 generations and 31,320 observations), we show that adaptation to multiple stressors shifts stressor interactions towards synergism. We show that trade-offs, where populations cannot optimally perform multiple tasks (i.e. adapting to multiple stressors), generate this bias towards synergism. We also show that removal of stressors from evolved populations does not necessarily increase fitness and that there is variation in the evolutionary trajectories of populations that experienced the same stressor regimes. Our results highlight outstanding questions at the interface between evolution and global change biology, and illustrate the importance of considering rapid adaptation when managing or restoring ecosystems subjected to multiple stressors under global change.

Warming and imidacloprid pulses determine macroinvertebrate community dynamics in experimental streams.

Sustainable management of freshwater and pesticide use is essential for mitigating the impacts of intensive agriculture in the context of a changing climate. To better understand how climate change will affect the vulnerability of freshwater ecosystems to chemical pollutants, more empirical evidence is needed on the combined effects of climatic and chemical stressors in environmentally realistic conditions. Our experiment provides the first empirical evaluation of stream macroinvertebrate community dynamics in response to one of the world's most widely used insecticides, imidacloprid, and increased water temperature. In a 7-week streamside experiment using 128 flow-through circular mesocosms, we investigated the effects of pulsed imidacloprid exposure (four environmentally relevant levels between 0 and 4.6 µg/L) and raised water temperature (ambient, 3°C above) on invertebrate communities representative of fast- and slow-flowing microhabitats. Invertebrate drift and insect emergence were monitored during three pesticide pulses (10 days apart), and benthic invertebrate communities were sampled after 24 days of heating and pesticide manipulations. All three manipulated factors strongly affected drift community composition. The first imidacloprid pulse and increased temperature had a greater impact on communities in fast-flowing mesocosms, which contained more pollution-sensitive EPT taxa (mayflies, stoneflies and caddisflies). Heating and imidacloprid caused increased emigration by drift, weak reductions in emergence, and negatively affected the benthic community. The combined effect of stressor manipulations and a 10-day natural heatwave drastically reduced relative abundances of EPT and insects overall and caused a shift to oligochaete-, crustacean- and gastropod-dominated communities. Contrary to our hypothesis, the very high yet realistic water temperatures reached in our experiment meant the negative effects of imidacloprid were clearest at ambient temperatures and fast flow. These findings demonstrate the potential combined impacts of imidacloprid contamination and heatwaves on freshwater invertebrate communities under future climate scenarios and highlight the need for more countries to take regulatory action to control neonicotinoid use.

Fine sediment and flow velocity impact bacterial community and functional profile more than nutrient enrichment.

Freshwater ecosystems face many simultaneous pressures due to human activities. Consequently, there has been a rapid loss of freshwater biodiversity and an increase in biomonitoring programs. Our study assessed the potential of benthic stream bacterial communities as indicators of multiple-stressor impacts associated with urbanization and agricultural intensification. We conducted a fully crossed four-factor experiment in 64 flow-through mesocosms fed by a pristine montane stream (21 d of colonization, 21 d of manipulations) and investigated the effects of nutrient enrichment, flow-velocity reduction and added fine sediment after 2 and 3 weeks of stressor exposure. We used high-throughput sequencing and metabarcoding techniques (16S rRNA genes), as well as curated biological databases (METAGENassit, MetaCyc), to identify changes in bacterial relative abundances and predicted metabolic functional profile. Sediment addition and flow-velocity reduction were the most pervasive stressors. They both increased α-diversity and had strong taxon-specific effects on community composition and predicted functions. Sediment and flow velocity also interacted frequently, with 88% of all bacterial response variables showing two-way interactions and 33% showing three-way interactions including nutrient enrichment. Changes in relative abundances of common taxa were associated with shifts in dominant predicted functions, which can be extrapolated to underlaying stream-wide mechanisms such as carbon use and bacterial energy production pathways. Observed changes were largely stable over time and occurred after just 2 weeks of exposure, demonstrating that bacterial communities can be well-suited for early detection of multiple stressors. Overall, added sediment and reduced flow velocity impacted both bacterial community structure and predicted function more than nutrient enrichment. In future research and stream management, a holistic approach to studying multiple-stressor impacts should include multiple trophic levels with their functional responses, to enhance our mechanistic understanding of complex stressor effects and promote establishment of more efficient biomonitoring programs.

Towards a unified study of multiple stressors: divisions and common goals across research disciplines.

