Diverging response patterns of terrestrial and aquatic species to hydromorphological restoration.

Although experiences with ecological restoration continue to accumulate, the effectiveness of restoration for biota remains debated. We complemented a traditional taxonomic analysis approach with information on 56 species traits to uncover the responses of 3 aquatic (fish, macroinvertebrates, macrophytes) and 2 terrestrial (carabid beetles, floodplain vegetation) biotic groups to 43 hydromorphological river restoration projects in Germany. All taxonomic groups responded positively to restoration, as shown by increased taxonomic richness (10-164%) and trait diversity (habitat, dispersal and mobility, size, form, life history, and feeding groups) (15-120%). Responses, however, were stronger for terrestrial than aquatic biota, and, contrary to our expectation, taxonomic responses were stronger than those of traits. Nevertheless, trait analysis provided mechanistic insights into the drivers of community change following restoration. Trait analysis for terrestrial biota indicated restoration success was likely enhanced by lateral connectivity and reestablishment of dynamic processes in the floodplain. The weaker response of aquatic biota suggests recovery was hindered by the persistence of stressors in the aquatic environment, such as degraded water quality, dispersal constraints, and insufficient hydromorphological change. Therefore, river restoration requires combined local- and regional-scale approaches to maximize the response of both aquatic and terrestrial organisms. Due to the contrasting responses of aquatic and terrestrial biota, the planning and assessment of river restoration outcomes should consider effects on both components of riverine landscapes.

Macroinvertebrate communities respond strongly but non-specifically to a toxicity gradient derived by effect-based methods.

Chemical pollution is one of the most important threats to freshwater ecosystems. The plethora of potentially occurring chemicals and their effects in complex mixtures challenge standard monitoring methods. Effect-based methods (EBMs) are proposed as complementary tools for the assessment of chemical pollution and toxic effects. To investigate the effects of chemical pollution, the ecological relevance of EBMs and the potential of macroinvertebrates as toxicity-specific bioindicators, ecological and ecotoxicological data were linked. Baseline toxicity, mutagenicity, dioxin-like and estrogenic activity of water and sediment samples from 30 river sites in central Germany were quantified with four in vitro bioassays. The responses of macroinvertebrate communities at these sites were assessed by calculating 16 taxonomic and functional metrics and by investigating changes in the taxonomic and trait composition. Principal component analysis revealed an increase in toxicity along a joint gradient of chemicals with different modes of action. This toxicity gradient was associated with a decrease in biodiversity and ecological quality, as well as significant changes in taxonomic and functional composition. The strength of the effects suggested a strong impact of chemical pollution and underlined the suitability of EBMs in detecting ecological relevant effects. However, the metrics, taxa, and traits associated with vulnerability or tolerance to toxicity were found to also respond to other stressors in previous studies and thus may have only a low potential as toxicity-specific bioindicators. Because macroinvertebrates respond integratively to all present stressors, linking both ecological and environmental monitoring is necessary to investigate the overall effects but also isolate individual stressors. EBMs have a high potential to separate the toxicity of chemical mixtures from other stressors in a multiple stressor scenario, as well as identifying the presence of chemical groups with specific modes of action.

Effects of multiple stressors on benthic invertebrates using Water Framework Directive monitoring data.

Multiple stressors affect freshwater systems and cause a deficient ecological status according to the European Water Framework Directive (WFD). To select effective mitigation measures and improve the ecological status, knowledge on the stressor hierarchy and individual and joined effects is necessary. However, compared to common stressors like nutrient enrichment and morphological degradation, the relative importance of micropollutants such as pesticides and pharmaceuticals is largely unaddressed. We used WFD monitoring data from Saxony (Germany) to investigate the importance of 85 environmental variables (including 34 micropollutants) for 18 benthic invertebrate metrics at 108 sites. The environmental variables were assigned to five groups (natural factors, nutrient enrichment, metals, micropollutants and morphological degradation) and were ranked according to their relative importance as group and individually within and across groups using Principal Component Analyses (PCAs) and Boosted Regression Trees (BRTs). Overall, natural factors contributed the most to the total explained deviance of the models. This variable group represented not only typological differences between sampling sites but also a gradient of human impact by strongly anthropogenically influenced variables such as electric conductivity and dissolved oxygen. These large-scale effects can mask the individual importance of the other variable groups, which may act more specifically at a subset of sites. Accordingly, micropollutants were not represented by a few dominant variables but rather a diverse palette of different chemicals with similar contribution. As a group, micropollutants contributed similarly as metals, nutrient enrichment and morphological degradation. However, the importance of micropollutants might be underestimated due to limitations of the current chemical monitoring practices.

