Generalist parasites persist in degraded environments: a lesson learned from microsporidian diversity in amphipods.

The present study provides new insight into suitable microsporidian­host associations. It relates regional and continental-wide host specialization in microsporidians infecting amphipods to degraded and recovering habitats across 2 German river catchments. It provides a unique opportunity to infer the persistence of parasites following anthropogenic disturbance and their establishment in restored rivers. Amphipods were collected in 31 sampling sites with differing degradation and restoration gradients. Specimens were morphologically (hosts) and molecularly identified (host and parasites). Amphipod diversity and abundance, microsporidian diversity, host phylogenetic specificity and continental-wide ß-specificity were investigated and related to each other and/or environmental variables. Fourteen microsporidian molecular operational taxonomic units (MOTUs), mainly generalist parasites, infecting 6 amphipod MOTUs were detected, expanding the current knowledge on the host range by 17 interactions. There was no difference in microsporidian diversity and host specificity among restored and near-natural streams (Boye) or between those located in urban and rural areas (Kinzig). Similarly, microsporidian diversity was generally not influenced by water parameters. In the Boye catchment, host densities did not influence microsporidian MOTU richness across restored and near-natural sites. High host turnover across the geographical range suggests that neither environmental conditions nor host diversity plays a significant role in the establishment into restored areas. Host diversity and environmental parameters do not indicate the persistence and dispersal of phylogenetic host generalist microsporidians in environments that experienced anthropogenic disturbance. Instead, these might depend on more complex mechanisms such as the production of resistant spores, host switching and host dispersal acting individually or conjointly.

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.

Long-term recovery of benthic food webs after stream restoration.

The assessment of restoration success often neglects trophic interactions within food webs, focusing instead on biodiversity and community structure. Here, we analysed the long-term recovery of food web structure based on stable isotopes (δ13C and δ15N) of benthic invertebrates and quantified responses of food web metrics to time since restoration. The samples derived from twelve restored sites with different restoration ages, sampled annually from 2012 to 2021, and covering an investigation period of up to 28 years after restoration for the whole catchment. Temporal developments of the restored sites were compared to the development of two near-natural sites. The restoration measures consisted of the cessation of sewage inflow and morphological restoration of the channels. As a clear and consistent result over almost all sites, trophic similarity (proportion of co-existing species occupying similar trophic niches) increased with time since restoration, and reached values of near-natural sites, suggesting an increase in the stability and resilience of the food webs. Surprisingly, resource diversity decreased at most restored sites within 10 years after restoration, probably due to the removal of wastewater-derived resources, and a shift towards leaf litter as the dominant resource following the regrowth of the riparian vegetation. Food chain length showed no consistent pattern over time at the different sites both increasing and decreasing with time since restoration. Overall, restoration had clear effects on the food web structure of stream ecosystems. While some effects such as the increase in trophic similarity were consistent at almost all sites, others such as response of the food chain length were context dependent. The study demonstrates the potential of utilizing food web metrics, particularly trophic similarity, in restoration research to achieve a more holistic understanding of ecosystem recovery.

How tolerances, competition and dispersal shape benthic invertebrate colonisation in restored urban streams.

Biotic communities often respond poorly to river restoration activities and the drivers of community recovery after restoration are not fully understood. According to the Asymmetric Response Concept (ARC), dispersal capacity, species tolerances to stressors, and biotic interactions are three key drivers influencing community recovery of restored streams. However, the ARC remains to be tested. Here we used a dataset on benthic invertebrate communities of eleven restored stream sections in a former open sewer system that were sampled yearly over a period of eleven years. We applied four indices that reflect tolerance against chloride and organic pollution, the community's dispersal capacity and strength of competition to the benthic invertebrate taxa lists of each year and site. Subsequently, we used generalised linear mixed models to analyse the change of these indices over time since restoration. Dispersal capacity was high directly after restoration but continuously decreased over time. The initial communities thus consisted of good dispersers and were later joined by more slowly dispersing taxa. The tolerance to organic pollution also decreased over time, reflecting continuous improvement of water quality and an associated increase of sensitive species. On the contrary, chloride tolerances did not change, which could indicate a stable chloride level throughout the sampling period. Lastly, competition within the communities, reflected by interspecific trait niche overlap, increased with time since restoration. We show that recovery follows a specific pattern that is comparable between sites. Benthic communities change from tolerant, fast dispersing generalists to more sensitive, slowly dispersing specialists exposed to stronger competition. Our results lay support to the ARC (increasing role of competition, decreasing role of dispersal) but also underline that certain tolerances may still shape communities a decade after restoration. Disentangling the drivers of macroinvertebrate colonisation can help managers to better understand recovery trajectories and to define more realistic restoration targets.