Anthropogenic environmental changes, or 'stressors', increasingly threaten biodiversity and ecosystem functioning worldwide. Multiple-stressor research is a rapidly expanding field of science that seeks to understand and ultimately predict the interactions between stressors. Reviews and meta-analyses of the primary scientific literature have largely been specific to either freshwater, marine or terrestrial ecology, or ecotoxicology. In this cross-disciplinary study, we review the state of knowledge within and among these disciplines to highlight commonality and division in multiple-stressor research. Our review goes beyond a description of previous research by using quantitative bibliometric analysis to identify the division between disciplines and link previously disconnected research communities. Towards a unified research framework, we discuss the shared goal of increased realism through both ecological and temporal complexity, with the overarching aim of improving predictive power. In a rapidly changing world, advancing our understanding of the cumulative ecological impacts of multiple stressors is critical for biodiversity conservation and ecosystem management. Identifying and overcoming the barriers to interdisciplinary knowledge exchange is necessary in rising to this challenge. Division between ecosystem types and disciplines is largely a human creation. Species and stressors cross these borders and so should the scientists who study them.

Advancing understanding and prediction in multiple stressor research through a mechanistic basis for null models.

Global environmental change is driven by multiple anthropogenic stressors. Conservation and restoration require understanding the individual and joint action of these stressors to evaluate and prioritize management measures. To date, most studies on multiple stressor effects have sought to identify potential stressor interactions, defined as deviations from null models, and related meta-analyses have focused on quantifying the relative proportion of stressor interactions across studies. These studies have provided valuable insights about the complexity of multiple stressor effects, but remain largely devoid of a theoretical framework for null model selection and prediction of effects. We suggest that multiple stressor research would benefit by (1) integrating and developing additional null models and (2) selecting null models based on their mechanistic assumptions of the stressor mode of action and organism sensitivities as well as stressor-effect relationships for individuals and populations. We present a range of null models and outline their underlying assumptions and application in multiple stressor research. Moving beyond mere description requires multiple stressor research to shift its focus from identifying statistically significant interactions to the use and development of mechanistic (null) models. Justified selection of the appropriate null model is a first step to achieve this.

Climate warming and agricultural stressors interact to determine stream macroinvertebrate community dynamics.

Global climate change is likely to modify the ecological consequences of currently acting stressors, but potentially important interactions between climate warming and land-use related stressors remain largely unknown. Agriculture affects streams and rivers worldwide, including via nutrient enrichment and increased fine sediment input. We manipulated nutrients (simulating agricultural run-off) and deposited fine sediment (simulating agricultural erosion) (two levels each) and water temperature (eight levels, 0-6°C above ambient) simultaneously in 128 streamside mesocosms to determine the individual and combined effects of the three stressors on macroinvertebrate community dynamics (community composition and body size structure of benthic, drift and insect emergence assemblages). All three stressors had pervasive individual effects, but in combination often produced additive or antagonistic outcomes. Changes in benthic community composition showed a complex interplay among habitat quality (with or without sediment), resource availability (with or without nutrient enrichment) and the behavioural/physiological tendency to drift or emerge as temperature rose. The presence of sediment and raised temperature both resulted in a community of smaller organisms. Deposited fine sediment strongly increased the propensity to drift. Stressor effects were most prominent in the benthic assemblage, frequently reflected by opposite patterns in individuals quitting the benthos (in terms of their propensity to drift or emerge). Of particular importance is that community measures of stream health routinely used around the world (taxon richness, EPT richness and diversity) all showed complex three-way interactions, with either a consistently stronger temperature response or a reversal of its direction when one or both agricultural stressors were also in operation. The negative effects of added fine sediment, which were often stronger at raised temperatures, suggest that streams already impacted by high sediment loads may be further degraded under a warming climate. However, the degree to which this will occur may also depend on in-stream nutrient conditions.

Climate warming and agricultural stressors interact to determine stream periphyton community composition.

