Abstract Urban landscapes can drastically alter light regimes through stream burial, and also modify water temperature patterns, both of which have profound implications for the degradation of dissolved organic matter (DOM) through photodegradation and biodegradation, respectively. Despite their likely significance, the dynamics of short-term biodegradation and photodegradation in urban environments remain poorly understood, with limited knowledge regarding the potential interplay between warming and stream burial effects. This study used a replicated flume experiment to investigate the effects of shading, warming (+ 4.5–6.6°C), and their interaction on DOM processing. We used optical techniques to characterize DOM quantity and composition, allowing us to assess photodegradation and biodegradation rates in urban stream analogues. Linear mixed effects models revealed that the degradation of the fluorescent DOM pool decreased under shaded conditions, accompanied by an increase in humic-like compounds. Additionally, shaded flumes exhibited a shift towards higher molecular weight organic matter, indicating the importance of photodegradation in DOM processing within urban rivers. Temperature effects on DOM processing rates were found to be relatively minor compared to shading, with no interaction with shading observed. Principal Component Analysis demonstrated clear distinctions between shaded and unshaded treatments. In contrast, no significant differences were observed between warmed and ambient temperature treatments. Our findings suggest that stream burial impedes DOM processing and alters DOM composition in urban headwaters by inhibiting the photodegradation of humic material. The temperature treatments examined had limited impacts on biodegradation over the relatively short timescales of this study. This study provides experimental support for daylighting interventions as a strategy to enhance DOM processing in urban streams and mitigate the flux of labile material to downstream ecosystems.

Freshwater habitats are drying more frequently and for longer under the combined pressures of climate change and overabstraction. Unsurprisingly, many aquatic species decline or become locally extinct as their benthic habitat is lost during stream droughts, but less is known about the potential 'winners': those terrestrial species that may exploit emerging niches in drying riverbeds. In particular, we do not know how these transient ecotones will respond as droughts become more extreme in the future. To find out we used a large-scale, long-term mesocosm experiment spanning a wide gradient of drought intensity, from permanent flows to full streambed dewatering, and analysed terrestrial invertebrate community assembly after 1 year. Droughts that caused stream fragmentation gave rise to the most diverse terrestrial invertebrate assemblages, including 10 species with UK conservation designations, and high species turnover between experimental channels. Droughts that caused streambed dewatering produced lower terrestrial invertebrate richness, suggesting that the persistence of instream pools may benefit these taxa as well as aquatic biota. Particularly intense droughts may therefore yield relatively few 'winners' among either aquatic or terrestrial species, indicating that the threat to riverine biodiversity from future drought intensification could be more pervasive than widely acknowledged.

River flow regimes face increasing pressure from human activities including water resource management operations and climate change. Consequently, extreme hydrological events are becoming more severe and commonplace, and there is a pressing need to understand and manage their ecological effects. Extreme low flows (ELFs)-those displaying significantly greater magnitudes and durations than typical low-flow conditions-are being increasingly experienced globally. Fish and macroinvertebrate responses to ELFs have been more widely researched relative to other organism groups in riverine environments, although such studies have employed variable methodological techniques. In this perspective piece, we identify field-based assessments and controlled experiments as two key research paradigms used to examine riverine faunal responses to ELFs. Field-based assessments are often explorative and can benefit from utilising large-scale and long-term datasets. Alternatively, controlled experiments typically employ more hypothesis-driven approaches and can establish strong cause and effect linkages through high replication and control over potentially confounding parameters. Each paradigm clearly possesses their respective strengths, which we highlight and discuss how these could be better harnessed to optimise scientific advancements. To date, studies examining faunal responses to ELFs in these two research paradigms have largely been undertaken in parallel. Here, we argue that future research should seek to develop closer synergies to optimise the quality and quantity of evidence to better understand riverine faunal responses to ELFs. Such scientific advances are of paramount importance given the vulnerability of riverine fauna, and the ecosystems they comprise, to a new era of ELFs in many global regions.

An understanding of how biotic communities are spatially organized is necessary to identify and prioritize habitats within landscape‐scale biodiversity conservation. Local contribution to beta diversity (LCBD) identifies individual habitats that make a significant contribution to beta diversity and may have important practical implications, particularly for conservation of habitat networks. In this study, we develop and apply a conservation prioritization approach based on LCBD in aquatic invertebrate communities from 132 ponds.