Stream macroinvertebrate communities in restored and impacted catchments respond differently to climate, land-use, and runoff over a decade.

Identifying which environmental drivers underlie degradation and improvements of ecological communities is a fundamental goal of ecology. Achieving this goal is a challenge due to diverse trends in both environmental conditions and ecological communities across regions, and it is constrained by the lack of long-term parallel monitoring of environmental and community data needed to study causal relationships. Here, we identify key environmental drivers using a high-resolution environmental - ecological dataset, an ensemble of the Soil and Water Assessment Tool (SWAT+) model, and ecological models to investigate effects of climate, land-use, and runoff on the decadal trend (2012-2021) of stream macroinvertebrate communities in a restored urban catchment and an impacted catchment with mixed land-uses in Germany. The decadal trends showed decreased precipitation, increased temperature, and reduced anthropogenic land-uses, which led to opposing runoff trends - with decreased runoff in the restored catchment and increased runoff in the impacted catchment. The two catchments also varied in decadal trends of taxonomic and trait composition and metrics. The most significant improvements over time were recorded in communities of the restored catchment sites, which have become wastewater free since 2007 to 2009. Within the restored catchment sites, community metric trends were primarily explained by land-use and evaporation trends, while community composition trends were mostly associated with precipitation and runoff trends. Meanwhile, the communities in the impacted catchment did not undergo significant changes between 2012 and 2021, likely influenced by the effects of prolonged droughts following floods after 2018. The results of our study confirm the significance of restoration and land-use management in fostering long-term improvements in stream communities, while climate change remains a prodigious threat. The coupling of long-term biodiversity monitoring with concurrent sampling of relevant environmental drivers is critical for preventative and restorative management in ecology.

Differential associations of five riverine organism groups with multiple stressors.

The decline of river and stream biodiversity results from multiple simultaneous occuring stressors, yet few studies explore responses explore responses across various taxonomic groups at the same locations. In this study, we address this shortcoming by using a coherent data set to study the association of nine commonly occurring stressors (five chemical, one morphological and three hydraulic) with five taxonomic groups (bacteria, fungi, diatoms, macro-invertebrates and fish). According to studies on single taxonomic groups, we hypothesise that gradients of chemical stressors structure community composition of all taxonomic groups, while gradients of hydraulic and morphological stressors are mainly related to larger organisms such as benthic macro-invertebrates and fish. Organisms were sampled over two years at 20 sites in two catchments: a recently restored urban lowland catchment (Boye) and a moderately disturbed rural mountainous catchment (Kinzig). Dissimilarity matrices were computed for each taxonomic group within a catchment. Taxonomic dissimilarities between sites were linked to stressor dissimilarities using multivariable Generalized Linear Mixed Models. Stressor gradients were longer in the Boye, but did in contrast to the Kinzig not cover low stress intensities. Accordingly, responses of the taxonomic groups were stronger in the Kinzig catchment than in the recently restored Boye catchment. The discrepancy between catchments underlines that associations to stressors strongly depend on which part of the stressor gradient is covered in a catchment. All taxonomic groups were related to conductivity. Bacteria, fungi and macro-invertebrates change with dissolved oxygen, and bacteria and fungi with total nitrogen. Morphological and hydraulic stressors had minor correlations with bacteria, fungi and diatoms, while macro-invertebrates were strongly related to fine sediment and discharge, and fish to high flow peaks. The results partly support our hypotheses about the differential associations of the different taxonomic groups with the stressors.

Multi-decadal improvements in the ecological quality of European rivers are not consistently reflected in biodiversity metrics.

