MARS papers & publications


Adams, J. L., Tipping, E., Feuchtmayr, H., Carter, H., Keenan, P. (2018). The contribution of algae to freshwater dissolved organic matter: Implications for UV spectroscopic analysis. Inland Waters 8, 01-12,

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The contribution of algae to freshwater dissolved organic matter: Implications for UV spectroscopic analysis

Dissolved organic matter (DOM) is an important constituent of freshwater that participates in a number of key ecological and biogeochemical processes but can be problematic during water treatment. Thus, the demand for rapid and reliable monitoring is growing, and spectroscopic methods are potentially useful. A model with 3 components—2 that absorb in the ultraviolet (UV) range and are present at variable concentrations and a third that does not absorb light and is present at a low constant concentration—was previously found to yield reliable predictions of dissolved organic carbon concentration [DOC]. The model underestimated [DOC] in shallow eutrophic lakes in the Yangtze Basin, China, however, raising the possibility that DOM derived from algae might be poorly estimated, an idea supported by new data reported here for eutrophic British lakes. We estimated the extinction coefficients in the UV range of algae-derived DOM from published data on algal cultures and from new data from outdoor mesocosm experiments in which high concentrations of DOC were generated under conditions comparable to those in eutrophic freshwaters. The results demonstrate the weak UV absorbance of DOM from algae compared to DOM from terrestrial sources. A modified model, in which the third component represents algae-derived DOM present at variable concentrations, allowed contributions of such DOM to be estimated by combining the spectroscopic data with [DOC] measured by laboratory combustion. Estimated concentrations of algae-derived DOC in 77 surface freshwater samples ranged from 0 to 8.6 mg L−1, and the fraction of algae-derived DOM ranged from 0% to 100%.

Amaral, S. D., Branco, P., Katopodis, C., Ferreira, M. T., Pinheiro, A. N., Santos, J. M. (2018). To swim or to jump? Passage behaviour of a potamodromous cyprinid over an experimental broad-crested weir. River Research and Applications 34, 174-182,

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To swim or to jump? Passage behaviour of a potamodromous cyprinid over an experimental broad-crested weir

Physical stressors, such as man-made obstacles, are considered one of the main causes that negatively affect freshwater fish. Even small weirs may impact fish populations, including potamodromous cyprinids, by partially or totally blocking upstream migratory movements. Some studies have addressed the effect of key hydraulic parameters on upstream movements past small weirs, but little is known on how these parameters interact to induce swimming or jumping behaviour in negotiating such obstacles. This study aims to evaluate the passage behaviour (swimming vs. jumping) of Iberian barbel (Luciobarbus bocagei), a potamodromous species, over an experimental small broad-crested weir, considering the effect of different plunge pool depths (D), waterfall heights (H), and weir discharges (Q). Results revealed that passage behaviour was highly dependent on combinations of plunge pool depths and waterfall heights. Barbel navigated most configurations by swimming (overall outcomes: 81.5% swimming vs. 18.5% jumping), except the ones with the higher waterfall heights (25 cm) tested. Therefore, higher waterfalls proved to be preponderant in the switching of passage behaviour from swimming to jumping. Regarding the discharge over the 1-m-wide weir (overall outcomes: 85% swimming vs. 15% jumping), contrary to what was expected, there was no evidence that passage behaviour was discharge-related, for the range of discharges tested (25–100 L/s). These results are useful to identify potential migration obstacles and should be taken into consideration in river restoration projects and when designing fishways for potamodromous cyprinids.

Baattrup-Pedersen, A., Garssen, A., Göthe, E., Hoffmann, C. C., Oddershede, A., Riis, T., van Bodegom, P. M., Larsen, S. E., Soons, M. (2018). Structural and functional responses of plant communities to climate change‐mediated alterations in the hydrology of riparian areas in temperate Europe. Ecology and Evolution, online,

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Structural and functional responses of plant communities to climate change‐mediated alterations in the hydrology of riparian areas in temperate Europe

The hydrology of riparian areas changes rapidly these years because of climate change‐mediated alterations in precipitation patterns. In this study, we used a large‐scale in situ experimental approach to explore effects of drought and flooding on plant taxonomic diversity and functional trait composition in riparian areas in temperate Europe. We found significant effects of flooding and drought in all study areas, the effects being most pronounced under flooded conditions. In near‐stream areas, taxonomic diversity initially declined in response to both drought and flooding (although not significantly so in all years) and remained stable under drought conditions, whereas the decline continued under flooded conditions. For most traits, we found clear indications that the functional diversity also declined under flooded conditions, particularly in near‐stream areas, indicating that fewer strategies succeeded under flooded conditions. Consistent changes in community mean trait values were also identified, but fewer than expected. This can have several, not mutually exclusive, explanations. First, different adaptive strategies may coexist in a community. Second, intraspecific variability was not considered for any of the traits. For example, many species can elongate shoots and petioles that enable them to survive shallow, prolonged flooding but such abilities will not be captured when applying mean trait values. Third, we only followed the communities for 3 years. Flooding excludes species intolerant of the altered hydrology, whereas the establishment of new species relies on time‐dependent processes, for instance the dispersal and establishment of species within the areas. We expect that altered precipitation patterns will have profound consequences for riparian vegetation in temperate Europe. Riparian areas will experience loss of taxonomic and functional diversity and, over time, possibly also alterations in community trait responses that may have cascading effects on ecosystem functioning.

Baattrup-Pedersen, A., Larsen, S. E., Andersen, D. K., Jepsen, N., Nielsen, J., Rasmussen, J. J. (2018). Headwater streams in the EU Water Framework Directive: Evidence-based decision support to select streams for river basin management plans. Science of the Total Environment 613-614, 1048-1054,

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Headwater streams in the EU Water Framework Directive: Evidence-based decision support to select streams for river basin management plans

Headwater streams are important contributors to aquatic biodiversity and may counteract negative impacts of anthropogenic stress on downstream reaches. In Denmark, the first river basin management plan (RBMP) included streams of all size categories, most being less than 2.5m wide (headwater streams). Currently, however, it is intensely debated whether the small size and low slopes, typical of Danish streams, in combination with degraded habitat conditions obstruct their ability to fulfill the ecological quality objectives required by the EU Water Framework Directive (WFD). The purpose of this study was to provide an analytically based framework for guiding the selection of headwater streams for RBMP. Specifically, the following hypotheses were addressed: i) stream slope, width, planform, and general physical habitat quality can act as criteria for selecting streams for the next generation of RBMPs, and ii) probability-based thresholds for reaching good ecological status can be established for some or all of these criteria, thus creating a sound, scientifically based, and clear selection process. The hypotheses were tested using monitoring data on Danish streams from the period 2004-2015. Significant linear relationships were obtained between the ecological quality ratio assessed by applying the Danish Stream Fauna Index (DSFIEQR) and stream slope, width, sinuosity, and DHI. The obtained models were used to produce pressure-response curves describing the probability of achieving good ecological status along gradients in these parameters. Next, threshold values for slope, width, sinuosity, and DHI were identified for selected probabilities of achieving minimum good ecological status. The obtained results can support managers and policy makers in prioritizing headwater streams for the 3rd RBMP. The approach applied is broadly applicable and can, for instance, help prioritization of restoration and conservation efforts in different types of ecosystems where the biota can be significantly linked to separate and quantifiable environmental characteristics.

Baattrup-Pedersen, A., Ovesen, N. B., Larsen, S. E., Andersen, D. K., Riis, T., Kronvang, B., Rasmussen, J. J. (2018). Evaluating effects of weed cutting on water level and ecological status in Danish lowland streams. Freshwater Biology, online,

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Evaluating effects of weed cutting on water level and ecological status in Danish lowland streams

At present, scientific evidence documenting effects of weed cutting in streams as a measure to improve flood protection and run‐off from agricultural land is scarce, which is surprising considering the huge effect that it has on stream ecology. Instead, weed cutting is performed under the assumption that removal of aquatic plant biomass improves runoff from agricultural land and prevents flooding of adjacent areas provided that it is performed regularly. • In this study, we examined linkages between weed cutting practice and water level reductions in 126 small‐ and medium‐sized Danish streams (catchment size 6 annual cuttings. Furthermore, we found that the ecological status was either moderate or poor/bad in streams with more than one annual cutting. • Our findings highlight that it is by no means certain that the current weed cutting practice is efficient for flood control since (1) regrowth is stimulated by frequent cuttings and a positive feedback loop may develop, necessitating even more frequent cuttings to maintain the discharge capacity of the streams, and (2) many species stimulated by weed cutting, like for instance Sparganium emersum, form dense canopy beds across the entire stream profile and therefore reduce the discharge capacity of the stream more than species growing in confined patches. We encourage more studies with the aim to identify how stream maintenance should be performed to optimise flood control without compromising the ability to reach good ecological stream quality.

Bechter, T., Baumann, K., Birk, S., Bolik, F., Graf, W., Pletterbauer, F. (2018). LaRiMo - A simple and efficient GIS-based approach for large-scale morphological assessment of large European rivers. Science of the Total Environment 628-629, 1191-1199,

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LaRiMo - A simple and efficient GIS-based approach for large-scale morphological assessment of large European rivers

Large rivers cover and function over large spatial extents. Accordingly, the detailed assessment of their morphology is complex. Here, a methodology is presented to assess large rivers’ morphology (LaRiMo) based on free datasets of geographic information systems. This approach could help to achieve a comparable, transboundary assessment of large river morphology to support the implementation of the European Water Framework Directive. The analyses are based on seven parameters describing processes and characteristics related to large river morphology. These parameters are evaluated for and compared between the rivers Danube, Elbe and Loire. A significantly higher amount of gravel and sand bars within the river bed strongly supported the overall higher morphological status of the Loire. A PCA highlighted that the parameters Wetlands, Active Riparian Zones and Free Flowing Sections similarly described a good morphological situation. In contrast, Canal, Dams and Impervious Surface indicated bad morphological conditions. Finally, the approach was successfully validated with data from a detailed, field-based morphological assessment for the Danube. LaRiMo represents an efficient and cost-effective approach to assess large river morphology across large extents. This method provides comparable results across countries and regions.

Bucak, T., Trolle, D., Tavşanoğlu, Ü. N., Çakıroğlu, A. İ., Özen, A., Jeppesen, E., Beklioğlu, M. (2018). Modeling the effects of climatic and land use changes on phytoplankton and water quality of the largest Turkish freshwater lake: Lake Beyşehir. Science of the Total Environment 621, 802-816,

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Modeling the effects of climatic and land use changes on phytoplankton and water quality of the largest Turkish freshwater lake: Lake Beyşehir

Climate change and intense land use practices are the main threats to ecosystem structure and services of Mediterranean lakes. Therefore, it is essential to predict the future changes and develop mitigation measures to combat such pressures. In this study, Lake Beyşehir, the largest freshwater lake in the Mediterranean basin, was selected to study the impacts of climate change and various land use scenarios on the ecosystem dynamics of Mediterranean freshwater ecosystems and the services that they provide. For this purpose, we linked catchment model outputs to the two different processed-based lake models: PCLake and GLM-AED, and tested the scenarios of five General Circulation Models, two Representation Concentration Pathways and three different land use scenarios, which enable us to consider the various sources of uncertainty. Climate change and land use scenarios generally predicted strong future decreases in hydraulic and nutrient loads from the catchment to the lake. These changes in loads translated into alterations in water level as well as minor changes in chlorophyll a (Chl-a) concentrations. We also observed an increased abundance of cyanobacteria in both lake models. Total phosphorus, temperature and hydraulic loading were found to be the most important variables determining cyanobacteria biomass. As the future scenarios revealed only minor changes in Chl-a due to the significant decrease in nutrient loads, our results highlight that reduced nutrient loading in a warming world may play a crucial role in offsetting the effects of temperature on phytoplankton growth. However, our results also showed increased abundance of cyanobacteria in the future may threaten ecosystem integrity and may limit drinking water ecosystem services. In addition, extended periods of decreased hydraulic loads from the catchment and increased evaporation may lead to water level reductions and may diminish the ecosystem services of the lake as a water supply for irrigation and drinking water.

Bussi, G,. Whitehead, P. G., Gutiérrez-Cánovas, C., Ledesma, J. L., Ormerod, S. J., Couture, R. M. (2018). Modelling the Effects of Climate and Land-use Change on the Hydrochemistry and Ecology of the River Wye (Wales). Science of the Total Environment 627, 733-743,

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Modelling the effects of climate and land-use change on the hydrochemistry and ecology of the River Wye (Wales)

Interactions between climate change and land use change might have substantial effects on aquatic ecosystems, but are still poorly understood. Using the Welsh River Wye as a case study, we linked models of water quality (Integrated Catchment - INCA) and climate (GFDL - Geophysical Fluid Dynamics Laboratory and IPSL - Institut Pierre Simon Laplace) under greenhouse gas scenarios (RCP4.5 and RCP8.5) to drive a bespoke ecosystem model that simulated the responses of aquatic organisms. The potential effects of economic and social development were also investigated using scenarios from the EU MARS project (Managing Aquatic Ecosystems and Water Resources under Multiple Stress). Longitudinal position along the river mediated response to increasing anthropogenic pressures. Upland locations appeared particularly sensitive to nutrient enrichment or potential re-acidification compared to lowland environments which are already eutrophic. These results can guide attempts to mitigate future impacts and reiterate the need for sensitive land management in upland, temperate environments which are likely to become increasingly important to water supply and biodiversity conservation as the effects of climate change intensify.

Couture, R.-M., Moe, S. J., Lin, Y., Kaste, Ø., Haande, S., Solheim, A. L. (2018). Simulating water quality and ecological status of Lake Vansjø, Norway, under land-use and climate change by linking process-oriented models with a Bayesian network. Science of the Total Environment 621, 713-724,

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Simulating water quality and ecological status of Lake Vansjø, Norway, under land-use and climate change by linking process-oriented models with a Bayesian network

Excess nutrient inputs and climate change are two of multiple stressors affecting many lakes worldwide. Lake Vansjø in southern Norway is one such eutrophic lake impacted by blooms of toxic blue-green algae (cyanobacteria), and classified as moderate ecological status under the EU Water Framework Directive. Future climate change may exacerbate the situation. Here we use a set of chained models (global climate model, hydrological model, catchment phosphorus (P) model, lake model, Bayesian Network) to assess the possible future ecological status of the lake, given the set of climate scenarios and storylines common to the EU project MARS (Managing Aquatic Ecosystems and Water Resources under Multiple Stress). The model simulations indicate that climate change alone will increase precipitation and runoff, and give higher P fluxes to the lake, but cause little increase in phytoplankton biomass or changes in ecological status. For the storylines of future management and land-use, however, the model results indicate that both the phytoplankton biomass and the lake ecological status can be positively or negatively affected. Our results also show the value in predicting a biological indicator of lake ecological status, in this case, cyanobacteria biomass with a BN model. For all scenarios, cyanobacteria contribute to worsening the status assessed by phytoplankton, compared to using chlorophyll-a alone.

Cremona, F., Tuvikene, L., Haberman, J., Nõges, P., Nõges, T. (2018). Factors controlling the three-decade long rise in cyanobacteria biomass in a eutrophic shallow lake. Science of the Total Environment 621, 352–359,

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Factors controlling the three-decade long rise in cyanobacteria biomass in a eutrophic shallow lake

We aimed at quantifying the importance of limnological variables in the decadal rise of cyanobacteria biomass in shallow hemiboreal lakes. We constructed estimates of cyanobacteria (blue-green algae) biomass in a large, eutrophic lake (Estonia, Northeastern Europe) from a database comprising 28 limnological variables and spanning more than 50 years of monitoring. Using a dual-model approach consisting in a boosted regression trees (BRT) followed by a generalized least squares (GLS) model, our results revealed that six variables were most influential for assessing the variance of cyanobacteria biomass. Cyanobacteria response to nitrate concentration and rotifer abundance was negative, whereas it was positive to pH, temperature, cladoceran and copepod biomass. Response to total phosphorus (TP) and total phosphorus to total nitrogen ratio was very weak, which suggests that actual in-lake TP concentration is still above limiting values. The most efficient GLS model, which explained nearly two thirds (r2 = 0.65) of the variance of cyanobacteria biomass included nitrate concentration, water temperature and pH. The very high number of observations (maximum n = 525) supports the robustness of the models. Our results suggest that the decadal rise of blue-green algae in shallow lakes lies in the interaction between cultural eutrophication and global warming which bring in-lake physical and chemical conditions closer to cyanobacteria optima.

Davidson, T. A., Audet, J., Jeppesen, E., Landkildehus, F., Lauridsen, T. L., Søndergaard, M., Syvaranta, J. (2018). Synergy between nutrients and warming enhances methane ebullition from shallow lakes. Nature Climate Change 8, 156-160,

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Synergy between nutrients and warming enhances methane ebullition from experimental lakes

Link to read only version of the paper "". Lakes and ponds are important natural sources of the potent greenhouse gas methane (CH4), with small shallow waters identified as particular hotspots1,2. Ebullition (bubbles) of CH4 makes up a large proportion of total CH4 flux3,4. However, difficulty measuring such episodic events5 makes prediction of how ebullition responds to nutrient enrichment and rising temperatures challenging. Here, the world’s longest running, mesocosm-based, shallow lake climate change experiment was used to investigate how the combination of warming and eutrophication (that is, nutrient enrichment) affects CH4 ebullition. Eutrophication without heating increased the relative contribution of ebullition from 51% to 75%. More strikingly the combination of nutrient enrichment and experimental warming treatments of +2–3 °C and +4–5 °C had a synergistic effect, increasing mean annual ebullition by at least 1900 mg CH4-C m−2 yr−1. In contrast, diffusive flux showed no response to eutrophication and only a small increase at higher temperatures (average 63 mg CH4–C m−2 yr−1). As shallow lakes are the most common lake type globally, abundant in highly climate sensitive regions6 and most vulnerable to eutrophication, these results suggest their current and future contributions to atmospheric CH4 concentrations may be significantly underestimated.

Davidson, T. A., Wetterich, S., Windirsch, T., Johansen, K. L., González-Bergonzoni, I., Grønnow, B., Jeppesen, E., Syvyränta, J., Olsen, J., Strunk, A., Larsen, N. K., Meyer, H., Søndergard, J., Dietz, R., Eulears, I., Mosbech, A. (2018). The history of seabird colonies and the North Water ecosystem: Contributions from palaeoecological and archaeological evidence. Ambio 47, 175-192,

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The history of seabird colonies and the North Water ecosystem: Contributions from palaeoecological and archaeological evidence

The North Water (NOW) polynya is one of the most productive marine areas of the Arctic and an important breeding area for millions of seabirds. There is, however, little information on the dynamics of the polynya or the bird populations over the long term. Here, we used sediment archives from a lake and peat deposits along the Greenland coast of the NOW polynya to track long-term patterns in the dynamics of the seabird populations. Radiocarbon dates show that the thick-billed murre (Uria lomvia) and the common eider (Somateria mollissima) have been present for at least 5500 cal. years. The first recorded arrival of the little auk (Alle alle) was around 4400 cal. years BP at Annikitsoq, with arrival at Qeqertaq (Salve Ø) colony dated to 3600 cal. years BP. Concentrations of cadmium and phosphorus (both abundant in little auk guano) in the lake and peat cores suggest that there was a period of large variation in bird numbers between 2500 and 1500 cal. years BP. The little auk arrival times show a strong accord with past periods of colder climate and with some aspects of human settlement in the area.

Feld, C. K., Fernandes, M. R. P., Ferreira, M. T., Hering, D., Ormerod, S. J., Venohr, M., Gutiérrez-Cánovas, C. (2018). Evaluating riparian solutions to multiple stressor problems in river ecosystems — A conceptual study. Water Research, in press,

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Evaluating riparian solutions to multiple stressor problems in river ecosystems — A conceptual study

Rivers are among the most sensitive of all ecosystems to the effects of global change, but options to prevent, mitigate or restore ecosystem damage are still inadequately understood. Riparian buffers are widely advocated as a cost-effective option to manage impacts, but empirical evidence is yet to identify ideal riparian features (e.g. width, length and density) which enhance ecological integrity and protect ecosystem services in the face of catchment-scale stressors. Here, we use an extensive literature review to synthesise evidence on riparian buffer and catchment management effects on instream environmental conditions (e.g. nutrients, fine sediments, organic matter), river organisms and ecosystem functions. We offer a conceptual model of the mechanisms through which catchment or riparian management might impact streams either positively or negatively. The model distinguishes scale-independent benefits (shade, thermal damping, organic matter and large wood inputs) that arise from riparian buffer management at any scale from scale-dependent benefits (nutrient or fine sediment retention) that reflect stressor conditions at broader (sub-catchment to catchment) scales. The latter require concerted management efforts over equally large domains of scale (e.g. riparian buffers combined with nutrient restrictions). The evidence of the relationships between riparian configuration (width, length, zonation, density) and scale-independent benefits is consistent, suggesting a high certainty of the effects. In contrast, scale-dependent effects as well as the biological responses to riparian management are more uncertain, suggesting that ongoing diffuse pollution (nutrients, sediments), but also sources of variability (e.g. hydrology, climate) at broader scales may interfere with the effects of local riparian management. Without concerted management across relevant scales, full biological recovery of damaged lotic ecosystems is unlikely. There is, nevertheless, sufficient evidence that the benefits of riparian buffers outweigh potential adverse effects, in particular if located in the upstream part of the stream network. This supports the use of riparian restoration as a no-regrets management option to improve and sustain lotic ecosystem functioning and biodiversity.

Gerner, N. V., Nafo, I., Winking, C., Wencki, K., Strehl, C., Wortberg, T., Niemann, A., Anzaldua, G., Lago, M., Birk, S. (2018). Large-scale river restoration pays off: A case study of ecosystem service valuation for the Emscher restoration generation project. Ecosystem Services 30, 327-338,

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Large-scale river restoration pays off: A case study of ecosystem service valuation for the Emscher restoration generation project

Though the Ecosystem Service (ESS) approach is considered promising for integrated ecosystem management, its operationalisation is hampered by the lack of agreed evaluation instruments. To demonstrate the suitability of a structured ESS evaluation, we conducted a case study estimating the impact of the restoration of the Emscher River and its tributaries on the provision, use and benefit of ESS. The Emscher restoration is a large-scale project with immense temporal and financial efforts. To assess the values generated by this restoration, we applied an ESS evaluation framework and quantified the regulation and maintenance ESS ‘self-purification capacity’, ‘maintaining nursery populations and habitats’ and ‘flood protection’ as well as cultural ESS such as aesthetic, recreational, educational and existence values. Final ESS were monetized using economic methods, e.g. ‘damage costs avoided’, ‘contingent valuation’ and ‘benefit transfer’. We estimated a market value/direct economic impact of 21,441,572 € per year as a result of the restoration. Furthermore, a non-market value for people who care about the local environment of 109,121,217 € per year was determined, representing the benefit with ‘non-use value’ from the Emscher restoration. Our case study demonstrated the successful application of the structured evaluation framework in practice. Its implications and limitations are discussed.