Lack of knowledge about how the various drivers of global climate change will interact with multiple stressors already affecting ecosystems is the basis for great uncertainty in projections of future biological change. Despite concerns about the impacts of changes in land use, eutrophication and climate warming in running waters, the interactive effects of these stressors on stream periphyton are largely unknown. We manipulated nutrients (simulating agricultural runoff), deposited fine sediment (simulating agricultural erosion) (two levels each) and water temperature (eight levels, 0-6 °C above ambient) simultaneously in 128 streamside mesocosms. Our aim was to determine the individual and combined effects of the three stressors on the algal and bacterial constituents of the periphyton. All three stressors had pervasive individual effects, but in combination frequently produced synergisms at the population level and antagonisms at the community level. Depending on sediment and nutrient conditions, the effect of raised temperature frequently produced contrasting response patterns, with stronger or opposing effects when one or both stressors were augmented. Thus, warming tended to interact negatively with nutrients or sediment by weakening or reversing positive temperature effects or strengthening negative ones. Five classes of algal growth morphology were all affected in complex ways by raised temperature, suggesting that these measures may prove unreliable in biomonitoring programs in a warming climate. The evenness and diversity of the most abundant bacterial taxa increased with temperature at ambient but not with enriched nutrient levels, indicating that warming coupled with nutrient limitation may lead to a more evenly distributed bacterial community as temperatures rise. Freshwater management decisions that seek to avoid or mitigate the negative effects of agricultural land use on stream periphyton should be informed by knowledge of the interactive effects of multiple stressors in a warming climate.

Heatwaves and carbon dioxide enrichment impact invertebrate drift and insect emergence patterns across time in experimental streams.

Climate change and human land use are considered key threats to freshwater invertebrates. Heatwaves can impact the phenology of insects and population dynamics, yet have been largely ignored in experiments compared to mean temperature changes. Another major anthropogenic stressor driving invertebrate community changes is deposited fine sediment; therefore, effects of key climate-change drivers on invertebrate drift and insect emergence rates may differ between sediment-impacted and non-impacted streams. However, this has never been tested in a realistic outdoor experiment. We investigated the individual and combined effects of two 7-day heatwaves, CO2 enrichment, flow velocity variability (periods of fast and slow) and fine sediment on stream drift and emergence responses, sampled four times during a 7-week experiment in 128 flow-through stream mesocosms. We examined invertebrate drift and insect emergence responses to the four stressors, and used these responses to help explain the benthic invertebrate community responses already assessed (sampled at the end of the experiment). Heatwave 1 strongly increased emergence (dominated by Chironomidae), causing an earlier emergence peak, an effect not repeated during heatwave 2, seven days later. During heatwave 1, emerged chironomids were larger in heated channels, but smaller in heated channels afterwards, suggesting a different effect on body size of short-term heatwaves to previous constant warming experiments. CO2 enrichment reduced drifting EPT and total and Chironomidae emergence on three sampling occasions each. After heatwave 1, total drift and total emergence were strongly reduced by heating in ambient-CO2 channels, whereas no reduction occurred in CO2-enriched channels. During heatwave 2, total drift increased in channels without sediment but not in channels with added sediment. Overall, our findings suggest heatwaves can shift the timing of stream insect emergence, regardless of longer-term mean temperatures. They also show that heatwaves, raised CO2, and fine sediment can modulate each others' effects on drift and emergence dynamics.

Effects of flow reduction and artificial light at night (ALAN) on litter decomposition and invertebrate communities in streams: A flume experiment.

River ecosystems are heavily impacted by multiple stressors, where effects can cascade downstream of point sources. However, a spatial approach to assess the effects of multiple stressors is missing. We assessed the local and downstream effects on litter decomposition, and associated invertebrate communities of two stressors: flow reduction and artificial light at night (ALAN). We used an 18-flow-through mesocosm system consisting of two tributaries, where we applied the stressors, merging in a downstream section. We assessed the changes in decomposition rate and invertebrate community structure in leaf bags. We found no effect of ALAN or its interaction with flow reduction on the litter decomposition or the invertebrate community in the tributaries. Flow reduction alone led to a 14.8 % reduction in decomposition rate in the tributaries. We recorded no effect of flow reduction on the macroinvertebrates community composition in the litter bags. We also observed no effects of the spatial arrangement of the stressors on the litter decomposition and macroinvertebrate community structure downstream. Overall, our results suggest the impact of stressors on litter decomposition and macroinvertebrate communities remained local in our experiment. Our work thus calls for further studies to identify the mechanisms and the conditions under which spatial effects dominate over local processes.

Assessing the response of an urban stream ecosystem to salinization under different flow regimes.