Habitat requirements of riparian arthropods on gravel bars: Implications for conservation and management of braided river floodplains.

In their pristine state, river landscapes consist of complex mosaics of aquatic and terrestrial habitats. They are highly dynamic and, with their harsh environments, offer living space for many specialists. In the present study, the habitat choice of specialists of the riparian arthropod community was studied on a near-natural stretch of the Upper Isar River. Study period was between May and July 2011. Araneae, Formicidae and Staphylinidae were the most common taxa. The dominant species was Pardosa wagleri with 1,092 individuals, followed by Arctosa cinerea with 184 and Paederidus rubrothoracicus with 154 individuals. These three species made up 54% of all located individuals and were considered as representatives for the invertebrate community. Remaining species had by far smaller proportions and were not determined further due to the low individual numbers. Habitat preferences for the three dominant species were analyzed using negative binomial regression. Common and important habitat features were non-silted and coarse gravel areas, which are neighboured by patches with an elevation 1m above the water. Furthermore, the absence of vegetation cover as well the absence of ants was crucial for the occurrence of the three model species. Habitat preferences were subject to seasonal influences due to various requirements of different life stages. Other influencing factors were competition and predation due to Formicidae and larger individuals of Lycosidae. This demonstrates the high importance of structurally rich riverbeds with a mosaic of distinct habitat patches for the three representative species. Our findings are a valuable contribution for the conservation and management of braided rivers and their characteristic gravel bar biocoenosis.

Decline in niche specialization and trait ß-diversity in benthic invertebrate communities of Central European low-mountain streams over 25 years.

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 ß-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 ß-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.

Aquatic macroinvertebrate assemblages in rivers influenced by mining activities.

Mining is one of the major pollution sources worldwide, causing huge disturbances to the environment. Industrial and artisanal mining activities are widespread in Mexico, a major global producer of various metals. This study aimed to assess the ecological impairments resulting from mining activities using aquatic macroinvertebrates assemblages (MA). A multiple co-inertia analysis was applied to determine the relationships between environmental factors, habitat quality, heavy metals, and aquatic macroinvertebrates in 15 study sites in two different seasons (dry and wet) along two rivers running across the Central Plateau of Mexico. The results revealed three contrasting environmental conditions associated with different MAs. High concentrations of heavy metals, nutrients, and salinity limit the presence of several families of seemingly sensitive macroinvertebrates. These factors were found to influence structural changes in MAs, showing that not only mining activities, but also agriculture and presence of villages in the basin, exert adverse effects on macroinvertebrate assemblages. Diversity indices showed that the lowest diversity matched both the most polluted and the most saline rivers. The rivers studied displayed high alkalinity and hardness levels, which can reduce the availability of metals and cause adverse effects on periphyton by inhibiting photosynthesis and damaging MAs. Aquatic biomonitoring in rivers, impacted by mining and other human activities, is critical for detecting the effect of metals and other pollutants to improve management and conservation strategies. This study supports the design of cost-effective and accurate water quality biomonitoring protocols in developing countries.

River restoration success depends on the species pool of the immediate surroundings.