Gonzalez-Bergonzoni, I., Kristensen, P. B., Baattrup-Pedersen, A., Alnøe, A. B., Astrup Kristensen, E. A., Riis, T. (2018). Riparian forest cover modifies fuelling sources for stream food webs but not trophic web length. Hydrobiologia 805, 291-310,

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Riparian forest modifies fuelling sources for stream food webs but not food-chain length in lowland streams of Denmark

Several studies have shown that the origin of carbon fuelling food webs in streams depends on riparian cover type. In forested stream sites allochthonous resources fuel food webs, whereas autochthonous resources support biomass in grassland (open-canopy) stream sites. However, some studies suggest that autochthonous carbon (of highest quality) is preferentially assimilated regardless of riparian cover and that the food-chain length (FCL) may be larger in grassland than in forested sites. We used stable isotopes of carbon and nitrogen in adjacent grassland and forested reaches to compare the contribution of autochthonous vs. allochthonous resources to the biomass of the whole macroinvertebrate assemblage and to the most abundant taxa. Moreover, we compared the FCL between forested and grassland sites by estimating the trophic position of brown trout, Salmo trutta. Autochthonous support to macroinvertebrate biomass was higher in grassland than in forested sites, often changing from a dominantly autochthonous to an allochthonous-generated biomass from grassland to forested. This held true for the whole macroinvertebrate assemblage and for specific species. FCL remained similar between reach types. Our study suggests that autochthonous resources are assimilated to a higher extent when their availability increases with canopy openness but allochthonous carbon sustain macroinvertebrate biomass in forested reaches.

Gu, J., He, H., Jin, H., Yu, J., Jeppesen, E., Nairn, R. W., Li, K. (2018). Synergistic negative effects of small-sized benthivorous fish and nitrogen loading on the growth of submerged macrophytes – Relevance for shallow lake restoration. Science of the Total Environment 610-611, 1572-1580,

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Synergistic negative effects of small-sized benthivorous fish and nitrogen loading on the growth of submerged macrophytes – Relevance for shallow lake restoration

Rapid recruitment of small fish after biomanipulation in warm lakes may delay the reestablishment of submerged macrophytes, not least at high nutrient concentrations. Success has recently been obtained in controlling phosphorus (P) loading to many lakes, but nitrogen (N) inputs often remain high. To determine the interactive effects of N loading and the abundance of small-sized fish on the growth of the submerged macrophyte Vallisneria natans, we conducted an outdoor mesocosm experiment with a factorial design on the north shore of Lake Taihu, China. The experiment involved two densities of small crucian carp – low (10 g m− 2) and high (40 g m− 2) – crossed with two levels of N loading – present-day external nutrient loading (P: 5 μg L− 1 day− 1, N: 130 μg L− 1 day− 1) and P: 5 μg L− 1 day− 1 with a three times higher N loading (N: 390 μg L− 1 day− 1). The results showed that nitrogen-fish interactions significantly hindered the growth of V. natans, particularly at the high N loading. At low N loading, high densities of fish decreased the relative growth rate, mean leaf length, leaf mass and root mass of V. natans by 16%, 5%, 8%, and 23%, respectively, compared with these measures at low fish densities. The effect of fish was even stronger when N loading was high, with decreases of 232%, 32%, 57%, and 47% for the respective plant growth measures. The stronger effect at high N loading was attributed to higher turbidity due to enhanced phytoplankton biomass and to increased consumption or damage of plants by the fish in response to the more nutrient-enriched plant tissue. Our results indicate that high abundance of small crucian carp in warm lakes may reduce the resilience of submerged macrophytes to external N loading, thereby lowering the chances of successful restoration by biomanipulation.

Guiterrez, M. F., Tavşanoğlu, Ü. N., Vidal, N., Yu, J., Teixeira-de Mello, F., Çakıroğlu, A. İ., He, H., Liu, Z., Jeppesen, E. (2018). Salinity shapes zooplankton communities and functional diversity and has complex effects on size structure in lakes. Hydrobiologia 813, 237-255,

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Salinity shapes zooplankton communities and functional diversity and has complex effects on size structure in lakes

Changes in zooplankton community structure and function were analyzed in 24 lakes covering a wide salinity gradient (from 0.5 to 115 g l-1) in a semiarid region in northwest China. We hypothesized that species richness (S), species diversity (H), functional diversity (FD), biomass, and size of zooplankton would decrease with increasing salinity. We found that S, H, and FD did decrease with increasing salinity, whereas zooplankton sizes, size range, and biomasses did not. In fact, the sizes of microcrustaceans were mainly regulated by the abundance of small fish. Besides the impoverishment of FD, the zooplankton functional groups also varied along the salinity gradient. A shift occurred from selective raptorial to more generalist microphagous rotifers, from selective to more generalist filter feeder cladocerans, and from dominance of microphagous herbivorous copepods to microphagous carnivores. Our study indicates that the ongoing salinization of lakes with climate warming will result in important changes in the zooplankton, affecting not only the structure but also the functioning of this community. A weakened top-down control by zooplankton on phytoplankton at moderate high salinities may be an indirect consequence, leading to a worsening of eutrophication symptoms. Loss of fish at high salinities may, however, counteract this effect.

Hein, T., Funk, A., Pletterbauer, F., Graf, W., Zsuffa, I., Haidvogl, G., Schinegger, R., Weigelhofer, G. (2018). Management challenges related to long-term ecological impacts, complex stressor interactions, and different assessment approaches in the Danube River Basin. River Research and Applications, 01-10,

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Management challenges related to long-term ecological impacts, complex stressor interactions, and different assessment approaches in the Danube River Basin

For centuries, rivers have experienced massive changes of their hydromorphic structures due to human activities. The Danube River, the second largest river in Europe, is a case in point for long-term societal imprint. Resulting human-induced pressures are a key issue for river management, aiming to improve the ecological conditions and guarantee the provision of ecosystem services. As the most international river basin in the world, the management of the Danube is particularly challenging and needs a well-organized cooperation of 19 nations. The recent river basin management plan has identified pollution and hydromorphological alterations as most pressing problems, but it has also acknowledged newly emerging issues. In this article, we present 3 specific examples of highly relevant issues for the future river basin management of the Danube: (a) long-term impacts in the catchment such as changes in flood patterns and potential ecological consequences; (b) complex feedback loops linking the spread of neozoa with intertwined stressor responses due to river engineering for different purposes; and (c) linkages between different assessment approaches based on European legal frameworks to analyse the specific pressures at different spatial scales. These examples highlight the need for a more integrated approach in future Danube River Basin management schemes. Furthermore, large-scale effects such as climate change and interactions of multiple pressures need to be addressed in future management to increase resilience of the river system and to allow a sustainable ecosystem-based management of rivers.

Hilt, S., Alirangues Nuñez, M. M., Bakker, E. S., Blindow, I., Davidson, T. A., Gillefalk, M., Hansson, L. A., Janse, J. H., Janssen, A. B. G., Jeppesen, E., Kabus, T., Kelly, A., Köhler, J., Lauridsen, T. L., Mooij, W. M., Noordhuis, R., Phillips, G., Rücker, J., Søndergaard, M., Teurlincx, S., van de Weyer, K., van Donk, E., Waterstraat, A., Willby, N., Sayer, C. D. (2018). Response of submerged macrophyte communities to external and internal restoration measures in north temperate shallow lakes. Frontiers in Plant Science 9, 194,

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Response of Submerged Macrophyte Communities to External and Internal Restoration Measures in North Temperate Shallow Lakes

Submerged macrophytes play a key role in north temperate shallow lakes by stabilizing clear-water conditions. Eutrophication has resulted in macrophyte loss and shifts to turbid conditions in many lakes. Considerable efforts have been devoted to shallow lake restoration in many countries, but long-term success depends on a stable recovery of submerged macrophytes. However, recovery patterns vary widely and remain to be fully understood. We hypothesize that reduced external nutrient loading leads to an intermediate recovery state with clear spring and turbid summer conditions similar to the pattern described for eutrophication. In contrast, lake internal restoration measures can result in transient clear-water conditions both in spring and summer and reversals to turbid conditions. Furthermore, we hypothesize that these contrasting restoration measures result in different macrophyte species composition, with added implications for seasonal dynamics due to differences in plant traits. To test these hypotheses, we analyzed data on water quality and submerged macrophytes from 49 north temperate shallow lakes that were in a turbid state and subjected to restoration measures. To study the dynamics of macrophytes during nutrient load reduction, we adapted the ecosystem model PCLake. Our survey and model simulations revealed the existence of an intermediate recovery state upon reduced external nutrient loading, characterized by spring clear-water phases and turbid summers, whereas internal lake Hilt et al. Response of Macrophytes to Restoration restoration measures often resulted in clear-water conditions in spring and summer with returns to turbid conditions after some years. External and internal lake restoration measures resulted in different macrophyte communities. The intermediate recovery state following reduced nutrient loading is characterized by a few macrophyte species (mainly pondweeds) that can resist wave action allowing survival in shallow areas, germinate early in spring, have energy-rich vegetative propagules facilitating rapid initial growth and that can complete their life cycle by early summer. Later in the growing season these plants are, according to our simulations, outcompeted by periphyton, leading to late-summer phytoplankton blooms. Internal lake restoration measures often coincide with a rapid but transient colonization by hornworts, waterweeds or charophytes. Stable clear-water conditions and a diverse macrophyte flora only occurred decades after external nutrient load reduction or when measures were combined.

Hutchins, M. G., Abesser, C., Prudhomme, C., Elliott, J. A., Bloomfield, J. P., Mansour, M. M., Hitt, O. E. (2018). Combined impacts of future land-use and climate stressors on water resources and quality in groundwater and surface waterbodies of the upper Thames river basin, UK. Science of the Total Environment 631, 962-986,

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Combined impacts of future land-use and climate stressors on water resources and quality in groundwater and surface waterbodies of the upper Thames river basin, UK

It is widely acknowledged that waterbodies are becoming increasingly affected by a wide range of drivers of change arising from human activity. To illustrate how this can be quantified a linked modelling approach was applied in the Thames river basin in southern UK. Changes to river flows, water temperature, river and reservoir quality were predicted under three contrasting future “storylines”; one an extension of present day rates of economic development, the others representing more extreme and less sustainable visions. Modelling revealed that lower baseflow conditions will arise under all storylines. For the less extreme storyline river water quality is likely to deteriorate but reservoir quality will improve slightly. The two more extreme futures could not be supported by current management strategies to meet water demand. To satisfy these scenarios, transfer of river water from outside the Thames river basin would be necessary. Consequently, some improvement over present day water quality in the river may be seen, and for most indicators conditions would be better than in the less extreme storyline. However, because phosphorus concentrations will rise, the invoked changes in water demand management would not be of a form suitable to prevent a marked deterioration in reservoir water quality.

Jeppesen, E., Appelt, M., Hastrup, K., Grønnow, B., Mosbech, A., Smol, J. P., Davidson, T. A. (2018). Living in an Oasis: Rapid Transformations, Resilience and Resistance in the North Water Area Societies and Ecosystems. Ambio 47, 296-309,

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Living in an oasis: Rapid transformations, resilience, and resistance in the North Water Area societies and ecosystems

Based on lake sediment data, archaeological findings, and historical records, we describe rapid transformations, resilience and resistance in societies and ecosystems, and their interactions in the past in the North Water area related to changes in climate and historical events. Examples are the formation of the polynya itself and the early arrival of people, ca. 4500 years ago, and later major human immigrations (different societies, cultural encounters, or abandonment) from other regions in the Arctic. While the early immigrations had relatively modest and localised effect on the ecosystem, the later-incoming culture in the early thirteenth century was marked by extensive migrations into and out of the area and abrupt shifts in hunting technologies. This has had long-lasting consequences for the local lake ecosystems. Large natural transformations in the ecosystems have also occurred over relatively short time periods related to changes in the polynya. Finally, we discuss the future perspectives for the North Water area given the many threats, but also opportunities.

Kaandorp, V. P., Molina-Navarro, E., Andersen, H. E., Bloomfield, J. P., Kuijper, M. J. M., de Louw, P. G. B. (2018). A conceptual model for the analysis of multi-stressors in linked groundwater–surface water systems. Science of the Total Environment 627, 880-895,

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A conceptual model for the analysis of multi-stressors in linked groundwater–surface water systems

Groundwater and surface water are often closely coupled and are both under the influence of multiple stressors. Stressed groundwater systems may lead to a poor ecological status of surface waters but to date no conceptual framework to analyse linked multi-stressed groundwater – surface water systems has been developed. In this paper, a framework is proposed showing the effect of groundwater on surface waters in multiple stressed systems. This framework will be illustrated by applying it to four European catchments, the Odense, Denmark, the Regge and Dinkel, Netherlands, and the Thames, UK, and by assessing its utility in analysing the propagation or buffering of multi-stressors through groundwater to surface waters in these catchments. It is shown that groundwater affects surface water flow, nutrients and temperature, and can both propagate stressors towards surface waters and buffer the effect of stressors in space and time. The effect of groundwater on drivers and states depends on catchment characteristics, stressor combinations, scale and management practises. The proposed framework shows how groundwater in lowland catchments acts as a bridge between stressors and their effects within surface waters. It shows water managers how their management areas might be influenced by groundwater, and helps them to include this important, but often overlooked part of the water cycle in their basin management plans. The analysis of the study catchments also revealed a lack of data on the temperature of both groundwater and surface water, while it is an important parameter considering future climate warming.

Larsen, S., Chase, J. M., Durance, I., Ormerod, S. J. (2018). Lifting the veil: Richness measurements fail to detect systematic biodiversity change over three decades. Ecology, in press.

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Lifting the veil: Richness measurements fail to detect systematic biodiversity change over three decades

While there is widespread recognition of human involvement in biodiversity loss globally, at smaller spatial extents the effects are less clear. One reason is that local effects are obscured by the use of summary biodiversity variables, such as species richness, that provide only limited insight into complex biodiversity change. Here, we use 30 years of invertebrate data from a metacommunity of ten streams in Wales (UK) combined with regional surveys, to examine temporal changes in multiple biodiversity measures at local, metacommunity and regional scales. There was no change in taxonomic or functional α and spatial β -diversity metrics at any scale over the 30-year time series, suggesting a relative stasis in the system and no evidence for on-going homogenisation. However, temporal changes in mean species composition were evident. Two independent approaches to estimate species niche breadth showed that compositional changes were associated with a systematic decline in mean community specialisation. Estimates of species-specific local extinction and immigration probabilities suggested that this decline was linked to lower re-colonisation rates of specialists, rather than greater local extinction rates. Our results reveal the need for caution in implying stasis from patterns in α and spatial β -diversity measures that might mask non-random biodiversity changes over time. We also show how different but complementary approaches to estimate niche breadth and functional distinctness of species can reveal long-term trends in community homogenisation likely to be important to conservation and ecosystem function.

Menezes, R. F., Attayde, J. L., Kosten, S., Lacerot, G. l., Coimbra e Sousa, L., Costa, L. S., Sternberg, L. D. S. L., dos Santos, A. C., de Medeiros Rodrigues, M., Jeppesen, E. (2018). Reduced precipitation to evaporation ratio increases eutrophication in tropical shallow lakes. Hydrobiologia, in press.

Molina-Navarro, E., Andersen, H. E., Nielsen, A., Thodsen, H., Trolle, D. (2018). Quantifying the combined effects of land use and climate changes on stream flow and nutrient loads: A modelling approach in the Odense Fjord catchment (Denmark). Science of the Total Environment 621, 253-264,

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Quantifying the combined effects of land use and climate changes on stream flow and nutrient loads: A modelling approach in the Odense Fjord catchment (Denmark)

Water pollution and water scarcity are among the main environmental challenges faced by the European Union, and multiple stressors compromise the integrity of water resources and ecosystems. Particularly in lowland areas of northern Europe, high population density, flood protection and, especially, intensive agriculture, are important drivers of water quality degradation. In addition, future climate and land use changes may interact, with uncertain consequences for water resources. Modelling approaches have become essential to address water issues and to evaluate ecosystem management. In this work, three multi-stressor future storylines combining climatic and socio-economic changes, defined at European level, have been downscaled for the Odense Fjord catchment (Denmark), giving three scenarios: High-Tech agriculture (HT), Agriculture for Nature (AN) and Market-Driven agriculture (MD). The impacts of these scenarios on water discharge and inorganic and organic nutrient loads to the streams have been simulated using the Soil and Water Assessment Tool (SWAT). The results revealed that the scenario-specific climate inputs were most important when simulating hydrology, increasing river discharge in the HT and MD scenarios (which followed the high emission 8.5 representative concentration pathway, RCP), while remaining stable in the AN scenario (RCP 4.5). Moreover, discharge was the main driver of changes in organic nutrients and inorganic phosphorus loads that consequently increased in a high emission scenario. Nevertheless, both land use (via inputs of fertilizer) and climate changes affected the nitrate transport. Different levels of fertilization yielded a decrease in the nitrate load in AN and an increase in MD. In HT, however, nitrate losses remained stable because the fertilization decrease was counteracted by a flow increase. Thus, our results suggest that N loads will ultimately depend on future land use and management in an interaction with climate changes, and this knowledge is of utmost importance for the achievement of European environmental policy goals.

Özen, A., Tavşanoğlu, Ü. N., Çakıroğlu, A. İ., Levi, E. E., Jeppesen, E., Beklioğlu, M. (2018). Patterns of microbial food webs in Mediterranean shallow lakes with contrasting nutrient levels and predation pressures. Hydrobiologia 806, 13-27,

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Patterns of microbial food webs in Mediterranean shallow lakes with contrasting nutrient levels and predation pressures

To elucidate the specific and combined effects of bottom-up and top-down control on the microbial community in warm lakes, we sampled microbial community along with physical–chemical and biological variables and performed in situ food web experiments, in 14 Turkish shallow lakes with contrasting nutrient levels and predation pressures. Our field results revealed that differences in microbial communities correlated with differences in zooplankton community structure, temperature (increasing nutrient concentrations, change in zooplankton composition), nutrient concentrations (increasing bacteria and heterotrophic nanoflagellate abundances with increasing nitrogen concentrations and temperature) and macrophyte coverage (ciliates as potential consumers of bacteria and HNF was strongest in macrophyte-dominated lakes). Our in situ experimental study revealed that the zooplankton not only affect the biomass and composition of microbial communities but also alter the microbial structure and trophic relationships. Our results therefore indicate that both bottom-up factors and top-down effects were important for the efficiency of the carbon transfer from bacteria to higher trophic levels in the study lakes. Due to an anticipated increase in eutrophication, temperature and alteration of the classical food web with climate warming, major changes in the microbial community of lakes are, therefore, expected in a warmer future in semi-arid Mediterranean climatic regions.

Piirsoo, K., Laas, A., Meinson, P., Nõges, P., Pall, P., Viik, M., Vilbaste, S., Nõges, T. (2018). Changes in particulate organic matter passing through a large shallow lowland lake. Proceedings of the Estonian Academy of Sciences 67, 93-105,

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Changes in particulate organic matter passing through a large shallow lowland lake

Different sources of particulate organic matter (POM) as well as its composition affect the biological food web and hence the self-purification potential and water quality of rivers. We studied the effect of a large shallow lake on the POM pool of the water passing through it. Over four years, we analysed monthly the amount and composition of POM and a set of environmental variables in the inflows and in the outflow of Lake Võrtsjärv (Estonia). In the inflows, the live pool of POM consisted of phytoplankton – small crypto-, dino-, and chlorophytes. The concentration of chlorophyll a (Chl a), as a proxy of phytoplankton biomass, was positively correlated with temperature and total phosphorus and negatively with dissolved silica, total nitrogen, and discharge. In the outflow, the share of the live component of POM was much larger than in the inflows but was also dominated by phytoplankton represented by grazing resistant filamentous cyanobacteria. Chl a was positively correlated with total phosphorus, temperature, pH, and precipitation, and negatively with dissolved silica, total nitrogen, and discharge in the outflow. The different amounts, composition, and seasonal dynamics of POM in the inflows and in the outflow have potentially substantial impacts on the food web with a predominating classical pathway in the inflows versus a detrital pathway in the outflow.

Pouso, S., Uyarra, M. C., Borja, A. (2018).The recovery of estuarine quality and the perceived increase of cultural ecosystem services by beach users: A case study from northern Spain. Journal of Environmental Management 212, 450-461,

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The recovery of estuarine quality and the perceived increase of cultural ecosystem services by beach users: A case study from northern Spain

In Europe, the quality of coastal bathing waters improved considerably in the last decades, mainly due to the more demanding legislation and the adoption of water sanitation plans. In the Nerbioi estuary (North Spain), the Wastewater Treatment Plan implemented between 1990 and 2001 resulted on an abrupt decrease in microbial concentration; thus, complying with bathing waters legislation and allowing recreational activities again in the three beaches of the estuary. However, little is known about how improvements in bathing waters influences the provision of cultural ecosystem services and human well-being. A questionnaire was used to study beach users' behaviour and perceptions and compared with environmental time-series data (microbial concentration and water transparency). Most respondents perceived an improvement in bathing waters quality and linked it to the estuarine sanitation. Nerbioi beaches are important recreational areas, mainly for local visitors, and water quality improvement was found to be a critical factor for deciding to visit these beaches. Furthermore, most visitors answered that they would not return if water conditions deteriorate. Significant differences existed between beaches, with the most inner beach presenting worse environmental conditions than the other two beaches; and matching user's perceptions. Our findings highlight that water sanitation actions are important for the recovery of degraded coastal environments and for the maintenance of ecosystem services. Also, that multidisciplinary research is necessary to better comprehend the links between environmental recovery and the provision of ecosystem services.

Romão, F. A., Santos, J. M., Katopodis, C., Pinheiro, A. N., Branco, P. (2018). How Does Season Affect Passage Performance and Fatigue of Potamodromous Cyprinids? An Experimental Approach in a Vertical Slot Fishway. Water 10, 395,

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How Does Season Affect Passage Performance and Fatigue of Potamodromous Cyprinids? An Experimental Approach in a Vertical Slot Fishway

Most fishway studies are conducted during the reproductive period, yet uncertainty remains on whether results may be biased if the same studies were performed outside of the migration season. The present study assessed fish passage performance of a potamodromous cyprinid, the Iberian barbel (Luciobarbus bocagei), in an experimental full-scale vertical slot fishway during spring (reproductive season) and early-autumn (non-reproductive season). Results revealed that no significant differences were detected on passage performance metrics, except for entry efficiency. However, differences between seasons were noted in the plasma lactate concentration (higher in early-autumn), used as a proxy for muscular fatigue after the fishway navigation. This suggests that, for potamodromous cyprinids, the evaluation of passage performance in fishways does not need to be restricted to the reproductive season and can be extended to early-autumn, when movements associated with shifts in home range may occur. The increased effort during the non-reproductive period suggests that adapting the operational regime of fishways, at biologically meaningful seasons in a year, should be assessed by considering the physiological state of the target species.

