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Aim: An intensively debated issue in macroecology is whether unicellular organisms show biogeographic patterns different from those of macroorganisms. One aspect of this debate addresses beta diversity, that is, do microbial organisms exhibit distance-decay patterns similar to those of macroorganisms? And if so, is the decay of community similarity caused by spatially limited dispersal or by niche-related factors? We studied the community similarity of stream diatoms, macroinvertebrates and bryophytes across the same set of sites in relation to environmental and geographic distance. Location: A geographical gradient of c. 1100 km in Finland. Methods: We first identified the subset of environmental variables that produced the highest correlation with community similarities for each taxonomic group. Based on these variables, we used partial Mantel tests to separate the independent influences of environmental and geographical distance for distance decay of community similarity, separately for diatoms, bryophytes and macroinvertebrates. Finally, macroinvertebrates were divided into three groups based on their different dispersal categories and a partial Mantel test was used to assess whether each of these groups were differently affected by environmental versus geographic distance, i.e. is dispersal a key factor in tests of niche versus neutral models. Results: The level of environmental control was by far the strongest for diatoms; however, all groups were controlled more by environmental factors than by limited dispersal. Macroinvertebrate species with low dispersal ability were significantly related to geographic distance, while more effective dispersers showed no relationship to geography but were instead strongly related to environmental distance. Main conclusions: Our results suggest that patterns between macro- and microorganisms are not fundamentally different, but the level of environmental control varies according to dispersal ability. The relative importance of niche versus dispersal processes is not simply a function of organism size but other traits (e.g. life-history type, dispersal capacity) may obscure this relationship.

Aim We examined the relative contributions of spatial gradients and local environmental conditions to macroinvertebrate assemblages of boreal headwater streams at three hierarchical extents: bioregion, ecoregion and drainage system. We also aimed to identify the environmental variables most strongly related to assemblage structure at each study scale, and to assess how the importance of these variables is related to regional context and spatial structuring at different scales. Location Northern Finland (62-68° N, 25-32° E). Methods Variation in macroinvertebrate data was partitioned using partial canonical correspondence analysis into components explained by spatial variables (nine terms from the cubic trend surface regression), local environmental variables (15 variables) and spatially structured environmental variation. Results The strength of the relationship between assemblage structure and local environmental variables increased with decreasing spatial extent, whereas assemblage variation related to spatial variables and spatially structured environmental variation showed the opposite pattern. At the largest extents, spatial variation was related to latitudinal gradients, whereas spatial autocorrelation among neighbouring streams was the likely mechanism creating spatial structure within drainage systems. Only stream size and water acidity were consistently important in explaining assemblage structure at all study scales, while the importance of other environmental variables was more context-dependent. Main conclusions The importance of local environmental factors in explaining macroinvertebrate assemblage structure increases with decreasing spatial extent. This scale-related pattern is not caused solely by changes in study extent, however, but also by variable sample sizes at different regional extents. The importance of environmental gradients is context-dependent and few factors are likely to be universally important correlates of macroinvertebrate assemblage structure. Finally, our results suggest that bioassessment should give due attention to spatial structuring of stream assemblages, because important assemblage gradients may not only be related to local factors but also to biogeographical constraints and neighbourhood dispersal processes.

To define whether the beta diversity of stream invertebrate communities in New Zealand exhibits geographical variation unexplained by variation in gamma diversity and, if so, what mechanisms (productivity, habitat heterogeneity, dispersal limitation, disturbance) best explain the observed broad‐scale beta diversity patterns. We sampled 120 streams across eight regions (stream catchments), spanning a north–south gradient of 12° of latitude, and calculated beta diversity (with both species richness and abundance data) for each region. We explored through a null model if beta diversity deviates from the expectation of stochastic assembly processes and whether the magnitude of the deviation varies geographically. We then performed multimodel inference analysis on the key environmental drivers of beta diversity, using Akaike's information criterion and model and predictor weights to select the best model(s) explaining beta diversity. Beta diversity was, unexpectedly, highest in the South Island. The null model analysis revealed that beta diversity was greater than expected by chance in all eight regions, but the magnitude of beta deviation was higher in the South Island, suggesting differences in environmental filtering and/or dispersal limitation between North and South Island. Habitat heterogeneity was the predominant driver of beta diversity of stream macroinvertebrates, with productivity having a secondary, and negative, contribution. This is one of the first studies accounting for stochastic effects while examining the ecological drivers of beta diversity. Our results suggest that local environmental heterogeneity may be the strongest determinant of beta diversity of stream invertebrates, more so than regional‐ or landscape‐scale variables. We explored the geographic distribution of β diversity of New Zealand stream invertebrates and studied whether their β diversity can be explained by one or more of the general mechanisms (productivity, habitat heterogeneity, dispersal limitation, disturbance) proposed to explain broad‐scale β diversity patterns. Our results suggest that local environmental heterogeneity may be the strongest determinant of β diversity of stream communities, more so than regional‐ or landscape‐scale variables.

