Inland navigation and land use interact to impact European freshwater biodiversity.

Inland navigation in Europe is proposed to increase in the coming years, being promoted as a low-carbon form of transport. However, we currently lack knowledge on how this would impact biodiversity at large scales and interact with existing stressors. Here we addressed this knowledge gap by analysing fish and macroinvertebrate community time series across large European rivers comprising 19,592 observations from 4,049 sampling sites spanning the past 32 years. We found ship traffic to be associated with biodiversity declines, that is, loss of fish and macroinvertebrate taxonomic richness, diversity and trait richness. Ship traffic was also associated with increases in taxonomic evenness, which, in concert with richness decreases, was attributed to losses in rare taxa. Ship traffic was especially harmful for benthic taxa and those preferring slow flows. These effects often depended on local land use and riparian degradation. In fish, negative impacts of shipping were highest in urban and agricultural landscapes. Regarding navigation infrastructure, the negative impact of channelization on macroinvertebrates was evident only when riparian degradation was also high. Our results demonstrate the risk of increasing inland navigation on freshwater biodiversity. Integrative waterway management accounting for riparian habitats and landscape characteristics could help to mitigate these impacts.

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.

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.

Bending Behavior of Hybrid Timber-Steel Beams.

Driven by climate change and the need for a more sustainable construction sector, policy is increasingly demanding and promoting timber hybrid construction methods. In the German state of Baden-Württemberg, every new public building has to be of timber or timber hybrid construction (Holzbauoffensive BW). The objective of multi-story buildings with large floor spans can only be achieved in a resource-efficient way by hybrid constructions combining timber and steel components. A research project recently completed at the Karlsruhe Institute of Technology was aimed at the development and systematic investigation of hybrid bending beams in which an advantageous combination of the materials steel and timber is used. For this purpose, steel profiles are integrated into timber cross-sections in a shear-resistant manner by adhesive bonding. As part of the experimental, numerical and analytical investigations, different cross-sections of steel and timber, as well as different construction materials, were considered (GL24h, LVL48p, LVL80p, S355 and S420). The results of large-scale four-point bending tests illustrate the potential of this new hybrid construction method. Depending on the geometry and material combinations tested, the bending stiffness could be increased by up to 250%, and the load-carrying capacity by up to 120%, compared to a glulam beam with identical dimensions.

Understanding temporal variability across trophic levels and spatial scales in freshwater ecosystems.

A tenet of ecology is that temporal variability in ecological structure and processes tends to decrease with increasing spatial scales (from locales to regions) and levels of biological organization (from populations to communities). However, patterns in temporal variability across trophic levels and the mechanisms that produce them remain poorly understood. Here we analyzed the abundance time series of spatially structured communities (i.e., metacommunities) spanning basal resources to top predators from 355 freshwater sites across three continents. Specifically, we used a hierarchical partitioning method to disentangle the propagation of temporal variability in abundance across spatial scales and trophic levels. We then used structural equation modeling to determine if the strength and direction of relationships between temporal variability, synchrony, biodiversity, and environmental and spatial settings depended on trophic level and spatial scale. We found that temporal variability in abundance decreased from producers to tertiary consumers but did so mainly at the local scale. Species population synchrony within sites increased with trophic level, whereas synchrony among communities decreased. At the local scale, temporal variability in precipitation and species diversity were associated with population variability (linear partial coefficient, ß = 0.23) and population synchrony (ß = -0.39) similarly across trophic levels, respectively. At the regional scale, community synchrony was not related to climatic or spatial predictors, but the strength of relationships between metacommunity variability and community synchrony decreased systematically from top predators (ß = 0.73) to secondary consumers (ß = 0.54), to primary consumers (ß = 0.30) to producers (ß = 0). Our results suggest that mobile predators may often stabilize metacommunities by buffering variability that originates at the base of food webs. This finding illustrates that the trophic structure of metacommunities, which integrates variation in organismal body size and its correlates, should be considered when investigating ecological stability in natural systems. More broadly, our work advances the notion that temporal stability is an emergent property of ecosystems that may be threatened in complex ways by biodiversity loss and habitat fragmentation.

The recovery of European freshwater biodiversity has come to a halt.

Owing to a long history of anthropogenic pressures, freshwater ecosystems are among the most vulnerable to biodiversity loss1. Mitigation measures, including wastewater treatment and hydromorphological restoration, have aimed to improve environmental quality and foster the recovery of freshwater biodiversity2. 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.

Long-term trends in abundances of non-native species across biomes, realms, and taxonomic groups in Europe.