Schinegger, R., Pucher, M., Aschauer, C., Schmutz, S. (2018). Configuration of multiple human stressors and their impacts on fish assemblages in Alpine river basins of Austria. Science of the Total Environment 616-617, 17-28,

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Configuration of multiple human stressors and their impacts on fish assemblages in Alpine river basins of Austria

This work addresses multiple human stressors and their impacts on fish assemblages of the Drava and Mura rivers in southern Austria. The impacts of single and multiple human stressors on riverine fish assemblages in these basins were disentangled, based on an extensive dataset. Stressor configuration, i.e. various metrics of multiple stressors belonging to stressor groups hydrology, morphology, connectivity and water quality were investigated for the first time at river basin scale in Austria. As biological response variables, the Fish Index Austria (FIA) and its related single as well as the WFD biological- and total state were investigated. Stressor-response analysis shows divergent results, but a general trend of decreasing ecological integrity with increasing number of stressors and maximum stressor is observed. Fish metrics based on age structure, fish region index and biological status responded best to single stressors and/or their combinations. The knowledge gained in this work provides a basis for advanced investigations in Alpine river basins and beyond, supports WFD implementation and helps prioritizing further actions towards multi-stressor restoration- and management.

Segurado, P., Almeida, C., Neves, R. J. J., Ferreira, M. T., Branco, P. (2018). Understanding multiple stressors in a Mediterranean basin: Combined effects of land use, water scarcity and nutrient enrichment. Science of the Total Environment 624, 1221-1233,

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Understanding multiple stressors in a Mediterranean basin: Combined effects of land use, water scarcity and nutrient enrichment

River basins are extremely complex hierarchical and directional systems that are affected by a multitude of interacting stressors. This complexity hampers effective management and conservation planning to be effectively implemented, especially under climate change. The objective of this work is to provide a wide scale approach to basin management by interpreting the effect of isolated and interacting factors in several biotic elements (fish, macroinvertebrates, phytobenthos and macrophytes). For that, a case study in the Sorraia basin (Central Portugal), a Mediterranean system mainly facing water scarcity and diffuse pollution problems, was chosen. To develop the proposed framework, a combination of process-based modelling to simulate hydrological and nutrient enrichment stressors and empirical modelling to relate these stressors - along with land use and natural background - with biotic indicators, was applied. Biotic indicators based on ecological quality ratios from WFD biomonitoring data were used as response variables. Temperature, river slope, % of agriculture in the upstream catchment and total N were the variables more frequently ranked as the most relevant. Both the two significant interactions found between single hydrological and nutrient enrichment stressors indicated antagonistic effects. This study demonstrates the potentialities of coupling process-based modelling with empirical modelling within a single framework, allowing relationships among different ecosystem states to be hierarchized, interpreted and predicted at multiple spatial and temporal scales. It also demonstrates how isolated and interacting stressors can have a different impact on biotic quality. When performing conservation or management plans, the stressor hierarchy should be considered as a way of prioritizing actions in a cost-effective perspective.

Sepp, M., Kõiv, T., Nõges, P., Nõges, T. (2018). Do organic matter metrics included in lake surveillance monitoring in Europe provide a broad picture of brownification and enrichment with oxygen consuming substances? Science of the Total Environment 610, 1288-1297,

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Do organic matter metrics included in lake surveillance monitoring in Europe provide a broad picture of brownification and enrichment with oxygen consuming substances?

Organic matter (OM) has numerous geochemical and ecological functions in inland waters and can affect water quality. Different parameters of aquatic OM are measured with various methods as no single analytical tool can provide definitive structural or functional information about it. In the present paper we review different OM metrics used in the European Union (EU) lake surveillance monitoring programmes and assess their suitability to provide sufficient data about the brownification and enrichment with oxygen consuming substances in European lakes. In the EU Water Framework Directive (WFD), metrics of OM are not mandatory physico-chemical parameters, but only recommended parameters to characterize water transparency, oxygenation conditions or acidification status. Our analysis shows that, as lake OM is monitored under the WFD in only 14 countries, no Europe-wide conclusions on the situation regarding brownification and organic enrichment can be drawn based on these data. Applied parameters in lake surveillance monitoring programmes are biochemical oxygen demand (BOD), chemical oxygen demand (COD), total organic carbon (TOC), dissolved organic carbon (DOC), water colour (WCol), and yellow substance. Different national OM metrics used avoid getting a broad picture of lake OM concentration changes in Europe over the last decades. Furthermore, our results demonstrate that the possibilities to convert different OM parameters to each other are limited because empirical relationships between them are region-specific. OM sensors for continuous measurements and remote sensing surveys could improve the effectiveness of lake OM monitoring, especially its temporal and spatial representativeness. It would be highly suggested to include in lake monitoring programmes also methods (e.g. absorbance or fluorescence spectroscopy) allowing to characterize the composition of OM as it influences strongly the biogeochemical role of OM in lakes.

Søndergaard, M., Lauridsen, T. L., Johansson, L. S., Jeppesen, E. (2018). Gravel pit lakes in Denmark: Chemical and biological state. Science of the Total Environment 612, 09-17,

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Gravel pit lakes in Denmark: Chemical and biological state

Mining of gravel and sand for construction purposes is big business and gravel pit lakes have become increasingly common all over the world. In Denmark, hundreds of gravel pit lakes have been created during the past decades. We investigated the chemical and biological status of 33–52 gravel pit lakes and compared the results with data from similar-sized natural Danish lakes. The area of the lakes ranged from 0.2 to 13 ha and their age from 0.5 to 26 years. Generally, the gravel pit lakes were clear with low nutrient concentrations, the median concentrations of total phosphorus and total nitrogen being 0.023 mg/l and 0.30 mg/l compared with 0.115 mg/l and 1.29 mg/l, respectively, in natural lakes. Correspondingly, median chlorophyll a was 5 μg/l in the gravel pit lakes and 36 μg/l in the natural lakes. Submerged macrophytes were found in all gravel pit lakes, with particularly high cover in the shallow ones. Most gravel pit lakes were deeper than the natural lakes, which may restrict the area potentially to be covered by submerged macrophytes, with implications also for the biological quality of the lakes. Fish were found in most of the gravel pit lakes, roach (Rutilus rutilus), perch (Perca fluviatilis) and rudd (Scardinius erythrophalmus) being the most frequently observed species. Fish stocking was common and included also non-native species such as carp (Cyprinus carpio) and rainbow trout (Oncorchynchus mykiss). Compared with the natural lakes, fish species richness and catch per gillnet were overall lower in the gravel pit lakes. Groundwater-fed gravel pit lakes add importantly to the number of high-quality lakes in Denmark and with an optimised design and by avoiding negative side effects, they can be positive for both nature and society.

Stefanidis, K., Panagopoulos,Y., Mimikou, M. (2018). Response of a multi-stressed Mediterranean river to future climate and socio-economic scenarios. Science of the Total Environment 627, 756-769,

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Response of a multi-stressed Mediterranean river to future climate and socio-economic scenarios

Streams and rivers are among the most threatened ecosystems in Europe due to the combined effects of multiple pressures related to anthropogenic activities. Particularly in the Mediterranean region, changes in hydromorphology along with increased nutrient loadings are known to affect the ecological functions and ecosystem services of streams and rivers with the anticipated climate change being likely to further impair their functionality and structure. In this study, we investigated the combined effects of agricultural driven stressors on the ecology and delivered services of the Pinios river basin in Greece under three future world scenarios developed within the EU funded MARS project. Scenarios are based on combinations of Representative Concentration Pathways and Shared Socioeconomic Pathways and refer to early century (2030) and mid-century (2060) representing future climate worlds with particular socioeconomic characteristics. To assess the responses of ecological and ecosystem service indicators to the scenarios we first simulated hydrology and water quality in Pinios with a process-based model. Simulated abiotic stressor parameters (predictors) were linked to two biotic indicators, the macroinvertebrate indicators ASPT and EPT, with empirical modelling based on boosted regression trees and general linear models. Our results showed that the techno world scenario driven by fast economic growth and intensive exploitation of energy resources had the largest impact on both the abiotic status (nutrient loads and concentrations in water) and the biotic indicators. In contrast, the predicted changes under the other two future worlds, consensus and fragmented, were more diverse and were mostly dictated by the projected climate. This work showed that the future scenarios, especially the mid-century ones, had significant impact on both abiotic status and biotic responses underpinning the need for implementing catchment management practices able to mitigate the ecological threat on waters in the long-term.

Venohr, M., Langhans, S. L., Peters, O., Holker, F., Arlinghaus, R., Mitchell, L., Wolter, C. (2018). The underestimated dynamics and impacts of water based recreational activities on freshwater ecosystems. Environmental Reviews, online,

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The underestimated dynamics and impacts of water-based recreational activities on freshwater ecosystems

Recreational activities on, in and along freshwaters (e.g., boating, bathing, angling) positively contribute to human well-being, but can concurrently stress aquatic ecosystems. While outdoor recreation, aquatic ecosystems and human well-being form coupled social-ecological systems, inherent fluxes and interactions between these have rarely been properly quantified. This paper synthesizes information on links between water-based recreational activities, effects on freshwater ecosystems integrity and recreational quality and proposes a novel framework for assessment and integrated management. This framework is based on understanding relationships between recreational quality, demand and use and recreational use-induced impacts on ecosystem state and function as well as ecological and social carrying capacities. Current management approaches of freshwater ecosystems addressing economic, environmental or recreational aspects are poorly linked and harmonized, and are further constrained by inadequate information on the dynamics and densities of recreational uses. Novel assessment and monitoring methods are needed to capture the short-term peak dynamics of water-based recreational uses, and we argue social media could play an increasingly important role here. An integrative recreation ecology management concept combined with peak usage information, has great potential to form the basis for next generation management approaches of freshwater and other ecosystems.

Volta, P., Jeppesen, E., Sala, P., Foglini, C., Galafassi, S., Puzzi, C., Winfield, I. J., (2018). Fish assemblages in deep Italian subalpine lakes: History and present status with an emphasis on non-native species. Hydrobiologia, in press.

Yu, Q., Wang, H.-Z., Jeppesen, E., Xu, C., Wang, H.-J. (2018). Reply to Cao et al.'s comment on “Does the responses of Vallisneria natans (Lour.) Hara to high nitrogen loading differ between the summer high-growth season and the low-growth season? Science of the Total Environment 601–602 (2017) 1513–1521”. Science of the Total Environment 615, 1093-1094,

Yu, Q., Wang, H.-Z., Li, Y., Ma, S.-N., Liang, X.-M., Jeppesen, E., Wang, H.-J. (2018). Higher tolerance of canopy-forming Potamogeton crispus than rosette-forming Vallisneria natans to high nitrogen concentration: A whole-ecosystem experiment. Frontiers in Plant Science, in press.

Zajicek, P., Radinger, J., Wolter, C. (2018). Disentangling multiple pressures on fish assemblages in large rivers. Science of the Total Environment 627, 1093-1105,

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Disentangling multiple pressures on fish assemblages in large rivers

European large rivers are exposed to multiple human pressures and maintained as waterways for inland navigation. However, little is known on the dominance and interactions of multiple pressures in large rivers and in particular inland navigation has been ignored in multi-pressure analyzes so far. We determined the response of ten fish population metrics (FPM, related to densities of diagnostic guilds and biodiversity) to 11 prevailing pressures including navigation intensity at 76 sites in eight European large rivers. Thereby, we aimed to derive indicative FPM for the most influential pressures that can serve for fish-based assessments. Pressures' influences, impacts and interactions were determined for each FPM using bootstrapped regression tree models. Increased flow velocity, navigation intensity and the loss of floodplains had the highest influences on guild densities and biodiversity. Interactions between navigation intensity and loss of floodplains and between navigation intensity and increased flow velocity were most frequent, each affecting 80% of the FPM. Further, increased sedimentation, channelization, organic siltation, the presence of artificial embankments and the presence of barriers had strong influences on at least one FPM. Thereby, each FPM was influenced by up to five pressures. However, some diagnostic FPM could be derived: Species richness, Shannon and Simpson Indices, the Fish Region Index and lithophilic and psammophilic guilds specifically indicate rhithralisation of the potamal region of large rivers. Lithophilic, phytophilic and psammophilic guilds indicate disturbance of shoreline habitats through both (i) wave action induced by passing vessels and (ii) hydromorphological degradation of the river channel that comes along with inland navigation. In European large rivers, inland navigation constitutes a highly influential pressure that adds on top of the prevailing hydromorphological degradation. Therefore, river management has to consider river hydromorphology and inland navigation to efficiently rehabilitate the potamal region of large rives.

Zeng, Q., Jeppesen, E., Gua, X., Mao, Z., Chen, H. (2018). Distribution, fate and risk assessment of PAHs in water and sediments from an aquaculture- and shipping-impacted subtropical lake, China. Chemosphere 201, 612-620,

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Distribution, fate and risk assessment of PAHs in water and sediments from an aquaculture- and shipping-impacted subtropical lake, China

The spatial-temporal distribution of polycyclic aromatic hydrocarbons (PAHs), their source, and potential health risks were determined in overlying water and surface sediments from Chinese Lake Guchenghu, adjacent commercial mitten crab ponds and the connected Wushen Canal to assess the contamination profile of the area. The total PAHs concentrations in sediment and water were 86.7-1790 ng g-1dry weight (dw) and 184-365 ng L-1in summer and 184-3140 ng g-1dw and 410-1160 ng L-1in winter. Two- and 3-ring PAHs were the predominant compounds in water, while PAHs with 4-6 rings dominated in the sediment at both upstream and downstream sites. PAHs concentrations in water and sediment correlated significantly. Diagnostic ratios and positive matrix factorization (PMF) analyses indicated a strong influence of pyrogenic sources, principally biomass combustion and vehicle emission, on the concentrations of PAHs. The distribution, source identification, and mean effects range median quotients (mERMQ) analyses suggested that the most contaminated area was located downstream and upstream of the Wushen Canal, followed by Lake Guchenghu and a commercial crab pond area. From an ecological point of view, PAHs posed a potential risk to drinking water sources as the concentrations exceeded the guideline value of 0.05 μg L-1. The risk posed by sediment PAHs appeared to be low except for the downstream sites, which showed a low to medium ecotoxicological risk. The total incremental lifetime cancer risks ranged between 10-7and 10-5, indicating a potential health risk for the local population when exposed to sediment from the area.

Zhen, W., Zhang, X., Guan, B., Yin, C., Yu, J., Jeppesen, E., Liu, Z. (2018). Stocking of herbivorous fish in eutrophic shallow clear-water lakes to reduce standing height of submerged macrophytes while maintaining their biomass. Ecological Engineering 113, 61-64,

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Stocking of herbivorous fish in eutrophic shallow clear-water lakes to reduce standing height of submerged macrophytes while maintaining their biomass

To balance the conservation value versus recreational use of shallow lakes, moderate herbivory may be needed in eutrophic lakes to avoid near surface growth while maintaining high vegetation biomass close to the sediment. However, over-grazing or even complete elimination of macrophytes by grass carp (Ctenopharyngodon idella) commonly used for control purposes has often been observed, leading to a shift from a clear to a turbid phy-toplankton-dominated state. We hypothesized that slow-growing and smaller-sized herbivorous fish species might be more suitable than grass carp to obtain the desired moderate control because they consume the top part of the vegetation without severely affecting the lower plant parts. To test the hypothesis, the effects of Wuchang bream (Megalobrama amblycephala), an endemic medium-sized herbivorous cyprinid, and grass carp on the biomass, density and trait of the macrophyte Vallisneria denseserrulata were compared in an enclosure experiment. We found that V. denseserrulata grew less tall but did not lose biomass under moderate herbivory by Wuchang bream due to increased plant density and leaf weight per length, whereas excessive herbivory by grass carp had strong negative effects on the plant biomass. Moreover, the plant had more and thicker leaves in the fish treatments than in the fishless controls. The growth of grass carp was much faster than that of Wuchang bream. Our findings suggest that stocking of Wuchang bream in proper densities may be more useful than grass carp for the management of V. denseserrulata and likely also other macrophyte species. More tests, especially at different fish densities are, however, needed to draw any firm conclusions regarding this hypothesis.

Zingel, P., Cremona, F., Nõges, T., Cao, Y., Neif, É. M., Coppens, J., Işkın, U., Lauridsen, T. L., Davidson, T. A., Søndergaard, M., Beklioglu, M., Jeppesen, E. (2018). Effects of warming and nutrients on the microbial food web in shallow lake mesocosms. European Journal of Protistology 64, 1-12,

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Effects of warming and nutrients on the microbial food web in shallow lake mesocosms

We analysed changes in the abundance, biomass and cell size of the microbial food web community (bacteria, heterotrophic nanoflagellates, ciliates) at contrasting nutrient concentrations and temperatures during a simulated heat wave. We used 24 mesocosms mimicking shallow lakes in which two nutrient levels (unenriched and enriched by adding nitrogen and phosphorus) and three different temperature scenarios (ambient, IPCC A2 scenario and A2 +%50) are simulated (4 replicates of each). Experiments using the mesocosms have been running un-interrupted since 2003. A 1-month heat wave was imitated by an extra 5 °C increase in the previously heated mesocosms (from 1st July to 1st August 2014). Changes in water temperature induced within a few days a strong effect on the microbial food web functioning, demonstrating a quick response of microbial communities to the changes in environment, due to their short generation times. Warming and nutrients showed synergistic effects. Microbial assemblages of heterotrophic nanoflagellates and ciliates responded positively to the heating, the increase being largest in the enriched mesocosms. The results indicate that warming and nutrients in combination can set off complex interactions in the microbial food web functioning.


Alahuhta, J., Kosten, S., Akasaka, M., Auderset, D., Azzella, M. M., Bolpagni, R., Bove, C. P., Chambers, P. A., Chappuis, E., Clayton, J., de Winton, M., Ecke, F., Gacia, E., Gecheva, G., Grillas, P., Hauxwell, J., Hellsten, S., Hjort, J., Hoyer, M. V., Ilg, C., Kolada, A., Kuoppala, M., Lauridsen, T., Li, E. H., Lukács, B. A., Mjelde, M., Mikulyuk, A., Mormul, R. P., Nishihiro, J., Oertli, B., Rhazi, L., Rhazi, M., Sass, L., Schranz, C., Søndergaard, M., Yamanouchi, T., Yu, Q., Wang, H., Willby, N., Zhang, X. K., Heino, J. (2017). Global variation in the beta diversity of lake macrophytes is driven by environmental heterogeneity rather than latitude. Journal of Biogeography 44, 1758–1769,

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Global variation in the beta diversity of lake macrophytes is driven by environmental heterogeneity rather than latitude

We studied global variation in beta diversity patterns of lake macrophytes using regional data from across the world. Specifically, we examined (1) how beta diversity of aquatic macrophytes is partitioned between species turnover and nestedness within each study region, and (2) which environmental characteristics structure variation in these beta diversity components. Global. We used presence–absence data for aquatic macrophytes from 21 regions distributed around the world. We calculated pairwise-site and multiple-site beta diversity among lakes within each region using Sørensen dissimilarity index and partitioned it into turnover and nestedness coefficients. Beta regression was used to correlate the diversity coefficients with regional environmental characteristics. Aquatic macrophytes showed different levels of beta diversity within each of the 21 study regions, with species turnover typically accounting for the majority of beta diversity, especially in high-diversity regions. However, nestedness contributed 30–50% of total variation in macrophyte beta diversity in low-diversity regions. The most important environmental factor explaining the three beta diversity coefficients (total, species turnover and nestedness) was elevation range, followed by relative areal extent of freshwater, latitude and water alkalinity range. Our findings show that global patterns in beta diversity of lake macrophytes are caused by species turnover rather than by nestedness. These patterns in beta diversity were driven by natural environmental heterogeneity, notably variability in elevation range (also related to temperature variation) among regions. In addition, a greater range in alkalinity within a region, likely amplified by human activities, was also correlated with increased macrophyte beta diversity. These findings suggest that efforts to conserve aquatic macrophyte diversity should primarily focus on regions with large numbers of lakes that exhibit broad environmental gradients.

Audet, J., Neif, E. M., Cao, Y., Hoffmann, C. C., Lauridsen, T. L., Larsen, S. E., Søndergaard, M., Jeppesen, E., Davidson, T. A. (2017). Heat-wave effects on greenhouse gas emissions from shallow lake mesocosms. Freshwater Biology 62, 1130–1142,

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Heat-wave effects on greenhouse gas emissions from shallow lake mesocosms

Shallow lakes are a key component of the global carbon cycle. It is, therefore, important to know how shallow lake ecosystems will respond to the current climate change. Global warming affects not only average temperatures, but also the frequency of heat waves (HW). The impact of extreme events on ecosystems processes, particularly greenhouse gas (GHG) emissions, is uncertain. Using the world's longest-running shallow lake experiment, we studied the effects of a simulated summer HW on the fluxes of carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O). The experimental mesocosms had been exposed to different temperature treatments and nutrient loading for 11 years prior to the artificial HW. In general, there was an increase in total GHG emissions during the 1-month artificial HW, with a significant increase in CO2, CH4 and N2O being observed in the shallow lake mesocosms. No significant effect of the HW on CO2 emissions could be traced, though, in the mesocosms with high nutrient levels. Furthermore, the data suggested that in addition to the direct effect of increased temperature on metabolic processes during the HW, biotic interactions exerted a significant control of GHG emissions. For example, at low nutrient levels, increased CO2 emissions were associated with low macrophyte abundance, whereas at high nutrient levels, decreased phytoplankton abundance was linked to increased emissions of CO2 and CH4. In contrast to the observable heat-wave effect, no clear general effect of the long-term temperature treatments could be discerned over the summer, likely because the potential effects of the moderate temperature increase, applied as a press disturbance, were overridden by biotic interactions. This study demonstrates that the role of biotic interactions needs to be considered within the context of global warming on ecosystem processes.

Baattrup-Pedersen, A., Göthe, E., Riis, T., Andersen, D. K., Larsen, S. E. (2017). A new paradigm for bio-monitoring - An example building on the Danish Stream Plant Index. Methods in Ecology and Evolution 8, 297-307,

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A new paradigm for biomonitoring: an example building on the Danish Stream Plant Index

1. Despite intensive efforts for more than a decade to develop Water Framework-compliant assessment systems, shortcomings continue to appear. In particular, the lack of reference conditions has hindered the development of assessment systems capturing the heart of the Water Framework Directive (WFD) – that ecological status should be set as the deviation from the natural, undisturbed condition. Recently, the Danish Stream Plant Index (DSPI) was developed. This system contrasts existing systems in that it builds on an expert interpretation of the normative definitions of ecological status classes in the WFD without taking pressure–impact relationships into account.

2. Here, we substantiate the approach taken in the development of DSPI and examine whether the DSPI class decreases with increasing level of anthropogenic stress and, additionally, whether the deviation from the natural undisturbed condition increases with decreasing DSPI class sensu WFD using trait composition of plant assemblages from Danish streams around year 1900 as a reference. We furthermore examine the trait composition of the vegetation in sites classified into different DSPI status classes to explore whether predictable patterns exist that can be used to identify the ultimate cause(s) of failure to meet ecological goals and help guide the selection of appropriate mitigation measures.

3. We observed that DSPI declined with several parameters indicative of environmental stress in Danish streams and, furthermore, that the deviation from the natural undisturbed condition regarding the trait composition of plant communities declined with increasing DSPI, implying that the trait composition of plant communities in the high DSPI status class was most similar to those occurring in Danish streams around year 1900. We also found that trait characteristics capable of disentangling important stressors in Danish streams varied consistently among sites classified into different DSPI classes.

4. Based on our findings, we call for new thinking. We suggest that more effort should be directed at describing reference conditions and interpreting the normative definitions of good, moderate, poor and bad instead of focusing solely on developing assessment systems using pressure–impact frameworks. We find this particularly important with respect to streams as these are seldom impacted by only a single stressor.

