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.N. Kaffenberger helped with initial data compilation. Funding for authors and data collection and processing was provided by the EU Horizon 2020 project eLTER PLUS (grant agreement no. 871128); the German Federal Ministry of Education and Research (BMBF; 033W034A); the German Research Foundation (DFG FZT 118, 202548816); Czech Republic project no. P505-20-17305S; the Leibniz Competition (J45/2018, P74/2018); the Spanish Ministerio de Economía, Industria y Competitividad—Agencia Estatal de Investigación and the European Regional Development Fund (MECODISPER project CTM 2017-89295-P); Ramón y Cajal contracts and the project funded by the Spanish Ministry of Science and Innovation (RYC2019-027446-I, RYC2020-029829-I, PID2020-115830GB-100); the Danish Environment Agency; the Norwegian Environment Agency; SOMINCOR—Lundin mining & FCT—Fundação para a Ciência e Tecnologia, Portugal; the Swedish University of Agricultural Sciences; the Swiss National Science Foundation (grant PP00P3_179089); the EU LIFE programme (DIVAQUA project, LIFE18 NAT/ES/000121); the UK Natural Environment Research Council (GLiTRS project NE/V006886/1 and NE/R016429/1 as part of the UK-SCAPE programme); the Autonomous Province of Bolzano (Italy); and the Estonian Research Council (grant no. PRG1266), Estonian National Program ‘Humanitarian and natural science collections’. The Environment Agency of England, the Scottish Environmental Protection Agency and Natural Resources Wales provided publicly available data. We acknowledge the members of the Flanders Environment Agency for providing data. This article is a contribution of the Alliance for Freshwater Life (www.allianceforfreshwaterlife.org).Peer reviewe

Biodiversity through time:coherence, stability and species turnover in boreal stream communities

Abstract Describing how and why species composition of ecological communities varies across spatial and temporal scales is a primary objective for ecological research. A key challenge is to distinguish changes in community composition resulting from external factors from the natural background variability. In this thesis I aimed to study: 1) the level of temporal variation in community composition of stream macroinvertebrates, 2) the role of different environmental factors to temporal variability, 3) the effect of temporal variability on bioassessment outcomes, and 4) comparability of different approaches to study community variability through time. A majority of the studied macroinvertebrate communities showed lower level of inter-annual variation, i.e. temporal turnover, than expected by chance. The observation of high community stability was further supported by the low level of inter-annual variation in taxonomic completeness (quotient of observed and expected number of species, O/E). Despite the low absolute variation in O/E, ecological status assessments varied annually. Thus the use of one year data may bias management decisions. Macroinvertebrate communities experienced similar dynamics across several spatial extents, from riffles within a stream to streams among regions, suggesting that large-scale extrinsic factors are the major driver of community dynamics. Especially climatically exceptional years may have a strong imprint on community variability. However, at the within-stream scale, coherence was lower than expected, indicating that community dynamics may be driven by different processes at different spatial extents. Stream macroinvertebrate community dynamics were strongly related to in-stream vegetation, temporal variability decreasing with increasing macrophyte cover. Importantly, the effect of in-stream vegetation on temporal turnover of macroinvertebrate communities was masked by the stochastic effect of habitat connectivity, suggesting that unless stochastic effects are controlled for, the role of deterministic processes may be obscured, thus affecting our ability to understand and predict community changes through time. In addition, different approaches to study temporal variability may disagree on estimates for the level of temporal turnover and factors explaining it – a fact that should be taken into account when planning and comparing studies.Tiivistelmä Yksi ekologisen tutkimuksen keskeisistä tavoitteista on kuvata, miten ja miksi eliöyhteisöjen koostumus muuttuu paikasta ja ajankohdasta toiseen. On tärkeää pystyä erottamaan erilaisten ulkoisten tekijöiden aiheuttamat muutokset luonnollisesta taustavaihtelusta. Väitöskirjani tavoitteena oli selvittää 1) miten paljon virtavesien pohjaeläinyhteisöissä tapahtuu ajallista vaihtelua 2) mitkä ympäristötekijät vaikuttavat yhteisöjen ajalliseen vaihteluun 3) miten ajallinen vaihtelu vaikuttaa ympäristön tilan arviointiin ja 4) kuinka vertailukelpoisia ovat eri lähestymistavat ajallista vaihtelua tutkittaessa. Valtaosa tutkituista pohjaeläinyhteisöistä vaihteli vuosien välillä vähemmän kuin olisi sattumalta odotettavissa osoittaen varsin suurta ajallista pysyvyyttä. Käsitystä yhteisöjen pysyvyydestä tuki myös vähäinen vuosittainen vaihtelu ekologista tilaa kuvaavassa taksonomisessa eheydessä (=havaitun ja odotetun lajiston suhde O/E). Huolimatta näennäisen pienestä vaihtelusta O/E suhteessa paikkakohtaiset tilaluokka-arviot saattoivat vaihtua vuodesta toiseen. Yhden vuoden aineistoon perustuvat tilan arvioinnit voivat siis johtaa virheellisiin johtopäätöksiin. Pohjaeläinyhteisöjen ajallinen vaihtelu oli samankaltaista eri mittakaavoilla niin peräkkäisten koskipaikkojen kuin eri alueilla sijaitsevien purojen välillä. Suuren mittakaavan ympäristötekijät näyttävät siis säätelevän eliöyhteisöjen ajallista vaihtelua. Erityisesti ilmastotekijöiltään poikkeukselliset vuodet säätelevät eliöyhteisöjä, ja niiden vaikutus voi näkyä vielä useiden vuosien kuluttua. Vaihtelun samankaltaisuus peräkkäisten koskipaikkojen välillä oli kuitenkin odotettua pienempää. Yhteisöjä voivat siis säädellä osittain eri tekijät eri mittakaavoilla. Tutkittujen pohjaeläinyhteisöjen ajallisen vaihtelun voimakkuus liittyi erityisesti vesikasvillisuuden määrään: vaihtelu väheni kasvillisuuden lisääntyessä. Kasvillisuuden määrän vaikutus peittyi kuitenkin satunnaisten tekijöiden alle. Jos satunnaisia tekijöitä ei huomioida, deterministiset syy-seuraussuhteet voivat jäädä huomaamatta heikentäen mahdollisuuksiamme ymmärtää ja ennustaa eliöyhteisöjen vaihtelua. Lisäksi eri lähestymistavat ajallista vaihtelua tutkittaessa voivat johtaa erilaisiin arvioihin vaihtelun suuruudesta ja siihen vaikuttavista tekijöistä, mikä tulisi ottaa huomioon tutkimuksia suunnitellessa ja niiden tuloksia vertailtaessa

