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Context: Community composition, environmental variation, and spatial structuring can influence ecosystem functioning, and ecosystem service delivery. While the role of space in regulating ecosystem functioning is well recognised in theory, it is rarely considered explicitly in empirical studies. Objectives: We evaluated the role of spatial structuring within and between regions in explaining the functioning of 36 reference and human-impacted streams. Methods: We gathered information on regional and local environmental variables, communities (taxonomy and traits), and used variance partitioning analysis to explain seven indicators of ecosystem functioning. Results: Variation in functional indicators was explained not only by environmental variables and community composition, but also by geographic position, with sometimes high joint variation among the explanatory factors. This suggests spatial structuring in ecosystem functioning beyond that attributable to species sorting along environmental gradients. Spatial structuring at the within-region scale potentially arose from movements of species and materials among habitat patches. Spatial structuring at the between-region scale was more pervasive, occurring both in analyses of individual ecosystem processes and of the full functional matrix, and is likely to partly reflect phenotypic variation in the traits of functionally important species. Characterising communities by their traits rather than taxonomy did not increase the total variation explained, but did allow for a better discrimination of the role of space. Conclusions: These results demonstrate the value of accounting for the role of spatial structuring to increase explanatory power in studies of ecosystem processes, and underpin more robust management of the ecosystem services supported by those processes.

River regulation for hydropower is undertaken worldwide, causing profound alterations to hydrological regimes and running water habitats. Regulated catchments are often subjected to additional stressors, arising inter alia from agriculture, forestry and industry, which are likely to interact with impacts of river regulation on fish and other biota. Such interactions are poorly understood, hindering planning of effective mitigation and restoration. We investigated fish responses to increased discharge (as a restoration measure) in regulated rivers in Sweden. We compiled electrofishing data from river channels downstream of hydropower dams, each of which either has or lacks a mandated minimum discharge corresponding to c. 5% of pre‐regulation discharge. We further analysed interactions between flow restoration and co‐occurring local and regional stressors. River channels without a mandated minimum discharge were characterized by a low diversity of fish species with traits favouring persistence under unpredictable environmental conditions, including omnivory, short life cycles and small size. Additional stressors further reduced diversity and increased dominance by broad‐niched, opportunistic species. Both the presence and magnitude of a mandated minimum discharge were positively related to fish diversity and density, and the relative density of three economically and recreationally valuable species. However, the size of these relationships frequently varied with the presence of additional stressors. Increasing levels of hydrological degradation and reduced connectivity at the catchment scale reduced positive flow–ecology relationships and hindered the restoration of fish communities towards reference conditions. However, application of a mandated minimum discharge also assisted in mitigating impacts of some co‐occurring stressors, especially reduced riparian integrity. Synthesis and applications. Additional stressors can strongly influence the outcomes of flow restoration for fish community diversity and composition. Our approach combining fish species and trait data from multiple flow restoration projects with information on additional stressors yielded valuable insights into factors affecting flow restoration success, useful for (a) identifying the systems most likely to benefit from mandated minimum flows, (b) modelling influences of multiple stressors on flow–ecology relationships, (c) prioritizing additional measures to manage co‐occurring stressors and enhance outcomes from flow restoration.

Summary The spatial organisation of biotic communities derives from factors operating at a wide range of spatial and temporal scales. Despite strong scientific evidence of prevalent spatial control of community composition in freshwater ecosystems, local environmental factors are often considered as the main drivers of community change. Furthermore, taxonomic approaches are most frequently used, and few studies have compared the relative importance of local and regional control of trait versus the taxonomic composition in stream ecosystems. Using a spatially dense data set covering all stream sizes in a lowland European region of c. 42 000 km2 and three organism groups (macrophytes, macroinvertebrates and fishes), we compared the relative importance of spatial and environmental determinants of species and trait composition in the study streams, classified into headwaters (stream order 1–2) and downstream sites (stream order >2). We hypothesised that (i) there is a higher correspondence between environmental conditions and trait composition than with species composition, (ii) dispersal limitation (pure spatial structuring) is greater in headwaters than in downstream sites and (iii) dispersal limitation (pure spatial structuring) is weakest for macroinvertebrates, intermediate for macrophytes and strongest for fishes. The most consistent pattern across organisms and stream order groups was a higher correspondence between environmental variation and trait composition as well as a higher number of environmental variables significantly related to trait composition than with species composition (hypothesis 1). Spatial structuring peaked in headwater macrophyte communities and downstream fish communities (hypotheses 2 & 3) – a pattern that was amplified when separate analyses of traits describing species dispersal potential were undertaken. Our study highlights the potential of traits to capture multiple environmental changes in stream ecosystems and illustrates how organism-specific and highly context-dependent patterns in community organisation can emerge as a consequence of interactions between habitat connectivity (i.e. top versus lower parts of the stream network) and organism dispersal potential.

