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Human impacts, particularly nutrient pollution and land-use change, have caused significant declines in the quality and quantity of freshwater resources. Most global assessments have concentrated on species diversity and composition, but effects on the multifunctionality of streams and rivers remain unclear. Here, we analyse the most comprehensive compilation of stream ecosystem functions to date to provide an overview of the responses of nutrient uptake, leaf litter decomposition, ecosystem productivity, and food web complexity to six globally pervasive human stressors. We show that human stressors inhibited ecosystem functioning for most stressor-function pairs. Nitrate uptake efficiency was most affected and was inhibited by 347% due to agriculture. However, concomitant negative and positive effects were common even within a given stressor-function pair. Some part of this variability in effect direction could be explained by the structural heterogeneity of the landscape and latitudinal position of the streams. Ranking human stressors by their absolute effects on ecosystem multifunctionality revealed significant effects for all studied stressors, with wastewater effluents (194%), agriculture (148%), and urban land use (137%) having the strongest effects. Our results demonstrate that we are at risk of losing the functional backbone of streams and rivers if human stressors persist in contemporary intensity, and that freshwaters are losing critical ecosystem services that humans rely on. We advocate for more studies on the effects of multiple stressors on ecosystem multifunctionality to improve the functional understanding of human impacts. Finally, freshwater management must shift its focus toward an ecological function-based approach and needs to develop strategies for maintaining or restoring ecosystem functioning of streams and rivers. Generalitat de Catalunya, Grant/Award Number: 2017SGR0976; Portuguese Foundation for Science and Technology, Grant/Award Number: CEEIND/02484/2018 and UIDP/04292/2020

[EN] Water diversion and pollution are two pervasive stressors in river ecosystems that often co-occur. Individual effects of both stressors on basal resources available to stream communities have been described, with diversion reducing detritus standing stocks and pollution increasing biomass of primary producers. However, interactive effects of both stressors on the structure and trophic basis of food webs remain unknown. We hypothesized that the interaction between both stressors increases the contribution of the green pathway in stream food webs. Given the key role of the high-quality, but less abundant, primary producers, we also hypothesized an increase in food web complexity with larger trophic diversity in the presence of water diversion and pollution. To test these hypotheses, we selected four rivers in a range of pollution subject to similar water diversion schemes, and we compared food webs upstream and downstream of the diversion. We characterized food webs by means of stable isotope analysis. Both stressors directly changed the availability of basal resources, with water diversion affecting the brown food web by decreasing detritus stocks, and pollution enhancing the green food web by promoting biofilm production. The propagation of the effects at the base of the food web to higher trophic levels differed between stressors. Water diversion had little effect on the structure of food webs, but pollution increased food chain length and trophic diversity, and reduced trophic redundancy. The effects at higher trophic levels were exacerbated when combining both stressors, as the relative contribution of biofilm to the stock of basal resources increased even further. Overall, we conclude that moderate pollution increases food web complexity and that the interaction with water abstraction seems to amplify this effect. Our study shows the importance of assessing the interaction between stressors to create predictive tools for a proper management of ecosystems. Ministerio de Economia, Industria y Competitividad, Gobierno de Espana, Grant/Award Number: GL2016-77487-R; European Social Fund; Diputacion Foral de Bizkaia; Serra Hunter Fellow; Labex, Grant/Award Number: ANR-10-LABX-41; H2020 European Research Council; Eusko Jaurlaritza; Consejo Nacional de Investigaciones Cientificas y Tecnicas; FRAGCLIM Consolidator, Grant/Award Number: 726176

Benthic biofilms are often assumed to control terrestrially-derived dissolved organic carbon (tDOC) uptake in streams. We tested this by comparing 13C-enriched ryegrass leachate uptake in an agricultural and a forest stream, hypothesizing that a greater abundance of autotrophic biofilms in the agricultural stream would cause its whole-stream tDOC uptake to be comparatively low. We measured whole-stream and biofilm tDOC tracer uptake, metabolism, bacterial and algal diversity, and nutrient status of benthic epilithic biofilms, and assessed whole-stream hydromorphology. Whole-stream uptake of tDOC was six times lower in the agricultural (3.0 mg m−2 day−1) than in the forest (19.0 mg m−2 day−1) stream, and tDOC uptake velocity indicated lower tDOC demand in the agricultural (1.2 mm min−1) than in the forest (1.9 mm min−1) stream. The agricultural stream differed from the forest stream by slightly lower transient storage capacity and higher benthic biofilm bacterial abundance and production, lower biofilm biomass and lower biofilm molar C:N, C:P, and N:P ratios. Changes in epilithic biofilms contributed little to the differences in whole-stream tDOC tracer uptake between streams, as biofilm tDOC uptake only amounted to 4% and 13% of whole-stream uptake in the forest and agricultural stream, respectively. This comparison of a forest and an agricultural stream suggests that agricultural stressors have the potential to diminish both whole-stream tDOC uptake and uptake efficiency. Furthermore, the weak link between biofilm and whole-stream tDOC uptake implies that benthic biofilms characteristics are poor predictors for human impacts on tDOC uptake in agricultural streams and that hot spots of tDOC uptake are likely situated in the hyporheic zone or in the stream water column.