Bækkelie, K. A. E., Schneider, S. C., Hagman, C. H. C., Petrin, Z. (2017). Effects of flow events and nutrient addition on stream periphyton and macroinvertebrates: An experimental study using flumes. Knowledgement and Management of Aquatic Ecosystems 418, 47,

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Effects of flow events and nutrient addition on stream periphyton and macroinvertebrates: an experimental study using flumes

We used flume experiments to study the effects of a temporary increase in stream flow on macroinvertebrates, leaf litter breakdown and soft-bodied benthic algae; both as a single stressor and eventually in combination with an increase in nutrient supply. In order to understand how well the flumes reflected the nearby stream ecosystem, we compared species composition of macroinvertebrates and benthic algae between the flumes and the nearby stream from which the flumes were supplied with water. As single stressors, nutrient addition and an increased flow velocity from 1.3 to 2.8 cm s-1 lead to an increase in the biomass of benthic algae, likely reflecting an improved transfer of nutrients into algal patches. However, the combined effect of flow and nutrient addition was smaller than the sum of both individual effects, likely because an increased biomass also was more susceptible to scouring. We found differences in macroinvertebrate and benthic algal taxon identity and abundance between stream and flumes. Since biodiversity is assumed to stabilize ecological functioning in response to disturbances and variation, we conclude that care should be taken in applying results from small scale experiments to stream ecosystems.

Beklioğlu, M., Bucak, T., Coppens, J., Bezirci, G., Tavşanoğlu, Ü. N., Çakıroğlu, A. İ., Levi, E. E., Erdoğan, Ş., Filiz, N., Özkan, K., Özen, A. (2017). Restoration of eutrophic lakes with fluctuating water levels: A 20 year monitoring study of 2 inter-connected lakes. Water 9, 127,

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Restoration of eutrophic lakes with fluctuating water levels: A 20 year monitoring study of 2 inter-connected lakes

Eutrophication continues to be the most important problem preventing a favorable environmental state and detrimentally impacting the ecosystem services of lakes. The current study describes the results of analyses of 20 year monitoring data from two interconnected Anatolian lakes, Lakes Mogan and Eymir, receiving sewage effluents and undergoing restoration. The first step of restoration in both lakes was sewage effluent diversion. Additionally, in hypertrophic Lake Eymir, biomanipulation was conducted, involving removal of benthi-planktivorous fish and prohibition of pike fishing. The monitoring period included high (H) and low (L) water levels (WL) enabling elucidation of the effects of hydrological changes on lake restoration. In shallower Lake Mogan, macrophyte abundance increased after the sewage effluent diversion in periods with low water levels even at turbid water. In comparatively deeper Lake Eymir, the first biomanipulation led to a clear water state with abundant macrophyte coverage. However, shortly after biomanipulation, the water clarity declined, coinciding with low water level (LWL) periods during which nutrient concentrations increased. A second biomanipulation was conducted, mostly during high water level (HWL) period, resulting in a major decrease in nutrient concentrations and clearer water, but without an expansion of macrophytes. We conclude that repetitive fish removal may induce recovery but its success may be confounded by high availability of nutrients and adverse hydrological conditions.

Borza, P., Huber, T., Leitner, P., Remund, N., Graf, W. (2017). Current velocity shapes co-existence patterns among invasive Dikerogammarus species. Freshwater Biology 62, 317-328,

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Current velocity shapes co-existence patterns among invasive Dikerogammarus species

RFacilitative interactions among co-evolved representatives of the endemic Ponto-Caspian fauna are regarded as a major factor of their invasion success. Nevertheless, the most renowned examples represent interactions between different trophic levels or functional groups, while ecologically similar species can be expected to show competition-based niche partitioning. Here, we test for differences in the realized niche of three invasive Dikerogammarus species (Crustacea: Gammaridae) in their co-occurring range. We sampled multiple habitats within sites distributed along the River Danube to test whether some environmental variables could reveal spatial niche differentiation among the three species of Dikerogammarus, and if so, to test a predictive model outside the zone of co-occurrence. Spatial niche differentiation was present among the species, primarily determined by current velocity (and associated substrate preference), likely reflecting a stress tolerance–competitive ability trade-off. Suspended matter concentration was also relevant, suggesting food resources (through filter feeding) might represent another important niche axis, somewhat loosening the terms of co-existence between D. haemobaphes and the other two species. Environmental variables could effectively explain the absence of D. bispinosus in the Lower Danube, implying that the co-existence of the three species is possible only along a sufficiently wide current velocity gradient, and the observed turnovers are the result of niche expansion in the absence of the stronger competitor. Hence, differences in invasion success may be attributed to a stress tolerance–competitive ability trade-off. Our results suggest the advantage of D. villosus is attributable to its competitive dominance, allowing it to monopolize lentic and/or structured habitats, which represents a fortunate pre-adaptation to anthropogenic alterations of aquatic ecosystems. The presence of D. villosus does not greatly affect the expansion of D. haemobaphes; however, the exclusion of D. bispinosus from lentic habitats by D. villosus probably strongly limits its potential to spread by active dispersal.

Borza, P., Huber, T., Leitner, P., Remund, N., Graf, W. (2017). Success factors and future prospects of Ponto–Caspian peracarid (Crustacea: Malacostraca) invasions: Is “the worst over”? Biological Invasions 19, 1517,

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Success factors and future prospects of Ponto–Caspian peracarid (Crustacea: Malacostraca) invasions: Is ‘the worst over’?

Ponto–Caspian peracarids (amphipods, isopods, mysids and cumaceans) represent one of the most successful groups of aquatic invaders comprising several high-impact species, such as Chelicorophium curvispinum, Dikerogammarus villosus, or Hemimysis anomala. In the present study we made the first attempt to compare biological traits and the environmental preferences of invasive and non-invasive members of the group based on both literature and field data (Joint Danube Survey 3, 2013) with the goal of identifying factors linked to invasion success and drawing conclusions on future invasion risks. Both datasets indicated substrate preference as an important factor in spontaneous range expansion; all invasive species are lithophilous, whereas the majority of non-invasives are psammo-pelophilous. The remaining seven presently non-invasive lithophilous species deserve special attention when considering potential future invaders; however, due to their rarity and possible negative interactions with earlier colonists we consider the probability of their expansion in the foreseeable future as low. Their potential expansion could most likely be of minor consequence anyway, since no considerable functional novelty can be attributed to them in addition to species already present. In this limited context (regarding habitats dominated by hard substrates and not considering the potential further spread of already invasive species) it might be justified to conclude that ‘the worst is over’. Nevertheless, impending navigation development projects both in the Danube–Main–Rhine and Dnieper–Pripyat–Bug–Vistula systems might favour the future spread of non-lithophilous species, which might imply a new invasion wave of Ponto–Caspian peracarids.

Brucet, S., Arranz, I., Mehner, T., Argillier, C., Beklioğlu, M., Benejam, L., Boll, T., Holmgren, K., Lauridsen, T. L., Svenning, J.-C., Winfield I. J., Jeppesen, E. (2017). Size diversity and species diversity relationships in fish assemblages of Western Palearctic lakes. Ecography 40, 01-12,

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Size diversity and species diversity relationships in fish assemblages of Western Palearctic lakes

Body size, coupled with abundance and taxonomy, may help to understand the mechanisms shaping community structure. Since the body size of fish is closely related to their trophic niche, size diversity (based on individual body size) of fish communities may capture intraspecific variations in fish trophic niches that are not detected by species diversity. Thus, the relationship between size diversity and species diversity may help to integrate variation at both intraspecific and interspecific levels. We studied the relationship between species diversity and size diversity as a measure of the degree of overlap in size among species and thereby the potential overlap in niches in a community. We hypothesized that the relationship between size diversity and species would be different across the European continent due to different levels of size overlap in fish communities. The data were derived from samplings of fish communities using standardised benthic gill nets in 363 lakes. At the continental scale, size diversity increased with species diversity; at the ecoregion scale, the slope of the relation changed across the continent, with the greatest mismatch occurring in northern Europe where communities comprised only one or a few species, but each of which exhibited a great range in size. There was an increase in slope towards the south with significant relations for four out of six ecoregions. The steeper size diversity-species diversity slope at lower latitudes is attributable to a lower overlap in fish size and thus likely to finer niche separation. Our results also suggest that size diversity is not a strong surrogate for species diversity in European lake fish communities. Thus, particularly in fish communities composed of few species, measuring size diversity may help to detect potential functional variation which may be neglected by measuring species diversity alone.

Brucet, S., Tavşanoğlu, Ü. N., Özen, A., Levi, E. E., Bezirci, G., Çakıroğlu, A. I., Jeppesen, E., Svenning, J.-C., Ersoy, Z., Beklioğlu, M. (2017). Size-based interactions across trophic levels in food webs of shallow Mediterranean lakes. Freshwater Biology 62, 1819–1830,

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Size-based interactions across trophic levels in food webs of shallow Mediterranean lakes

1. Body size is a key trait of an organism which determines the dynamics of predator–prey interactions. Most empirical studies on the individual size distribution of the aquatic community have focused on the variations in body size of a single trophic level as a response to certain environmental variables or biotic factors. Few studies, however, have evaluated how individual size structure is altered simultaneously across interacting trophic levels and locations. Such comparative examinations of the size distribution in predator and prey communities may bring insight into the strength of the interactions between adjacent trophic levels. 2. We assessed the potential predation effect of size-structured predators (i.e. predation by individuals of different sizes) on prey size structure using data from 30 shallow Turkish lakes spanning over five latitudinal degrees. We correlated size diversity and size evenness of predator and prey assemblages across the planktonic food web after accounting for the confounding effects of temperature and resource availability which may also affect size structure. We expected to find a negative relationship between size diversity of predators and prey due to the enhanced strength of top-down control with increasing predator size diversity. We also hypothesised that competitive interactions for resources in less productive systems would promote a higher size diversity. We further expected a shift towards reduced size diversity and evenness at high temperatures. 3. In contrast to our hypothesis, we found a positive correlation between size structures of two interacting trophic levels of the planktonic food web; thus, highly size-diverse fish assemblages were associated with highly size-diverse zooplankton assemblages. The size evenness of fish and phytoplankton assemblages was negatively and positively related to temperature, respectively. Phytoplankton size diversity was only weakly predicted by the resource availability. 4. Our results suggest that size structure within a trophic group may be controlled by the size structure at adjacent trophic levels, as well as by temperature and resource availability. The positive relationship between the size diversity of fish and zooplankton suggests that higher diversity of the resources drives a higher size diversity of consumers or vice versa, and these effects are beyond those mediated by taxonomic diversity. In contrast, the size diversity and size evenness of phytoplankton are mainly influenced by physical factors in this region and perhaps in warm shallow lakes in general.

Bucak, T., Trolle, D., Andersen, H. E., Thodsen, H., Erdogan, S., Levi, E. E., Filiz, N., Jeppesen, E., Beklioglu, M. (2017). Future water availability in the largest freshwater Mediterranean lake is at great risk as evidenced from simulations with the SWAT model. Science of the Total Environment 581-582, 413-425,

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Future water availability in the largest freshwater Mediterranean lake is at great risk as evidenced from simulations with the SWAT model

Inter- and intra-annual water level fluctuations and changes in water flow regime are intrinsic characteristics of Mediterranean lakes. Additionally, considering climate change projections for the water-limited Mediterranean region, increased air temperatures and decreased precipitation are anticipated, leading to dramatic declines in lake water levels as well as severe water scarcity problems. The study site, Lake Beyşehir, the largest freshwater lake in the Mediterranean basin, is – like other Mediterranean lakes – threatened by climatic changes and over-abstraction of water for irrigated crop farming. Therefore, implementation of strict water level management policies is required. In this study, an integrated modeling approach was used to predict the future water levels of Lake Beyşehir in response to potential future changes in climate and land use. Water level estimation was performed by linking the catchment model Soil and Water Assessment Tool (SWAT) with a Support Vector Regression model (ε-SVR). The projected increase in temperature and decrease in precipitation based on the climate change models led to an enhanced potential evapotranspiration and reduced total runoff. On the other hand, the effects of various land use scenarios within the catchment appeared to be comparatively insignificant. According to the ε-SVR model results, changes in hydrological processes caused a water level reduction for all scenarios. Moreover, the MPI-ESM-MR General Circulation Model outputs produced the most dramatic results by predicting that Lake Beyşehir may dry out by the 2040s with the current outflow regime. The results indicate that shallow Mediterranean lakes may face a severe risk of drying out and losing their ecosystem values in the near future if the current intensity of water abstraction is not reduced. In addition, the results also demonstrate that outflow management and sustainable use of water sources are vital to sustain lake ecosystems in water-limited regions.

Calapez, A. R., Branco, P., Santos, J. M., Ferreira, M. T., Hein, T., Brito, A. G., Feio, M. J. (2017). Macroinvertebrate short-term responses to flow variation and oxygen depletion: A mesocosm approach. Science of the Total Environment 599-600, 1202-1212,

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Macroinvertebrate short-term responses to flow variation and oxygen depletion: A mesocosm approach

In Mediterranean rivers, water scarcity is a key stressor with direct and indirect effects on other stressors, such as water quality decline and inherent oxygen depletion associated with pollutants inputs. Yet, predicting the responses of macroinvertebrates to these stressors combination is quite challenging due to the reduced available information, especially if biotic and abiotic seasonal variations are taken under consideration. This study focused on the response of macroinvertebrates by drift to single and combined effects ofwater scarcity and dissolved oxygen (DO) depletion over two seasons (winter and spring). A factorial design of two flowvelocity levels - regular and low (vL) - with three levels of oxygen depletion - normoxia, medium depletion (dM) and higher depletion (dH) - was carried out in a 5-artificial channels system, in short-term experiments. Results showed that both stressors individually and together had a significant effect on macroinvertebrate drift ratio for both seasons. Single stressor effects showed that macroinvertebrate drift decreased with flow velocity reduction and increased with DO depletion, in both winter and spring experiments. Despite single stressors opposing effects in drift ratio, combined stressors interaction (vL × dM and vL × dH) induced a positive synergistic drift effect for both seasons, but only in winter the drift ratio was different between the levels of DO depletion. Stressors interaction in winter seemed to intensify drift response when reached lower oxygen saturation. Also, drift patterns were different between seasons for all treatments, which may depend on individual's life stage and seasonal behaviour. Water scarcity seems to exacerbate the oxygen depletion conditions resulting into a greater drifting of invertebrates. The potential effects of oxygen depletion should be evaluated when addressing the impacts of water scarcity on river ecosystems, since flow reductions will likely contribute to a higher oxygen deficit, particularly in Mediterranean rivers.

Cao Y., Olsen, S., Gutierrez, M. F., Brucet, S., Davidson, T. A., Li, W., Lauridsen, T. L., Søndergaard, M., Jeppesen, E. (2017). Temperature effects on periphyton, epiphyton and epipelon under a nitrogen pulse in low-nutrient experimental freshwater lakes. Hydrobiologia 795, 267-279,

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Temperature effects on periphyton, epiphyton and epipelon under a nitrogen pulse in low-nutrient experimental freshwater lakes

The ongoing global climate change involves not only increased temperatures but may also produce more frequent extreme events, such as severe rainfall that could trigger a pulse of nutrients to lakes. In shallow lakes, this may affect primary producers through a number of direct and indirect mechanisms. We conducted a six-month mesocosm experiment to elucidate how periphyton (on inert substrata), epiphyton and epipelon biomass responded to a nitrogen (N) pulse, an approximately tenfold enrichment of the NO3-pool, under three contrasting warming scenarios: ambient temperature and ca. +3°C and ca. +4.5°C elevated temperatures (hereafter T1, T2 and T3). After the N pulse, we found a higher periphyton biomass at elevated than at ambient temperatures but no change in epiphyton biomass. Epipelon biomass was lower in T3 than in T1. Both periphyton and epiphyton biomasses correlated negatively with snail biomass, while epiphyton biomass correlated positively with light. Different responses to higher temperatures under short-term extreme nutrient loading conditions may be attributed to differences in the access to nutrient sources and light. Our data suggest that the biomass of periphyton in oligotrophic clear-water lakes will increase significantly under conditions exhibiting short-term extreme nutrient loading in a warmer climate.

Carbonell, J. A., Velasco, J., Millán, A., Green, A. J., Coccia, C., Guareschi, S., Gutiérrez-Cánovas, C. (2017). Biological invasion modifies the co-occurrence patterns of insects along a stress gradient. Functional Ecology 31, 1957–1968,

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Biological invasion modifies the co-occurrence patterns of insects along a stress gradient

1. Biological invasions have become one of the most important drivers of biodiversity loss and ecosystem change world-wide. However, it is still unclear how invasions may interact with local abiotic stressors, which are expected to increase as global change intensifies. Furthermore, we know little about the response to biological invasions of insects, despite their disproportionate contribution to global animal biodiversity. 2. The aim of the present work is to investigate the impact of an invasive aquatic insect on the co-occurrence patterns of native species of insects along a salinity gradient, and determine which assembly rules are driving these patterns. 3. First, we characterised the habitat specialisation and functional niches of each species from physiological and biological traits, respectively, and their degree of overlap. Second, we used field data to compare the co-occurrence patterns of native and invasive species in invaded and non-invaded areas of southern Iberia and northern Morocco. Finally, we tested if habitat filtering or niche differentiation assembly rules mediate their co-occurrence. 4. In non-invaded areas, habitat filtering drives habitat segregation of species along the salinity gradient, with a lower contribution of niche differentiation. The presence of the invasive insect modifies the distribution and co-occurrence patterns of native species. In invaded areas, niche differentiation seems to be the main mechanism to avoid competition among the invasive and native species, enabling coexistence and resource partitioning. 5. The combined study of functional niche similarity and abiotic stressor tolerance of invasive and native species can improve our understanding of the effects of invasive species along abiotic stress gradients. This approach may increase our capacity to predict the outcomes of biological invasion in a global change context.

Cremona, F., Vilbaste, S., Couture, R.M., Nõges, P., Nõges, T. (2017). Is the future of large shallow lakes blue-green? Comparing the response of a catchment-lake model chain to climate predictions. Climatic Change 141(2), 347-361,

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Is the future of large shallow lakes blue-green? Comparing the response of a catchment-lake model chain to climate predictions

We constructed a model chain into which regional climate-related variables (air temperature, precipitation) and a lake’s main tributary hydrological indicators (river flow, dissolved inorganic carbon) were employed for predicting the evolution of planktonic blue-green algae (cyanobacteria) and zooplankton (rotifer) biomass in that lake for the mid-21st century. Simulations were based on the future climate predicted under both the Representative Concentration Pathways 4.5 and 8.5 scenarios which, combined with three realistic policy-making and basin land-use evolution lead to six scenarios for future water quality. Model outputs revealed that mean annual river flow is expected to decline between 3 to 20%, depending of the scenario. Concentration of river dissolved inorganic carbon is predicted to follow the opposite trend and might soar up to twice the 2005-2014 average concentration. Lake planktonic primary producers will display quantitative changes in the future decades whereas zooplankters will not. A 2 to 10% increase in mean cyanobacteria biomass is accompanied by a stagnation (-3 to +2%) of rotifer biomass. Changes in cyanobacteria and rotifer phenology are expected: a surge of cyanobacteria biomass in winter and a shortening of the rotifer biomass spring peak. The expected quantitative changes on the biota were magnified in those scenarios where forested area conversion to cropland and water abstraction were the greatest.

Ding, N., Yang, W., Zhou, Y., Gonzalez-Bergonzoni, I., Zhang, J., Chen, K., Vidal, N., Jeppesen, E., Liu, Z., Wang, B. (2017). Different responses of functional traits and diversity of stream macroinvertebrates to environmental and spatial factors in the Xishuangbanna watershed of the upper Mekong River Basin, China. Science of the Total Environment 574, 288-299,

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Different responses of functional traits and diversity of stream macroinvertebrates to environmental and spatial factors in the Xishuangbanna watershed of the upper Mekong River Basin, China

Functional traits and diversity indices have provided new insights into community responses to stressors. Most traits of aquatic organisms have frequently been tested for predictability and geographical stability in response to environmental variables, but such tests of functional diversity indices are rare. We sampled macroinvertebrates at 18 reference sites (RS) and 35 disturbed sites (DS) from headwater streams in the upper Mekong River Basin, Xishuangbanna (XSBN), China. We selected 29 qualitative categories of eight traits and then calculated five functional diversity indices, namely functional richness (FRic), functional evenness (FEve), functional dispersion (FDis), functional divergence (FDiv) and Rao's Quadratic Entropy (RaoQ), and two trait diversity indices, namely trait richness (TR) and trait diversity (TD). We used combination of RLQ and fourth-corner to examine the response of traits and functional diversity to the disturbance and environmental variables. We used variance partitioning to explore the relative role of environmental variables and spatial factors in constraining trait composition and functional diversity. We found that the relative frequency of ten trait categories, and the values of TD, TR, FRic and FDis in RS were significantly different (p<0.05) from DS. In addition, the seven traits (except for "habit") demonstrated a predictable response of trait patterns along the integrative environmental gradients. Environmental variables significantly contributed to most of the traits, functional diversity and trait diversity. However, spatial variables were mainly significant in shaping ecological traits, FRic and FEve. Our results confirm the dominant role of environmental variables in the determination of community trait composition and functional diversity, and substantiate the contribution of spatial vectors in explaining the variance of functional traits and diversity. We conclude that the traits "Refuge", "External protection", "Respiration" and "Body shape", and diversity indices FDis, TD, and TR are promising indicators of stream conditions at XSBN.

Elosegi, A., Gessner, M. O., Young, R. G. (2017). River doctors: Learning from medicine to improve ecosystem management. Science of the Total Environment 595, 294-302,

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River doctors: Learning from medicine to improve ecosystem management

Effective ecosystem management requires a robust methodology to analyse, remedy and avoid ecosystem damage. Here we propose that the overall conceptual framework and approaches developed over millennia in medical science and practice to diagnose, cure and prevent disease can provide an excellent template. Key principles to adopt include combining well-established assessment methods with new analytical techniques and restricting both diagnosis and treatment to qualified personnel at various levels of specialization, in addition to striving for a better mechanistic understanding of ecosystem structure and functioning, as well as identifying the proximate and ultimate causes of ecosystem impairment. In addition to applying these principles, ecosystem management would much benefit from systematically embracing how medical doctors approach and interview patients, diagnose health condition, select treatments, take follow-up measures, and prevent illness. Here we translate the overall conceptual framework from medicine into environmental terms and illustrate with examples from rivers how the systematic adoption of the individual steps proven and tested in medical practice can improve ecosystem management.

Ersoy, Z., Jeppesen, E., Sgarzi, S., Arranz, I., Cañedo-Argüelles, M., Quintana, X, D., Landkildehus, F., Lauridsen, T. L., Bartrons, M., Brucet, S. (2017). Size-based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn, Iceland. Freshwater Biology 62, 1942–1952.