Humans impact terrestrial, marine and freshwater ecosystems, yet many broad-scale studies have found no systematic, negative biodiversity changes (for example, decreasing abundance or taxon richness). Here we show that mixed biodiversity responses may arise because community metrics show variable responses to anthropogenic impacts across broad spatial scales. We first quantified temporal trends in anthropogenic impacts for 1,365 riverine invertebrate communities from 23 European countries, based on similarity to least-impacted reference communities. Reference comparisons provide necessary, but often missing, baselines for evaluating whether communities are negatively impacted or have improved (less or more similar, respectively). We then determined whether changing impacts were consistently reflected in metrics of community abundance, taxon richness, evenness and composition. Invertebrate communities improved, that is, became more similar to reference conditions, from 1992 until the 2010s, after which improvements plateaued. Improvements were generally reflected by higher taxon richness, providing evidence that certain community metrics can broadly indicate anthropogenic impacts. However, richness responses were highly variable among sites, and we found no consistent responses in community abundance, evenness or composition. These findings suggest that, without sufficient data and careful metric selection, many common community metrics cannot reliably reflect anthropogenic impacts, helping explain the prevalence of mixed biodiversity trends.

Taxonomic and functional reorganization in Central European stream macroinvertebrate communities over 25 years.

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.

Stressors affecting the ecological status of temporary rivers in the Mediterranean region.

Temporary rivers are widespread in the Mediterranean region and impose a challenge for the implementation of the Water Framework Directive (WFD) and other environmental regulations. Surprisingly, an overarching analysis of their ecological status and the stressors affecting them is yet missing. We compiled data on the ecological status of 1504 temporary rivers in seven European Mediterranean region countries and related their ecological status (1) to publicly available data on pressures from the European WISE-WFD dataset, and (2) to seven more specific stressors modelled on a sub-catchment scale. More than 50 % of the temporary water bodies in the Mediterranean countries reached good or even high ecological status. In general, status classes derived from phytobenthos and macrophyte assessment were higher than those derived from the assessment of benthic invertebrates or fish. Of the more generally defined pressures reported to the WISE-WFD database, the most relevant for temporary rivers were 'diffuse agricultural' and 'point urban waste water'. Of the modelled more specific stressors, agricultural land use best explained overall ecological status, followed by total nitrogen load, and urban land use, while toxic substances, total phosphorus load and hydrological stressors were less relevant. However, stressors differed in relevance, with total nitrogen being most important for macrophytes, and agricultural land use for phytobenthos, benthic invertebrates and fish. For macrophytes, ecological quality increased with stressor intensity. The results underline the overarching effect of land use intensity for the ecological status of temporary water bodies. However, assessment results do not sufficiently reflect hydrological stress, most likely as the biological indicators used to evaluate these systems were designed for perennial water bodies and thus mainly target land use and nutrient impacts. We conclude that biomonitoring systems need to be updated or newly developed to better account for the specific situation of temporary water bodies.

The Asymmetric Response Concept explains ecological consequences of multiple stressor exposure and release.

Our capacity to predict trajectories of ecosystem degradation and recovery is limited, especially when impairments are caused by multiple stressors. Recovery may be fast or slow and either complete or partial, sometimes result in novel ecosystem states or even fail completely. Here, we introduce the Asymmetric Response Concept (ARC) that provides a basis for exploring and predicting the pace and magnitude of ecological responses to, and release from, multiple stressors. The ARC holds that three key mechanisms govern population, community and ecosystem trajectories. Stress tolerance is the main mechanism determining responses to increasing stressor intensity, whereas dispersal and biotic interactions predominantly govern responses to the release from stressors. The shifting importance of these mechanisms creates asymmetries between the ecological trajectories that follow increasing and decreasing stressor intensities. This recognition helps to understand multiple stressor impacts and to predict which measures will restore communities that are resistant to restoration.

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.

The interplay of nutrients, dissolved inorganic carbon and algae in determining macrophyte occurrences in rivers.