Rates of biological invasion have increased over recent centuries and are expected to increase in the future. Whereas increasing rates of non-native species incursions across realms, taxonomic groups, and regions are well-reported, trends in abundances within these contexts have lacked analysis due to a paucity of long-term data at large spatiotemporal scales. These knowledge gaps impede prioritisation of realms, regions, and taxonomic groups for management. We analysed 180 biological time series (median 15 ± 12.8 sampling years) mainly from Long-Term Ecological Research (LTER) sites comprising abundances of marine, freshwater, and terrestrial non-native species in Europe. A high number (150; 83,3 %) of these time series were invaded by at least one non-native species. We tested whether (i) local long-term abundance trends of non-native species are consistent among environmental realms, taxonomic groups, and regions, and (ii) if any detected trend can be explained by climatic conditions. Our results indicate that abundance trends at local scales are highly variable, with evidence of declines in marine and freshwater long-term monitoring sites, despite non-native species reports increasing rapidly since the late 1970s. These declines were driven mostly by abundance trends in non-native fish, birds, and invertebrate species in three biogeographic regions (Continental, Atlantic, and the North Sea). Temperature and precipitation were important predictors of observed abundance trends across Europe. Yet, the response was larger for species with already declining trends and differed among taxa. Our results indicate that trends in biological invasions, especially across different taxonomic groups, are context-dependent and require robust local data to understand long-term trends across contexts at large scales. While the process of biological invasion is spatiotemporally broad, economic or ecological impacts are generally realised on the local level. Accordingly, we urge proactive and coordinated management actions from local to large scales, as invasion impacts are substantial and dynamics are prone to change.

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.

Multidecadal data indicate increase of aquatic insects in Central European streams.

In recent years, declining insect biodiversity has sparked interest among scientists and drawn the attention of society and politicians. However, our understanding of the extent of this decline is incomplete, particularly for freshwater insects that provide a key trophic link between aquatic and terrestrial ecosystems, but that are also especially vulnerable to climate change. To investigate the response of freshwater insects to climate change, we quantified shifts in insect abundance and diversity across 7264 samples covering Central Europe during 1990-2018 and related these changes to annual data on temperature and precipitation. We observed both increases in richness (10.6 %) and abundance (9.5 %) of freshwater insects over the past three decades. These changes were related to increases in summer temperature and summer precipitation, which had negative effects on species richness, and to increases in winter temperature and precipitation, which had positive effects. Further we found that increased temperature was generally related to increased abundance, whereas increased precipitation was associated with declines, thus highlighting the particularly varying impacts on differing insect orders. Given that freshwater insects have been more severely affected by global change than marine and terrestrial species, the observed increases are a positive sign, but the overall situation of freshwater invertebrates is still critical.

Long-term trends and drivers of biological invasion in Central European streams.

Rates of biological invasion continue to accelerate and threaten the structure and function of ecosystems worldwide. High habitat connectivity, multiple pathways, and inadequate monitoring have rendered aquatic ecosystems vulnerable to species introductions. Past riverine invasion dynamics were largely restricted to large rivers, leaving out smaller rivers that commonly harbour high freshwater biodiversity. Moreover, biodiversity time series have rarely been used to investigate invasions across larger spatial-temporal scales, limiting our understanding of aquatic invasion dynamics. Here, we used 6067 benthic invertebrate samples from streams and small rivers from the EU Water Framework Directive monitoring program collected across Central Europe between 2000 and 2018 to assess temporal changes to benthic invertebrate communities as well as non-native species. We assessed invasion rates according to temperature, precipitation, elevation, latitude, longitude, and stream type. Overall, average daily temperatures significantly increased by 0.02 °C per annum (0.34 °C in total) while annual precipitation significantly decreased by 0.01 mm per annum (-67.8 mm over the study period), paralleled with significant increases in overall species richness (12.3 %) and abundance (14.9 %); water quality was relatively stable. Non-native species richness increased 5-fold and abundance 40-fold, indicating an ongoing community shift from native to non-native species. The observed increase in invasions was stronger in low mountain rivers compared to low mountain streams, with the share of non-native species abundance and richness declining with increasing elevation and latitude but increasing with temperature. We found thermophilic non-native species invasion success was greatest in larger sized streams, at lower latitudes, lower elevations and higher temperatures. These results indicate that widespread environmental characteristics (i.e., temperature) could heighten invasion success and confer refuge effects (i.e., elevation and latitude) in higher sites. High altitude and latitude environments should be prioritised for prevention efforts, while biosecurity and management should be improved in lowland areas subject to greater anthropogenic pressure, where non-native introductions are more likely.

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.

Alien species and climate change drive shifts in a riverine fish community and trait compositions over 35 years.