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Size-based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn, Iceland

1. Trophic cascade studies have so far mostly focused on changes in the abundance, biomass, or average size of prey and predators. In contrast, individual size-based interactions, playing a key role in the trophic structure and functioning of aquatic ecosystems, have been less explored. 2. We conducted a 3-month in situ experiment in Lake Myvatn, Iceland, with two fish treatments (with and without fish, Gasterosteus aculeatus). After the first month of the experiment, Anabaena blooms appeared in the lake. We studied the effects of fish predation and occurrence of cyanobacteria blooms on the individual size structure (i.e. the distribution of the number of organisms over a size range) of zooplankton and phytoplankton. We also assessed the potential consequences for trophic transfer efficiency (TTE) (measured as the predator to prey biomass ratio) in the planktonic food web. 3. Our results showed that fish predation and cyanobacteria bloom had a negative relationship with size diversity of zooplankton, which became dominated by small-sized individuals in both cases. The phytoplankton size diversity changed over time particularly due to the blooming of large-sized Anabaena, and its increase was apparently mainly driven by changes in resources. 4. Low zooplankton size diversity related to fish predation reduced TTE, particularly in the enclosures with fish. This may be because low zooplankton size diversity represents a lower partition of resources among consumers, thereby decreasing the trophic energy transfer. With the occurrence of Anabaena bloom, high phytoplankton size diversity coincided with a lower energy transfer in all enclosures likely due to reduced zooplankton grazing when large-sized colony-forming Anabaena dominated. 5. In conclusion, our results indicate that both top-down and bottom-up forces significantly influence the size structure of planktonic communities. The changes in size structure were related to shifts in the energy transfer efficiency of the Lake Myvatn food web. Thus, our study underpins the importance of taking into account size-based interactions in the study of trophic cascades, particularly in a warming climate where strong planktivorous fish predation and frequent cyanobacteria blooms may occur.

Forsius, M., Räike, A., Poutanen, H., Mattsson, T., Kankaanpää, S., Kortelainen, P., Vuorilehto, V.-P. (2017). Observed and predicted future changes of total organic carbon in the lake Päijänne catchment (southern Finland): Implications for water treatment of the Helsinki metropolitan area. Boreal Environment Research 22, 317-336,

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Observed and predicted future changes of total organic carbon in the lake Päijänne catchment (southern Finland): Implications for water treatment of the Helsinki metropolitan area.

Lake Päijänne (1083 km2) is the main drinking-water source for more than one million people of the Helsinki metropolitan area and the lake is important also for fisheries and recreation. We carried out a detailed study on observed and modelled future changes of total organic carbon (TOC) in this large lake catchment (26 459 km2), and assessed the implications for current water treatment processes. Concentrations of TOC/CODMn in the lake and its sub-catchments increased during the period 2000–2014 (CODMn was used as a TOC proxy in the statistical analysis because of long-term data availability). Stepwise multiple regression analysis on the relationships between CODMn, water quality parameters, air temperature and runoff in selected 18 catchments indicated that the explaining variables are site-specific and that the observed recent changes are mainly due to a combination of the simultaneously interacting factors of climate and atmospheric deposition. Linear regression analysis revealed no relationships between CODMn increases and landuse related factors or specific catchment characteristics. A catchment-scale dynamic model system (VEMALA) was used to predict future concentrations and fluxes of TOC, using three different climate change scenarios. TOC concentrations were predicted to decrease by 19% in the Dry scenario and increase by 13% in the Wet scenario by the end of the century. The current treatment processes in the city of Helsinki can likely meet the challenges set by climate change in form of changed raw water TOC and quality. Integrated carbon (C) balance calculations indicated large changes in future C fluxes and lake retention time. These changes would markedly affect several key ecosystem processes such as transport of organically bound nutrients to sea areas and the importance of surface waters in the overall landscape C budgets.

Gao J., Zhong, P., Ning, J., Liu, Z., Jeppesen, E. (2017). Herbivory of omnivorous fish shapes the food web structure of a Chinese tropical eutrophic lake: Evidence from stable isotope and fish gut content analyses. Water 9(1), 69,

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Herbivory of omnivorous fish shapes the food web structure of a Chinese tropical eutrophic lake: Evidence from stable isotope and fish gut content analyses

Studies suggest that, unlike the situation in temperate lakes, high biomasses of omnivorous fish are maintained in subtropical and tropical lakes when they shift from a turbid phytoplankton-dominated state to a clear water macrophyte-dominated state, and the predation pressure on large-bodied zooplankton therefore remains high. Whether this reflects a higher degree of herbivory in warm lakes than in temperate lakes is debatable. We combined food web studies using stable isotopes with gut content analyses of the most dominant fish species to elucidate similarities and differences in food web structure between a clear water macrophyte-dominated basin (MDB) and a turbid phytoplankton-dominated basin (PDB) of Huizhou West Lake, a shallow tropical Chinese lake. The δ 13 C–δ 15 N biplot of fish and invertebrates revealed community-wide differences in isotope-based metrics of the food webs between MDB and PDB. The range of consumer δ 15 N (NR) was lower in MDB than in PDB, indicating shorter food web length in MDB. The mean nearest neighbor distance (MNND) and standard deviation around MNND (SDNND) were higher in MDB than in PDB, showing a markedly low fish trophic overlap and a more uneven packing of species in niches in MDB than in PDB. The range of fish δ 13 C (CR) of consumers was more extensive in MDB than in PDB, indicating a wider feeding range for fish in MDB. Mixing model results showed that macrophytes and associated periphyton constituted a large fraction of basal production sources for the fish in MDB, while particulate organic matter (POM) contributed a large fraction in PDB. In MDB, the diet of the dominant fish species, crucian carp (Carassius carassius), consisted mainly of vegetal matter (macrophytes and periphyton) and zooplankton, while detritus was the most important food item in PDB. Our results suggest that carbon from macrophytes with associated periphyton may constitute an important food resource for omnivorous fish, and this may strongly affect the feeding niche and the strength of the top-down trophic cascade between fish and zooplankton in the restored, macrophyte-dominated basin of the lake. This dual effect (consumption of macrophytes and zooplankton) may reduce the chances of maintaining the clear water state at the prevailing nutrient levels in the lake, and regular removal of large crucian carp may therefore be needed to maintain a healthy ecosystem state.

Gieswein, A., Hering, D., Feld, C. K. (2017). Additive effects prevail: The response of biota to multiple stressors in an intensively monitored watershed. Science of the Total Environment 593-594, 27-35,

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Additive effects prevail: The response of biota to multiple stressors in an intensively monitored watershed

Freshwater ecosystems are impacted by a range of stressors arising from diverse human-caused land and water uses. Identifying the relative importance of single stressors and understanding how multiple stressors interact and jointly affect biology is crucial for River Basin Management. This study addressed multiple human-induced stressors and their effects on the aquatic flora and fauna based on data from standard WFD monitoring schemes. For altogether 1095 sites within a mountainous catchment, we used 12 stressor variables covering three different stressor groups: riparian land use, physical habitat quality and nutrient enrichment. Twenty-one biological metrics calculated from taxa lists of three organism groups (fish, benthic invertebrates and aquatic macrophytes) served as response variables. Stressor and response variables were subjected to Boosted Regression Tree (BRT) analysis to identify stressor hierarchy and stressor interactions and subsequently to Generalised Linear Regression Modelling (GLM) to quantify the stressors standardised effect size. Our results show that riverine habitat degradation was the dominant stressor group for the river fauna, notably the bed physical habitat structure. Overall, the explained variation in benthic invertebrate metrics was higher than it was in fish and macrophyte metrics. In particular, general integrative (aggregate) metrics such as % Ephemeroptera, Plecoptera and Trichoptera (EPT) taxa performed better than ecological traits (e.g. % feeding types). Overall, additive stressor effects dominated, while significant and meaningful stressor interactions were generally rare and weak. We concluded that given the type of stressor and ecological response variables addressed in this study, river basin managers do not need to bother much about complex stressor interactions, but can focus on the prevailing stressors according to the hierarchy identified.

Globevnik, L., Koprivsek, M., Snoj, L. (2017). Metadata to the MARS spatial database. Freshwater Metadata Journal 21, 1-7,

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Metadata to the MARS spatial database

The MARS spatial database (MARSgeoDB) supports analyses of European waters, providing common reference spatial layers and selected data on indicators of pressures, state and impacts of European waters. It is developed within the European research project MARS (Managing Aquatic ecosystems and water Resources under multiple Stress) in accordance with the WISE (Water Information System in Europe) concept. It is built on the ECRINS (European Catchments and Rivers Network System) spatial database (from the European Environment Agency), consisting of river segments, lakes and functional elementary catchments (FECs). It includes other available European spatial layers, such as River Basin Districts (RBDs), RBD sub-units, coastlines, regions, water bodies as reported under the WFD (Water Framework Directive) in 2010 and WISE SoE (State of Environment) locations. For spatial objects representing waters in the MARSgeoDB we compiled indicators of pressure, state and impact: physical-chemical indicators, ecological quality ratio, ecological status, chemical status, hydromorphological status, land use, population, nitrogen and phosphorus diffuse pollution, Eurostat agricultural data, UWWTD (Urban Waste Water Treatment Directive) point sources of organic pollution, E-PRTR (The European Pollutant Release and Transfer Register) point sources of large emissions to water, hydro-morphological changes/naturalness of rivers, meteorological and hydrological characteristics. To calculate pressures acting on selected locations on waters we derived surface water receiving areas (polygons representing catchments/hinterlands). We assigned broad ecological types to rivers (20 types) and lakes (15 types) objects in the MARSgeoDB using abiotic criteria as proposed by EEA ETC/ICM (European Topic Centre on Inland, Coastal and Marine waters) in 2015. A corresponding water body code and national ecological types were assigned as well. Spatial and associated attribute data were quality checked, unified when needed, harmonised and interlinked.

Gonzalez-Bergonzoni, I., Johansen, K. L., Mosbech, A., Landkildehus, F., Jeppesen, E., Davidson, T. A. (2017). Small birds, big effects: The little auk (Alle alle) transforms high Arctic ecosystems. Proceedings of the Royal Society B: Biological Sciences 284(1849), 20162572,

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Small birds, big effects: The little auk (Alle alle) transforms high Arctic ecosystems

In some arctic areas, marine-derived nutrients (MDN) resulting from fish migrations fuel freshwater and terrestrial ecosystems, increasing primary production and biodiversity. Less is known, however, about the role of seabird-MDN in shaping ecosystems. Here, we examine how the most abundant seabird in the North Atlantic, the little auk (Alle alle), alters freshwater and terrestrial ecosystems around the North Water Polynya (NOW) in Greenland. We compare stable isotope ratios (δ15N and δ13C) of freshwater and terrestrial biota, terrestrial vegetation indices and physical–chemical properties, productivity and community structure of fresh waters in catchments with and without little auk colonies. The presence of colonies profoundly alters freshwater and terrestrial ecosystems by providing nutrients and massively enhancing primary production. Based on elevated δ15N in MDN, we estimate that MDN fuels more than 85% of terrestrial and aquatic biomass in bird influenced systems. Furthermore, by using different proxies of bird impact (colony distance, algal δ15N) it is possible to identify a gradient in ecosystem response to increasing bird impact. Little auk impact acidifies the freshwater systems, reducing taxonomic richness of macroinvertebrates and truncating food webs. These results demonstrate that the little auk acts as an ecosystem engineer, transforming ecosystems across a vast region of Northwest Greenland.

Göthe, E., Baattrup-Pedersen, A., Wiberg-Larsen, P., Graeber, D., Kristensen, E. A., Friberg, N. (2017). Environmental and spatial controls of taxonomic versus trait composition of stream biota. Freshwater Biology 62(2), 397-413,

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Environmental and spatial controls of taxonomic versus trait composition of stream biota

The spatial organisation of biotic communities derives from factors operating at a wide range of spatial and temporal scales. Despite strong scientific evidence of prevalent spatial control of community composition in freshwater ecosystems, local environmental factors are often considered as the main drivers of community change. Furthermore, taxonomic approaches are most frequently used, and few studies have compared the relative importance of local and regional control of trait versus the taxonomic composition in stream ecosystems. Using a spatially dense data set covering all stream sizes in a lowland European region of c. 42 000 km2 and three organism groups (macrophytes, macroinvertebrates and fishes), we compared the relative importance of spatial and environmental determinants of species and trait composition in the study streams, classified into headwaters (stream order 1–2) and downstream sites (stream order >2). We hypothesised that (i) there is a higher correspondence between environmental conditions and trait composition than with species composition, (ii) dispersal limitation (pure spatial structuring) is greater in headwaters than in downstream sites and (iii) dispersal limitation (pure spatial structuring) is weakest for macroinvertebrates, intermediate for macrophytes and strongest for fishes. The most consistent pattern across organisms and stream order groups was a higher correspondence between environmental variation and trait composition as well as a higher number of environmental variables significantly related to trait composition than with species composition (hypothesis 1). Spatial structuring peaked in headwater macrophyte communities and downstream fish communities (hypotheses 2 & 3) – a pattern that was amplified when separate analyses of traits describing species dispersal potential were undertaken. Our study highlights the potential of traits to capture multiple environmental changes in stream ecosystems and illustrates how organism-specific and highly context-dependent patterns in community organisation can emerge as a consequence of interactions between habitat connectivity (i.e. top versus lower parts of the stream network) and organism dispersal potential.

Graeber, D., Jensen, T. M., Rasmussen, J., Riis, T., Wiberg-Larsen, P., Baattrup-Pedersen, A. (2017). Multiple stress response of lowland stream benthic macroinvertebrates is dependent on habitat type. Science of the Total Environment 599, 1517-1523,

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Multiple stress response of lowland stream benthic macroinvertebrates depends on habitat type

Worldwide, lowland stream ecosystems are exposed to multiple anthropogenic stress due to the combination of water scarcity, eutrophication, and fine sedimentation. The understanding of the effects of such multiple stress on stream benthic macroinvertebrates has been growing in recent years. However, the interdependence of multiple stress and stream habitat characteristics has received little attention, although single stressor studies indicate that habitat characteristics may be decisive in shaping the macroinvertebrate response. We conducted an experiment in large outdoor flumes to assess the effects of low flow, fine sedimentation, and nutrient enrichment on the structure of the benthic macroinvertebrate community in riffle and run habitats of lowland streams. For most taxa, we found a negative effect of low flow on macroinvertebrate abundance in the riffle habitat, an effect which was mitigated by fine sedimentation for overall community composition and the dominant shredder species (Gammarus pulex) and by nutrient enrichment for the dominant grazer species (Baetis rhodani). In contrast, fine sediment in combination with low flow rapidly affected macroinvertebrate composition in the run habitat, with decreasing abundances of many species. We conclude that the effects of typical multiple stressor scenarios on lowland stream benthic macroinvertebrates are highly dependent on habitat conditions and that high habitat diversity needs to be given priority by stream managers to maximize the resilience of stream macroinvertebrate communities to multiple stress.

Grizzetti, B., Pistocchi, A., Liquete, C., Udias, A., Bouraoui, F., van de Bund, W. (2017). Human pressures and ecological status of European rivers. Scientific Reports 7(1), 205,

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Human pressures and ecological status of European rivers

Humans have increased the discharge of pollution, altered water flow regime and modified the morphology of rivers. All these actions have resulted in multiple pressures on freshwater ecosystems, undermining their biodiversity and ecological functioning. The European Union has adopted an ambitious water policy to reduce pressures and achieve a good ecological status for all water bodies. However, assessing multiple pressures on aquatic ecosystems and understanding their combined impact on the ecological status is challenging, especially at the large scale, though crucial to the planning of effective policies. Here, for the first time, we quantify multiple human pressures and their relationship with the ecological status for all European rivers. We considered ecological data collected across Europe and pressures assessed by pan-European models, including pollution, hydrological and hydromorphological alterations. We estimated that in one third of EU’s territory rivers are in good ecological status. We found that better ecological status is associated with the presence of natural areas in floodplains, while urbanisation and nutrient pollution are important predictors of ecological degradation. We explored scenarios of improvement of rivers ecological status for Europe. Our results strengthen the need to halt urban land take, curb nitrogen pollution and maintain and restore nature along rivers.

Hanganu, J., Constantinescu, A., Doroftei, M., Tošić, K. (2017). Investigation of the effects of multiple pressures in the lower Danube basin. Water and Environment Journal, online,

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Investigation of the effects of multiple pressures in the lower Danube basin: Effects of multiple pressures in the lower Danube basin

The main scope of the paper is the implementation of future scenarios in the Lower Danube basin, supporting the development of River Basin Management Plans and future environmental policies. The 800 000 km² Danube River Basin extends across 19 countries, of which 14 are contracting parties of the ICPDR. The Lower Danube is occupied mostly by Romania, including the Danube Delta. The Danube Delta hydrological complex covers 5800 km² comprising over 300 lakes, three main branches and a vast network of channels. Changes impacting significant ecosystem services such as recreation, fishing, flows, ecotourism, flood regulation, navigation and reed/wood resources have to be evaluated and the quantification of those elements must be performed in order to incorporate them into the modelling process.

Hao, B., Wu, H., Cao, Y., Xing, W., Jeppesen, E., Li, W. (2017). Comparison of periphyton communities on natural and artificial macrophytes with contrasting morphological structures. Freshwater Biology 62, 1783-1793,

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Comparison of periphyton communities on natural and artificial macrophytes with contrasting morphological structures

It remains an open question whether or not artificial macrophytes are good alternatives to natural macrophytes in studies of periphyton abundance and composition in lakes. Here, a mesocosm experiment was conducted in winter (when plant growth is low) to compare simultaneously the periphyton community on three submerged macrophytes (Potamogeton lucens, Vallisneria sp. and Cabomba caroliniana) with contrasting leaf structural complexities (leaf fractal dimension = 1.12, 1.17 and 1.37, respectively) and on three types of artificial macrophytes with similar morphologies as the natural plants. We also compared intertreatment differences in phytoplankton sampled from mesocosms. Both for natural and artificial macrophytes, the periphyton chlorophyll a (Chl-a) was positively associated with leaf fractal dimension. Although the morphological structure of natural and artificial plants and the physicochemical characteristics of the water were similar, the periphyton community differed between natural and artificial macrophytes, with the difference being dependent on the leaf structural complexity of the macrophytes. For leaves with a simple structural complexity, the abundance and composition of periphyton on natural and artificial plants were not statistically different. In addition, periphyton Chl-a, density and biovolume were higher on the adaxial side than on the abaxial side of natural P. lucens leaves, but no differences were found between sides of the artificial leaves. For leaves with a medium structural complexity, the abundance of periphyton was lower on the natural than artificial plants, and the proportion of diatoms to the total community differed. For leaves with a high structural complexity, periphyton Chl-a of the artificial plants was notably higher than on the natural plants, while no significant differences were found for periphyton density, biovolume, and the proportion of diatoms and green algae. Permutational multivariate analysis of periphyton genus composition confirmed that periphyton composition on the artificial plants (medium and high leaf structural complexities) was different overall from that on the natural plants. Phytoplankton Chl-a, density, biovolume, and diversity did not show any pronounced differences among treatments. Our results suggest that artificial macrophytes cannot fully substitute for natural plants even when they are morphologically similar. Artificial macrophytes should therefore be used with caution when investigating the periphyton community on macrophytes.

He, H., Hu, E., Yu, J., Luo, X., Li, K., Liu, Z., Jeppesen, E. (2017). Does turbidity induced by crucian carp (Carassius auratus) limit phytoplankton growth? A mesocosm study. Environmental Science and Pollution Research 24(5), 5012-5018,

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Does turbidity induced by crucian carp (Carassius auratus) limit phytoplankton growth? A mesocosm study

It is well established that benthivorous fish in shallow lakes can create turbid conditions that influence phytoplankton growth both positively, as a result of elevated nutrient concentration in the water column, and negatively, due to increased attenuation of light. The net effect depends upon the degree of turbidity induced by the benthivores. Stocked Carassius carassius dominate the benthivorous fish fauna in many nutrient-rich Chinese subtropical and tropical shallow lakes, but the role of the species as a potential limiting factor in phytoplankton growth is ambiguous. Clarification of this relationship will help determine the management strategy and cost of restoring eutrophic lakes in China and elsewhere. Our outdoor mesocosm experiment simulating the effect of high density of crucian carp on phytoplankton growth and community structure in eutrophic shallow lakes suggests that stocking with this species causes resuspension of sediment, thereby increasing light attenuation and elevating nutrient concentrations. However, the effect of light attenuation was insufficient to offset the impact of nutrient enhancement on phytoplankton growth, and significant increases in both phytoplankton biomass and chlorophyll a concentrations were recorded. Crucian carp stocking favored the dominance of diatoms and led to lower percentages (but not biomass) of buoyant cyanobacteria. The dominance of diatoms may be attributed to a competitive advantage of algal cells with high sedimentation velocity in an environment subjected to frequent crucian carp-induced resuspension and entrainment of benthic algae caused by the fish foraging activities. Our study demonstrates that turbidity induced by stocked crucian carp does not limit phytoplankton growth in eutrophic waters. Thus, removal of this species (and presumably other similar taxa) from subtropical or tropical shallow lakes, or suspension of aquaculture, is unlikely to boost phytoplankton growth, despite the resulting improvements in light availability.

He, H., Luo, X., Jin, H., Gu, J., Jeppesen, E., Liu, Z., Li, K. (2017). Effects of exposed artificial substrata on the competition between phytoplankton and benthic algae: Implications for shallow lake restoration. Water 9, 24,

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Effects of exposed artificial substrata on the competition between phytoplankton and benthic algae: Implications for shallow lake restoration

Phytoplankton and benthic algae coexist in shallow lakes and the outcome of the competition between these two photoautotrophs can markedly influence water clarity. It is well established that exposed artificial substrate in eutrophic waters can remove nutrients and fine particles from the water column via the attached periphyton canopy. However, the effects of the introduction of artificial substrate on the competition between planktonic and benthic primary producers remain to be elucidated. We conducted a short-term outdoor mesocosm experiment to test the hypothesis that the nutrient and light changes induced by exposed artificial substrate (polythene nets) would benefit the benthic algae. Artificial substrate significantly reduced total nitrogen and phosphorus concentrations and water clarity improved, the latter due to the substrate-induced reduction of both organic and inorganic suspended solids. Consequently, as judged from changes in chlorophyll a (Chl-a) concentrations in water and sediment, respectively, exposed artificial substrate significantly reduced the phytoplankton biomass, while benthic algae biomass increased. Our results thus indicate that exposed artificial substrate may be used as a tool to re-establish benthic primary production in eutrophic shallow lakes after an external nutrient loading reduction, paving the way for a benthic-or a macrophyte-dominated system. Longer term and larger scale experiments are, however, needed before any firm conclusions can be drawn on this.

