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Climate warming can lead to a replacement of species that favour cold temperatures by species that favour warm temperatures. However, the implications of such thermic shifts for the functioning of ecosystems remain poorly understood. Here, we used stream macroinvertebrate biological and ecological traits to quantify the relative contribution of cold, intermediate and warm temperature-adapted taxa to changes in community functional diversity (FD) using a dataset of 3781 samples collected in Central Europe over 25 years, from 1990 to 2014. Our analyses indicated that functional diversity of stream macroinvertebrate communities increased over the study period. This gain was driven by a net 39 % increase in the richness of taxa that favour intermediate temperatures, which comprise the highest share in the community, and to a 97 % increase in the richness of taxa that favour warm temperatures. These warm temperature-adapted taxa displayed a distinct and more diverse suite of functional traits compared to the cold temperature-adapted group and thus contributed disproportionately to local FD on a per-taxon basis. At the same time, taxonomic beta-diversity declined significantly within each thermal group, in association with increasing local taxon richness. This study shows that over recent decades, small low-mountain streams in Central Europe have experienced a process of thermophilization and increasing functional diversity at local scales. However, a progressive homogenisation occurred at the regional scale, with communities converging towards similar taxonomic composition. As the reported increase in local functional diversity can be attributed mostly to the intermediate temperature-adapted taxa and a few expanding warm temperature-adapted taxa, these patterns could mask more subtle loss of sensitive cold temperature-adapted taxa with irreplaceable functional traits. In light of increasing climate warming, preservation of cold habitat refuges, should be considered a priority in river conservation., Competing Interests: Declaration of competing interest This manuscript has not been submitted to, nor is under review at, another journal or other publishing venue. The authors have no affiliation with any organization with a direct or indirect financial interest in the subject matter discussed in the manuscript., (Copyright © 2023 Elsevier B.V. All rights reserved.)

Owing to a long history of anthropogenic pressures, freshwater ecosystems are among the most vulnerable to biodiversity loss 1 . Mitigation measures, including wastewater treatment and hydromorphological restoration, have aimed to improve environmental quality and foster the recovery of freshwater biodiversity 2 . Here, using 1,816 time series of freshwater invertebrate communities collected across 22 European countries between 1968 and 2020, we quantified temporal trends in taxonomic and functional diversity and their responses to environmental pressures and gradients. We observed overall increases in taxon richness (0.73% per year), functional richness (2.4% per year) and abundance (1.17% per year). However, these increases primarily occurred before the 2010s, and have since plateaued. Freshwater communities downstream of dams, urban areas and cropland were less likely to experience recovery. Communities at sites with faster rates of warming had fewer gains in taxon richness, functional richness and abundance. Although biodiversity gains in the 1990s and 2000s probably reflect the effectiveness of water-quality improvements and restoration projects, the decelerating trajectory in the 2010s suggests that the current measures offer diminishing returns. Given new and persistent pressures on freshwater ecosystems, including emerging pollutants, climate change and the spread of invasive species, we call for additional mitigation to revive the recovery of freshwater biodiversity., (© 2023. The Author(s).)

Biotic homogenization is one of the key aspects of the current biodiversity crisis. Here we analyzed the trends of three facets of niche homogenization, i.e. niche specialization, trait α-diversity and spatial β-diversity, over a period of 25 years (1990-2014) using a large dataset of 3782 stream benthic invertebrate samples collected from central European low-mountain streams. We studied a set of traits describing the ecological niche of species and their functions: body size, feeding groups, substrate preferences, flow preferences, stream zonation preferences and saprobity. Trait composition changed significantly during the study period, and we identified an overall increase in niche homogenization. Specifically, community niche specialization significantly decreased by 20.3% over the 25-year period, with declines ranging from -16.0 to -40.9% for zonation-, flow-, substrate-preferences, body size and feeding traits. Trait diversity did not change significantly, although we recorded significant decreases by -14.2% and -10.2% for flow- and substrate-preference and increases by 5.8% and 22.6% for feeding traits and zonation preference over the study period. Trait spatial β-diversity significantly decreased by -53.0%, with substrate-preference, feeding groups and flow-preference traits declining from -61.9% to -75.3% over the study period. This increased niche homogenization is likely driven by the increase of down-stream typical taxa, which are favored by warming temperatures. Further, it is in apparent contradiction with the recorded increase in abundance (+35.9%) and taxonomic richness (+39.2%) over the same period. Even such increases do not safeguard communities from undergoing niche homogenization, indicating that recovery processes may differ with regard to community taxonomic composition and traits. Our results emphasize the complexity of community responses to global change and warrant caution when founding conclusions based solely on single community metrics., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 The Authors. Published by Elsevier B.V. All rights reserved.)