Nitrogen and phosphorous concentrations are widely considered to drive macrophyte assemblages in rivers. However, Dissolved Inorganic Carbon (DIC) - available for plants as CO2 and HCO3- - is also of major relevance. Based on literature, we present a conceptual model on the interaction between algae, macrophytes, DIC, pH, light, N, P and the surface water and sedimental compartment. Analysing two separate datasets (i) on river physico-chemistry and chlorophyll-a, and (ii) on river physico-chemistry and macrophytes we quantify three connections within this concept: (1) the correlation of chlorophyll-a versus pH, (2) the correlation of TP versus chlorophyll-a and (3) the occurrence of HCO3-users and CO2-only-users among macrophytes along the DIC gradient. Chlorophyll-a correlated positively with pH (R-squared = 77%, p < .001) due to increased carbon dioxide uptake of phytoplankton. Surface water TP did not linearly correlate with chlorophyll-a concentrations. Obligate and optionally submerged macrophyte species that utilise HCO3- were separated from CO2-only-users by HCO3- concentrations, with an area under the curve (AUC) of 68% and 70% (both p < .001) between groups. Obligate and optionally submerged macrophyte assemblages only composed of HCO3-users and those exclusively composed of CO2-only-users showed an even stronger separation based on the HCO3- concentration, with both an AUC of 82% and 78% (both p < .001). Our results underline that DIC can greatly affect riverine macrophytes. However, absolute concentrations of HCO3- are less relevant, while the connection to pH is more important, reflecting CO2 concentrations. River monitoring and management should consider the interaction between nutrients DIC, surface water and sedimental compartment as important factors affecting macrophyte occurrence, rather than solely focussing on surface water nutrients.

The effect of lateral connectedness on the taxonomic and functional structure of fish communities in a lowland river floodplain.

In river floodplains many conservation programs focus on the main river channel as the richest in species. Lateral floodplain waterbodies, which contribute largely to functional processes in river systems, often remain overlooked and exposed to anthropogenic pressures. Although the role of hydrological connectedness between lateral waterbodies and the main river on taxonomic composition of fish communities is well understood, effects on functional community composition is much less studied. Abundance data of fish communities were gathered from 152 electrofishing sites in the main channel and lateral floodplain waterbodies of the river Lippe (Germany), over 18 years. These data were used to compare taxonomic, functional, conservation and recreational fishing aspects along the floodplain lateral connectedness gradient. Fish species richness decreased along the lateral continuum from the main river channel to isolated floodplain waterbodies. In contrast, the relative abundance of endangered and also of non-native species increased along this gradient, highlighting the ecological and conservational importance of floodplain waterbodies. Species composition in floodplain waterbodies differed across the connectedness gradient showing distinct assemblages which were not merely subsets of the main channel. The variability of life-history and feeding strategists among classes of lateral connectedness confirmed the importance of each connectivity class in contributing to the overall floodplain functional diversity. This study highlights the need of preserving fish taxonomic and functional biodiversity across the floodplain as one integrated hydrosystem. Conservation and restoration measures should therefore extend to include the whole floodplain area and the complete spectrum of differently connected floodplain waterbodies in addition to the main channel of the river.

Re-meandering German lowland streams: qualitative and quantitative effects of restoration measures on hydromorphology and macroinvertebrates.

We investigated the effects of two river restoration projects on hydromorphology and macroinvertebrate fauna in two German lowland rivers, the Schwalm and the Gartroper Mühlenbach. The stream channels were re-meandered and the floodplain levels were lowered to better connect the streams to their floodplains. The restoration was performed 10 years ago in the Schwalm and 2 years ago in the Gartroper Mühlenbach. We compared the restored reaches to nearby anthropogenically straightened reaches. Twenty-five hydromorphological parameters were recorded on twenty transects; between nine and 23 substrate-specific macroinvertebrate samples were taken per reach. Several hydromorphological parameters, such as the number and width of channel features and the number of substrate types, were significantly higher in the restored reaches compared to nearby anthropogenically straightened reaches. Total numbers of invertebrate families, genera, and taxa were also higher in the restored reaches than in the anthropogenically straightened reaches. Biotic substrates like dead wood or macrophytes were more abundant in the restored reaches, and these substrates hosted 28 taxa not found in the straightened reaches. While diversity was high in both restored reaches, overall abundance increased only in the river that was restored 10 years ago. Using NMS-analysis, substrate-specific faunistic samples of the restored reaches were compared to those of the straightened reaches. Our results revealed different invertebrate communities on the same substrates in the recently restored river. In the 10-year-old restoration, however, the same substrates were similarly inhabited. This comparable colonization of substrates may reflect succession in the macroinvertebrate community. The results are discussed according to the re-colonization potentials of the upstream and downstream reaches and the dispersal capacity of taxa.

Development and validation of a macroinvertebrate-based biomonitoring tool to assess fine sediment impact in small mountain streams.