The science of the total environment 769, 144324 (2021). doi:10.1016/j.scitotenv.2020.144324, Published by Elsevier Science, Amsterdam [u.a.]

The composition of littoral macroinvertebrate communities in lakes is governed by multiple natural and anthropogenic environmental influences interacting at different spatial scales. Since ecological assessment methods using littoral macroinvertebrates should respond specifically to a single stressor, knowledge on the unique effects of a given stressor is necessary. To effectively disentangle the effects of hydromorphology and trophic state requires analysing macroinvertebrate communities at lake sites with the full range of both stressors. We used a dataset of 98 lakes encompassing the entire gradient of geographical locations, lake types, hydromorphological degradation and trophic states in Central European lakes. We studied the unique and joint effects of hydromorphology and trophic state on macroinvertebrate richness, community composition and the Littoral Invertebrate Multimetric Index based on Composite Sampling (LIMCO). Variation partitioning analyses were conducted to test the importance of hydromorphology relative to trophic state across and within hydromorphological states (natural shorelines, hard and soft shore modifications) and trophic states (oligotrophic to hypertrophic states). At natural, hard and soft modification sites, hydromorphology explained 10, 16 and 19%, respectively, of the average unique variation of diversity, community composition and the LIMCO index, whereas trophic state explained on average 2, 5 and 5%, respectively. Similarly, in low, medium and high trophic state lakes, hydromorphology explained 10, 15 and 7%, respectively, of the average unique variation of diversity, community composition and the LIMCO index, whereas trophic state explained on average 0.3, 3 and 6%, respectively. Our results demonstrate that littoral hydromorphology was a more important driver of macroinvertebrate diversity, community composition and LIMCO than trophic state across hydromorphological states and trophic states. This indicates that multiple stressors in lakes act hierarchically on littoral macroinvertebrate communities and that the hydromorphological degradation of littoral zones is the primary driver for altered communities.

Accidental spills or illegal discharges of pesticides in aquatic ecosystems can lead to exposure levels that strongly exceed authorized pesticide concentrations, causing major impacts on aquatic ecosystems. Such short-term events often remain undetected in regular monitoring programs with infrequent sampling. In early spring 2015, we identified a catastrophic pesticide spill with the insecticide cypermethrin in the Holtemme River, Germany. Based on existing pre-event macroinvertebrate community data, we monitored the effects and recovery of the macroinvertebrate community for more than two years after the spill. Strong short-term effects were apparent for all taxa with the exception of Chironomidae and Tubificidae. Effects could also be observed on the community level as total abundance, taxa number and biomass strongly decreased. Total abundance and taxa number showed a fast recovery. Regarding long-term effects, the total biomass remained substantially below the pre-contamination level (76%) until the end of the study. Also the abundances of three taxa (Gammarus, Leuctra, Limnius Ad.) did not return to levels prior to the spill even after 26 months. This lack of the taxon-specific recovery was likely due to their long generation time and a low migration ability due to a restricted connectivity between the contaminated site and uncontaminated stream sections. These factors proved to be stronger predictors for the recovery than the pesticide tolerance. We revealed that the biological indicators SPEARpesticides and share of Ephemeroptera, Plecoptera and Trichoptera (EPT) are not suitable for the identification of such extreme events, when nearly all taxa are eradicated. Both indicators are functioning only when repeated stressors initiate long-term competitive replacement of sensitive by insensitive taxa. We conclude that pesticide spills can have significant long-term effects on stream macroinvertebrate communities. Regular ecological monitoring is imperative to identify such ecosystem impairments, combined with analytical chemistry methods to identify the potential sources of spills.