Climate warming often results in species range shifts, biodiversity loss and accumulated climatic debts of biota (i.e. slower changes in biota than in temperature). Here, we analyzed the changes in community composition and temperature signature of stream invertebrate communities over 25 years (1990-2014), based on a large set of samples (n = 3782) over large elevation, latitudinal and longitudinal gradients in central Europe. Although warming was moderate (average 0.5 °C), we found a strong reorganization of stream invertebrate communities. Total abundance (+35.9%) and richness (+39.2%) significantly increased. The share of abundance (TA) and taxonomic richness (TR) of warm-dwelling taxa (TA: +73.2%; TR: +60.2%) and medium-temperature-dwelling taxa (TA: +0.4%; TR: +5.8%) increased too, while cold-dwelling taxa declined (TA: -61.5%; TR: -47.3%). The community temperature index, representing the temperature signature of stream invertebrate communities, increased at a similar pace to physical temperature, indicating a thermophilization of the communities and, for the first time, no climatic debt. The strongest changes occurred along the altitudinal gradient, suggesting that stream invertebrates use the spatial configuration of river networks to track their temperature niche uphill. Yet, this may soon come to an end due to the summit trap effect. Our results indicate an ongoing process of replacement of cold-adapted species by thermophilic species at only 0.5 °C warming, which is particularly alarming in the light of the more drastic climate warming projected for coming decades., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Long-term observations on riverine benthic invertebrate communities enable assessments of the potential impacts of global change on stream ecosystems. Besides increasing average temperatures, many studies predict greater temperature extremes and intense precipitation events as a consequence of climate change. In this study we examined long-term observation data (10-32years) of 26 streams and rivers from four ecoregions in the European Long-Term Ecological Research (LTER) network, to investigate invertebrate community responses to changing climatic conditions. We used functional trait and multi-taxonomic analyses and combined examinations of general long-term changes in communities with detailed analyses of the impact of different climatic drivers (i.e., various temperature and precipitation variables) by focusing on the response of communities to climatic conditions of the previous year. Taxa and ecoregions differed substantially in their response to climate change conditions. We did not observe any trend of changes in total taxonomic richness or overall abundance over time or with increasing temperatures, which reflects a compensatory turnover in the composition of communities; sensitive Plecoptera decreased in response to warmer years and Ephemeroptera increased in northern regions. Invasive species increased with an increasing number of extreme days which also caused an apparent upstream community movement. The observed changes in functional feeding group diversity indicate that climate change may be associated with changes in trophic interactions within aquatic food webs. These findings highlight the vulnerability of riverine ecosystems to climate change and emphasize the need to further explore the interactive effects of climate change variables with other local stressors to develop appropriate conservation measures., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Although ecological restoration generally succeeds in increasing physical heterogeneity, many projects fail to enhance biota. Researchers have suggested several possible explanations, including insufficient restoration intensity, or time-lags in ecological responses that prevent detection of significant changes in short-term monitoring programs. This study aims to evaluate whether benthic macroinvertebrate communities responded to an expanded set of stream restoration measures within a study period of one to five years after completion of the restoration project. We studied 10 forest streams in northern Sweden that were channelized in the past for timber floating. Managers subjected six of these streams to habitat restoration, on each of these we selected two reaches, located in close proximity but differing in restoration intensity. In "basic" restored reaches, the restoration managers broke up the channelized banks and returned cobbles and small boulders to the main channel. In "enhanced" restoration reaches, they added additional large wood and boulders to reaches previously subjected to basic restoration, and rehabilitated gravel beds. The remaining four streams were not restored, and thus represent the baseline impacted (channelized) condition. We surveyed stream benthic assemblages before the enhanced restoration (year 2010) and three times afterward between 2011 and 2015. Five years after restoration, macroinvertebrate assemblages at the enhanced restored reaches were more differentiated from channelized conditions than those at basic-restored reaches. This reflected increased relative abundances of the insect orders Ephemeroptera and Trichoptera and the bivalve molluscs Sphaeriidae and decreased relative abundances of Chironomidae (Diptera). Analysis of functional traits provided further insights on the mechanistic explanations driving the recovery, e.g., indicating that the augmented channel retention capacity at enhanced restored reaches favored taxa adapted to slow flow conditions and more effectively retained passive aquatic dispersers. The increased restoration intensity in enhanced restored reaches has resulted in shifts in the composition of benthic macroinvertebrate assemblages, including increases in more sensitive taxa. These shifts became fully apparent five years after the enhanced restoration. Our results emphasize the value of longer-term monitoring to assess ecological responses following restoration, and of undertaking additional restoration as a valuable management option for previously restored sites that failed to achieve biotic recovery., (© 2017 by the Ecological Society of America.)