Urban streams are exposed to a variety of anthropogenic stressors. Freshwater salinization is a key stressor in these ecosystems that is predicted to be further exacerbated by climate change, which causes simultaneous changes in flow parameters, potentially resulting in non-additive effects on aquatic ecosystems. However, the effects of salinization and flow velocity on urban streams are still poorly understood as multiple-stressor experiments are often conducted at pristine rather than urban sites. Therefore, we conducted a mesocosm experiment at the Boye River, a recently restored stream located in a highly urbanized area in Western Germany, and applied recurrent pulses of salinity along a gradient (NaCl, 9 h daily of +0 to +2.5 mS/cm) in combination with normal and reduced current velocities (20 cm/s vs. 10 cm/s). Using a comprehensive assessment across multiple organism groups (macroinvertebrates, eukaryotic algae, fungi, parasites) and ecosystem functions (primary production, organic-matter decomposition), we show that flow velocity reduction has a pervasive impact, causing community shifts for almost all assessed organism groups (except fungi) and inhibiting organic-matter decomposition. Salinization affected only dynamic components of community assembly by enhancing invertebrate emigration via drift and reducing fungal reproduction. We caution that the comparatively small impact of salt in our study can be due to legacy effects from past salt pollution by coal mining activities >30 years ago. Nevertheless, our results suggest that urban stream management should prioritize the continuity of a minimum discharge to maintain ecosystem integrity. Our study exemplifies a holistic approach for the assessment of multiple-stressor impacts on streams, which is needed to inform the establishment of a salinity threshold above which mitigation actions must be taken.

Assessing remote sensing as a tool to monitor hydrological stress in Irish catchments with Freshwater Pearl Mussel populations.

The West Coast of Ireland hosts many of the few populations of Freshwater Peal Mussels (FPM) left in Europe. The decline of this keystone species is strongly related to deteriorating hydrological conditions, specifically to the threat of low flows during dry summers. Populations still capable of reproducing require a minimum discharge and flow velocity to support juvenile mussels, or else stress builds up and an entire generation may be lost. Monitoring environmental and hydrological conditions in small and remote FPM catchments is difficult due to the lack of infrastructure. Indices derived from remote sensing imagery can be used to assess hydrological variables at the catchment scale. Here, five indices are tested as possible surrogates for soil moisture and evapotranspiration, based on two relevant land-cover types: open peat habitats (OPH) and forestry. Selected indices are then assessed in their ability to reproduce seasonal patterns and in their response to a severe drought event. The moisture stress index (MSI) and normalized difference vegetation index (NDVI) were found to be the best surrogates for soil moisture and evapotranspiration respectively. Both indices showed seasonality patterns in the two land-cover types, although the variability of MSI was significantly higher. During the 2018 drought, MSI visibly increased only in OPH, while NDVI rose only for forestry. The results suggest that OPH enhances the long-term hydrological resilience of a catchment by conserving water in the peat substrate, while industrial forestry plantations exacerbate the pressure on water during drier periods. This has consequences for river discharge, freshwater biodiversity and specifically for FPM. Implementing these surrogates have the potential to identify land-use management strategies that reduce and even avert the effects of drought on FPM. Such strategies are increasingly necessary in a climate change context, as recurring summer droughts are expected in most of Europe.

Using weighted expert judgement and nonlinear data analysis to improve Bayesian belief network models for riverine ecosystem services.

Rivers are a key part of the hydrological cycle and a vital conduit of water resources, but are under increasing threat from anthropogenic pressures. Linking pressures with ecosystem services is challenging because the processes interconnecting the physico-chemical, biological and socio-economic elements are usually captured using heterogenous methods. Our objectives were, firstly, to advance an existing proof-of-principle Bayesian belief network (BBN) model for integration of ecosystem services considerations into river management. We causally linked catchment stressors with ecosystem services using weighted evidence from an expert workshop (capturing confidence among expert groups), legislation and published literature. The BBN was calibrated with analyses of national monitoring data (including non-linear relationships and ecologically meaningful breakpoints) and expert judgement. We used a novel expected index of desirability to quantify the model outputs. Secondly, we applied the BBN to three case study catchments in Ireland to demonstrate the implications of changes in stressor levels for ecosystem services in different settings. Four out of the seven significant relationships in data analyses were non-linear, highlighting that non-linearity is common in ecosystems, but rarely considered in environmental modelling. Deficiency of riparian shading was identified as a prevalent and strong influence, which should be addressed to improve a broad range of societal benefits, particularly in the catchments where riparian shading is scarce. Sediment load had a lower influence on river biology in flashy rivers where it has less potential to settle out. Sediment interacted synergistically with organic matter and phosphate where these stressors were active; tackling these stressor pairs simultaneously can yield additional societal benefits compared to the sum of their individual influences, which highlights the value of integrated management. Our BBN model can be parametrised for other Irish catchments whereas elements of our approach, including the expected index of desirability, can be adapted globally.