Estimates of benthic invertebrate community variability and its environmental determinants differ between snapshot and trajectory designs

Abstract Long-term data sets are essential for biodiversity research and monitoring. Researchers use 2 major approaches in the study of temporal variability of biological communities: 1) the trajectory approach (monitoring sites across several consecutive years) and 2) the snapshot approach (comparing sites among few sampling events several years apart). We used data on benthic macroinvertebrate communities in 23 near-pristine forested streams to compare these 2 approaches for different study periods ranging from 3 to 14 y. We asked whether the level of temporal turnover and the identity of the best explanatory variables underlying it were comparable across studies based on differing approaches, study periods, or total duration. The 2 approaches yielded partly different stories about the level of community variability and its environmental correlates. With the snapshot approach, variation in community similarity and factors explaining it reflected short-term (e.g., year-specific) conditions, which could be misinterpreted as long-term trends, the difference being most evident for periods that began or ended in an extreme drought year. Our results imply that snapshot studies may lead to ambiguous conclusions, whereas the trajectory approach yielded more consistent results. Trajectory data of differing length showed minor differences, apart from studies with the shortest durations. Overall, our results suggest that time sequences of ∼6 y of trajectory data (i.e., 6 generations for most benthic invertebrates in boreal streams) may be needed for the among-year similarity of macroinvertebrate communities in near-pristine streams to stabilize. If temporal replication is limited (snapshots/very short time sequences) the outcome depends strongly on the particular years included in a comparison. Based on our results, we advise caution when basing conclusions on a comparison of a few (e.g., just 2) occasions several years apart or on very short time sequences

Bacterial communities in a subarctic stream network:spatial and seasonal patterns of benthic biofilm and bacterioplankton