Previous studies evaluating the success of river restorations have rarely found any consistent effects on benthic invertebrate assemblages. In this study, we analyzed data from 24 river restoration projects in Germany dating back 1 to 12 years and 1231 data sets from adjacent river reaches that lie within 0-5, 5-10, and 10-15 km rings centered on the restored sites. We calculated restoration success and recolonization potential of adjacent river reaches based on stream-type-specific subsets of taxa indicative for good or bad habitat quality. On average, the restorations did not improve the benthic invertebrate community quality. However, we show that restoration success depends on the presence of source populations of desired taxa in the surrounding of restored sites. Only where source populations of additional desired taxa existed within a 0-5 km ring around the restored sites were benthic invertebrate assemblages improved by the restoration. Beyond the 5-km rings, this recolonization effect was no longer detected. We present here the first field results to support the debated argument that a lack of source populations in the areas surrounding restored sites may play an important role in the failure to establish desired invertebrate communities by the means of river restorations. In contrast, long-range dispersal of invertebrates seems to play a subordinate role in the recolonization of restored sites. However, because the surroundings of the restored sites were far from good ecological quality, the potential for improvement of restored sites was limited.

Linking environmental with biological data: Low sampling frequencies of chemical pollutants and nutrients in rivers reduce the reliability of model results.

Linking environmental and biological data using ecological models can provide crucial knowledge about the effects of water quality parameters on freshwater ecosystems. However, a model can only be as reliable as its input data. Here, the influence of sampling frequency of temporal variable environmental input data on the reliability of model results when linked to biological data was investigated using Threshold Indicator Taxa Analysis (TITAN) and species sensitivity distributions (SSDs). Large-scale biological data from benthic macroinvertebrates and matching water quality data including four metals and four nutrients of up to 559 site-year combinations formed the initial data sets. To compare different sampling frequencies, the initial water quality data sets (n = 12 samples per year, set as reference) were subsampled (n = 10, 8, 6, 4, 2 and 1), annual mean values calculated and used as input data in the models. As expected, subsampling significantly reduced the reliability of the environmental input data across all eight substances. For TITAN, the use of environmental input data with a reduced reliability led to a considerable (1) loss of information because valid taxa were no longer identified, (2) gain of unreliable taxon-specific change points due to false positive taxa, and (3) bias in the change point estimation. In contrast, the reliability of the SSD results appeared to be much less reduced. However, closer examination of the SSD input data indicated that existing effects were masked by poor model performance. The results confirm that the sampling frequency of water quality data significantly influences the reliability of model results when linked with biological data. For studies limited to low sampling frequencies, the discussion provides recommendations on how to deal with low sampling frequencies of temporally variable water quality data when using them in TITAN, in SSDs, and in other ecological models.

Moderate warming over the past 25 years has already reorganized stream invertebrate communities.

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.

Towards stressor-specific macroinvertebrate indices: Which traits and taxonomic groups are associated with vulnerable and tolerant taxa?

Monitoring of macroinvertebrate communities is frequently used to define the ecological health status of rivers. Ideally, biomonitoring should also give an indication on the major stressors acting on the macroinvertebrate communities supporting the selection of appropriate management measures. However, most indices are affected by more than one stressor. Biological traits (e.g. size, generation time, reproduction) could potentially lead to more stressor-specific indices. However, such an approach has rarely been tested. In this study we classify 324 macroinvertebrate taxa as vulnerable (decreasing abundances) or tolerant (increasing abundances) along 21 environmental gradients (i.e. nutrients, major ions, oxygen and micropollutants) from 422 monitoring sites in Germany using Threshold Indicator Taxa Analysis (TITAN). Subsequently, we investigate which biological traits and taxonomic groups are associated with taxa classified as vulnerable or tolerant with regard to specific gradients. The response of most taxa towards different gradients was similar and especially high for correlated gradients. Traits associated with vulnerable taxa across most gradients included: larval aquatic life stages, isolated cemented eggs, reproductive cycle per year <1, scrapers, aerial and aquatic active dispersal and plastron respiration. Traits associated with tolerant taxa included: adult aquatic life stages, polyvoltinism, ovoviviparity or egg clutches in vegetation, food preference for dead animals or living microinvertebrates, substrate preference for macrophytes, microphytes, silt or mud and a body size >2-4cm. Our results question whether stressor-specific indices based on macroinvertebrate assemblages can be achieved using single traits, because we observed that similar taxa responded to different gradients and also similar traits were associated with vulnerable and tolerant taxa across a variety of water quality gradients. Future studies should examine whether combinations of traits focusing on specific taxonomic groups achieve higher stressor specificity.