Perceived multiple stressor effects depend on sample size and stressor gradient length.

Multiple stressors are continuously deteriorating surface waters worldwide, posing many challenges for their conservation and restoration. Combined effect types of multiple stressors range from single-stressor dominance to complex interactions. Identifying prevalent combined effect types is critical for environmental management, as it helps to prioritise key stressors for mitigation. However, it remains unclear whether observed single and combined stressor effects reflect true ecological processes unbiased by sample size and length of stressor gradients. Therefore, we examined the role of sample size and stressor gradient lengths in 158 paired-stressor response cases with over 120,000 samples from rivers, lakes, transitional and marine ecosystems around the world. For each case, we split the overall stressor gradient into two partial gradients (lower and upper) and investigated associated changes in single and combined stressor effects. Sample size influenced the identified combined effect types, and stressor interactions were less likely for cases with fewer samples. After splitting gradients, 40 % of cases showed a change in combined effect type, 30 % no change, and 31 % showed a loss in stressor effects. These findings suggest that identified combined effect types may often be statistical artefacts rather than representing ecological processes. In 58 % of cases, we observed changes in stressor effect directions after the gradient split, suggesting unimodal stressor effects. In general, such non-linear responses were more pronounced for organisms at higher trophic levels. We conclude that observed multiple stressor effects are not solely determined by ecological processes, but also strongly depend on sampling design. Observed effects are likely to change when sample size and/or gradient length are modified. Our study highlights the need for improved monitoring programmes with sufficient sample size and stressor gradient coverage. Our findings emphasize the importance of adaptive management, as stress reduction measures or further ecosystem degradation may change multiple stressor-effect relationships, which will then require associated changes in management strategies.

Time-cumulative effects of neonicotinoid exposure, heatwaves and food limitation on stream mayfly nymphs: A multiple-stressor experiment.

The global intensification of agriculture has resulted in pesticides playing an increasingly important role as anthropogenic stressors and drivers of environmental change. There is also a growing need to determine if other environmental stressors, especially those predicted to worsen with climate change, interact with pesticides to alter their effects on non-target biota. Two such stressors are increased extreme temperature events and periods of food limitation. This study is the first to investigate the combined effects of the world's most widely used insecticide, imidacloprid, with heatwaves and food limitation on a freshwater animal. A 6-week, full-factorial laboratory experiment with Deleatidium spp. mayfly nymphs was performed to investigate the potential for direct and delayed interactive effects of simulated heatwaves and starvation with chronic exposure to a field-realistic concentration of imidacloprid (0.4 µg/L). The experiment included two 6-day simulated heatwaves, one during a starvation period prior to imidacloprid addition, and one during the first 6 days of imidacloprid exposure. The simulated heatwaves alone caused such drastic negative effects on Deleatidium survival and mobility that mainly antagonistic interactions were observed with the other stressors, though delayed synergisms between imidacloprid and the second heatwave also affected mayfly mobility. Time-cumulative toxicity of imidacloprid was evident, with imidacloprid first affecting mayfly mobility after 12 days but eventually causing the strongest effects of all manipulated stressors. However, lethal effects of imidacloprid could only be detected in the absence of heatwaves and starvation, possibly as a result of selection for stronger individuals due to prior exposure to these stressors. Our findings demonstrate that heatwaves of increasing severity will critically affect sensitive freshwater organisms such as mayflies, and that the impacts of widespread pesticide use on freshwater ecosystems under global climate change cannot be ignored.

Imidacloprid dominates the combined toxicities of neonicotinoid mixtures to stream mayfly nymphs.