Protected areas are a principal conservation tool for addressing biodiversity loss. Such protection is especially needed in freshwaters, given their greater biodiversity losses compared to terrestrial and marine ecosystems. However, broad-scale evaluations of protected area effectiveness for freshwater biodiversity are lacking. Here, we provide a continental-scale analysis of the relationship between protected areas and freshwater biodiversity using 1,754 river invertebrate community time series sampled between 1986 and 2022 across ten European countries. Protected areas primarily benefited poor-quality communities (indicative of higher human impacts) that were protected, or that gained protection, across a substantial proportion of their upstream catchment. Protection had little to no influence on moderate- and high-quality communities, although high-quality communities potentially provide less scope for effect. Our results reveal the overall limited effectiveness of current protected areas for freshwater biodiversity, likely because they are typically designed and managed to achieve terrestrial conservation goals. Broadly improving effectiveness for freshwater biodiversity requires catchment-scale management approaches involving larger and more continuous upstream protection, and efforts to address remaining stressors. These approaches would also benefit connected terrestrial and coastal ecosystems, thus generally helping bend the curve of global biodiversity loss. The effects of current protected areas on freshwater biodiversity are poorly understood. Here, the authors show that European protected areas have overall limited influence on changes in river biodiversity, underscoring the urgent need for improved effectiveness.

Increasing nutrient loads are impairing water quality and ecological status of aquatic ecosystems globally, but their effects on Arctic freshwaters remain less studied. Nutrients in Arctic freshwaters are increasing due to expanding anthropogenic land use and climate change induced alterations in nutrient leaching and transport. Also increased occurrences of non-native semelparous Pacific pink salmon ( Oncorhynchus gorbuscha) in the northern European rivers can enhance nutrient availability. These additional nutrients may be readily available for aquatic primary producers in often inherently (ultra-)oligotrophic Arctic freshwaters. We used an experimental field set-up with nutrient diffusing substrates to study temporal and spatial differences in the response of primary producers to increased phosphorus (P) and nitrogen (N) levels in Arctic rivers. Additional NO 3 -N enhanced algal growth (chlorophyll -a (Chl- a) amount) of epilithic biofilm. PO 4 -P alone had no effect, indicating N as the primary limiting factor for algal growth. This was particularly evident in autumn while early summer showed evidence for secondary P limitation. The response to nutrient additions was mostly dictated by the background availability of inorganic N. Our results suggest that Arctic rivers are highly sensitive to nutrient additions, especially N. Thus, any increase in nutrients can boost algal growth causing cascading effects throughout Arctic ecosystems.

1. Multiscale determinants of diversity and the relationship between regional (RSR) and local richness (LSR) have recently attracted increased attention, yet such studies on stream organisms remain scarce. We studied the relationships among RSR, β-diversity, LSR and local environmental variables in 120 headwater streams in Finland. Approximately similar-sized areas of eight drainage systems were defined as regions, and 15 stream riffles (= locality) per region were sampled. 2. RSR showed a strong positive relationship with mean LSR (R2= 0·686), and there was no sign of curvilinearity within the observed range of RSR. RSR was also positively, although non-significantly, related to β-diversity (r = 0·662). 3. In stepwise regression, RSR was the first variable to enter the model, and a model incorporating RSR and stream width explained 32·5% of variation in LSR. If RSR was omitted from the model, then stream width emerged as the most important variable, followed by water pH, which together accounted for 20·6% of variation in LSR. 4. At the within-region scale, different variables were important in accounting for variation in LSR. Factors correlated with LSR reflected either stream size, spatial heterogeneity, adverse water chemistry conditions (pH), or a limiting resource base for macroinvertebrates (nutrient concentrations). 5. The major role of RSR in setting the upper limit to LSR suggests that macroinvertebrate assemblages of boreal headwater streams are unsaturated. This finding is supported by evidence from experimental studies, where it has been shown that competitive interactions among stream macroinvertebrates are effective only at very small spatial scales, and competitive exclusion is prevented typically by frequent disturbances. However, although RSR was generally the most influential variable contributing to LSR, it is far from clear whether RSR consistently sets the limits to LSR, or vice versa. For instance, uniformly adverse water chemistry conditions across a region may lead to low levels of local richness and low species turnover among sites, leading eventually to an impoverished regional species pool. 6. These findings do not deny the importance of local factors, but emphasize that understanding the organization of stream benthic communities requires simultaneous examination of factors prevailing at multiple spatial scales.