Alien fish substantially impact aquatic communities. However, their effects on trait composition remain poorly understood, especially at large spatiotemporal scales. Here, we used long-term biomonitoring data (1984-2018) from 31 fish communities of the Rhine river in Germany to investigate compositional and functional changes over time. Average total community richness increased by 49 %: it was stable until 2004, then declined until 2010, before increasing until 2018. Average abundance decreased by 9 %. Starting from 198 individuals/m2 in 1984 abundance largely declined to 23 individuals/m2 in 2010 (-88 %), and then consequently increased by 678 % up to 180 individuals/m2 until 2018. Increases in abundance and richness starting around 2010 were mainly driven by the establishment of alien species: while alien species represented 5 % of all species and 0.1 % of total individuals in 1993, it increased to 30 % (7 species) and 32 % of individuals in 2018. Concomitant to the increase in alien species, average native species richness and abundance declined by 26 % and 50 % respectively. We identified increases in temperature, precipitation, abundance and richness of alien fish driving compositional changes after 2010. To get more insights on the impacts of alien species on fish communities, we used 12 biological and 13 ecological traits to compute four trait metrics each. Ecological trait dispersion increased before 2010, probably due to diminishing ecologically similar native species. No changes in trait metrics were measured after 2010, albeit relative shares of expressed trait modalities significantly changing. The observed shift in trait modalities suggested the introduction of new species carrying similar and novel trait modalities. Our results revealed significant changes in taxonomic and trait compositions following alien fish introductions and climatic change. To conclude, our analyses show taxonomic and functional changes in the Rhine river over 35 years, likely indicative of future changes in ecosystem services.

Evaluating the biological validity of European river typology systems with least disturbed benthic macroinvertebrate communities.

Humans have severely altered freshwater ecosystems globally, causing a loss of biodiversity. Regulatory frameworks, like the Water Framework Directive, have been developed to support actions that halt and reverse this loss. These frameworks use typology systems that summarize freshwater ecosystems into environmentally delineated types. Within types, ecosystems that are minimally impacted by human activities, i.e., in reference conditions, are expected to be similar concerning physical, chemical, and biological characteristics. This assumption is critical when water quality assessments rely on comparisons to type-specific reference conditions. Lyche Solheim et al. (2019) developed a pan-European river typology system, the Broad River Types, that unifies the national Water Framework Directive typology systems and is gaining traction within the research community. However, it is unknown how similar biological communities are within these individual Broad River Types. We used analysis of similarities and classification strength analysis to examine if the Broad River Types delineate distinct macroinvertebrate communities across Europe and whether they outperform two ecoregional approaches: the European Biogeographical Regions and Illies' Freshwater Ecoregions. We determined indicator and typical taxa for the types of all three typology systems and evaluated their distinctiveness. All three typology systems captured more variation in macroinvertebrate communities than random combinations of sites. The results were similar among typology systems, but the Broad River Types always performed worse than either the Biogeographic Regions or Illies' Freshwater Ecoregions. Despite reaching statistical significance, the statistics of analysis of similarity and classification strength were low in all tests indicating substantial overlap among the macroinvertebrate communities of different types. We conclude that the Broad River Types do not represent an improvement upon existing freshwater typologies when used to delineate macroinvertebrate communities and we propose future avenues for advancement: regionally constrained types, better recognition of intermittent rivers, and consideration of biotic communities.

Invasion impacts and dynamics of a European-wide introduced species.

Globalization has led to the introduction of thousands of alien species worldwide. With growing impacts by invasive species, understanding the invasion process remains critical for predicting adverse effects and informing efficient management. Theoretically, invasion dynamics have been assumed to follow an "invasion curve" (S-shaped curve of available area invaded over time), but this dynamic has lacked empirical testing using large-scale data and neglects to consider invader abundances. We propose an "impact curve" describing the impacts generated by invasive species over time based on cumulative abundances. To test this curve's large-scale applicability, we used the data-rich New Zealand mud snail Potamopyrgus antipodarum, one of the most damaging freshwater invaders that has invaded almost all of Europe. Using long-term (1979-2020) abundance and environmental data collected across 306 European sites, we observed that P. antipodarum abundance generally increased through time, with slower population growth at higher latitudes and with lower runoff depth. Fifty-nine percent of these populations followed the impact curve, characterized by first occurrence, exponential growth, then long-term saturation. This behaviour is consistent with boom-bust dynamics, as saturation occurs due to a rapid decline in abundance over time. Across sites, we estimated that impact peaked approximately two decades after first detection, but the rate of progression along the invasion process was influenced by local abiotic conditions. The S-shaped impact curve may be common among many invasive species that undergo complex invasion dynamics. This provides a potentially unifying approach to advance understanding of large-scale invasion dynamics and could inform timely management actions to mitigate impacts on ecosystems and economies.