Impacts on biodiversity and the functioning of ecosystems are often overlooked in small waterbodies as they are not regularly monitored as part of the Water Framework Directive (WFD). These small systems are, however, often essential for freshwater biodiversity and are frequently characterized by high beta-diversity. In this study we examined shredder diversity and ecosystem functioning (leaf litter breakdown) as indicators of environmental stress along a gradient of urbanisation. The native isopod Asellus aquaticus and the non-native shrimp Crangonyx pseudogracilis (Amphipoda) were dominant species of the shredder communities in 26 study ponds. Variation in shredder community composition among ponds was explained by pond surface area (P < 0.05) and the coverage of impermeable surface within 500 m (P < 0.05), indicative of urban impact, which was strongly, and negatively correlated with shredder richness. Site mean total leaf breakdown was 0.0020 k d−1 (range 0.0003 – 0.0053), whilst breakdown rates owing to microbial and fungal activity were lower and averaged 0.0010 k d−1 (range 6.72−5 − 0.0036). Both total and microbial activity driven breakdown rates showed relatively weak relationships with the environmental gradient reflecting degrees of urbanization. A model-averaging procedure suggested that total litter breakdown rates could be attributed to activity of shredders with A. aquaticus and C. pseudogracilis contributing most, explaining 65% of the variance (R2 = 0.65, F = 13.7, P < 0.001), which both significantly and similarly predicted total leaf breakdown rates (P < 0.05). Total leaf breakdown rates were significantly correlated with the densities of both C. pseudogracilis (P < 0.001) and A. aquaticus (P < 0.001), however partial correlations indicated that the former showed the strongest association.The process of urbanisation can have significant, and negative impacts upon freshwater ecosystems, including small water bodies such as ponds both regarding biodiversity and ecosystem functioning. The findings of the current study suggest that ecosystem functioning indicators such as leaf litter decomposition should be paired with structural measures. This is because function sustained by tolerant invading species would not otherwise be detected, and their presence is increasingly likely due to climate change and urbanization.

Climate change is expected to intensify the effect of environmental stressors on riverine ecosystems. Extreme events, such as low flow and heatwaves, could have profound consequences for stream ecosystem functioning, but research on the impact of these stressors and their interaction across multiple processes, remains scarce. Here, we report the results of a two-month stream mesocosm experiment testing the effect of low flow (66% water level reduction, without gravel exposure) and heatwaves (three 8-d episodes of +5 degrees C above ambient with 10-15 days recovery between each episode) on a suite of ecosystem processes (i.e. detrital decomposition, biofilm accrual, ecosystem metabolism and DOC quantity and quality). Low flow reduced whole system metab-olism, suppressing the rates of gross primary production (GPP) and ecosystem respiration (ER), but elevated DOC concentration. Overall, habitat contraction was the main driver of reduced ecosystem functioning in the low flow treatment. By contrast, heatwaves increased decomposition, algal accrual, and humic-like DOC, but reduced leaf decomposition efficiency. Net ecosystem production (NEP) generally decreased across the experiment but was most pronounced for low flow and heatwaves when occurring independently. Assessment of NEP responses to the three successive heatwave events revealed that responses later in the sequence were more reduced (i.e. more similar to controls), suggesting biofilm communities may acclimate to autumn heatwaves. However, when heatwaves co-occurred with low flow, a strong reduction in both ER and GPP was observed, suggesting in-creased microbial mortality and reduced acclimation. Our study reveals autumn heatwaves potentially elongate the growth season for primary producers and stimulate decomposers. With climate change, river ecosystems may become more heterotrophic, with faster processing of recalcitrant carbon. Further research is required to identify the impacts on higher trophic levels, meta-community dynamics and the potential for legacy effects gen-erated by successive low flows and heatwaves. (c) 2021 Elsevier B.V. All rights reserved.