An ability to understand and predict invasions is elemental for controlling the detrimental effects of introduced organisms on native biota. In eastern North America, European brown trout generally dominates over, and eventually replaces, the native brook trout. We show here that in northern Europe the pattern of replacement between these two species is reversed: when transferred to North European streams, brook trout spread extensively and partially replaced the native brown trout. The effect of brook trout on brown trout was habitat-specific: brook trout excluded the native species only in small tributary streams where the reproduction of brown trout was severely reduced, whereas in larger streams brown trout was largely unaffected. Thus, the pattern of coexistence among the two salmonids in our study area is approaching that typically observed in North American streams. In both areas, brook trout ultimately settles in small headwater streams, but the process of replacement differs profoundly: in North Europe, brook trout replaces brown trout in headwater streams, whereas in North America these same streams are the ultimate refuge area for brook trout under the invasion pressure by brown trout. Our results underline the importance of knowing species' niche characteristics to explain and predict biological invasions.

Concentrations of dissolved organic carbon (DOC) have increased over the past few decades, causing freshwater browning. Impacts of browning on biodiversity have been little studied, despite many of the individual stressors associated with browning being known to control freshwater communities. We explored the responses of benthic invertebrates along a wide gradient of DOC concentrations (3.6–27 mg L−1) in 63 boreal streams variously impacted by peatland drainage or peat production. DOC was a prime determinant of macroinvertebrate diversity and abundance, with the strongest negative response in algal scrapers. Threshold indicator taxa analysis indicated an abrupt community change at 12–13 mg DOC L−1, with only four taxa increasing, while 13 taxa decreased along the gradient. Our findings of both a gradual loss and abrupt change of biodiversity along a browning gradient provide a benchmark against which changes to stream biodiversity relative to the predicted browning trend can be gauged.

1. There is compelling evidence that anthropogenic disturbance can decrease biodiversity and impair ecological functioning. A major challenge to biodiversity–ecosystem function research is to disentangle the effects of biodiversity loss on ecosystem functions from the direct effects of human disturbance. 2. We studied the influence of human disturbance (acidification and eutrophication) and a natural stressor (low pH due to bedrock geology) on leaf-shredding macroinvertebrates, fungal decomposers and leaf decomposition rates in boreal streams. We used pyrosequencing techniques to determine fungal richness and assemblage structure. 3. Decomposition rates were higher in anthropogenically disturbed than in circumneutral reference or naturally acidic sites, but did not differ between the latter two groups. Macroinvertebrate richness was higher in circumneutral than in human-impacted or naturally acidic sites, and shredder evenness was highest in circumneutral sites. Fungal evenness was also lower in human-disturbed than in reference sites, whereas fungal richness did not vary among site groups. 4. Decomposition rate in fine-mesh bags was related positively to current velocity and fungal dominance, while in coarse-mesh bags, it was related positively to total phosphorus. In anthropogenically disturbed streams, the effects of low pH were overridden by eutrophication, and increased decomposition rates resulted from disturbance-induced increase in species dominance rather than richness. Furthermore, decomposition rates were positively correlated with abundances of dominant taxa, suggesting that ecosystem processes may be driven by a few key species. 5. Synthesis and applications. Our results suggest that leaf decomposition rates are insensitive to natural background variation, supporting the use of decomposition assays, preferably accompanied by molecular analysis of fungal assemblages, to assess stream ecosystem health. Instead of focusing solely on diversity, however, more emphasis should be placed at changes in dominance patterns, particularly if management aims are to improve stream ecosystem functioning.

1. There is controversy over how the success of ecological restoration should be measured. Traditionally, emphasis has been placed on species diversity and other community attributes, whereas the restoration of ecosystem processes has received less attention. Here, we combine replicated field experiments and a field survey to provide an ecosystem-level measure of stream restoration success. 2. Numerous headwater streams in Finland, and in many other parts of the world, have been channelized for timber transport, resulting in channels with simplified structure and flow. Recently, programmes have been launched to restore these streams to their pre-channelization condition. While the efficacy of restoration in improving fish habitat has been tested, little is known about effects on other stream biota or on the retention of leaf litter, despite its importance in trophic dynamics of forested headwater streams. Using a before-after-control-intervention (BACI) designed experiment with multiple reference and experimental streams, we examined restoration-induced changes in retention efficiency by conducting leaf-release experiments before (1993) and after (1996) restoration. 3. Substrate heterogeneity increased, but moss cover decreased dramatically following restoration. Retention efficiency in restored streams was higher than in channelized, but lower than in natural, streams. Algae-feeding scrapers were the only macroinvertebrate group whose density increased significantly after restoration. 4. Aquatic mosses were a key retentive feature in both channelized and natural streams, but their importance to retention was strikingly reduced by restoration. During restoration work, mosses are detached from large areas of the stream bed, exposing bare stone surfaces for colonization by periphytic algae. 5. A more effective restoration technique would involve the use of moss transplants, or the addition of large woody debris, to increase retentiveness and thus enhance the availability of organic material to benthic consumers. This case study on rivers illustrates how restoration projects benefit from an ecosystem perspective and from measures of ecosystem processes in assessing restoration success.