Hermoso, V., Filipe, A. F., Segurado, P., Beja, P. (2017). Freshwater conservation in a fragmented world: Dealing with barriers in a systematic planning framework. Aquatic Conservation: Marine and Freshwater Ecosystems 28,17-25,

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Freshwater conservation in a fragmented world: Dealing with barriers in a systematic planning framework

1. Disruption of longitudinal connectivity poses one of the most important threats to the persistence of freshwater biodiversity worldwide. Longitudinal connectivity plays a key role by facilitating ecological processes, such as migrations or transfer of energy along river networks. For this reason, effective conservation of freshwater biodiversity is highly dependent on our capacity to maintain all processes associated with connectivity. Freshwater protected areas are commonly affected by disruptions of connectivity due to anthropogenic activities and recent approaches to addressing connectivity when identifying priority areas have overlooked the limitations that human perturbations pose to connectivity. 2. Here, a novel approach is presented to address this issue by accounting for the spatial distribution of barriers in Marxan, a commonly used tool for conservation planning. This approach is first tested on a simulated example and then applied to the identification of priority areas for the conservation of freshwater vertebrates in the Iberian Peninsula (Spain and Portugal). 3. When using this new approach, the number of disrupted connections within priority areas can be significantly reduced at no additional cost in terms of area needed, which would help maintain connectivity among populations of species with low-medium migratory needs. 4. Given the widespread occurrence of barriers in the study region, the improvement in connectivity within priority areas also resulted in the selection of river reaches closer to the headwaters and the river mouth. Focusing on both extremes of the longitudinal gradient might compromise the effectiveness of conservation efforts for long migratory species, such as the European eel. This inevitably means that additional management measures, like barrier removal or construction of fish ladders, would be necessary to ensure these migratory species may complete their life cycles. 5. The method demonstrated here could be applied to other regions where connectivity is compromised.

Iglesias, C., Meerhoff, M., Johanson, L. S., Gonzalez-Bergonzoni, I., Mazzeo, N., Pacheco, J. P., Teixeira de Mello, F., Goyenola, G., Lauridsen, T., Sondergard, M., Davidson, T., Jeppesen, E. (2016). Stable isotopes analysis confirms substantial differences between subtropical and temperate shallow lakes food webs. Hydrobiologia 784, 111-123,

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Stable isotope analysis confirms substantial differences between subtropical and temperate shallow lake food webs

Differences in trophic web structure in otherwise similar ecosystems as a consequence of direct or indirect effects of ambient temperature differences can lead to changes in ecosystem functioning. Based on nitrogen and carbon stable isotope analysis, we compared the food-web structure in a series of subtropical (Uruguay, 30–35°S) and temperate (Denmark, 55–57°N) shallow lakes. The food-web length was on average one trophic position shorter in the subtropical shallow lakes compared with their temperate counterparts. This may reflect the fact that the large majority of subtropical fish species are omnivores (i.e., feed on more than one trophic level) and have a strong degree of feeding niche overlap. The shapes of the food webs of the subtropical lakes (truncated and trapezoidal) suggest that they are fuelled by a combination of different energy pathways. In contrast, temperate lake food webs tended to be more triangular, likely as a result of more simple pathways with a top predator integrating different carbon sources. The effects of such differences on ecosystem functioning and stability, and the connection with ambient temperature as a major underlying factor, are, however, still incipiently known.

Jeppesen, E., Lauridsen, T. L., Christoffersen, K. S., Landkildehus, F., Geertz-Hansen, P., Amsinck, S. L., Søndergaard, M., Davidson, T. A., Rigét, F. (2017). The structuring role of fish in Greenland lakes: An overview based on contemporary and paleoecological studies of 87 lakes from the low and the high Arctic. Hydrobiologia 800, 99-113,

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The structuring role of fish in Greenland lakes: an overview based on contemporary and paleoecological studies of 87 lakes from the low and the high Arctic

Lakes in Greenland are species-poor ecosystems and many are fishless. We studied the structuring role of fish in lakes in high- and low-Arctic Greenland. Major differences were observed in the trophic structure of the 87 lakes studied. Pelagic zooplankton biomass was on average 3–4-fold higher in the fishless lakes and dominated by large-bodied taxa such as Daphnia, the phyllopod Branchinecta and the tadpole shrimp Lepidurus. In contrast, small-bodied crustaceans dominated the lakes with fish. Analysis of microcrustacean remains in the surface sediment and contemporary benthic invertebrates also showed a marked influence of fish on community structure and the size of the taxa present. The cascading effect of fish on the microbial communities was modest, and no differences were observed for chlorophyll a. The cascading effect of fish on invertebrates depended, however, on the species present, being largest between fishless lakes and lakes hosting only sticklebacks (Gasterosteus aculeatus), while lakes with both Arctic charr (Salvelinus arcticus) and stickleback revealed a more modest response, indicating that presence of charr modulates the predation effect of sticklebacks. It is predicted that more lakes in Greenland will be colonised by fish in a future warmer climate, and this will substantially alter these vulnerable ecosystems.

Jeppesen, E., Søndergaard, M. (2017). Lake Restoration and Management in a Climate Change Perspective: An introduction. Water 9, 122,

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Lake Restoration and Management in a Climate Change Perspective: An introduction

Lakes all around the globe are under severe pressure due to an increasing anthropogenic impact from a growing population in a more developed world. Accordingly, today, many lakes are highly eutrophic and suffer from severe blooms of often toxic cyanobacteria and may become even more eutrophic in the future unless strong lake management actions are taken. Recent research has further shown that global warming and subsequent changes in water use will further exacerbate the eutrophication process in lakes. There is therefore a growing demand for lake restoration and insight into sustainable lake management. The measures to be taken, however, depend on the climate and other local conditions. This special issue addresses lake restoration and management with special emphasis on the restoration of eutrophicated lakes within a climate change perspective. The papers included collectively highlight that the ongoing climate change affects lake water quality by (1) changes in external and internal nutrient loading; (2) higher frequency of extreme events (such as hurricanes); (3) temperature‐induced changes in biota, biotic interactions; and (4) water level. Lower nutrient loading is therefore needed in a future warmer world to achieve the same ecological state as today. Several papers discuss lake restoration methods within a climate change perspective and show practical results, notably of various attempts of biomanipulation. Finally, some papers discuss the effects of other anthropogenic stressors and their interaction with climate.

Lemm, J. U., Feld, C. K. (2017). Identification and interaction of multiple stressors in central European lowland rivers. Science of the Total Environment 603-604, 148-154,

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Identification and interaction of multiple stressors in central European lowland rivers

Interactions of multiple stressors in lotic systems have received growing interest and have been analysed in a growing number of studies using experiment and survey data. In this study, we present a protocol to identify, display and analyse stressors of rivers and their interactions (additive, synergistic or antagonistic). We used a dataset of 125 samples of central European lowland rivers comprising hydromorphological, physico-chemical and land use stressor and pressure variables as well as benthic macroinvertebrate traits as biological response variables. To identify and visualise multiple stressor combinations jointly operating in the data set, we applied social network analysis. The main co-occurring stressor combination was fine sediment accumulation (hydromorphological stress) and enhanced phosphorus concentration (nutrient stress). Agricultural (cropland) and urban land use were identified as the main large scale environmental pressures. Stressor interactions were analysed using generalised linear regression modelling (GLM) including pairwise interaction terms. Altogether, 14 macroinvertebrate response variables were tested on six stressor combinations and revealed predominantly additive effects (80% of all significant models with absolute standardised effect sizes > 0.1). Significant antagonistic and synergistic interactions occurred in almost 20% of the models. Fine sediment stress was more influential and frequent than nutrient stress. The methodology presented here is standardisable and thus could help inform practitioners in aquatic ecosystem monitoring about prominent combinations of multiple stressors and their interactions. Yet, further understanding of the mechanisms behind the biological responses is required to be able to derive appropriate guidance for management. This applies to rather complex stressors and pressures, such as land use, for which more detailed data (e.g. nutrient concentrations, fine sediment entry, pesticide pollution) is often missing.

Lin, Q., Xu, L., Hou, J., Liu, Z., Jeppesen, E., Han, B.-P. (2017). Responses of trophic structure and zooplankton community to salinity and temperature in Tibetan lakes: Implication for the effect of climate warming. Water Research 124, 618-629,

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Responses of trophic structure and zooplankton community to salinity and temperature in Tibetan lakes: Implication for the effect of climate warming

Warming has pronounced effects on lake ecosystems, either directly by increased temperatures or indirectly by a change in salinity. We investigated the current status of zooplankton communities and trophic structure in 45 Tibetan lakes along a 2300 m altitude and a 76 g/l salinity gradient. Freshwater to hyposaline lakes mainly had three trophic levels: phytoplankton, small zooplankton and fish/Gammarus, while mesosaline to hypersaline lakes only had two: phytoplankton and large zooplankton. Zooplankton species richness declined significantly with salinity, but did not relate with temperature. Furthermore, the decline in species richness with salinity in lakes with two trophic levels was much less abrupt than in lakes with three trophic levels. The structural variation of the zooplankton community depended on the length of the food chain, and was significantly explained by salinity as the critical environmental variable. The zooplankton community shifted from dominance of copepods and small cladoceran species in the lakes with low salinity and three trophic levels to large saline filter-feeding phyllopod species in those lakes with high salinity and two trophic levels. The zooplankton to phytoplankton biomass ratio was positively related with temperature in two-trophic-level systems and vice versa in three-trophic-level systems. As the Tibetan Plateau is warming about three times faster than the global average, our results imply that warming could have a considerable impact on the structure and function of Tibetan lake ecosystems, either via indirect effects of salinization/desalinization on species richness, composition and trophic structure or through direct effects of water temperature on trophic interactions.

Liu, H., Cao, Y., Li, W., Zhang, Z., Jeppesen, E., Wang, W. (2017). The effects of cadmium pulse dosing on physiological traits and growth of the submerged macrophyte Vallisneria spinulosa and phytoplankton biomass: A mesocosm study. Environmental Science and Pollution Research 24, 15308-15314,

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The effects of cadmium pulse dosing on physiological traits and growth of the submerged macrophyte Vallisneria spinulosa and phytoplankton biomass: a mesocosm study

Pulse inputs of heavy metals are expected to increase with a higher frequency of extreme climate events (heavy rain), leading to stronger erosion of contaminated and fertilized farmland soils to freshwaters, with potentially adverse effects on lake ecosystems. We conducted a 5-month mesocosm study to elucidate the responses of the submerged macrophyte Vallisneria spinulosa and phytoplankton to four different doses of cadmium (Cd): 0 (control), 0.05, 0.5, and 5 g m(-2) (CK, I, II, and III, respectively) under mesotrophic conditions. We found that total phosphorus concentrations were larger in the three Cd pulse treatments, whereas total nitrogen concentrations did not differ among the four treatments. The contents of chlorophyll a and soluble sugar in macrophyte leaves decreased in III, and total biomass, ramet number, plant height, and total stolon length of macrophytes were lower in both II and III. In contrast, abundances of the three main phytoplankton taxa-Cyanophyta, Chlorophyta, and Bacillariophyta-did not differ among treatments. Total phytoplankton biomass was, however, marginally lower in CK than in the Cd treatments. We conclude that exposure to strong Cd pulses led to significantly reduced growth of macrophytes, while no obvious effect appeared for phytoplankton.

Melcher, A. H., Bakken, T. H., Friedrich, T., Greimel, F., Humer, N., Schmutz, S., Zeiringer, B., Webb, J. A. (2017). Drawing together multiple lines of evidence from assessment studies of hydropeaking pressures in impacted rivers. Freshwater Science 36 (1), 220-230,

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Drawing together multiple lines of evidence from assessment studies of hydropeaking pressures in impacted rivers

Hydropeaking has negative effects on aquatic biota, but the causal relationships have not been studied extensively, especially when hydropeaking occurs in combination with other environmental pressures. The available evidence comes mainly from case studies demonstrating river-specific effects of hydropeaking that result in modified microhabitat conditions and lead to declines in fish populations. We used multiple lines of evidence to attempt to strengthen the evidence base for models of ecological response to flow alteration from hydropeaking. First, we synthesized evidence of ecological responses from relevant studies published in the scientific literature. We found considerable evidence of the ecological effects of hydropeaking, but many causal pathways are poorly understood, and we found very little research on the interactive effects of hydropeaking and other pressures. As a 2nd line of evidence, we used results from analyses of large-scale data sets. These results demonstrated the extent to which hydropeaking occurs with other pressures, but did not elucidate individual or interactive effects further. Thus, the multiple lines of evidence complemented each other, but the main result was to identify knowledge gaps regarding hydropeaking and a consequent pressing need for novel approaches, new questions, and new ways of thinking that can fill them.

Molina-Navarro, E., Andersen, H. E., Nielsen, A., Thodsen, H., Trolle, D. (2017). The impact of the objective function in multi-site and multi-variable calibration of the SWAT model. Environmental Modelling and Software 93, 255-267,

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The impact of the objective function in multi-site and multi-variable calibration of the SWAT model

Automatic calibration of complex hydro-ecological models is an increasingly important issue which involves making decisions. One of the most relevant is the choice of the objective function, but its effects have been scarcely studied in complex models. We have used the SWAT model to assess the impact of the objective function for a multi-site (4 stations) and multi-variable (OrgP, OrgN, NO3−, PO43−) calibration of the Odense catchment (Denmark). Six calibration schemes were tested, varying the objective function and the nutrient fractions targeted. The best performance metrics (R2, NSE, PBIAS) were obtained when using NSE as objective function and targeting N-fractions and P-fractions separately. The scheme was validated in another SWAT set-up in northern Denmark. Although NSE is often questioned, we found it as an adequate objective function when addressing a multi-site and multi-variable calibration. Our findings may serve as guideline for hydro-ecological modellers, being useful to achieve watershed management goals.

Neif, É. M., Graeber, D., Rodrigues, L., Rosenhøj-Leth, S., Jensen, T. M., Wiberg-Larsen, P., Landkildehus, F., Riis, T., Baattrup-Pedersen, A. (2017). Responses of benthic algal communities and their traits to experimental changes in fine sediments, nutrients and flow. Freshwater Biology 62, 1539-1550,

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Responses of benthic algal communities and their traits to experimental changes in fine sediments, nutrients and flow

1. Lowland stream ecosystems are subjected to multiple anthropogenic stressors, usually nutrient enrichment in combination with sedimentation of fine particles and low flow periods in summer. Here, we investigated the temporal development of the benthic algae community in response to these three stressors and linkages to the trait characteristics of the community to explore the mechanisms responsible for stress-induced community changes. 2. We investigated the response of benthic algae species composition, traits (life forms, cell size categories), biovolume and chlorophyll a (Chl-a) concentration to low flow in combination with nutrient enrichment and fine sedimentation in twelve large outdoor stream flumes (12 m long) resembling small streams in size and habitat characteristics. The experiment consisted of two phases: a normalflow phase followed by a low-flow phase (90% current velocity reduction), each spanning 4 weeks. We applied a eutrophication scenario (mean increases of 1.14–5.48 mg N/L and 0.01–0.06 mg P/L in the flumes for dissolved inorganic nitrogen and phosphate respectively) throughout the experiment. Under low flow, we supplemented this with a fine sedimentation scenario (>90% stream bed cover). We took samples once in the normal-flow phase and every week during the low-flow phase. 3. We observed strong responses in the benthic algae community to sudden changes in low flow and fine sedimentation, mediating rapid species turnover with a decreased algal biovolume and increased abundance of large, motile species. However, we did not observe any pronounced responses to nutrient enrichment. In contrast to the observations for other variables, we found a continuous increase in Chl-a concentration during low flow. This was likely due to continuous fine sedimentation during this phase, reducing light availability which probably resulted in an increase of cell-level Chl-a concentration in response to light limitation and lower rates of light-induced Chl-a degradation. 4. The rapid response of the benthic algal community to the applied stressors suggests that even short periods of major stressor exposure may significantly affect benthic algae in lowland systems. We suggest that short-term stress events may have cascading effects on several important ecosystem processes given the importance of benthic algae for the productivity of these systems.

Nõges, T., Anneville, O., Guillard, J., Haberman, J., Järvalt, A., Manca, M., Morabito, G., Rogora, M., Thackeray, S. J., Volta, P., Winfield, I. J., Nõges, P. (2017). Fisheries impacts on lake ecosystem structure in the context of a changing climate and trophic state. Journal of Limnology 77, 46-61,

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Fisheries impacts on lake ecosystem structure in the context of a changing climate and trophic state

Through cascading effects within lake food webs, commercial and recreational fisheries may indirectly affect the abundances of organisms at lower trophic levels, such as phytoplankton, even if they are not directly consumed. So far, interactive effects of fisheries, changing trophic state and climate upon lake ecosystems have been largely overlooked. Here we analyse case studies from five European lake basins of differing trophic states (Lake Võrtsjärv, two basins of Windermere, Lake Geneva and Lake Maggiore) with long-term limnological and fisheries data. Decreasing phosphorus concentrations (re-oligotrophication) and increasing water temperatures have been reported in all five lake basins, while phytoplankton concentration has decreased only slightly or even increased in some cases. To examine possible ecosystem-scale effects of fisheries, we analysed correlations between fish and fisheries data, and other food web components and environmental factors. Re-oligotrophication over different ranges of the trophic scale induced different fish responsesIn the deeper lakes Geneva and Maggiore, we found a stronger link between phytoplankton and planktivorous fish and thus a more important cascading top-down effect than in other lakes. This connection makes careful ecosystem-based fisheries management extremely important for maintaining high water quality in such systems. We also demonstrated that increasing water temperature might favour piscivores at low phosphorus loading, but suppresses them at high phosphorus loading and might thus either enhance or diminish the cascading top-down control over phytoplankton with strong implications for water quality.

Olsen, S., Cao, Y., Gutierrez, M. F., Brucet, S., Landkildehus, F., Lauridsen, T. L., Davidson, T. A., Søndergaard, M., Jeppesen, E., Risgaard-Petersen, N. (2017). Effect of a nitrogen pulse on ecosystem N processing at different temperatures: A mesocosm experiment with 15NO3- addition. Freshwater Biology 62, 1232-1243,

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Effect of a nitrogen pulse on ecosystem N processing at different temperatures: A mesocosm experiment with 15NO3- addition

1. Shallow lakes may play an important role for the nitrogen (N) balance in drainage basins by processing, transferring and retaining N inputs. An increase in the frequency of storm-induced short-term N pulses and increased water temperatures are both likely outcomes of climate change, potentially affecting the N processing in lakes. 2. An experiment with a K 15 NO 3 À pulse addition (increase in NO 3 À concentration from c. 0.1 to 2 mg/L) was carried out in 12 mesocosms with relatively low (applies to Danish lakes) total N (TN) and total phosphorus (TP) concentrations (c. 0.3 mg N L -1 and 0.04 mg P L -1) to assess the effects of an N pulse on N processing and storage in shallow lake ecosystems. The mesocosms have a hydraulic retention time of approximately two and a half months, and at the time of the experiment, they had been adapted to contrasting temperatures for a period of 10 years: ambient, T3 (heat-ing according to the Intergovernmental Panel on Climate Change 2007 A2 scenario, +3.7–4.5°C, depending on season) and T5 (heating with A2 + 50%, +4.9–6.6°C). 3. Macrophytes and filamentous algae retained up to 40% and 30% of the added 15 N, respectively, reflecting their high biomass in the mesocosms. Macrophytes and filamentous algae constituted between 70% and 80% of the biomass of all primary producers during the experiment in the T3 and ambient treatments and between 20% and 40% in T5. By comparison, less than 1% of the added 15 N diffused to the sediment and less than 5% was lost to the atmosphere as N 2 gas. Snails represented the long-term storage of 15 N, retaining up to 6% of the tracer and with detectable enrichment 100 days after tracer addition. 4. We found no significant differences among the temperature treatments in the 15 N turnover after pulse dosing. However, a larger percentage of 15 N was stored in macrophytes in the ambient and T3 mesocosms, reflecting higher biomasses than in T5 where filamentous algae were more abundant. Macrophytes and filamentous algae rather than temperature were therefore key controllers of N processing during the summer N pulse in these shallow, relatively low TP lakes.

Poikane, S., Ritterbusch, D., Argillier, C., Bialokoz, W., Blabolil, P., Breine, J., Jaarsma, N., Krause, T., Kubecka, J., Lauridsen, T., Nõges, P., Peirson, G., Virbickas, T. (2017). Response of fish communities to multiple pressures: Development of a total anthropogenic pressure intensity index. Science of the Total Environment 586, 502-511,

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Response of fish communities to multiple pressures: Development of a total anthropogenic pressure intensity index

Lakes in Europe are subject to multiple anthropogenic pressures, such as eutrophication, habitat degradation and introduction of alien species, which are frequently inter-related. Therefore, effective assessment methods addressing multiple pressures are needed. In addition, these systems have to be harmonised (i.e. intercalibrated) to achieve common management objectives across Europe. Assessments of fish communities inform environmental policies on ecological conditions integrating the impacts of multiple pressures. However, the challenge is to ensure consistency in ecological assessments through time, across ecosystem types and across jurisdictional boundaries. To overcome the serious comparability issues between national assessment systems in Europe, a total anthropogenic pressure intensity (TAPI) index was developed as a weighted combination of the most common pressures in European lakes that is validated against 10 national fish-based water quality assessment systems using data from 556 lakes. Multi-pressure indices showed significantly higher correlations with fish indices than single-pressure indices. The best-performing index combines eutrophication, hydromorphological alterations and human use intensity of lakes. For specific lake types also biological pressures may constitute an important additional pressure. The best-performing index showed a strong correlation with eight national fish-based assessment systems. This index can be used in lake management for assessing total anthropogenic pressure on lake ecosystems and creates a benchmark for comparison of fish assessments independent of fish community composition, size structure and fishing-gear. We argue that fish-based multiple-pressure assessment tools should be seen as complementary to single-pressure tools offering the major advantage of integrating direct and indirect effects of multiple pressures over large scales of space and time.

Ren, L., He, D., Chen, Z., Jeppesen, E., Lauridsen, T. L., Søndergaard, M., Liu, Z., Wu, Q. L. (2017). Warming and nutrient enrichment in combination increase stochasticity and beta diversity of bacterioplankton assemblages across freshwater mesocosms. The ISME Journal 11(3), 613-625,

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Warming and nutrient enrichment in combination increase stochasticity and beta diversity of bacterioplankton assemblages across freshwater mesocosms

The current climate warming and eutrophication are known to interactively threaten freshwater biodiversity; however, the interactive effects on lacustrine bacterioplankton diversity remain to be determined. Here, we analyzed the spring bacterioplankton community composition (BCC) in 24 outdoor, flow-through mesocosms (mimicking shallow lake environments) under 3 temperature scenarios and 2 nutrient regimes. Our results revealed that neither long-term warming (8.5 years) nor nutrient enrichment had significant effects on bacterioplankton alpha diversity, whereas long-term enhanced warming (elevated 50% above the IPCC A2 climate scenario) and nutrient enrichment in combination increased bacterioplankton beta diversity. We also found that BCC shifted significantly under enhanced warming and nutrient-enriched conditions towards decreased relative abundances of Actinobacteria, Bacteroidetes and Betaproteobacteria, whereas the percentages of Cyanobacteria, total rare phyla and unclassified phyla significantly increased. Null-model tests indicated that deterministic processes played a more important role than stochastic processes in determining BCC. However, the relative importance of stochasticity, primarily ecological drift, was enhanced and contributed to the increased beta diversity of BCC under enhanced warming and nutrient-enriched conditions. Overall, our study suggests that the synergetic effects of warming and nutrient enrichment may result in high variability in the composition of bacterioplankton communities in lacustrine water bodies.