Predicted trends towards more intense droughts are of particular significance for running water ecosystems, as the loss of critical stream habitat can provoke sudden changes in biodiversity and shifts in community structure. However, analysing ecological responses to the progressive loss of stream habitat requires a continuous disturbance gradient that can only be generated through large-scale manipulations of streamflow. In the first experiment of its kind, we used large artificial stream channels (mesocosms) as analogues of spring-fed headwaters and simulated a gradient of drought intensity that encompassed flowing streams, disconnected pools, and dry streambeds. We used breakpoint analysis to analyse macroinvertebrate community responses to intensifying drought, and identify the taxa and compositional metrics sensitive to small changes in drought stress. We detected breakpoints for >60% of taxa, signalling sudden population crashes or irruptions as drought intensified. Abrupt changes were most pronounced where riffle dewatering isolated pools. In the remnant wetted habitat, we observed a shift to larger body sizes across the community, primarily driven by irruptions of predatory midge larvae and coincident population collapses among prey species (worms and smaller midges). Our results suggest that intense predation in confined, fragmented stream habitat can lead to unexpected changes in body sizes, challenging the conventional wisdom that droughts favour the small. Pool fragmentation might thus be the most critical stage of habitat loss during future droughts, as the point at which impacted rivers and streams begin to exhibit major shifts in fundamental food web properties.

Aim In fresh waters, most biogeographical understanding of how extreme events such as drought modify biodiversity and ecosystem functioning derives from static, spatial comparisons of ecological communities, between intact and disturbed sites or along stress gradients. Impacts of drought on the development of ecological communities over time remain poorly resolved, with information on parallel trends in community structure and function particularly scarce. In theory, drought could progressively eliminate both species and functional traits, rendering communities increasingly taxonomically and functionally nested subsets of their pre-existing counterparts. Alternatively, drought could create new niche opportunities, producing a continuous turnover of species and traits, or simply constrain natural community succession. Location Dorset, UK. Taxon Aquatic invertebrates. Methods We studied temporal changes in community structure and function in artificial streams over 2 years, comparing drought (frequent drying) with control (constant flow) conditions. Temporal beta diversity was partitioned into turnover and nestedness components, calculated using both presence-absence and abundance data, and analysed using time-lag and null modelling approaches. Results Community development was comparable taxonomically under control and drought conditions, driven primarily by temporal turnover of species. Under control conditions, corresponding trends in functional composition were not apparent, and species turnover was characterized by the progressive replacement of some species by others of equivalent abundance. By contrast, species turnover in disturbed communities was accompanied by both functional turnover and greater loss of individuals, indicating that new colonists were not equivalent, either functionally or numerically, to those they replaced. Furthermore, functional dissimilarities between time points were greatest under drought, and more similar in magnitude to taxonomic dissimilarities, implying that drying reduced the stability and redundancy of functional attributes. Main conclusion A shift to drier climate could disrupt the natural development of stream community structure, and undermine functional stability, at local and biogeographical scales, with potentially significant consequences for ecosystem services provisioning in fresh waters.

Functional traits are increasingly being used to predict extinction risks and range shifts under long‐term climate change scenarios, but have rarely been used to study vulnerability to extreme climatic events, such as supraseasonal droughts. In streams, drought intensification can cross thresholds of habitat loss, where marginal changes in environmental conditions trigger disproportionate biotic responses. However, these thresholds have been studied only from a structural perspective, and the existence of functional nonlinearity remains unknown. We explored trends in invertebrate community functional traits along a gradient of drought intensity, simulated over 18 months, using mesocosms analogous to lowland headwater streams. We modelled the responses of 16 traits based on a priori predictions of trait filtering by drought, and also examined the responses of trait profile groups (TPGs) identified via hierarchical cluster analysis. As responses to drought intensification were both linear and nonlinear, generalized additive models (GAMs) were chosen to model response curves, with the slopes of fitted splines used to detect functional thresholds during drought. Drought triggered significant responses in 12 (75%) of the a priori‐selected traits. Behavioural traits describing movement (dispersal, locomotion) and diet were sensitive to moderate‐intensity drought, as channels fragmented into isolated pools. By comparison, morphological and physiological traits showed little response until surface water was lost, at which point we observed sudden shifts in body size, respiration mode and thermal tolerance. Responses varied widely among TPGs, ranging from population collapses of non‐aerial dispersers as channels fragmented to irruptions of small, eurythermic dietary generalists upon extreme dewatering. Our study demonstrates for the first time that relatively small changes in drought intensity can trigger disproportionately large functional shifts in stream communities, suggesting that traits‐based approaches could be particularly useful for diagnosing catastrophic ecological responses to global change.