Historically, close attention has been paid to negative impacts associated with nutrient loads to streams and rivers, but today hydromorphological alterations are considered increasingly implicated when lowland streams do not achieve good ecological status. Here, we explore if trait-abundance patterns of aquatic plants change along gradients in hydromorphological degradation and eutrophication in lowland stream sites located in Denmark. Specifically, we hypothesised that: i) changes in trait-abundance patterns occur along gradients in hydromorphological degradation and ii) trait-abundance patterns can serve to disentangle effects of eutrophication and hydromorphological degradation in lowland streams reflecting that the mechanisms behind changes differ. We used monitoring data from a total of 147 stream reaches with combined data on aquatic plant species abundance, catchment land use, hydromorphological alterations (i.e. planform, cross section, weed cutting) and water chemistry parameters. Traits related to life form, dispersal, reproduction and survival together with ecological preference values for nutrients and light (Ellenberg N and L) were allocated to 41 species representing 79% of the total species pool. We found clear evidence that habitat degradation (hydromorphological alterations and eutrophication) mediated selective changes in the trait-abundance patterns of the plant community. Specific traits could distinguish hydromorphological degradation (free-floating, surface; anchored floating leaves; anchored heterophylly) from eutrophication (free-floating, submerged; leaf area). We provide a conceptual framework for interpretation of how eutrophication and hydromorphological degradation interact and how this is reflected in trait-abundance patterns in aquatic plant communities in lowland streams. Our findings support the merit of trait-based approaches in biomonitoring as they shed light on mechanisms controlling structural changes under environmental stress. The ability to disentangle several stressors is particularly important in lowland stream environments where several stressors act in concert since the impact of the most important stressor can be targeted first, which is essential to improve the ecological status.

Most river restoration projects have applied relatively small-scale measures focused on improving specific instream conditions, with only limited outcomes for biodiversity in rivers and their adjacent riparian habitats. Here, we investigate the effects of both small- and large-scale restoration projects on floodplain vegetation across 20 European catchments. We focused on the roles of different restoration parameters (i.e., the number, spatial extent and type of restoration measure applied and restoration age) and specific environmental characteristics in regulating changes in plant diversity and trait composition following restoration. Among restoration characteristics, restoration type was the only significant determinant of plant community responses, with stream channel widening having the strongest effects, particularly on the diversity and composition of species traits favoured by increases in physical disturbance (e.g. flooding) and open habitat patch availability (e.g. plant growth form, life strategy and life span). Of the environmental variables, altitude and discharge were positively and most strongly related to responses of both species and trait diversity. Our results emphasise the value of (i) choosing relevant restoration measures that affect environmental conditions of importance for the target organism group and (ii) conducting restoration projects in environmental settings where the likelihood of restoration "success" is maximised.

Despite intensive efforts for more than a decade to develop Water Framework-compliant assessment systems, shortcomings continue to appear. In particular, the lack of reference conditions has hindered the development of assessment systems capturing the heart of the WFD – that ecological status should be set as the deviation from the natural, undisturbed condition. Recently, the Danish Stream Plant Index (DSPI) was developed. This system contrasts existing systems in that it builds on an expert interpretation of the normative definitions of ecological status classes in the WFD without taking pressure-impact relationships into account. Here, we substantiate the approach taken in the development of DSPI and examine if the DSPI class decreases with increasing level of anthropogenic stress and, additionally, if the deviation from the natural undisturbed condition increases with decreasing DSPI class sensu WFD using trait composition of plant assemblages from Danish streams around year 1900 as a reference. We furthermore examine the trait composition of the vegetation in sites classified into different DSPI status classes to explore if predictable patterns exist that can be used to identify the ultimate cause(s) of failure to meet ecological goals and help guide the selection of appropriate mitigation measures. We observed that DSPI declined with several parameters indicative of environmental stress in Danish streams and, furthermore, that the deviation from the natural undisturbed condition regarding the trait composition of plant communities declined with increasing DSPI, implying that the trait composition of plant communities in the high DSPI status class was most similar to those occurring in Danish streams around year 1900. We also found that trait characteristics capable of disentangling important stressors in Danish streams varied consistently among sites classified into different DSPI classes. Based on our findings, we call for new thinking. We suggest that more effort should be directed at describing reference conditions and interpreting the normative definitions of good, moderate, poor and bad instead of focusing solely on developing assessment systems using pressure-impact frameworks. We find this particularly important with respect to streams since these are seldom impacted by only a single stressor. This article is protected by copyright. All rights reserved.

Stream assemblages are structured by a combination of local (environmental filtering and biotic interactions) and regional factors (e.g., dispersal related processes). The relative importance of environmental and spatial (i.e., regional) factors structuring stream assemblages has been frequently assessed in previous large-scale studies, but biotic predictors (potentially reflecting local biotic interactions) have rarely been included. Diatoms may be useful for studying the effect of trophic interactions on community structure since: (1) a majority of experimental studies shows significant grazing effects on diatom species composition, and (2) assemblages can be divided into guilds that have different susceptibility to grazing. We used a dataset from boreal headwater streams in south-central Sweden (covering a spatial extent of ∼14000 km(2)), which included information about diatom taxonomic composition, abundance of invertebrate grazers (biotic factor), environmental (physicochemical) and spatial factors (obtained through spatial eigenfunction analyses). We assessed the relative importance of environmental, biotic, and spatial factors structuring diatom assemblages, and performed separate analyses on different diatom guilds. Our results showed that the diatom assemblages were mainly structured by environmental factors. However, unique spatial and biological gradients, specific to different guilds and unrelated to each other, were also evident. We conclude that biological predictors, in combination with environmental and spatial variables, can reveal a more complete picture of the local vs. regional control of species assemblages in lotic environments. Biotic factors should therefore not be overlooked in applied research since they can capture additional local control and therefore increase accuracy and performance of predictive models. The inclusion of biotic predictors did, however, not significantly influence the unique fraction explained by spatial factors, which suggests low bias in previous assessments of unique regional control of stream assemblages.