Riis, T., Levi, P. S., Baattrup-Pedersen, A., Jeppesen, K. G., Rosenhøj Leth, S. (2017). Experimental drought changes ecosystem structure and function in a macrophyte-rich stream. Aquatic Sciences 79, 841-853,

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Experimental drought changes ecosystem structure and function in a macrophyte-rich stream

Water abstraction continues to increase worldwide, causing periods with extreme low-flow in many streams, which will likely intensify in the future due to climate change. Extreme low-flow may have major effects on in-stream habitats, organisms, and consequently ecosystem functions. We investigated the effects of a 2 months experimentally induced extreme low-flow scenario on the physical, biological, and functional characteristics in a macrophyte-rich lowland stream using a before-after, control-impact (BACI) approach. We quantified nutrient dynamics, including inorganic nitrogen and phosphorus concentrations, ammonium uptake, and whole-stream metabolism. We found a significant decline in the stream wetted habitat area, an increase in water temperature, and an increase in the accumulation of fine organic matter with reduced flow, but no significant changes in dissolved oxygen or benthic chlorophyll a concentrations. Furthermore, the relative demand and overall uptake of ammonium was lower in the low-flow reach relative to the control reach, whereas the relative demand and uptake of phosphate were higher at low-flow. Our results demonstrate that low-flow conditions cause resource limitation in stream biota most likely due to increased thickness of the diffusive boundary layers and an enhanced heterotrophic activity in the accumulated fine organic matter. Our results imply that the basal resources for productivity shift from autotrophic towards more heterotrophic resources causing a shift at higher trophic levels towards more detritivore based and less herbivore based food webs with implications for the invertebrate community composition and the distribution of functional feeding groups. Based on the strong links observed between low-flow and nutrient uptake, we suggest that functional metrics are suitable to assess the effects of low-flow conditions in small streams.

Romão, F., Quaresma, A. L., Branco, P. J., Santos, J. M., Amaral, S., Ferreira, M. T., Katopodis, C., Pinheiro, A. N. (2017). Passage performance of two cyprinids with different ecological traits in a fishway with distinct vertical slot configurations. Ecological Engineering 105, 180-188,

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Passage performance of two cyprinids with different ecological traits in a fishway with distinct vertical slot configurations

Adequately designed vertical slot fishways (VSF) mitigate the impact of anthropogenic obstructions onfish migrations. Until now, most studies on VSF were conducted focusing on high priority species, partic-ularly salmonids, while other species, such as cyprinids, have received less attention. In Mediterraneanrivers, where water availability is a problem, effective low discharge fishways are desirable. Attemptingto contribute towards filling this gap, the present study focuses on the behaviour and passage perfor-mance of two Iberian cyprinids with different ecological traits, the Iberian barbel (Luciobarbus bocagei,Steindachner, 1864) and the Southern Iberian chub (Squalius pyrenaicus, Günther, 1868). Fish passageperformance was investigated in a full-scale experimental VSF under two different slot configurations(C1 and C2), which require different discharges for equal mean water depths in the pools (Q = 110 L s−1, forC1, and Q = 81 L s−1, for C2). Results show that the chub, a small-bodied fish that utilises the upper portionof the water column, performed a higher number of upstream movements in C2, while for the barbel, alarge-bodied potamodromous bottom-oriented fish, the performance was similar in both configurations.With similar overall passage success, slot configuration C2 requires a lower discharge to operate, makingit the more cost-effective geometry, especially in regions affected by water scarcity.

Santos, J. M., Rivaes, R., Boavida, I., Branco, P. J. (2017). Structural microhabitat use by endemic cyprinids in a Mediterranean-type river: Implications for restoration practices. Aquatic Conservation: Marine and Freshwater Ecosystems 28, 01–11,

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Structural microhabitat use by endemic cyprinids in a Mediterranean-type river: Implications for restoration practices

Endemic freshwater fish from the Mediterranean region are among the most threatened species in the world owing to increasing river degradation. Because of such threats, the number of river restoration projects has greatly increased. However, they are seldom planned with consideration of the species' life history, often resulting in erroneous practices that compromise their success. This study assessed the seasonal and size-related microhabitat use by three endemic cyprinids (Iberian barbel, Luciobarbus bocagei; Iberian straight-mouth nase, Pseudochondrostoma polylepis; and calandino, Squalius alburnoides) using a modified point electrofishing procedure in a Mediterranean river. A multivariate approach was then employed to analyse both structural resource use and availability data. All species showed non-random microhabitat use. The barbel and nase shifted to faster-flowing positions (>25 cm s−1) with a coarser substratum (>150 mm particle size) during spring and to sheltered positions (50–100% instream cover) during autumn. Calandino selected more covered areas in autumn (>60% cover) and shifted to shallower positions from this season (>40 cm) to summer (

Schinegger, R., Aschauer, C., Mühlmann, H., Schmutz, S. (2017). Metadata: MARS multiple stressors and biological dataset of Drava & Mura Basins. Freshwater Metadata Journal 24, 1-8,

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Metadata: MARS multiple stressors and biological dataset of Drava & Mura Basins

This work/dataset addresses human stressors and their impacts on fishassemblages in the Austrian Drava and Mura River Basins. It supports theEU-project MARS (Managing Aquatic ecosystems and water Resourcesunder multiple Stress) by analysing single and multiple stressors,environmental effects and stressor combinations/interactions. Data sourcesare mainly shape files and MS ACCESS databases.With the help of point data on connectivity disruptions (barriers) and linedata on hydromorphological & water quality stressors (on water body level),six mainly hydromorphological stressors from the national inventoryassessment of the EU Water Framework Directive were recoded andaggregated into new variables, i.e. stressor metrics. These then werecompared with point data (fish sampling sites) and related information onfish assemblages (Fish Index Austria and related single metrics as well asthe WFD biological and total status).

Søndergaard, M., Lauridsen, T. L., Johansson, L. S., Jeppesen, E. (2017). Nitrogen or phosphorus limitation in lakes and its impact on phytoplankton biomass and submerged macrophyte covers. Hydrobiologia 795, 35-48,

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Nitrogen or phosphorus limitation in lakes and its impact on phytoplankton biomass and submerged macrophyte covers

We used data on nutrients, chlorophyll a (Chla) and submerged macrophyte cover from up to 817 Danish lakes to elucidate seasonal variations in nitrogen (N) and phosphorus (P) concentrations and to study the impact of N or its role in combination with P. In both deep and shallow lakes, we found marked seasonality in the ratio between total N and total P (TN:TP) and in the inorganic concentrations of nitrogen (DIN), indicating that N more easily becomes a limiting nutrient as summer proceeds. TN:TP reached its lowest values of <7 (by mass) in August in 25% of the shallow lakes. Chla generally related more strongly to TP than to TN, but at high TP concentrations TN explained more of the variability in Chla than TP. Macrophyte cover tended to decrease at increasing TN when TP was between 0.1 and 0.4 mg/l. At macrophyte cover above 20%, Chla was considerably lower compared with lakes with low macrophyte cover. We conclude that P is of key importance for the ecological quality of Danish lakes but that increased N concentrations, particularly in shallow lakes with moderate to high TP, may have significantly adverse effects on lake water quality and ecological status in summer.

Søndergaard, M., Lauridsen, T. L., Johansson, L. S., Jeppesen, E. (2017). Repeated fish removal to restore lakes: Case study Lake Væng, Denmark - Two biomanipulations during 30 years of monitoring. Water 9(1), 43,

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Repeated fish removal to restore lakes: Case study Lake Væng, Denmark - Two biomanipulations during 30 years of monitoring

Biomanipulation by fish removal has been used in many shallow lakes as a method to improve lake water quality. Here, we present and analyse 30 years of chemical and biological data from the shallow and 16 ha large Lake Vaeng, Denmark, which has been biomanipulated twice with a 20-year interval by removing roach (Rutilus rutilus) and bream (Abramis brama). After both biomanipulations, Lake Vaeng shifted from a turbid, phytoplankton-dominated state to a clear, water macrophyte-dominated state. Chlorophyll a was reduced from 60–80 µg·L −1 to 10–30 µg·L −1 and the coverage of submerged macrophytes, dominated by Elodea canadensis, increased from <0.1% to 70%–80%. Mean summer total phosphorus was reduced from about 0.12 to 0.07 mg·L −1 and total nitrogen decreased from 1.0 to 0.4 mg·L −1. On a seasonal scale, phosphorus and chlorophyll concentrations changed from a summer maximum during turbid conditions to a winter maximum under clear conditions. The future of Lake Vaeng is uncertain and a relatively high phosphorus loading via the groundwater, and the accumulation of a mobile P pool in the sediment make it likely that the lake eventually will return to turbid conditions. Repeated fish removals might be a relevant management strategy to apply in shallow lakes with a relatively high external nutrient loading.

Spears, B. M., Futter, M. N., Jeppesen, E., Huser, B. J., Ives, S., Davidson, T. A., Adrian, R., Angeler, D. G., Burthe, S. J., Carvalho, L., Daunt, F., Gsell, A. S., Hessen, D. O., Janssen, A. B. G., Mackay, E. B., May, L., Moorhouse, H., Olsen, S., Søndergaard, M., Woods, H., Thackeray, S. J. (2017). Ecological resilience in lakes and the conjunction fallacy. Nature Ecology & Evolution 1, 1616-1624,

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Ecological resilience in lakes and the conjunction fallacy

There is a pressing need to apply stability and resilience theory to environmental management to restore degraded ecosystems effectively and to mitigate the effects of impending environmental change. Lakes represent excellent model case studies in this respect and have been used widely to demonstrate theories of ecological stability and resilience that are needed to underpin preventative management approaches. However, we argue that this approach is not yet fully developed because the pursuit of empirical evidence to underpin such theoretically grounded management continues in the absence of an objective probability framework. This has blurred the lines between intuitive logic (based on the elementary principles of probability) and extensional logic (based on assumption and belief) in this field.

Tavşanoğlu, U.N, Šorf, M., Stefanidis, K., Brucet, S., Agasild, H., Boho, D., Scharfenberger, U., Beklioğlu, M., Hejzlar, J., Papastergiadou, E., Adrian, R., Angeler, D., Nõges, T., Çakıroğlu, A. I., Özen, A., Drakare, S., Søndergaard, M., Jeppesen, E. (2017). Effects of nutrient and water level changes on the composition and size structure of zooplankton communities under different climatic conditions: A Pan-European mesocosm experiment. Aquatic Ecology 51, 257-273,

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Effects of nutrient and water level changes on the composition and size structure of zooplankton communities under different climatic conditions: A Pan-European mesocosm experiment

Lentic ecosystems act as sentinels of climate change, and evidence exists that their sensitivity to warming varies along a latitudinal gradient. We assessed the effects of nutrient and water level variability on zooplankton community composition, taxonomic diversity and size structure in different climate zones by running a standardised controlled 6-months (May to November) experiment in six countries along a European north–south latitudinal temperature gradient. The mesocosms were established with two different depths and nutrient levels. We took monthly zooplankton samples during the study period and pooled a subsample from each sampling to obtain one composite sample per mesocosm. We found a significant effect of temperature on the community composition and size structure of the zooplankton, whereas no effects of water depth or nutrient availability could be traced. The normalised size spectrum became flatter with increasing temperature reflecting higher zooplankton size diversity due to higher abundance of calanoid copepods, but did not differ among depths or nutrient levels. Large-bodied cladocerans such as Daphnia decreased with temperature. Taxonomic diversity was positively related to size diversity, but neither of the two diversity measures demonstrated a clear pattern along the temperature gradient nor with nutrient and water levels. However, genus richness decreased at the warm side of the temperature gradient. Our experiment generally supports recent empirically based findings that a continuing temperature increase may result in lower genus richness and lower abundance of large-sized zooplankton grazers, the latter likely resulting in reduced control of phytoplankton.

Teichert, N., Pasquaud, S., Borja, A., Chust, G., Uriarte, A., Lepage, M. (2017). Living under stressful conditions: Fish life history strategies across environmental gradients in estuaries. Estuarine, Coastal and Shelf Science 188, 18-26,

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Living under stressful conditions: Fish life history strategies across environmental gradients in estuaries

The life history strategies of fishes can be defined by specific combinations of demographic traits that influence species performances depending on environmental features. Hence, the constraints imposed by the local conditions restrict the range of successful strategies by excluding species poorly adapted. In the present study, we compared the demographic strategies of fish caught in 47 estuaries of the North East Atlantic coast, aiming to determine the specific attributes of resident species and test for changes in trait associations along the environmental gradients. Eight demographic traits were considered to project our findings within a conceptual triangular model, composed on three endpoint strategies: (i) periodic (large size, long generation time, high fecundity); (ii) opportunistic (small size, short generation time, high reproductive effort); and (iii) equilibrium (low fecundity, large egg size, parental care). We demonstrated that various life history strategies co-exist in estuaries, but equilibrium species were scarce and restricted to euhaline open-water. Resident species form a specialised assemblage adapted to high spatiotemporal variability of estuarine conditions, i.e. opportunistic attributes associated with parental care. Even with these singular attributes, our findings revealed changes in distribution of resident species across the estuarine gradients linked to their life history traits. Among other patterns, the diversity of life history strategies significantly decreased from euhaline to oligohaline areas and along gradient of human disturbances. These trends were associated with a convergence of species traits toward short generation times, suggesting that long-lived species with late maturation are more severely impacted by disturbance and environmental stress.

Teichert, N., Lepage, M., Sagouis, A., Borja, A., Chust, G., Ferreira, M. T., Pasquaud, S., Schinegger, R., Segurado, P., Argillier, C. (2017). Functional redundancy and sensitivity of fish assemblages in European rivers, lakes and estuarine ecosystems. Scientific Reports 7, 17611,

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Functional redundancy and sensitivity of fish assemblages in European rivers, lakes and estuarine ecosystems

The impact of species loss on ecosystems functioning depends on the amount of trait similarity between species, i.e. functional redundancy, but it is also influenced by the order in which species are lost. Here we investigated redundancy and sensitivity patterns across fish assemblages in lakes, rivers and estuaries. Several scenarios of species extinction were simulated to determine whether the loss of vulnerable species (with high propensity of extinction when facing threats) causes a greater functional alteration than random extinction. Our results indicate that the functional redundancy tended to increase with species richness in lakes and rivers, but not in estuaries. We demonstrated that i) in the three systems, some combinations of functional traits are supported by non-redundant species, ii) rare species in rivers and estuaries support singular functions not shared by dominant species, iii) the loss of vulnerable species can induce greater functional alteration in rivers than in lakes and estuaries. Overall, the functional structure of fish assemblages in rivers is weakly buffered against species extinction because vulnerable species support singular functions. More specifically, a hotspot of functional sensitivity was highlighted in the Iberian Peninsula, which emphasizes the usefulness of quantitative criteria to determine conservation priorities.

Tolonen, K. T., Vilmi, A., Karjalainen, S. M., Hellsten, S., Heino, J. (2017). Do different facets of littoral macroinvertebrate diversity show congruent patterns in a large lake system? Community Ecology 18(1), 109-116,

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Do different facets of littoral macroinvertebrate diversity show congruent patterns in a large lake system?

Modern biodiversity research focuses on multiple diversity facets because different indices may describe different ecological and environmental processes, as well as the effects of varied disturbances of natural and anthropogenic origins. We investigated littoral macroinvertebrate diversity in a large boreal lake system and specifically explored congruence of indices within and between the three diversity facets: species diversity, functional diversity and taxonomic distinctness. First, we found that the indices of taxonomic distinctness were the most sensitive indicators of eutrophication. Second, we observed that most correlations between the indices within the same diversity facet, and between the indices of functional and species diversity, were relatively strong. However, the indices of taxonomic distinctness (?+ and ?+) were weakly associated with other metrics of diversity, emphasising the importance of taxonomic distinctness as a complementary dimension of biodiversity. Therefore, our observations support the importance to examine multiple facets for mapping biodiversity or for assessing the effects of anthropogenic disturbances on biological communities.

Trochine, C., Brucet, S., Argillier, C., Arranz, I., Beklioğlu, M., Benejam, L., Ferreira, T., Hesthagen, T., Holmgren, K., Jeppesen, E., Kelly, F., Krause, T., Rask, M., Volta, P., Winfield, I. J., Mehner, T. (2017). Non-native Fish Occurrence and Biomass in 1943 Western Palearctic Lakes and Reservoirs and their Abiotic and Biotic Correlates. Ecosystems 21, 395-409,

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Non-native Fish Occurrence and Biomass in 1943 Western Palearctic Lakes and Reservoirs and their Abiotic and Biotic Correlates

Invasion of non-native species is considered a major threat to global biodiversity. Here we present a comprehensive overview of the occurrence, richness and biomass contribution of non-native fish species in 1943 standing water bodies from 14 countries of the Western Palearctic, based on standardised fish catches by multi-mesh gillnetting. We expected strong geographical gradients to emerge in the occurrence of non-natives. We further hypothesised that the contribution by nonnatives to the local fish community biomass was correlated with local richness and the trophic level of native and non-native species. Non-native fish species occurred in 304 of 1943 water bodies (16%). If the average number of occupied water bodies per country was weighted by number of water bodies per country, the grand mean occurrence of non-natives in Western Palearctic water bodies was 10%. Exotic (non-native to the Palearctic) and translocated (non-native only to parts of the Palearctic) species were found in 164 (8.4%) or 235 (12.1%) of the water bodies, respectively. The occurrence and local richness of non-native fish species increased with temperature, precipitation and lake area and were substantially higher in reservoirs than in natural lakes. High local biomass contributions of non-native species were strongly correlated with low richness of native species and high richness of non-native species, whereas the trophic level of the fish species had only a weak effect. Single non-native species rarely dominated community biomass, but high biomass contributions and thus strong community and ecosystem impacts can be expected if several non-native species accumulate in a water body.

Wang, H. J., Xiao, X. C., Wang, H. Z., Li, Y., Yu, Q., Liang, X. M., Feng, W. S., Shao, J. C., Rybicki, M., Jungmann, D., Jeppesen, E. (2017). Effects of high ammonia concentrations on three cyprinid fish: Acute and whole-ecosystem chronic tests. Science of the Total Environment 598, 900-909,

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Effects of high ammonia concentrations on three cyprinid fish: Acute and whole-ecosystem chronic tests

A number of studies have revealed ammonia to be toxic to aquatic organisms; however, little is known about its effects under natural conditions. To elucidate the role of ammonia, we conducted 96-h acute toxicity tests as well as a whole-ecosystem chronic toxicity test for one year in ten 600-m² ponds. Three common cyprinids, silver carp Hypophthalmichthys molitrix Val. (H.m.), bighead carp Aristichthys nobilis Richardson (A.n.), and gibel carp Carassius auratus gibelio Bloch (C.g.), were used as test organisms. The 96-h LC50 values of un-ionized ammonia (NH3) for H.m., A.n., and C.g. were 0.35, 0.33, and 0.73 mg L− 1, respectively. In the ponds, annual mean NH3 ranged between 0.01 and 0.54 mg L− 1, with 4 ponds having a NH3 higher than the LC50 of A.n. (lowest LC50 in this study). No fish were found dead in the high-nitrogen ponds, but marked histological changes were found in livers and gills. Despite these changes, the specific growth rate of H.m. and A.n. increased significantly with NH3. Our pond results suggest that fish might be more tolerant to high ammonia concentrations in natural aquatic ecosystems than under laboratory conditions. Our finding from field experiments thus suggests that the existing regulatory limits for reactive nitrogen (NH3) established from lab toxicity tests might be somewhat too high at the ecosystem conditions. Field-scale chronic toxicity tests covering full life histories of fish and other aquatic organisms are therefore encouraged in order to optimize determination of the effects of ammonia in natural environments.

Wengrat, S., Padial, A. A., Jeppesen, E., Davidson, T. A., Fontana, L., Costa-Böddeker, S., Bicudo, D. C. (2017). Paleolimnological records reveal biotic homogenization driven by eutrophication in tropical reservoirs. Journal of Paleolimnology, 01-11,

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Paleolimnological records reveal biotic homogenization driven by eutrophication in tropical reservoirs

Biodiversity changes in response to eutrophication, climate variability and species invasions. These pressures have been shown to reduce community heterogeneity at various scales; however, how productivity drives homogenization patterns in a community of primary producers, such as diatoms, has not been studied. Using a dataset with good temporal resolution, obtained from cores collected from seven tropical reservoirs, we evaluated patterns of spatial and temporal homogenization, i.e. the trends in temporal α-diversity and spatial β-diversity (change in community composition), of diatom assemblages over the past 60–100 years. The paleolimnological records allowed us to study biodiversity trends since the initial community (reservoir construction) in those systems with low anthropogenic impact and also those undergoing eutrophication. No clear trend of spatial β-diversity change over time was found when all reservoirs were analyzed together. However, when only eutrophic reservoirs were considered, a marked decrease in the spatial β-diversity occurred, suggesting that eutrophication leads to homogenization of the diatom assemblage. These findings were reinforced by the lack of change in β-diversity when the age of the reservoirs was standardized, indicating that the reservoirs’ ontogeny did not influence the spatial β-diversity trend and β-diversity did not increase even in the reservoirs with low anthropogenic impact. In addition, the results showed a decrease of α-diversity over time for almost all the eutrophic reservoirs, as well as a decrease in the total species pool for the reservoirs, although periphytic diatoms may be favored by the appearance and sometimes mass development of floating macrophytes in warm, shallow eutrophic reservoirs. This study supports the role of eutrophication as one of the main drivers of diatom assemblage homogenization in tropical reservoirs, with a significant loss of species over time.

Wright, R. F., Couture, R.-M., Christiansen, A. B., Guerrero, J.-L., Kaste, Ø., Barlaup. B. T. (2017). Effects of multiple stresses hydropower, acid deposition and climate change on water chemistry and salmon populations in the River Otra, Norway. Science of the Total Environment 574, 128-138,

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Effects of multiple stresses hydropower, acid deposition and climate change on water chemistry and salmon populations in the River Otra, Norway

Many surface waters in Europe suffer from the adverse effects of multiple stresses. The Otra River, southernmost Norway, is impacted by acid deposition, hydropower development and increasingly by climate change. The river holds a unique population of land-locked salmon and anadromous salmon in the lower reaches. Both populations have been severely affected by acidification. The decrease in acid deposition since the 1980s has led to partial recovery of both populations. Climate change with higher temperatures and altered precipitation can potentially further impact fish populations. We used a linked set of process-oriented models to simulate future climate, discharge, and water chemistry at five sub-catchments in the Otra river basin. Projections to year 2100 indicate that future climate change will give a small but measureable improvement in water quality, but that additional reductions in acid deposition are needed to promote full restoration of the fish communities. These results can help guide management decisions to sustain key salmon habitats and carry out effective long-term mitigation strategies such as liming. The Otra River is typical of many rivers in Europe in that it fails to achieve the good ecological status target of the EU Water Framework Directive. The programme of measures needed in the river basin management plan necessarily must consider the multiple stressors of acid deposition, hydropower, and climate change. This is difficult, however, as the synergistic and antagonistic effects are complex and challenging to address with modelling tools currently available.