Contamination of the environment with toxic chemicals such as pesticides has become a global problem. Understanding the role of chemical contaminants as stressors in ecological systems is therefore an important research need in the 21st century. In surface freshwaters, mixtures of neonicotinoid insecticides are being detected around the world as more monitoring data become available. Combinations of imidacloprid, clothianidin and thiamethoxam are commonly found, but studies testing their combined toxicities to freshwater invertebrates are rare. Taking a multiple-stressor approach, we employed a full-factorial design to investigate the individual and combined chronic toxicities of these three neonicotinoids in a 28-day laboratory experiment using Deleatidium spp. mayfly nymphs. Imidacloprid (1.2 µg/L achieved concentration) reduced mayfly survival (by 50% on Day 28) and mobility (~100%) more than clothianidin (1.1 µg/L, affecting about 25% of individuals across the responses measured) and thiamethoxam (2.9 µg/L, affecting 12%). Imidacloprid interacted with the other two neonicotinoids to cause a greater-than-additive negative effect when combined until 25 days of exposure, after which the strong negative overall effects of imidacloprid prevented these interactions from being observed. Our findings represent a novel contribution to multiple-stressor research by demonstrating the combined effects of chronic exposure to environmentally relevant neonicotinoid concentrations on an ecologically important stream insect taxon. These results emphasise the higher toxicity of imidacloprid to non-target freshwater insects compared to clothianidin and thiamethoxam, implying that stricter regulation to control the use of imidacloprid may need to be prioritised to protect vulnerable aquatic insect populations that provide key links to terrestrial food webs. Finally, our study provides an ecological, multiple-stressor comparison for related ecotoxicological investigations indicating neonicotinoid mixtures can deviate from additive toxicity.

Impacts of multiple anthropogenic stressors on stream macroinvertebrate community composition and functional diversity.

Ensuring the provision of essential ecosystem services in systems affected by multiple stressors is a key challenge for theoretical and applied ecology. Trait-based approaches have increasingly been used in multiple-stressor research in freshwaters because they potentially provide a powerful method to explore the mechanisms underlying changes in populations and communities. Individual benthic macroinvertebrate traits associated with mobility, life history, morphology, and feeding habits are often used to determine how environmental drivers structure stream communities. However, to date multiple-stressor research on stream invertebrates has focused more on taxonomic than on functional metrics. We conducted a fully crossed, 4-factor experiment in 64 stream mesocosms fed by a pristine montane stream (21 days of colonization, 21 days of manipulations) and investigated the effects of nutrient enrichment, flow velocity reduction and sedimentation on invertebrate community, taxon, functional diversity and trait variables after 2 and 3 weeks of stressor exposure. 89% of the community structure metrics, 59% of the common taxa, 50% of functional diversity metrics, and 79% of functional traits responded to at least one stressor each. Deposited fine sediment and flow velocity reduction had the strongest impacts, affecting invertebrate abundances and diversity, and their effects translated into a reduction of functional redundancy. Stressor effects often varied between sampling occasions, further complicating the prediction of multiple-stressor effects on communities. Overall, our study suggests that future research combining community, trait, and functional diversity assessments can improve our understanding of multiple-stressor effects and their interactions in running waters.

Making waves. Bridging theory and practice towards multiple stressor management in freshwater ecosystems.

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.

Multiple stressors and stream macroinvertebrate community dynamics: Interactions between fine sediment grain size and flow velocity.

Agricultural development has resulted in the degradation of freshwater ecosystems worldwide. Two key stressors impacting streams and rivers draining agricultural catchments are deposited fine sediment (e.g. due to erosion) and reduced flows (e.g. due to water abstraction, dams, or climate change). Past studies have identified fine sediment as a 'master stressor' in streams, but the effects of different sediment grain sizes in combination with reduced flow velocity are poorly understood. We manipulated deposited fine sediment (no added sediment; silt: 0-0.125 mm; fine sand: 0.125-0.250 mm; coarse sand: 1-2 mm) and flow velocity (fast: 26.5 cm/s; medium: 13.9 cm/s; slow: 0.0 cm/s) simultaneously in 60 outdoor stream mesocosms. We determined the individual and combined effects of these stressors on the benthic, drifting, and emerging stream macroinvertebrate communities. Both fine sediment and reduced flow velocity had pervasive detrimental impacts on stream invertebrate communities. Negative effects of sediment were worse at the smaller two grain sizes for some responses (abundance of Chironomidae, Copepoda, Psilochorema spp.); however, for several sediment-sensitive common taxa or community-level invertebrate metrics, effects were negative regardless of grain size. Although their combined effects were mainly additive, sediment impacts were worsened by reduced flow velocities in several cases. Our findings imply that (a) especially for sediment-sensitive species, all fine sediment <2 mm has profound negative effects, (b) sediment grain size matters for some invertebrate taxa, where severity of impacts increased as particle size decreased, and (c) negative effects of sedimentation can become worse when combined with reduced flow velocity.