Land use and climate change are driving widespread modifications to the biodiverse and functionally unique headwaters of rivers. In temperate and boreal regions, many headwaters drain peatlands where land management and climate change can cause significant soil erosion and peat deposition in rivers. However, effects of peat deposition in river ecosystems remain poorly understood. We provide two lines of evidence-derived from sediment deposition gradients in experimental mesocosms (0-7.5 g/m2 ) and headwaters (0.82-9.67 g/m2 )-for the adverse impact of peat deposition on invertebrate community biodiversity. We found a consistent negative effect of sediment deposition across both the experiment and survey; at the community level, decreases in density (1956 to 56 individuals per m2 in headwaters; mean 823 ± 129 (SE) to 288 ± 115 individuals per m2 in mesocosms) and richness (mean 12 ± 1 to 6 ± 2 taxa in mesocosms) were observed. Sedimentation increased beta diversity amongst experimental replicates and headwaters, reflecting increasing stochasticity amongst tolerant groups in sedimented habitats. With increasing sedimentation, the density of the most common species, Leuctra inermis, declined from 290 ± 60 to 70 ± 30 individuals/m2 on average in mesocosms and >800 individuals/m2 to 0 in the field survey. Traits analysis of mesocosm assemblages suggested biodiversity loss was driven by decreasing abundance of invertebrates with trait combinations sensitive to sedimentation (longer life cycles, active aquatic dispersal of larvae, fixed aquatic eggs, shredding feeding habit). Functional diversity metrics reinforced the idea of more stochastic community assembly under higher sedimentation rates. While mesocosm assemblages showed some compositional differences to surveyed headwaters, ecological responses were consistent across these spatial scales. Our results suggest short-term, small-scale stressor experiments can inform understanding of "real-world" peatland river ecosystems. As climate change and land-use change are expected to enhance peatland erosion, significant alterations to invertebrate biodiversity can be expected where these eroded soils are deposited in rivers.

Climatic extremes are becoming more frequent and intense across much of the globe, potentially transforming the biodiversity and functioning of affected ecosystems. In freshwaters, hydrological extremes such as drought can regulate beta diversity, acting as powerful environmental filters to dictate the complement of species and functional traits found at local and landscape scales. New methods that enable beta diversity and its functional equivalent to be partitioned into turnover (replacement of species/functions) and nestedness‐resultant (gain/loss of species/functions) components may offer novel insights into the parallel impacts of drought on ecosystem structure and function. Using a series of artificial channels (mesocosms) designed to mimic perennial headwater streams, we experimentally manipulated streamflows to simulate a gradient of drought intensity. We then modelled taxonomic and functional turnover and nestedness of macroinvertebrate communities along this gradient, validating direct gradient approaches (bootstrapping, Mantel tests) against null models of nestedness. Drought intensification produced significant environmental distance decay trends (i.e. communities became increasingly taxonomically and functionally dissimilar the more differentially disturbed by drought they were). Taxonomic distance decay was primarily driven by turnover, while the functional trend reflected a combination of richness differences and turnover at different points along the gradient. Taxonomic and functional distance decay slopes were not significantly different, implying that communities were functionally vulnerable to drying. The increased frequency and intensity of droughts predicted under most climate change scenarios could thus profoundly modify not only the structure of running water invertebrate communities, but also the ecosystem functions they underpin.

Urbanisation represents a growing threat to natural communities across the globe. Small aquatic habitats such as ponds are especially vulnerable and are often poorly protected by legislation. Many ponds are threatened by development and pollution from the surrounding landscape, yet their biodiversity and conservation value remain poorly described. Here we report the results of a survey of 30 ponds along an urban land-use gradient in the West Midlands, UK. We outline the environmental conditions of these urban ponds to identify which local and landscape scale environmental variables determine the biodiversity and conservation value of the macroinvertebrate assemblages in the ponds. Cluster analysis identified four groups of ponds with contrasting macroinvertebrate assemblages reflecting differences in macrophyte cover, nutrient status, riparian shading, the nature of the pond edge, surrounding land-use and the availability of other wetland habitats. Pond conservation status varied markedly across the sites. The richest macroinvertebrate assemblages with high conservation value were found in ponds with complex macrophyte stands and floating vegetation with low nutrient concentrations and little surrounding urban land. The most impoverished assemblages were found in highly urban ponds with hard-engineered edges, heavy shading and nutrient rich waters. A random forest classification model revealed that local factors usually had primacy over landscape scale factors in determining pond conservation value, and constitute a priority focus for management.