Revisiting restored river reaches - Assessing change of aquatic and riparian communities after five years.

Hydromorphological restructuring of river sections, i.e. river restoration measures, often has little effects on aquatic biota, even in case of strong habitat alterations. It is often supposed that the biotic response is simply delayed as species require additional time to recolonize the newly generated habitats and to establish populations. To identify and specify the supposed lag time between restoration and biotic response, we investigated 19 restored river reaches twice in a five-year interval. The sites were restored one to ten years prior to the first sampling. We sampled three aquatic (fish, benthic invertebrates, macrophytes) and two riparian organism groups (ground beetles and riparian vegetation) and analyzed changes in assemblage composition and biotic metrics. With the exception of ground beetle assemblages, we observed no significant changes in richness and abundance metrics or metrics used for biological assessment. However, indicator taxa for near-natural habitat conditions in the riparian zone (indicators for regular inundation in plants and river bank specialists in beetles) improved significantly in the five-year interval. Contrary to general expectations in river restoration planning, we neither observed a distinct succession of aquatic communities nor a general trend towards "good ecological status" over time. Furthermore, multiple linear regression models revealed that neither the time since restoration nor the morphological status had a significant effect on the biological metrics and the assessment results. Thus, the stability of aquatic assemblages is strong, slowing down restoration effects in the aquatic zone, while riparian assemblages improve more rapidly. When defining restoration targets, the different timelines for ecological recovery after restoration should be taken into account. Furthermore, restoration measures should not solely focus on local habitat conditions but also target stressors acting on larger spatial scales and take other measures (e.g. species reintroduction) into consideration.

Effects of changing climate on European stream invertebrate communities: A long-term data analysis.

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.

Water quality variables and pollution sources shaping stream macroinvertebrate communities.

In 2015, over 90 percent of German rivers failed to reach a good ecological status as demanded by the EU Water Framework Directive (WFD). Deficits in water quality, mainly from diffuse pollution such as agricultural run-off, but also from wastewater treatment plants (WWTPs), have been suggested as important drivers of this decline in ecological quality. We modelled six macroinvertebrate based metrics indicating ecological quality for 184 streams in response to a) PCA-derived water quality gradients, b) individual water quality variables and c) catchment land use and wastewater exposure indices as pollution drivers. The aim was to evaluate the relative importance of key water quality variables and their sources. Indicator substances (i.e. carbamazepine and caffeine indicating wastewater exposure; herbicides indicating agricultural run-off) represented micropollutants in the analyses and successfully related water quality variables to pollution sources. Arable and urban catchment land covers were strongly associated with reduced ecological quality. Electric conductivity, oxygen concentration, caffeine, silicate and toxic units with respect to pesticides were identified as the most significant in-stream predictors in this order. Our results underline the importance to manage diffuse pollution, if ecological quality is to be improved. However, we also found a clear impact of wastewater on ecological quality through caffeine. Thus, improvement of WWTPs, especially preventing the release of poorly treated wastewater, will benefit freshwater communities.

Cross-realm assessment of climate change impacts on species' abundance trends.

Climate change, land-use change, pollution and exploitation are among the main drivers of species' population trends; however, their relative importance is much debated. We used a unique collection of over 1,000 local population time series in 22 communities across terrestrial, freshwater and marine realms within central Europe to compare the impacts of long-term temperature change and other environmental drivers from 1980 onwards. To disentangle different drivers, we related species' population trends to species- and driver-specific attributes, such as temperature and habitat preference or pollution tolerance. We found a consistent impact of temperature change on the local abundances of terrestrial species. Populations of warm-dwelling species increased more than those of cold-dwelling species. In contrast, impacts of temperature change on aquatic species' abundances were variable. Effects of temperature preference were more consistent in terrestrial communities than effects of habitat preference, suggesting that the impacts of temperature change have become widespread for recent changes in abundance within many terrestrial communities of central Europe.