Fluvial networks are globally relevant for the processing of dissolved organic matter (DOM). To investigate the change in molecular DOM diversity along the river course, high-field FTICR mass spectrometry and NMR spectroscopy of riverine DOM as well as bacterial abundance and activity were measured in a third order stream along a land-use gradient from pristine, agricultural to urban landscapes. DOM composition showed a clear evolution along the river course with an initial decrease of average oxidation and unsaturation followed by an increased relative abundance of CHNO and CHOS compounds introduced by agriculture and waste water, respectively. DOM composition was dominated by rather unsaturated CHO compounds (H/C ≤ 1) in headwaters and by more aliphatic molecules at downstream sites. Oxygenated functional groups shifted from aromatic ethers and hydroxyl groups to aliphatic carboxylic acids and aliphatic hydroxyl groups. This massive dislocation of oxygen significantly increased the diversity of atomic environments in branched aliphatic groups from headwater to downstream DOM. Mass spectra of DOM enabled the detection of compositional relationships to bacterial abundance and activity which was positively related to more aliphatic components (H/C > 1) and negatively related to unsaturated components. FTICR mass and NMR spectra corroborated the initial decline in DOM molecular diversity predicted by the River Continuum Concept (RCC) but demonstrated an anthropogenic increase in the molecular diversity of DOM further downstream. While the high DOM molecular diversity in first order headwater streams was the result of small scale ecosystem plurality, agriculture and waste water treatment introduced many components in the lower reaches. These anthropogenic influences together with massive bacterial oxidation of DOM contributed to a growth of molecular diversity of downstream DOM whose composition and structure differed entirely from those found in pristine headwaters.

Agricultural and urban land use has dramatically increased over the last century and one consequence is the release of anthropogenic chemicals into aquatic ecosystems. One of the rarely studied consequences is the effect of land use change on internal concentrations of organic micropollutants (OMPs) in aquatic invertebrates and its effects on their genotype diversity. Here, we applied population genetic and internal concentrations of OMPs analyses to determine evolutionary implications of chemical pollution on Gammarus pulex populations from a natural and two agricultural streams. Along 14 consecutive months sampled, 26 different OMPs were quantified in G. pulex extracts with the highest number, concentration, and toxic pressure in the anthropogenically stressed stream ecosystems. Our results indicate distinct internal OMP profiles and changes in both genetic variation and genetic structure in streams affected by anthropogenic activity. Genetic variation was attributed to chemical pollution whereas changes in the genetic structure were attributed to environmental disturbances, such as changes in discharge in the impacted stream ecosystems, which worked both independently and in tandem. Finally, we conclude that human-impacted streams are subjected to severe alterations in their population genetic patterns compared to nonimpacted stream ecosystems.

Length–mass relationships are widely used to estimate body mass from body dimensions for freshwater macroinvertebrates. The relationships are influenced by environmental conditions and should be applied within ecosystems and geographic regions similar to those for which they were estimated. However, very few relationships exist for littoral macroinvertebrates, and thus we provide length–mass relationships for macroinvertebrates from lakes of the Central European lowlands. We compared log-linear and nonlinear methods for fitting length–mass relationships and tested the smearing factor for removing bias in mass predictions from log-linear models. We also estimated conversion factors to correct for mass changes during ethanol preservation and assessed the transferability of our results to different geographical regions. We showed that the log-linear approach gave better results in fitting length–mass relationships, while residuals showed that nonlinear models over-predict the mass of small individuals. The smearing correction factor successfully removed bias introduced by log transformation, and relationships transferred well between lakes in the same and different geographical regions. In total, 52 bias-corrected length–mass relationships are provided for littoral macroinvertebrates that are applicable also to lakes in geographic regions with similar environmental conditions, such as the Central European lowlands or the temperate lowland zone of America.