Abstract Water-column bacterial communities are assembled by different mechanisms at different stream network positions, with headwater communities being controlled by mass effects (advection of bacteria from terrestrial soils) while downstream communities are mainly driven by environmental sorting. Conversely, benthic biofilms are colonized largely by the same set of taxa across the entire network. However, direct comparisons of biofilm and bacterioplankton communities along whole stream networks are rare. We used 16S rRNA gene amplicon sequencing to explore the spatiotemporal variability of benthic biofilm (2 weeks old vs. mature biofilm) and water-column communities at different network positions of a subarctic stream from early summer to late autumn. Amplicon sequence variant (ASV) richness of mature biofilm was about 2.5 times higher than that of early biofilm, yet the pattern of seasonality was the same, with the highest richness in midsummer. Biofilm bacterial richness was unrelated to network position whereas bacterioplankton diversity was negatively related to water residence time and distance from the source. This pattern of decreasing diversity along the network was strongest around midsummer and diminished greatly as water level increased towards autumn. Biofilm communities were phylogenetically clustered at all network positions while bacterioplankton assemblages were phylogenetically clustered only at the most downstream site. Both early and mature biofilm communities already differed significantly between upstream (1st order) and midstream (2nd order) sections. Network position was also related to variation in bacterioplankton communities, with upstream sites harbouring substantially more unique taxa (44% of all upstream taxa) than midstream (20%) or downstream (8%) sites. Some of the taxa that were dominant in downstream sections were already present in the upmost headwaters, and even in riparian soils, where they were very rare (relative abundance <0.01%). These patterns in species diversity and taxonomic and phylogenetic community composition of the riverine bacterial metacommunity were particularly strong for water-column communities, whereas both early and mature biofilm exhibited weaker spatial patterns. Our study demonstrated the benefits of studying bacterioplankton and biofilm communities simultaneously to allow testing of ecological hypotheses about biodiversity patterns in freshwater bacteria

Partitioning of benthic biodiversity in boreal streams:contributions of spatial, inter-annual, and seasonal variability

Abstract 1. Studies on biodiversity patterns should optimally relate different scales of temporal community variability to spatial variability. Although temporal biodiversity variability is often negligible compared to spatial variation, it may still constitute a substantial source of overall community variability in stream ecosystems. Boreal streams exhibit seasonally recurring environmental periodicity, which can be expected to induce synchronous dynamics of abiotic variables among sites, and consequently, to produce spatial synchrony of deterministically controlled biological communities with higher intra- than inter-annual community variability. 2. We sampled benthic macroinvertebrates in 10 near-pristine boreal streams on three different seasons (spring, summer, autumn) across 4 consecutive years in northern Finland. We aimed to identify the relative contributions of spatial, inter-annual, and seasonal variability to overall benthic biodiversity; and relate variation in benthic invertebrate communities to key environmental factors, particularly in-stream habitat diversity. 3. Among-site spatial variability was clearly the most important source of variation for both species richness and community dissimilarity. Of the two temporal scales, inter-annual variability contributed more to variation in taxonomic richness and seasonal variability slightly more to variability in community composition. 4. Only inter-annual variation differed systematically from random expectation, indicating strong stability (low variability) of stream macroinvertebrate communities across years, with less variation at sites with higher substrate heterogeneity. Considering the distinct seasonality of the boreal stream environment, seasonal variability accounted for an unexpectedly low amount of total community variability. 5. Although differences between seasons were small, autumnal sampling is likely to be the least susceptible to climatic vagaries, thus providing the most consistent and predictable conditions for benthic sampling in boreal streams, particularly for bioassessment purposes. Exceptional climatic conditions are becoming more frequent in northern Europe, probably causing substantial and largely unpredictable changes in benthic community composition. As a result, the importance of temporal (relative to spatial) community variability may increase

Excess of nitrogen reduces temporal variability of stream diatom assemblages

Abstract Nutrient enrichment degrades water quality and threatens aquatic biota. However, our knowledge on (dis)similarities in temporal patterns of biota among sites of varying level of nutrient stress is limited. We addressed this gap by assessing temporal (among seasons) variation in algal biomass, species diversity and composition of diatom assemblages in three streams that differ in nutrient stress, but are otherwise similar and share the same regional species pool. We monitored three riffle sections in each stream bi-weekly from May to October in 2014. Temporal variation in water chemistry and other environmental variables was mainly synchronous among riffles within streams and often also among streams, indicating shared environmental forcing through time. We found significant differences in diatom assemblage composition among streams and, albeit less so, also among riffles within streams. Diatom assemblages in the two nutrient-enriched streams were more similar to each other than to those in the nutrient-poor stream. Taxa richness did not differ consistently among the streams, and did not vary synchronously at any spatial scale. Temporal variation in diatom assemblage composition decreased with increasing DIN:TotP ratio, likely via a negative effect on sensitive taxa while maintaining favorable conditions for certain tolerant taxa, irrespective of season. This relationship weakened but remained significant even after controlling for stochastic effects, suggesting deterministic mechanisms between nutrient levels and diatom assemblage stability. After controlling for stochastic effects temporal variability was best explained by DIN suggesting that excess of nitrogen reduces temporal variability(intra-annual beta diversity) of diatom assemblages. The high temporal variation, and especially the lack of temporal synchrony at the within streams scale, suggests that single sampling at a single site may be insufficient to reliably assess and monitor a complete stream water body. Our results also showed that measures including species identity outperform traditional diversity metrics in detecting nutrient stress in streams

Drainage-induced browning causes both loss and change of benthic biodiversity in headwater streams