Field data reveal low critical chemical concentrations for river benthic invertebrates.

River ecosystems are of immense ecological and social importance. Despite the introduction of wastewater treatment plants and advanced chemical authorization procedures in Europe, chemical pollution is still a major threat to freshwater ecosystems. Here, large-scale monitoring data was exploited to identify taxon-specific chemical concentrations beyond which benthic invertebrate taxa are unlikely to occur using Threshold Indicator Taxa Analysis (TITAN). 365 invertebrate taxa and 25 organic chemicals including pesticides, pharmaceuticals, plasticisers, flame retardants, complexing agents, a surfactant and poly- and monocyclic aromatic hydrocarbons from a total of 399 sites were analysed. The number of taxa that responded to each of these chemicals varied between 0% and 21%. These sensitive taxa belonged predominantly to the groups Plecoptera, Coleoptera, Trichoptera, Ephemeroptera, Turbellaria, Megaloptera, Crustacea, and Diptera. Strong effects were observed in response to wastewater-associated compounds, confirming that wastewater is an important cause of biological degradation. The majority of change points identified for each compound were well below predicted no-effect concentrations derived from laboratory toxicity studies. Thus, the results show that chemicals are likely to induce effects in the environment at concentrations much lower than expected based on laboratory experiments. Overall, it is confirmed that chemical pollution is still an important factor shaping the distribution of invertebrate taxa, suggesting the need for continued efforts to reduce chemical loads in rivers.

A newly developed dispersal metric indicates the succession of benthic invertebrates in restored rivers.

Dispersal capacity plays a fundamental role in the riverine benthic invertebrate colonization of new habitats that emerges following flash floods or restoration. However, an appropriate measure of dispersal capacity for benthic invertebrates is still lacking. The dispersal of benthic invertebrates occurs mainly during the aquatic (larval) and aerial (adult) life stages, and the dispersal of each stage can be further subdivided into active and passive modes. Based on these four possible dispersal modes, we first developed a metric (which is very similar to the well-known and widely used saprobic index) to estimate the dispersal capacity for 802 benthic invertebrate taxa by incorporating a weight for each mode. Second, we tested this metric using benthic invertebrate community data from a) 23 large restored river sites with substantial improvements of river bottom habitats dating back 1 to 10years, b) 23 unrestored sites very close to the restored sites, and c) 298 adjacent surrounding sites (mean±standard deviation: 13.0±9.5 per site) within a distance of up to 5km for each restored site in the low mountain and lowland areas of Germany. We hypothesize that our metric will reflect the temporal succession process of benthic invertebrate communities colonizing the restored sites, whereas no temporal changes are expected in the unrestored and surrounding sites. By applying our metric to these three river treatment categories, we found that the average dispersal capacity of benthic invertebrate communities in the restored sites significantly decreased in the early years following restoration, whereas there were no changes in either the unrestored or the surrounding sites. After all taxa had been divided into quartiles representing weak to strong dispersers, this pattern became even more obvious; strong dispersers colonized the restored sites during the first year after restoration and then significantly decreased over time, whereas weak dispersers continued to increase. The successful application of our metric to river restoration might be promising in further applications of this metric, for example, in analyzing metacommunity structure or community's recovery from extreme events such as floods, droughts or catastrophic pollution episodes.

Time is no healer: increasing restoration age does not lead to improved benthic invertebrate communities in restored river reaches.