A better management of nutrient inflows into lakes has led to an improvement in their conditions (i.e. reversal of eutrophication) and the effects of this on macroinvertebrate communities that inhabit different lake-depth zones is largely unknown. This paper reports a comparison of macroinvertebrate communities living in the eulittoral, infralittoral and sublittoral/profundal zones of Lake Nemi (Central Italy) before and after its natural recovery from eutrophication following the deviation of domestic wastewater. The infralittoral zone responded more rapidly than the other two depth-zones to the improved ecological conditions, as shown by larger differences in community composition between the two periods. In the eulittoral sand, the combined effects of hydromorphological pressures and reversal of eutrophication hindered the biotic response. In the eulittoral and infralittoral zones, typical taxa of mesotrophic waters appeared or increased their abundances after the eutrophication reversal. Benthic invertebrate response was slower in the sublittoral/profundal zone due to deoxygenation that continued to prevail in the deepest area of the lake during summer. However, both tolerant and more sensitive taxa were collected there for the first time. After the reversal of eutrophication, the percentage of molluscan+large crustaceans increased in the infralittoral zone, whereas the oligochaete/chironomid ratio decreased in both sublittoral/profundal and infralittoral zones. Functional feeding metrics (percentages of filter-feeders, collector-gatherers, miners and scrapers/grazers) differently tracked the reversal of eutrophication in the three depth-zones probably according to the effects of the reduction of nutrients on food-web structure influencing macroinvertebrates. Biological Monitoring Working Party (BMWP) and the Average Score Per Taxon (ASPT) seemed to respond to eutrophication reversal only in the sublittoral/profundal zone, where deoxygenation plays a major role as a structuring agent of the community. Our results suggest that the effects of reversal of eutrophication can be better assessed by examining the response of the communities belonging to each zone individually., (Copyright © 2016 Elsevier B.V. All rights reserved.)

Large wood (LW), defined as pieces of wood greater than 10 cm in diameter and 1 m long, is well known to alter river hydromorphology and the availability of potential food resources for consumers. However, there has been a lack of studies investigating whether these can cause shifts in the trophic base, which may explain alterations to the total abundance and taxonomic structure of the macroinvertebrate assemblage., We aimed to determine how the presence of LW altered the trophic base of the macroinvertebrate consumer assemblage in a lowland river, and to provide a methodological comparison of two assimilation-based food web methods: stable-isotope analysis (SIA) and fatty-acid biomarker profiles (FA). To do so, we quantified the contribution of trophic resources to the diets of macroinvertebrates colonising the surface of LW, present in this study as single logs, and surrounding bed sediments with those from bed sediments of a nearby control site with minimal amounts of LW., SIA showed that the macroinvertebrate food web, even for non-filter feeding taxa, was mostly sustained by seston exported from a lake 1 km upstream, highlighting a high degree of lake-river coupling. The presence of wood altered the trophic base from being predominantly seston-supported to one with increased support from epixylic autochthonous production (i.e. periphyton and bryophytes on wood). Terrestrial matter (i.e. leaves and grass) and organic sediments were a relatively unimportant fraction of the trophic base (<10%) in all locations., FA did not directly track the influence of seston, but instead differentiated between overall allochthonous (terrestrial) and autochthonous (aquatic) components of the trophic base. In particular, FA analysis demonstrated the higher nutritional value of autochthonous primary producers, and provided supporting evidence that most consumers, even seston-feeders, were primarily supported by autochthonous resources and not by allochthonous matter. FA indicated shifts in some taxa-specific diets not detected by stable isotopes alone., Our study demonstrated that the combined use of stable isotopes and fatty acids provides new insights into determining the trophic base of a complex food web with trophic resources of both terrestrial/aquatic and lacustrine/riverine origins. In addition, directly comparing results from both stable-isotope and fatty-acid analyses provided additional information on selective feeding by seston-feeding taxa on autochthonous and allochthonous fractions of the seston., The presence of LW in the river channel decreased lake-river coupling by providing alternative basal resources, primarily through increasing high-quality autochthonous production on wood and by providing a superior substratum for net-spinning caddisflies to feed on a fraction of the seston richer in essential fatty acids. River management strategies that incorporate instream LW therefore have the potential to alter energy flows and enhance ecosystem productivity by increasing the quantity and quality of available basal food resources. [ABSTRACT FROM AUTHOR]