Summary 1. Shoreline development and the associated loss of littoral habitats represent a pervasive alteration of the ecological integrity of lakes and have been identified as major drivers for the loss of littoral biodiversity world-wide. Little is known about the effects of shoreline development on the structure of, and energy transfer in, littoral food webs, even though this information is urgently needed for management and mitigation measures. 2. We measured macroinvertebrate biomass and analysed potential food resources using stable isotopes (δ13C, δ15N) and mixing models to compare the complexity and the trophic base of littoral food webs between undeveloped and developed shorelines in three North German lowland lakes. 3. The lower diversity of littoral habitats found at developed shorelines was associated with lower diversity of food resources and consumers. Consequently, the number of trophic links in food webs at developed shorelines was up to one order of magnitude lower as compared with undeveloped shorelines. 4. Mixing model analysis showed that consumer biomass at undeveloped shorelines was mainly derived from fine particulate organic matter (FPOM) and coarse particulate organic matter of terrestrial origin (CPOM). The contribution of CPOM to consumer biomass was twofold lower at developed shorelines, and consumer biomass was mainly derived from FPOM and suspended particulate organic matter. 5.Synthesis and application. Shoreline development impacts the flow of organic matter within littoral food webs primarily through the reduction in littoral habitat diversity. These effects are exacerbated by clearcutting of the riparian vegetation, which disrupts cross-boundary couplings between the riparian and the littoral zone. Lakeshore conservation should focus on preserving the structural integrity of the littoral zone, while restoration of coarse woody debris, reed and root habitats can be a cost-efficient measure to improve degraded lakeshores. The local effects of shoreline development demonstrated in this study might lead to whole-lake effects, but future studies are needed to derive thresholds at which shoreline development has consequences for the structure and functioning of the entire ecosystem.

Ecosystems are generally linked via fluxes of nutrients and energy across their boundaries. For example, freshwater ecosystems in temperate regions may receive significant inputs of terrestrially derived carbon via autumnal leaf litter. This terrestrial particulate organic carbon (POC) is hypothesized to subsidize animal production in lakes, but direct evidence is still lacking. We divided two small eutrophic lakes each into two sections and added isotopically distinct maize litter to the treatment sections to simulate increased terrestrial POC inputs via leaf litter in autumn. We quantified the reliance of aquatic consumers on terrestrial resources (allochthony) in the year subsequent to POC additions by applying mixing models of stable isotopes. We also estimated lake-wide carbon (C) balances to calculate the C flow to the production of the major aquatic consumer groups: benthic macroinvertebrates, crustacean zooplankton, and fish. The sum of secondary production of crustaceans and benthic macroinvertebrates supported by terrestrial POC was higher in the treatment sections of both lakes. In contrast, total secondary and tertiary production (supported by both autochthonous and allochthonous C) was higher in the reference than in the treatment sections of both lakes. Average aquatic consumer allochthony per lake section was 27–40%, although terrestrial POC contributed less than about 10% to total organic C supply to the lakes. The production of aquatic consumers incorporated less than 5% of the total organic C supply in both lakes, indicating a low ecological efficiency. We suggest that the consumption of terrestrial POC by aquatic consumers facilitates a strong coupling with the terrestrial environment. However, the high autochthonous production and the large pool of autochthonous detritus in these nutrient-rich lakes make terrestrial POC quantitatively unimportant for the C flows within food webs.

Secondary and tertiary wastewater treatment is common in developed countries, but little is known about the responses of lotic ecosystems to contemporary wastewater treatment plant (WWTP) discharge. We examined the effects of WWTP discharge on various ecosystem components and functions of 2 morphologically and chemically impacted lowland streams near Berlin, Germany. We sampled one reach upstream and one reach downstream of a WWTP in each stream during each of 5 sampling campaigns. Discharge of treated wastewater resulted in increased concentrations of total organic C, total N, and total P in the sediments and in elevated macrophyte and benthic invertebrate biomasses. However, adverse effects of the WWTPs on the benthic invertebrate communities were small compared to effects reported in previous studies. This difference was a result of the higher purification efficiency of modern WWTPs, but also of significant structural degradation and eutrophication of the streams that already had impoverished the invertebrate community upstream of the WWTPs. Whole-stream community respiration (CR24) and gross primary production (GPP) were both enhanced by WWTP discharge. WWTP discharge generally caused diminished NH4- and PO4-uptake efficiencies, but did not necessarily lead to diminished NO3- uptake efficiencies of streams. Increases in areal NO3-uptake rates caused by the discharge of a large municipal WWTP were high enough to result in increased load-specific NO3-uptake efficiencies. Our study shows that the effects of present-day WWTPs on stream ecosystem functioning clearly differ from the former impacts of poorly treated wastewater. Present-day WWTP discharges mainly cause eutrophication and subsequent side effects and low nutrient-retention efficiencies relative to the high nutrient concentrations and loads of impacted streams. Our results highlight the need for efficient tertiary treatment of wastewater and for the refinement of agricultural practices to reduce diffuse nutrient loadings. We found evidence that even efficiently treated wastewater can have extensive effects on stream ecosystem structure and function. Therefore, adequate dilution rates always should be considered when routing treated wastewater through lotic networks. Our findings on the response of key ecosystem variables to present-day WWTP loading underline the importance of scientifically based stream management.