Evidence for successful restoration of riverine communities is scarce, particularly for benthic invertebrates. Among the multitude of reasons discussed so far for the lack of observed effects is too short of a time span between implementation and monitoring. Yet, studies that explicitly focus on the importance of restoration age are rare. We present a comprehensive study based on 44 river restoration projects in Germany, focusing on standardized benthic invertebrate sampling. A broad gradient ranging from 1 to 25years in restoration age was available. In contrast to clear improvements in habitat heterogeneity, benthic community responses to restoration were inconsistent when compared to control sections. Taxon richness increased in response to restoration, but abundance, diversity and various assessment metrics did not respond clearly. Restoration age was a poor predictor of community composition and community change, as no significant linear responses could be detected using 34 metrics. Moreover, only 5 out of 34 tested metrics showed non-linear shifts at restoration ages of 2 to 3years. This might be interpreted as an indication of a post-restoration disturbance followed by a re-establishment of pre-restoration conditions. BIO-ENV analysis and fourth-corner modeling underlined the low importance of restoration age, but revealed high importance of catchment-scale characteristics (e.g., ecoregion, catchment size and land use) in controlling community composition and community change. Overall, a lack of time for community development did not appear to be the ultimate reason for impaired benthic invertebrate communities. Instead, catchment-scale characteristics override the effectiveness of restoration. To enhance the ecological success of future river restoration projects, we recommend improving water quality conditions and catchment-scale processes (e.g., connectivity and hydrodynamics) in addition to restoring local habitat structure.

Disentangling environmental drivers of benthic invertebrate assemblages: The role of spatial scale and riverscape heterogeneity in a multiple stressor environment.

It is broadly acknowledged that freshwater ecosystems are affected by multiple stressors, but the relative importance of individual stressors in impairing riverine communities remains unclear. We investigated the impacts of multiple stressors, incorporating in-stream water quality, riparian and catchment land use and stream morphology, on riverine benthic invertebrate communities, while considering the spatial scales of factors and the heterogeneity of riverscapes. We performed a stepwise regression procedure linking 21 abiotic and 20 community metrics using Generalized Linear Models on data from 1018 river sites spread across Germany. High impact stressors (e.g., nutrients and water temperature) were identified for various community metrics. Both the combination of relevant stressors and their explanatory value differed significantly across streams of different sizes and ecoregions. In large rivers, the riparian land use was less important in determining community structure compared to lower order streams. Thus, possible mitigating effects of revegetated riparian buffer strips are likely to be overwhelmed by the influence of catchment-wide land use. Our results indicated substantial variability in stressors for the range of metrics studied, providing insight into potential target parameters for effective ecosystem management. To achieve long lasting successes in managing, protecting and restoring running waters, it is of vital importance to recognize the heterogeneity of riverscapes and to consider large-scale influences.

Environmental Controls on River Assemblages at the Regional Scale: An Application of the Elements of Metacommunity Structure Framework.

Understanding factors that structure regional biodiversity is important for linking ecological and biogeographic processes. Our objective was to explore regional patterns in riverine benthic invertebrate assemblages in relation to their broad positioning along the river network and examine differences in composition, biodiversity (alpha and beta diversity), and environmental drivers. We up-scaled methods used to examine patterns in metacommunity structure (Elements of Metacommunity Structure framework) to examine faunal distribution patterns at the regional extent for 168 low-mountain stream invertebrate assemblages in central Germany. We then identified the most influential environmental factors using boosted regression trees. Faunal composition patterns were compartmentalised (Clementsian or quasi-Clementsian), with little difference from headwaters to large rivers, potentially reflecting the regional scale of the study, by crossing major catchment boundaries and incorporating different species pools. While idealised structures did not vary, environmental drivers of composition varied considerably between river sections and with alpha diversity. Prediction was substantially weaker, and the importance of space was greater, in large rivers compared to other sections suggesting a weakening in species sorting downstream. Further, there was a stronger transition in composition than for alpha diversity downstream. The stronger links with regional faunal composition than with richness further emphasises the importance of considering the alternative ways in which anthropogenic stressors are operating to affect biodiversity patterns. Our approach allowed bridging the gap between local (or metacommunity) and regional scales, providing key insights into drivers of regional biodiversity patterns.