Increased fine sediment deposition is recognised as one of the major causes of biological impairment of rivers and streams influencing all components of aquatic communities. Notably, stream macroinvertebrates are affected showing changes in abundance and community composition. This makes macroinvertebrates an attractive choice for biomonitoring fine sediment stress. However, there are substantial knowledge gaps regarding the quantification of deposited fine sediment and the identification of taxa sensitive to fine sediment deposition, which could serve as indicators. In this study, we developed a stream type-specific index based on the taxon-specific response of macroinvertebrates to deposited fine sediment in small, coarse substrate-dominated mountain streams. We sampled fine sediment at 73 sampling sites in Western Germany (Europe) in spring 2014 and 2015 using a sediment remobilization technique. Macroinvertebrate taxalists originating from WFD monitoring surveys were available for all sites. We applied Threshold Indicator Taxa ANalysis (TITAN) on the fine sediment mass of the sampling sites and the corresponding macroinvertebrate taxalists to identify indicator taxa, which were then used for index development. In total, TITAN identified 95 reliable indicator taxa, of which some taxa tolerated large amounts of fine sediment (e.g., Gammarus roeselii and Tubificidae Gen. sp.), while others were found to be highly sensitive to increased fine sediment mass (e.g., Elodes sp. and Limnius perrisi). The newly developed index was tested on an independent data set and performed well in detecting fine sediment stress (Spearman's r = 0.63). Furthermore, the index was better related to the deposited fine sediment mass as compared to other fine sediment indices and standard metrics used for monitoring purposes under the Water Framework Directive (WFD). The diagnostic index can be a cost-effective biomonitoring tool for stream managers and can be used as a proxy for the impact of deposited fine sediment on the reach scale.

Effect of river restoration on life-history strategies in fish communities.

Assessments of river restoration outcomes are mostly based on taxonomic identities of species, which may not be optimal because a direct relationship to river functions remains obscure and results are hardly comparable across biogeographic borders. The use of ecological species trait information instead of taxonomic units may help to overcome these challenges. Abundance data for fish communities were gathered from 134 river restoration projects conducted in Switzerland, Germany and Finland, monitored for up to 15 years. These data were related to a dataset of 22 categories of ecological traits describing fish life-history strategies to assess the outcome of the restoration projects. Restoration increased trait functional diversity and evenness in projects that were situated in the potamal zone of rivers. Restoration effect increased with the length of the restored river reaches. In areas with low levels of anthropogenic land use, the peak of the restoration effect was reached already within one to five years after the restoration and effect receded thereafter, while communities responded later in areas with higher levels of anthropogenic land use. In the lower potamal zone, a shift towards opportunistic life-history strategists was observed. In the upper rhithral zone, in contrast, species with an opportunistic life-history strategy increased only in the first five years of restoration, followed by a shift towards equilibrium strategists at restorations older than 5 years. This pattern was more pronounced in rivers with higher level of anthropogenic land use and longer restored river reaches. Restoration reduced the variability in community trait composition between river reaches suggesting that community trait composition within these zones converges when rivers are restored. This study showed how ecological traits are suitable to analyse restoration outcomes and how such an approach can be used for the evaluation and comparison of environmental management actions across geographical regions.

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.

Reintroduction of freshwater macroinvertebrates: challenges and opportunities.

Species reintroductions - the translocation of individuals to areas in which a species has been extirpated with the aim of re-establishing a self-sustaining population - have become a widespread practice in conservation biology. Reintroduction projects have tended to focus on terrestrial vertebrates and, to a lesser extent, fishes. Much less effort has been devoted to the reintroduction of invertebrates into restored freshwater habitats. Yet, reintroductions may improve restoration outcomes in regions where impoverished regional species pools limit the self-recolonisation of restored freshwaters. We review the available literature on macroinvertebrate reintroductions, focusing on identifying the intrinsic and extrinsic factors that determine their success or failure. Our study reveals that freshwater macroinvertebrate reintroductions remain rare, are often published in the grey literature and, of the attempts made, approximately one-third fail. We identify life-cycle complexity and remaining stressors as the two factors most likely to affect reintroduction success, illustrating the unique challenges of freshwater macroinvertebrate reintroductions. Consideration of these factors by managers during the planning process and proper documentation - even if a project fails - may increase the likelihood of successful outcomes in future reintroduction attempts of freshwater macroinvertebrates.

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.

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.