Lake shores are characterised by a high natural variability, which is increasingly threatened by a multitude of anthropogenic disturbances including morphological alterations to the littoral zone. The European Water Framework Directive (EU WFD) calls for the assessment of lake ecological status by monitoring biological quality elements including benthic macroinvertebrates. To identify cost- and time-efficient sampling strategies for routine lake monitoring, we sampled littoral invertebrates in 32 lakes located in different geographical regions in Europe. We compared the efficiency of two sampling methodologies, defined as habitat-specific and pooled composite sampling protocols. Benthic samples were collected from unmodified and morphologically altered shorelines. Variability within macroinvertebrate communities did not differ significantly between sampling protocols across alteration types, lake types and geographical regions. Community composition showed no significant differences between field composite samples and artificially generated composite samples, and correlation coefficients between macroinvertebrate metrics calculated with both methods and a predefined morphological stressor index were similar. We conclude that proportional composite sampling represents a time- and cost-efficient method for routine lake monitoring as requested under the EU WFD, and may be applied across various European geographical regions. [ABSTRACT FROM AUTHOR]

Coarse woody debris ( CWD) in the littoral zone of lakes constitutes a preferred habitat for macroinvertebrates and fish. CWD differs in the surface complexity depending on its decay status. Therefore, CWD may provide distinct types of shelters and thus modify the structure of the macroinvertebrate community as well as its susceptibility to fish predation., We ran an enclosure experiment in a lake littoral zone to test the effect of surface complexity of CWD on the interactions between the predator, Eurasian perch ( Perca fluviatilis) and its potential macroinvertebrate prey. We deployed 10 enclosures containing fresh wood with a smooth surface and 10 enclosures containing decayed wood with a more complex, rough surface and allowed colonisation by macroinvertebrates. Five enclosures of each type were then stocked by perch and exposed to fish predation., The abundance and biomass of macroinvertebrates were significantly higher on decayed wood with greater surface complexity than on fresh wood; however, the type of CWD did not strongly influence the taxonomic composition and diversity of invertebrates., The direct effect of perch predation on the macroinvertebrate community was weak. Perch reduced only the abundance of adult Dikerogammarus villosus, while other potential prey, such as chironomids, was more abundant in the presence of the fish. The impact of perch consumption of these larvae was probably obscured by interspecific interactions among chironomids and D. villosus, which were impaired in the fish enclosures., We found no clear evidence that the influence of perch on macroinvertebrates was mediated by the complexity of the wood surface. However, fish diet analysis showed that on decayed wood, perch preferentially consumed chironomids, and consumption of D. villosus was much lower, while on fresh wood, the preferential consumption of chironomids decreased with increasing consumption of gammarids. This suggests that such differences in fish diet could be an effect of complex interactions between wood microstructure, prey density and its ability to find refuge in CWD., The effect of CWD microstructure on predator-prey interactions was visible with respect to interspecific relationships between chironomids and gammarids, which on more complex decayed wood were moderated in the absence of perch. [ABSTRACT FROM AUTHOR]

Abstract: Implementation of the EU Water Framework Directive (WFD) has drawn much attention to hydromorphological alterations of surface waters. The Lake Habitat Survey (LHS) protocol provides a method for characterising and assessing the physical habitats of lakes and reservoirs. Two metrics were developed based on this method: the Lake Habitat Modification Score (LHMS) and the Lake Habitat Quality Assessment (LHQA), as measures of lake modification and habitat value, respectively. However, the use of these metrics to predict measures of ecological quality remains largely untested. Thus, we assessed the relationships between LHS metrics and the littoral macroinvertebrate community in 42 lakes across Europe. A significant relationship was found between littoral macrophyte descriptors and riparian natural land cover variables of the LHQA score and macroinvertebrate community composition in 2 out of 4 European regions. No relationship was found between macroinvertebrate community composition and the LHMS. Some significant correlations were found between selected macroinvertebrate metrics and the LHS scores, but this pattern was not consistent across regions, and no relationship was found with the overall LHMS or LHQA scores. This demonstrates that the LHS metrics do not consistently predict the quality of littoral macroinvertebrate communities across Europe, and a region specific approach may be necessary. However, we could demonstrate a relationship between the site specific LHS variables and the macroinvertebrate community at the site level, and in some cases at the regional level. Therefore, although the LHS metrics do not appear to be a useful for relating habitat quality and pressure to littoral macroinvertebrate communities, selected LHS variables may exhibit stronger relationships with the biota. [Copyright &y& Elsevier]

Highlights: [•] We developed an assessment tool for morphological alterations of lake shores. [•] Macroinvertebrates in natural and morphologically degraded lake sites were sampled. [•] Macrozoobenthos communities differed among European geographical regions. [•] Biological indication scores correlated well with the morphological stressor. [•] Two biotic multimetric indices are available based on habitat and composite samples. [ABSTRACT FROM AUTHOR]