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[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

The sum of benthic autotrophic and bacterial production often exceeds the sum of pelagic autotrophic and bacterial production, and hence may contribute substantially to whole-lake carbon fluxes, especially in shallow lakes. Furthermore, both benthic and pelagic autotrophic and bacterial production are highly edible and of sufficient nutritional quality for animal consumers. We thus hypothesised that pelagic and benthic transfer efficiencies (ratios of production at adjacent trophic levels) in shallow lakes should be similar. We performed whole ecosystem studies in two shallow lakes (3.5ha, mean depth 2m), one with and one without submerged macrophytes, and quantified pelagic and benthic biomass, production and transfer efficiencies for bacteria, phytoplankton, epipelon, epiphyton, macrophytes, zooplankton, macrozoobenthos and fish. We expected higher transfer efficiencies in the lake with macrophytes, because these provide shelter and food for macrozoobenthos and may thus enable a more efficient conversion of basal production to consumer production. In both lakes, the majority of the whole-lake autotrophic and bacterial production was provided by benthic organisms, but whole-lake primary consumer production mostly relied on pelagic autotrophic and bacterial production. Consequently, transfer efficiency of benthic autotrophic and bacterial production to macrozoobenthos production was an order of magnitude lower than the transfer efficiency of pelagic autotrophic and bacterial production to rotifer and crustacean production. Between-lake differences in transfer efficiencies were minor. We discuss several aspects potentially causing the unexpectedly low benthic transfer efficiencies, such as the food quality of producers, pelagic-benthic links, oxygen concentrations in the deeper lake areas and additional unaccounted consumer production by pelagic and benthic protozoa and meiobenthos at intermediate or top trophic levels. None of these processes convincingly explain the large differences between benthic and pelagic transfer efficiencies. Our data indicate that shallow eutrophic lakes, even with a major share of autotrophic and bacterial production in the benthic zone, can function as pelagic systems with respect to primary consumer production. We suggest that the benthic autotrophic production was mostly transferred to benthic bacterial production, which remained in the sediments, potentially cycling internally in a similar way to what has previously been described for the microbial loop in pelagic habitats. Understanding the energetics of whole-lake food webs, including the fate of the substantial benthic bacterial production, which is either mineralised at the sediment surface or permanently buried, has important implications for regional and global carbon cycling.

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.

Summary 1. Ship-induced waves disturb benthic invertebrate assemblages colonizing littoral zones of lakes and rivers. However, the impact of ship-induced waves on invertebrates has rarely been quantified, and the influencing factors have not been addressed. 2. In an experimental wave tank, five benthic invertebrate species, Bithynia tentaculata, Calopteryx splendens, Dikerogammarus villosus, Gammarus roeseli and Laccophilus hyalinus, were exposed to waves of increasing shear stress (0.43–2.19 N m−2). Mean number of detached individuals was recorded for five littoral habitats [coarse woody debris (CWD), reeds, sand, stones and tree roots], representing different levels of structural complexity as quantified by their fractal dimensions (FD). 3. Results showed that detachment of invertebrates was significantly related to shear stress in all habitats except tree roots. Detachments averaged for the five species were significantly lower in habitats with a high degree of structural complexity, decreasing in the habitat sequence: sand, CWD, stones, reeds and tree roots. 4. Consistent with their different morphologies and methods of attachment to substrates, the five species displayed differences in their response to hydraulic stress that were dependent on habitat. 5. The increasing sheltering effect of structural habitat complexity was mirrored by increasing dissipation of the kinetic energy of waves; i.e. the FD of the habitat was positively correlated with shear stress reduction due to the flow resistance of the habitat. 6. Network habitats such as tree roots provided the best sheltering conditions against hydraulic disturbance, because they combined good refuge availability for all studied invertebrate species and maximal dissipation of kinetic wave energy. Consequently, persistent anthropogenic impacts, such as lakeshore modification or long-term exposure to ship-induced waves, which cause disappearance of complex littoral habitats such as tree roots or dense reed belts, will drastically increase the adverse effects of boating and ship traffic on littoral invertebrate assemblages.

Summary 1 The shores of many lakes have been substantially altered by human developments such as erosion control structures or recreational beaches. Such alterations are likely to increase in the future, yet almost nothing is known about their impacts on the littoral macroinvertebrate community. 2 Macroinvertebrates were studied in seven German lowland lakes exhibiting natural shorelines (reference), retaining walls, ripraps and recreational beaches to examine impacts on the eulittoral (0–0·2 m water depth) and infralittoral (0·2–1·2 m water depth) communities associated with the three types of shoreline development. 3 Among sites, eulittoral species richness and abundance of Coleoptera, Gastropoda, Trichoptera, shredders and xylophagous species were lowest on beaches and retaining walls but ripraps did not differ significantly from natural shorelines. Retaining walls and ripraps had no significant impact on the infralittoral macroinvertebrate community. Conversely, beaches had significantly lower infralittoral species richness and abundance of Ephemeroptera, Trichoptera and shredders than natural shorelines. Furthermore, species richness was correlated positively with habitat heterogeneity expressed as number of habitat types. 4 Among lakes, whole-lake littoral macroinvertebrate density increased with increasing proportion of developed shorelines due to increasing abundances of Chironomidae. The remaining macroinvertebrate major groups decreased with increasing proportion of shoreline development. 5 Synthesis and applications. The biological impacts of shoreline development in lowland lakes depend upon the extent to which structural complexity and heterogeneity of littoral habitats are reduced. Hence, we recommend that management programmes focus upon the conservation of littoral habitat complexity and habitat heterogeneity. The biological effects of shoreline development may be assessed efficiently by combining an assessment of the morphological status of lakeshores and information on macroinvertebrate indicator species with a defined response to the loss of their preferred habitats.

Summary 1. Nutrient inputs from urban and agricultural land use often result in shifts in species composition of pelagic and profundal invertebrate communities. Here, we test if nutrient enrichment affects the composition of eulittoral macroinvertebrate communities, and, if so, if macroinvertebrate communities of five different habitat types reflect differences in trophic state. 2. Macroinvertebrate community composition of 36 lakes was significantly correlated with total phosphorus (TP) concentration, the proportion of coarse woody debris (CWD) and root habitats and the proportion of grassland. 3. However, macroinvertebrate communities of five major habitat types from eight lakes were more dissimilar among habitats than among trophic states. Community composition of reed and stone habitats was significantly correlated with wind exposure but not TP concentration, while macroinvertebrate composition of sand habitats was related to TP concentration and coarse sediments. In CWD and root habitats, both TP concentration and a predominance of invasive species covaried, which made it difficult to relate the observed compositional differences to either trophic state or to the effects of competition between native and invasive species. 4. Trophic state influenced the composition of eulittoral macroinvertebrate communities but to a lesser extent than has been previously reported for profundal habitats. Moreover, the effects of trophic state were nested within habitat type and were partially superseded by biotic interactions and small-scaled habitat complexity. Although eulittoral macroinvertebrate communities were not strong indicators of the trophic state of lowland lakes, they may be used to assess other anthropogenic impacts on lakeshores.