Sedimentation is a pervasive environmental pressure affecting rivers globally. Headwaters draining catchments rich in organic soils (i.e., peat) are particularly vulnerable to enhanced sedimentation caused by land management and environmental change, yet many of the ecological consequences of peat deposition are poorly understood. We conducted a before‐after‐control‐impact experiment in two rivers draining blanket peatland in Northern England to test the effect of sediment inputs on water quality, macroinvertebrate drift, macroinvertebrate community structure, and ecosystem metabolism. Sediment addition increased concentrations of dissolved organic carbon, total oxidised nitrogen and suspended sediment concentration in rivers, and intensified the total drift of macroinvertebrates particularly at night. By contrast, the abundance and richness of benthic macroinvertebrates were unaffected, except for declines in Coleoptera abundance in one river. The gross primary production of both rivers was strongly suppressed as the benthos was smothered by sediment. Community respiration also declined, albeit by different extents in the two rivers. Our experiment revealed that short‐term pulses of organic sediment in rivers can have broad effects on water quality and biota, from influences on the dispersal of individual organisms to the modification of ecosystem processes. Organic sediments therefore warrant further examination, to include longer observation periods and more sites. It is particularly important to clarify the extent to which impacts extend from peatland streams into larger rivers downstream. Such studies are necessary to inform global management efforts to restore the integrity of river ecosystems under a range of water and biodiversity policy mechanisms.

Pond networks support high levels of biodiversity when compared to other freshwater ecosystems such as rivers, lakes and streams. The persistence of species in these small, sometimes ephemeral, aquatic habitats depends on the dispersal of individuals among ponds in the landscape. However, the number of ponds across the landscape is at a historical low as urbanisation and intensified agricultural practices have led to a substantial loss of ponds (nodes in the pond network) over more than a century. Here, we examine the extent and drivers of pond loss in a heavily urbanised landscape (Birmingham, UK) over 105 years and determine how pond loss influences key structural properties of the pond network using graph theoretic approaches. Specifically, we calculated minimum spanning trees (MST) and performed percolation analyses to determine changes in both the spatial configuration and resilience of the pond network through time. Pond numbers declined by 82% between ca1904 and 2009, such that pond density decreased from 7.1 km-2 to 1.3 km-2. The MST analyses revealed increased distance between ponds in the network (i.e. edge length increased) by up to 49% over the 105-year period, indicating that ponds in the modern landscape (2009) were considerably more isolated, with fewer neighbours. This study demonstrates that graph theory has an excellent potential to inform the management of pond networks in order to support ecological communities that are less vulnerable to environmental change.

The biotic resistance hypothesis provides one of several explanations for the limited biological recovery of streams recovering chemically from acidification. The hypothesis proposes that acidification has changed the presence, abundance and interactions among species in acidified streams to the extent that acid-sensitive colonists cannot re-invade even where acidity has ameliorated. As a first step in testing for biotic resistance in streams, we conducted a field experiment to determine whether the success (growth rate) of acid-sensitive recolonists (mayfly nymphs, Baetis rhodani) is reduced by competition with abundant acid-tolerant residents (stonefly nymphs, Leuctra inermis) in a chemically recovering Welsh stream (UK). Gut contents analysis revealed a marked overlap in resource use between the two species. However, when Baetis was exposed to several (0, 0.25, 0.5 and 1 times ambient) densities of its putative competitor, Leuctra, growth rates of the colonist were not affected by the residents at any of the densities tested. These results do not support the hypothesis that resident species constrain colonist populations by affecting growth rates through competition for limited resources or interference. Further work is required to assess whether independent and/or interactive ecological effects of other common residents might affect colonists in ecosystems recovering from past stressors.

Talk and workshop given at 11th INTECOL Congress, London, 22nd August 2013.