The aim of this study was to assess land use effects on the density, biomass, and instantaneous secondary production (IP) of benthic invertebrates in a fifth-order tropical river. Invertebrates were sampled at 11 stations along the Rio das Mortes (upper Rio Grande, Southeast Brazil) in the dry and the rainy season 2010/2011. Invertebrates were counted, determined, and measured to estimate their density, biomass, and IP. Water chemical characteristics, sediment heterogeneity, and habitat structural integrity were assessed in parallel. Total invertebrate density, biomass, and IP were higher in the dry season than those in the rainy season, but did not differ significantly among sampling stations along the river. However, taxon-specific density, biomass, and IP differed similarly among sampling stations along the river and between seasons, suggesting that these metrics had the same bioindication potential. Variability in density, biomass, and IP was mainly explained by seasonality and the percentage of sandy sediment in the riverbed, and not directly by urban or agricultural land use. Our results suggest that the consistently high degradation status of the river, observed from its headwaters to mouth, weakened the response of the invertebrate community to specific land use impacts, so that only local habitat characteristics and seasonality exerted effects.

Reduction of flow constitutes one of the most severe human alterations to rivers, as it affects the key abiotic feature of these ecosystems. While there has been considerable progress in understanding the effects of reduced flow on benthic macroinvertebrates, cascading effects of flow reduction on dissolved oxygen concentrations (DO) have not yet received much attention. We compared the macroinvertebrate composition between reference conditions and a situation after several years of discharge reduction in the Spree River (Brandenburg, Germany). Community composition shifted from rheophilic species to species indifferent to flow conditions. Filter feeders were partially replaced by collector/gatherers, which likely reduces the retention of organic matter, and thus the self-purification capacity of the river section. These shifts were associated with low discharge during summer, cascading into daily DO concentration minima of less than 5 mg l−1 which prevailed 74% of the days in summer. This depletion of DO after flow reduction presumably caused the observed species turnover. Hence, flow reduction in lowland rivers may not only directly impair the ecological functions provided by benthic macroinvertebrates but may also act indirectly by depleting DO concentrations.

Lake ecosystems are strongly linked to their terrestrial surroundings by material and energy fluxes across ecosystem boundaries. However, the contribution of terrestrial particulate organic carbon (tPOC) from annual leaf fall to lake food webs has not yet been adequately traced and quantified. In this study, we conducted whole-lake experiments to trace artificially added tPOC through the food webs of two shallow lakes of similar eutrophic status, but featuring alternative stable regimes (macrophyte rich vs. phytoplankton dominated). Lakes were divided with a curtain, and maize (Zea mays) leaves were added, as an isotopically distinct tPOC source, into one half of each lake. To estimate the balance between autochthonous carbon fixation and allochthonous carbon input, primary production and tPOC and tDOC (terrestrial dissolved organic carbon) influx were calculated for the treatment sides. We measured the stable isotope ratios of carbon (d13C) of about 800 samples from all trophic consumer levels and compared them between lake sides, lakes, and three seasons. Leaf litter bag experiments showed that added maize leaves were processed at rates similar to those observed for leaves from shoreline plants, supporting the suitability of maize leaves as a tracer. The lake-wide carbon influx estimates confirmed that autochthonous carbon fixation by primary production was the dominant carbon source for consumers in the lakes. Nevertheless, carbon isotope values of benthic macroinvertebrates were significantly higher with maize additions compared to the reference side of each lake. Carbon isotope values of omnivorous and piscivorous fish were significantly affected by maize additions only in the macrophytedominated lake and d13C of zooplankton and planktivorous fish remained unaffected in both lakes. In summary, our results experimentally demonstrate that tPOC in form of autumnal litterfall is rapidly processed during the subsequent months in the food web of shallow lakes and is channeled to secondary and tertiary consumers predominantly via the benthic pathways. A more intense processing of tPOC seems to be connected to a higher structural complexity in littoral zones, and hence may differ between shallow lakes of alternative stable states. peerReviewed