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

Dark matters:contrasting responses of stream biofilm to browning and loss of riparian shading

Abstract Concentrations of terrestrial-derived dissolved organic carbon (DOC) in freshwater ecosystems have increased consistently, causing freshwater browning. The mechanisms behind browning are complex, but in forestry-intensive regions browning is accelerated by land drainage. Forestry actions in streamside riparian forests alter canopy shading, which together with browning is expected to exert a complex and largely unpredictable control over key ecosystem functions. We conducted a stream mesocosm experiment with three levels of browning (ambient vs. moderate vs. high, with 2.7 and 5.5-fold increase, respectively, in absorbance) crossed with two levels of riparian shading (70% light reduction vs. open canopy) to explore the individual and combined effects of browning and loss of shading on the quantity (algal biomass) and nutritional quality (polyunsaturated fatty acid and sterol content) of the periphytic biofilm. We also conducted a field survey of differently colored (4.7 to 26.2 mg DOC L−1) streams to provide a ‘reality check’ for our experimental findings. Browning reduced greatly the algal biomass, suppressed the availability of essential polyunsaturated fatty acids, especially eicosapentaenoic acid (EPA), and sterols, but increased the availability of terrestrial-derived long-chain saturated fatty acids (LSAFA). In contrast, loss of shading increased primary productivity, which resulted in elevated sterol and EPA contents of the biofilm. The field survey largely repeated the same pattern: biofilm nutritional quality decreased significantly with increasing DOC, as indicated particularly by a decrease of the ω-3:ω-6 ratio and increase in LSAFA content. Algal biomass, in contrast, was mainly controlled by dissolved inorganic nitrogen (DIN) concentration, while DOC concentration was of minor importance. The ongoing browning process is inducing a dramatic reduction in the nutritional quality of the stream biofilm. Such degradation of the major high-quality food source available for stream consumers may reduce the trophic transfer efficiency in stream ecosystems, potentially extending across the stream-forest ecotone

Meta-analysis of multidecadal biodiversity trends in Europe

Abstract Local biodiversity trends over time are likely to be decoupled from global trends, as local processes may compensate or counteract global change. We analyze 161 long-term biological time series (15–91 years) collected across Europe, using a comprehensive dataset comprising ~6,200 marine, freshwater and terrestrial taxa. We test whether (i) local long-term biodiversity trends are consistent among biogeoregions, realms and taxonomic groups, and (ii) changes in biodiversity correlate with regional climate and local conditions. Our results reveal that local trends of abundance, richness and diversity differ among biogeoregions, realms and taxonomic groups, demonstrating that biodiversity changes at local scale are often complex and cannot be easily generalized. However, we find increases in richness and abundance with increasing temperature and naturalness as well as a clear spatial pattern in changes in community composition (i.e. temporal taxonomic turnover) in most biogeoregions of Northern and Eastern Europe

The global EPTO database:worldwide occurrences of aquatic insects

Abstract Motivation: Aquatic insects comprise 64% of freshwater animal diversity and are widely used as bioindicators to assess water quality impairment and freshwater ecosystem health, as well as to test ecological hypotheses. Despite their importance, a comprehensive, global database of aquatic insect occurrences for mapping freshwater biodiversity in macroecological studies and applied freshwater research is missing. We aim to fill this gap and present the Global EPTO Database, which includes worldwide geo-referenced aquatic insect occurrence records for four major taxa groups: Ephemeroptera, Plecoptera, Trichoptera and Odonata (EPTO). Main type of variables contained: A total of 8,368,467 occurrence records globally, of which 8,319,689 (99%) are publicly available. The records are attributed to the corresponding drainage basin and sub-catchment based on the Hydrography90m dataset and are accompanied by the elevation value, the freshwater ecoregion and the protection status of their location. Spatial location and grain: The database covers the global extent, with 86% of the observation records having coordinates with at least four decimal digits (11.1 m precision at the equator) in the World Geodetic System 1984 (WGS84) coordinate reference system. Time period and grain: Sampling years span from 1951 to 2021. Ninety-nine percent of the records have information on the year of the observation, 95% on the year and month, while 94% have a complete date. In the case of seven sub-datasets, exact dates can be retrieved upon communication with the data contributors. Major taxa and level of measurement: Ephemeroptera, Plecoptera, Trichoptera and Odonata, standardized at the genus taxonomic level. We provide species names for 7,727,980 (93%) records without further taxonomic verification. Software format: The entire tab-separated value (.csv) database can be downloaded and visualized at https://glowabio.org/project/epto_database/. Fifty individual datasets are also available at https://fred.igb-berlin.de, while six datasets have restricted access. For the latter, we share metadata and the contact details of the authors