Most research on the effects of environmental change in freshwaters has focused on incremental changes in average conditions, rather than fluctuations or extreme events such as heatwaves, cold snaps, droughts, floods or wildfires, which may have even more profound consequences. Such events are commonly predicted to increase in frequency, intensity and duration with global climate change, with many systems being exposed to conditions with no recent historical precedent. We propose a mechanistic framework for predicting potential impacts of environmental fluctuations on running-water ecosystems by scaling up effects of fluctuations from individuals to entire ecosystems. This framework requires integration of four key components: effects of the environment on individual metabolism, metabolic and biomechanical constraints on fluctuating species interactions, assembly dynamics of local food webs, and mapping the dynamics of the meta-community onto ecosystem function. We illustrate the framework by developing a mathematical model of environmental fluctuations on dynamically assembling food webs. We highlight (currently limited) empirical evidence for emerging insights and theoretical predictions. For example, widely supported predictions about the effects of environmental fluctuations are: high vulnerability of species with high per capita metabolic demands such as large-bodied ones at the top of food webs; simplification of food web network structure and impaired energetic transfer efficiency; and reduced resilience and top-down relative to bottom-up regulation of food web and ecosystem processes. We conclude by identifying key questions and challenges that need to be addressed to develop more accurate and predictive bio-assessments of the effects of fluctuations, and implications of fluctuations for management practices in an increasingly uncertain world.

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Droughts are intensifying under climate change. Research into the resilience of stream food webs to drought now shows that ‘rewiring’ of food web structure in the face of species losses helps to buffer changes to the overall network structure. Droughts are intensifying across the globe1,2, with potentially devastating implications for freshwater ecosystems3,4. We used new network science approaches to investigate drought impacts on stream food webs and explored potential consequences for web robustness to future perturbations. The substructure of the webs was characterized by a core of richly connected species5 surrounded by poorly connected peripheral species. Although drought caused the partial collapse of the food webs6, the loss of the most extinction-prone peripheral species triggered a substantial rewiring of interactions within the networks’ cores. These shifts in species interactions in the core conserved the underlying core/periphery substructure and stability of the drought-impacted webs. When we subsequently perturbed the webs by simulating species loss in silico, the rewired drought webs were as robust as the larger, undisturbed webs. Our research unearths previously unknown compensatory dynamics arising from within the core that could underpin food web stability in the face of environmental perturbations.

Summary Pesticides can have strong deleterious impacts in fresh waters, but understanding how these effects cascade through natural ecosystems, from microbes to apex predators, is limited because research that spans multiple levels of biological organisation is rare. We report how an accidental insecticide spill altered the structure and functioning of a river across levels ranging from genes to ecosystems. We quantified the impacts on assemblages of microbes, diatoms, macroinvertebrates and fish and measured leaf‐litter decomposition rates and microbial functional potential at upstream control and downstream impacted sites 2 months after the spill. Both direct and indirect impacts were evident across multiple levels of organisation and taxa, from the base of the food web to higher trophic levels. At the molecular level, differences in functional gene abundance within the impacted sites reflected a combination of direct and indirect effects of the pesticide, via elevated abundances of microbial populations capable of using chlorpyrifos as a resource (i.e. direct effect) and oxidising ammonia released by decaying macroinvertebrate carcasses (i.e. indirect effect). At the base of the food chains, diatom taxa found only in the impacted sites were an order‐of‐magnitude larger in cell size than the largest comparable taxa in control communities, following the near extirpation of their consumers. Population biomass of the key detritivore Gammarus pulex was markedly lower, as was the rate of litter decomposition in the impacted sites. This was partially compensated for, however, by elevated microbial breakdown, suggesting another indirect food‐web effect of the toxic spill. Although many species exhibited population crashes or local extirpation, total macroinvertebrate biomass and abundance were largely unaffected due to a compensatory elevation in small tolerant taxa such as oligochaetes, and/or taxa which were in their adult aerial life stage at the time of the spill (e.g. chironomids) and thus avoided contact with the polluted waters and were able to repopulate the river quickly. Mass–abundance scaling of trophic links between consumers and resources revealed extensive restructuring within the food web. This case study shows that pesticides can affect food‐web structure and ecosystem functioning, both directly and indirectly across levels of biological organisation. It also demonstrates how an integrated assessment approach, as adopted here, can elucidate links between microbiota, macroinvertebrates and fish, for instance, thus improving our understanding of the range of biological consequences of chemical contamination in natural ecosystems.