Sampling and modelling rare species: Conceptual guidelines for the neglected majority.

Biodiversity conservation faces a methodological conundrum: Biodiversity measurement often relies on species, most of which are rare at various scales, especially prone to extinction under global change, but also the most challenging to sample and model. Predicting the distribution change of rare species using conventional species distribution models is challenging because rare species are hardly captured by most survey systems. When enough data are available, predictions are usually spatially biased towards locations where the species is most likely to occur, violating the assumptions of many modelling frameworks. Workflows to predict and eventually map rare species distributions imply important trade-offs between data quantity, quality, representativeness and model complexity that need to be considered prior to survey and analysis. Our opinion is that study designs need to carefully integrate the different steps, from species sampling to modelling, in accordance with the different types of rarity and available data in order to improve our capacity for sound assessment and prediction of rare species distribution. In this article, we summarize and comment on how different categories of species rarity lead to different types of occurrence and distribution data depending on choices made during the survey process, namely the spatial distribution of samples (where to sample) and the sampling protocol in each selected location (how to sample). We then clarify which species distribution models are suitable depending on the different types of distribution data (how to model). Among others, for most rarity forms, we highlight the insights from systematic species-targeted sampling coupled with hierarchical models that allow correcting for overdispersion and spatial and sampling sources of bias. Our article provides scientists and practitioners with a much-needed guide through the ever-increasing diversity of methodological developments to improve the prediction of rare species distribution depending on rarity type and available data.

A global agenda for advancing freshwater biodiversity research.

Global freshwater biodiversity is declining dramatically, and meeting the challenges of this crisis requires bold goals and the mobilisation of substantial resources. While the reasons are varied, investments in both research and conservation of freshwater biodiversity lag far behind those in the terrestrial and marine realms. Inspired by a global consultation, we identify 15 pressing priority needs, grouped into five research areas, in an effort to support informed stewardship of freshwater biodiversity. The proposed agenda aims to advance freshwater biodiversity research globally as a critical step in improving coordinated actions towards its sustainable management and conservation.

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.

Multidecadal changes in functional diversity lag behind the recovery of taxonomic diversity.

While there has been increasing interest in how taxonomic diversity is changing over time, less is known about how long-term taxonomic changes may affect ecosystem functioning and resilience. Exploring long-term patterns of functional diversity can provide key insights into the capacity of a community to carry out ecological processes and the redundancy of species' roles. We focus on a protected freshwater system located in a national park in southeast Germany. We use a high-resolution benthic macroinvertebrate dataset spanning 32 years (1983-2014) and test whether changes in functional diversity are reflected in taxonomic diversity using a multidimensional trait-based approach and regression analyses. Specifically, we asked: (i) How has functional diversity changed over time? (ii) How functionally distinct are the community's taxa? (iii) Are changes in functional diversity concurrent with taxonomic diversity? And (iv) what is the extent of community functional redundancy? Resultant from acidification mitigation, macroinvertebrate taxonomic diversity increased over the study period. Recovery of functional diversity was less pronounced, lagging behind responses of taxonomic diversity. Over multidecadal timescales, the macroinvertebrate community has become more homogenous with a high degree of functional redundancy, despite being isolated from direct anthropogenic activity. While taxonomic diversity increased over time, functional diversity has yet to catch up. These results demonstrate that anthropogenic pressures can remain a threat to biotic communities even in protected areas. The differences in taxonomic and functional recovery processes highlight the need to incorporate functional traits in assessments of biodiversity responses to global change.

Long-term data reveal unimodal responses of ground beetle abundance to precipitation and land use but no changes in taxonomic and functional diversity.

While much of global biodiversity is undoubtedly under threat, the responses of ecological communities to changing climate, land use intensification, and long-term changes in both taxonomic and functional diversity over time, has still not been fully explored for many taxonomic groups, especially invertebrates. We compiled time series of ground beetles covering the past two decades from 40 sites located in five regions across Germany. We calculated site-based trends for 21 community metrics representing taxonomic and functional diversity of ground beetles, activity density (a proxy for abundance), and activity densities of functional groups. We assessed both overall and regional temporal trends and the influence of the global change drivers of temperature, precipitation, and land use on ground beetle communities. While we did not detect overall temporal changes in ground beetle taxonomic and functional diversity, taxonomic turnover changed within two regions, illustrating that community change at the local scale does not always correspond to patterns at broader spatial scales. Additionally, ground beetle activity density had a unimodal response to both annual precipitation and land use. Limited temporal change in ground beetle communities may indicate a shifting baseline, where community degradation was reached prior to the start of our observation in 1999. In addition, nonlinear responses of animal communities to environmental change present a challenge when quantifying temporal trends.