Wu, N., Dong, X., Baattrup-Pedersen, A., Riis, T. (2017). Using river microalgae as indicators for freshwater biomonitoring: review of published research and future directions. Ecological Indicators 81, 124-131,

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Using river microalgae as indicators for freshwater biomonitoring: Review of published research and future directions

Trait-based approaches may give insights into underlying mechanisms of relationships between biological communities and environmental stressors, and are increasingly used in ecological studies, but are only very recently considered for freshwater riverine microalgae. Here, we i) review the research trend in riverine microalgae during the past 26 years in order to conduct a quantitative and qualitative analysis for global trends in the research field, ii) summarize the use of algae traits in riverine biomonitoring and iii) propose future research perspectives. The bibliometric analysis showed that the annual number of publications on microalgae increased significantly from 1991 to 2016, although their proportions to total numbers of scientific articles remained steady. The studies have become increasingly concerned on issues arisen from global environmental changes such as “eutrophication”, “pollution”, “land use”, “biomonitoring”, “biodiversity”, “functional group”, etc. The use of algae traits in biomonitoring has become popular and includes e.g. functional diversity, cell size, guild, life form, eco-morphology, spore formation as well as algal quality. Here we collate all relevant algal traits, their different categories and propose their responses to resource supply and disturbance frequency in a conceptual model, which should be validated in future studies. In order to expand the knowledge and future use of microalgae in biomonitoring research efforts should also include: i) description of relationships between algal traits and ecosystem functions (e.g., nutrient uptake, metabolism, energy transfer across the food web) and underlying mechanisms; ii) selection of robust traits reflecting and disentangling the effects of multiple stressors; iii) water resource management in an interdisciplinary manner linking risk assessment and management scenarios by an integrated modelling system using microalgae.

Xu, M., Dong, X., Yang, X., Wang, R., Zhang, K., Zhao, Y., Davidson, T. A., Jeppesen, E. (2017). Using palaeolimnological data and historical records to assess long-term dynamics of ecosystem services in typical Yangtze shallow lakes (China). Science of the Total Environment 584, 791–802,

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Using palaeolimnological data and historical records to assess long-term dynamics of ecosystem services in typical Yangtze shallow lakes (China)

Long-term patterns of lake ecosystem services (ESs) are crucial for lake management. • Both palaeolimnological and documentary records revealed ES variations in Yangtze lakes. • Provisioning and regulating services have exhibited both tradeoff and syner-gy since 1900s. Understanding the dynamics of ecosystem services (ESs) is crucial for sustainable resource management. Palaeolimnological records have a great potential to reveal long-term variations and dynamic interactions in ESs, especially supporting/regulating services, which are not easily quantified by documentary records. To elucidate the variations between eight important ESs in shallow lake ecosystems, we combined documentary records with palaeolimnological proxies (covering the past 100 years) from two typical lakes (Lakes Taibai and Zhangdu) of the Yangtze River basin. Although all supporting services and some provisioning services have increased, the regulating services of the two lakes have markedly declined, in particular since the 1950s. Human activities, including hydrological intervention, nutrient input and land-use change, were identified as the main factors behind the observed variations. Both in Lake Taibai and Zhangdu, primary production and biodiversity (supporting services) have increased (synergies), whereas climate and water purification (regulating services) have significantly decreased (tradeoffs) since the 1950s when attempts were made by the local population to reach a higher land/ fish ESs level. By considering long-term records, dynamic tradeoff and synergy relationship between various ESs relative to different types of human " modification " in a temporal perspective, we suggest valuable information can be gained in future lake management initiatives.

Yu, Q., Wang, H.-J., Wang, H.-Z., Li, Y., Liang, X.-M., Xu, C., Jeppesen, E. (2017). Does the responses of Vallisneria natans (Lour.) Hara to high nitrogen loading differ between the summer high-growth season and the low-growth season? Science of the Total Environment 601-602, 1513-1521,

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Does the responses of Vallisneria natans (Lour.) Hara to high nitrogen loading differ between the summer high-growth season and the low-growth season?

Yu et al.'s paper showed very interesting effects of high nitrogen (N) on the submerged macrophytes Vallisneria natans: active growth in the growing season enabled the macrophytes partly to overcome the ammonium stress. This result was evident in an experiment using ten pond ecosystems; however, their conclusion that shading induced by high phytoplankton biomass together with the toxicity of high ammonium contributed to the decrease of macrophytes growth was not strongly supported by the data provided in the paper. Three factors influencing how submerged macrophytes respond to high ammonium, not addressed by Yu et al.'s paper, are toxicity of ammonium/ammonia (NH4(+)/NH3), the precise extent of shading in water and species-specific characteristics of macrophytes. In conclusion, a comprehensive consideration of abiotic and biotic factors that involve in the responses of submerged macrophytes to high N is urged in future studies of the role of high N on the growth of submerged macrophytes.

Zajicek, P., Wolter, C. (2017). The gain of additional sampling methods for the fish-based assessment of large rivers. Fisheries Research 197, 15-24,

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The gain of additional sampling methods for the fish-based assessment of large rivers

Fishes serve as indicators in ecological assessments of European large rivers. Electrofishing is the standard fishing method although it is restricted to the shallow littoral shoreline. Fish occurring in the open water zone of the main channel remain consequently underestimated. Additional sampling methods that cover the midchannel of rivers could close the electrofishing gap, but strengths’, weaknesses and gains of both electrofishing and additional sampling methods for fish-based assessments of large rivers have not been contrasted yet. We analyzed a unique dataset consisting of 2693 fish samplings in European large rivers and compared electrofishing with the additional sampling methods trawling, seining, and drift-netting. We compiled fish metrics commonly used in fish-based assessments yielded by the different gears and highlight the differences in fish species, biodiversity metrics (Shannon Index, Evenness, Simpson Index), the Fish Region Index (FRI) and densities of fish in selected guilds (eurytopic, rheophilic, lithophilic, phytophilic, psammophilic, potamal) that are considered indicative for the degradation of habitats in large rivers. Electrofishing yielded overall highest numbers of species, biodiversity metrics and densities of fish guilds, except for the number of migratory and Habitat Directive species, the FRI and densities of potamal fish. The additional gears, predominantly trawling, captured additional rheophilic and lithophilic species. Trawling also assessed most migratory and Habitat Directive species and yielded higher densities of potamal fish as well as larger fish than electrofishing. Trawl catches further estimated higher biodiversity compared to seining, while the latter yielded higher densities of eurytopic, rheophilic, lithophilic and phytophilic fish. Drift-netting yielded the lowest estimates overall but sample size was very low. We suggest that electrofishing is an appropriate method to assess and evaluate the effects of hydromorphological degradation and rehabilitation on fish, and to guide river management. It sufficiently well represents the typical fish assemblage of large rivers despite its restriction to the shoreline. In contrast, assessing specifically Habitat Directive, migratory and rare species, as well as obtaining complete species inventories, e.g., for biodiversity assessments, requires complementary sampling of the mid-channel of large rivers by additional gears such as trawling.

Zhang, Y., Jeppesen, E., Liu, X., Qin, B., Shi, K., Zhou, Y., Thomaz, S. M., Deng, J. (2017). Global loss of aquatic vegetation in lakes. Earth-Science Reviews 173, 259-265,

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Global loss of aquatic vegetation in lakes

Quantitative global assessments of aquatic vegetation dynamics in lakes are lacking despite reports of the losses of submerged aquatic vegetation. We conducted a comprehensive global assessment of aquatic vegetation at 155 study sites. We also included ≥2 yr of information on the absolute or relative area of aquatic vegetation from the literature. We calculated the difference between initial and final observed aquatic vegetation area (or cover) to represent the overall trends over time. We classified the study sites of aquatic vegetation into the categories “increasing”,“decreasing” or “no change” using a threshold of 10%. Aquatic vegetation area (or cover) decreased in 101 study sites, particularly in China (35 study sites), increased in 43 study sites, and showed nomarked changes in 11 study sites. Our results revealed an accelerating decrease rate (vegetation loss in terms of area or cover) over time: 13.5 ± 16.9%/yr (1900–1980), 21.8 ± 28.9%/yr (1980–2000) and 33.6 ± 59.8%/yr (after 2000). Moreover, the area (or cover) increase rate in lakes where aquatic vegetation showed recoverydecreased from 23.5 ± 29.9%/yr (1980–2000) to 16.8 ± 13.2%/yr (after 2000). We conclude that aquaticvegetation loss is accelerating, especially that of submerged aquatic vegetation and particularly in lakes with an area larger than 50 km2. The predominance of decreasing vegetation found in our study is likely caused by multiple stressors such as eutrophication, algal blooms, land reclamation, aquaculture cultivation and global climate changes.

Zhang, W., Jeppesen, E., Wang, M., Xu, X., Wang, L. (2017). Allelopathic effect boosts Chrysosporum ovalisporum dominance in summer at the expense of Microcystis panniformis in a shallow coastal water body. Environmental Science and Pollution Research 24(5), 4666-4675,

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Allelopathic effect boosts Chrysosporum ovalisporum dominance in summer at the expense of Microcystis panniformis in a shallow coastal water body.

The increased occurrence of harmful cyanobacterial species and, with this, higher frequency of cyanobacteria blooms, closely associated with eutrophication and climate change, have attracted increasing attention worldwide. However, competition mechanisms between the different bloom-forming cyanobacteria species remain to be elucidated. In this paper, for the first time, the allelopathic effect of the cyanobacterium Chrysosporum ovalisporum on the cyanobacterium Microcystis panniformis is reported. The results of our study conducted in a Chinese shallow coastal water body demonstrated that the biomass of M. panniformis was relatively low during the C. ovalisporum blooming period. Co-cultivation of a C. ovalisporum strain with a M. panniformis strain showed strong inhibition of the growth of M. panniformis but stimulation of C. ovalisporum. Thus, filtrate of C. ovalisporum culture had a strong inhibitory effect on the performance of M. panniformis by decreasing the maximum optical quantum yield (Fv/Fm), the electron transport rate (ETR) of PS II and the onset of light saturation (Ik) and by increasing the alkaline phosphatase (ALP) activity and superoxide dismutase (SOD) activity of M. panniformis. Our results suggest that the interspecific allelopathic effect plays an important role in the competition between different cyanobacteria species. We foresee the importance of C. ovalisporum to intensify in a future warmer world, not least in small- to medium-sized, warm and high conductivity coastal water bodies.

Zhang X., Tang, Y., Jeppesen E., Liu, Z. (2017). Biomanipulation-induced reduction of sediment phosphorus release in a tropical shallow lake. Hydrobiologica 794(1), 49-57,

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Biomanipulation-induced reduction of sediment phosphorus release in a tropical shallow lake

Biomanipulation via fish regulation combined with submerged plant introduction is an effective measure to restore eutrophic shallow lakes. Improved water quality and clarity promote growth of benthic algae, which with submerged plants may limit sediment phosphorus (P) release, thereby reinforce lake recovery. Our study sought to evaluate the effect of such a biomanipulation on water quality, benthic algal development and sediment P release in a shallow, tropical lake by (1) comparing porewater and lake water quality, light intensity and benthic algal development in restored and unrestored sections; (2) conducting a 32P radiotracer experiment to track P release from sediment cores sampled from both sections. The biomanipulation led to lower total P, total dissolved P, and soluble reactive P concentrations in lake water, lower phytoplankton biomass, and increased light intensity at sediment surface, stimulating benthic algal development. Moreover, sediment 32P release was lower in the restored than unrestored section. Concurrently, dissolved oxygen levels in upper layers of the sediment cores were higher in the restored section. Our study indicates that the biomanipulation improved water quality and enhanced growth of benthic algae, thereby reducing sediment P release, which may be one of the main mechanisms to create successful restoration.

Zhou, Y., Ma, J., Zhang, Y., Qin, B., Jeppesen, E., Shi, K., Brookes, J. D., Spencer, R. G. M., Zhu, G., Gao, G. (2017). Improving water quality in China: Environmental investment pays dividends. Water Research 118, 152-159,

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Improving water quality in China: Environmental investment pays dividends

This study highlights how Chinese economic development detrimentally impacted water quality in recent decades and how this has been improved by enormous investment in environmental remediation funded by the Chinese government. To our knowledge, this study is the first to describe the variability of surface water quality in inland waters in China, the affecting drivers behind the changes, and how the government-financed conservation actions have impacted water quality. Water quality was found to be poorest in the North and the Northeast China Plain where there is greater coverage of developed land (cities + cropland), a higher gross domestic product (GDP), and higher population density. There are significant positive relationships between the concentration of the annual mean chemical oxygen demand (COD) and the percentage of developed land use (cities + cropland), GDP, and population density in the individual watersheds (p < 0.001). During the past decade, following Chinese government-financed investments in environmental restoration and reforestation, the water quality of Chinese inland waters has improved markedly, which is particularly evident from the significant and exponentially decreasing GDP-normalized COD and ammonium (NH4⁺-N) concentrations. Our results demonstrate that the increasing GDP in China over the past decade did not occur at the continued expense of its inland water ecosystems. This offers hope for the future, also for other industrializing countries, that with appropriate environmental investments a high GDP can be reached and maintained, while simultaneously preserving inland aquatic ecosystems, particularly through management of sewage discharge.

Zhou, Y., Shi, K., Liu, X., Jeppesen, E., Zhang, Y., Zhou, Q., Wu, H., Tang, X. Zhu, G. (2017). Fluorescence peak integration ratio IC:IT as a new potential real-time tracer differentiating dissolved organic matter of allochthonous and autochthonous origin in waters. Science of the Total Environment 574, 1588-1598,

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Fluorescence peak integration ratio IC:IT as a new potential indicator tracing the compositional changes in chromophoric dissolved organic matter

The present study demonstrates that the ratio of fluorescence integration of peak C to peak T (IC:IT) can be used as an indicator tracing the compositional dynamics of chromophoric dissolved organic matter (CDOM). CDOM absorption and fluorescence spectroscopy and stable isotope δ13C were determined on a seasonal basis in seventeen Chinese inland waters as well as in a series of mixing and photodegradation experiments in the lab. A strong positive linear correlation was recorded between IC:IT and the ratio of terrestrial humic-like C1 to tryptophan-like C4 (C1:C4) derived by parallel factor analysis. The r2 for the linear fitting between IC:IT and C1:C4 (r2 = 0.80) was notably higher than between C1:C4 and other indices tested, including the ratio of CDOM absorption at 250 nm to 365 nm, i.e. a(250):a(365) (r2 = 0.09), spectral slope (S275–295) (r2 = 0.26), spectral slope ratio (SR) (r2 = 0.31), the humification index (HIX) (r2 = 0.47), the recent autochthonous biological contribution index (BIX) (r2 = 0.27), and a fluorescence index (FI370) (r2 = 0.07). IC:IT exhibited larger variability than the remaining six indices and a closer correlation with stable isotope δ13C than that observed for a(250):a(365), S275–295, SR, FI370, and BIX during field campaigns. Confirming our field observations, significant correlations were recorded between IC:IT and the remaining six indices, and IC:IT also demonstrated notably larger variability than the six other indices during our wastewater addition experiment. Compared with HIX, eutrophic water addition and photobleaching substantially decreased IC:IT but had no pronounced effect on a(250):a(365), S275–297, SR, FI370, and BIX, further suggesting that IC:IT is the most efficient indicator of the CDOM compositional dynamics.

Zhou, Y., Yao, X., Zhang, Y., Shi, K., Zhang, Y., Jeppesen, E., Gao, G., Zhu, G., Qin, B. (2017). Potential rainfall-intensity and pH-driven shifts in the apparent fluorescent composition of dissolved organic matter in rainwater. Environmental Pollution 224, 638-648,

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Potential rainfall-intensity and pH-driven shifts in the apparent fluorescent composition of dissolved organic matter in rainwater

Perturbations of rainwater chromophoric dissolved organic matter (CDOM) fluorescence induced by changes in rainfall intensity and pH were investigated by field observations and laboratory pH titrations. Microbial humic-like fluorophores dominated the rainwater CDOM pool, followed by tryptophan-like and tyrosine-like substances. Increased rainfall intensity had notable dilution effects on all six fluorescent components (C1-C6) identified using parallel factor (PARAFAC) analysis, the effect being especially pronounced for the microbial humic-like C1, tryptophan-like C3, and tyrosine-like C5. The results also indicated that increasing pH from 7 to 9 led to decreased fluorescence intensity (Fmax) of all the six components, while a pH increase from 5 to 7, resulted in increasing Fmax of terrestrial humic-like C2, tyrosine-like C5, and tryptophan-like C6 and decreasing microbial humic-like C1, tryptophan-like C3, and fulvic-like C4. Two-dimensional correlation spectroscopy (2D-COS) demonstrated that synchronous fluorescence responded first to pH modifications at fulvic-like wavelength (λEx/λEm = ∼316/416 nm), followed by tyrosine-like wavelength (λEx/λEm = ∼204/304 nm), tryptophan-like wavelength (λEx/λEm = ∼226/326 nm), microbial humic-like wavelength (∼295/395 nm), and finally terrestrial humic-like wavelength (∼360/460 nm). Our results suggest that a decrease in areas affected by acid rain in South China occurring at present may possibly result in apparent compositional changes of CDOM fluorescence. The decreased rainfall in South-West China and increased rainfall in North-West China during the past five decades may possibly accordingly result in increased and decreased Fmax of all the six components identified in South-West and North-West China, respectively.

Zhou, L., Zhou, Y., Hu, Y., Cai, J., Bai, C., Shao, K., Gao, G., Zhang, Y., Jeppesen, E., Tang, X. (2017). Hydraulic connectivity and evaporation control the water quality and sources of chromophoric dissolved organic matter in Lake Bosten in arid northwest China. Chemosphere 188, 608-617,

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Hydraulic connectivity and evaporation control the water quality and sources of chromophoric dissolved organic matter in Lake Bosten in arid northwest China

Lake Bosten is the largest oligosaline lake in arid northwestern China, and water from its tributaries and evaporation control the water balance of the lake. In this study, water quality and chromophoric dissolved organic matter (CDOM) absorption and fluorescence were investigated in different seasons to elucidate how hydraulic connectivity and evaporation may affect the water quality and variability of CDOM in the lake. Mean suspended solids and turbidity were significantly higher in the upstream tributaries than in the lake, the difference being notably more pronounced in the wet than in the dry season. A markedly higher mean first principal component (PC1) score, which was significantly positively related to protein-like components, and a considerably lower fluorescence peak integration ratio - IC:IT, indicative of the terrestrial humic-like CDOM contribution percentage, were observed in the lake than in the upstream tributaries. Correspondingly, notably higher contribution percentages of terrestrial humic-like components were observed in the river mouth areas than in the remaining lake regions. Furthermore, significantly higher mean turbidity, and notably lower mean conductivity and salinity, were recorded in the southwestern Kaidu river mouth than in the remaining lake regions in the wet season. Notably higher mean salinity is recorded in Lake Bosten than in upstream tributaries. Autochthonous protein-like associated amino-acids and also PC1 scores increased significantly with increasing salinity. We conclude that the dynamics of water quality and CDOM composition in remote arid Lake Bosten are strongly driven by evaporation and also the hydraulic connectivity between the upstream tributaries and the downstream lake.


Alahuhta, J., Hellsten, S., Kuoppala, M., Riihimäki, J. (2016). Regional and local determinants of macrophyte community composition in high-latitude lakes in Finland. Hydrobiologia 812, 99-114,

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Regional and local determinants of macrophyte community compositions in high-latitude lakes of Finland

Species distributions are structured by regional and local determinants, which operate at multiple spatial and temporal scales. The purpose of our work was to distinguish the relative roles of local variables, climate, geographical location and post glaciation condition (i.e. delineation between supra- and subaquatic lakes during the post-glacial Anculys Lake) in explaining variation in macrophyte community composition of all taxa, helophytes and hydrophytes. In addition, we investigated how these four explanatory variable groups affected macrophyte strategy groups based on Grime’s classification. Using partial linear regression and variation partitioning, we found that macrophyte communities are primarily filtered by local determinants together with regional characteristics at the studied spatial scale. We further evidenced that post glaciation condition indirectly influenced on local water quality variables, which in turn directly contributed to the macrophyte communities. We thus suggest that regional determinants interact with local-scale abiotic factors in explaining macrophyte community patterns and examining only regional or local factors is not sufficient for understanding how aquatic macrophyte communities are structured locally and regionally.

Alahuhta, J., Aroviita, J. (2016). Quantifying the relative importance of natural variables, human disturbance and spatial processes in ecological status indicators of boreal lakes. Ecological Indicators 63, 240–248,

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Quantifying the relative importance of natural variables, human disturbance and spatial processes in ecological status indicators of boreal lakes

Spatial processes are increasingly associated with species distributions in freshwaters. However, these processes are usually neglected in bioassessment techniques, which may introduce uncontrolled variation in ecological indicators used to express human disturbance. We used partial linear regression to quantify the relative importance of natural variables, human disturbance and spatial variables in structuring variation in boreal lake status indicators based on six biological indicator groups (phytoplankton, macrophytes, diatoms, littoral and profundal macroinvertebrates and fish). We found that, of the pure fractions, human disturbance explained most variation (7-32%) of the Ecological Quality Ratios (EQRs) for all groups, with the exception of littoral macroinvertebrate metric, which was most controlled by natural and spatial variables (15% and 16%, respectively). In addition, pure fractions of natural and spatial variables and joint fractions of different explanatory variable groups structured all biological metrics to various degrees. Phytoplankton, diatom and profundal macroinvertebrate EQRs responded purest to human disturbance but only weakly to pure natural or spatial variation. Our work demonstrates that spatial processes and spatial structuring of the environment can bias bioassessment techniques and hinder the detection of human impact. Thus, it is important to acknowledge spatial autocorrelation, context of metacommunity dynamics, species dispersal traits and variable spatial extent when defining reference conditions and bioassessment techniques for different biological groups. More research is needed to better understand the relative role of spatial processes on ecological metrics originated from different freshwater ecosystems. To this end, our work provides an example of how sources of variation can be identified to increase accuracy in freshwater bioassessment.

Alnøe, A. B., Riis, T., Baattrup-Pedersen, A. (2016). Comparison of metabolic rates among macrophyte and nonmacrophyte habitats in streams. Freshwater Science 35, 909-921,

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Comparison of metabolic rates among macrophyte and nonmacrophyte habitats in streams

Little is known about the relative contribution of different stream habitats to reach-scale metabolism. We measured in situ metabolism in sand, gravel, stone, and macrophyte habitats to compare metabolic rates among these habitat types and to compare habitat-weighted measurements with reach-scale measurements. We used open-bottom chambers in sand, gravel, and macrophyte habitats and closed-bottom chambers in stones, and we estimated reach-scale metabolism from 2-station O2 budgets. Macrophyte habitats had a significantly higher gross primary production (GPP) and community respiration (CR) than stone, gravel, and sand habitats. A large part of this difference was associated with epiphytic biofilm: 28% of net ecosystem production (NEP), 20% of CR, and 24% of GPP. Macrophyte habitats contributed proportionally more to reach-scale metabolism than did the other habitat types. Forty-one percent of reach-scale NEP, 60% of ecosystem respiration (ER), and 50% of GPP were associated with this habitat type even though only 14% of the reach was covered by macro-phytes. We found significant linear relationships between GPP and CR and the amount of autotrophic biomass in the streams. The rates reported in 11 literature studies fit into our observed relationships, showing the generality of our findings. The rates we obtained expand the range of reported metabolic values in relation to auto-trophic biomass for both low and high biomass. The importance of macrophyte habitat can be ascribed to the macrophytes themselves, the associated epiphytic biofilm, and the fine organic material trapped in the dense stands. We conclude that besides having an effect on the structural elements in streams, macrophytes contribute significantly to stream ecosystem functions.

Arora, R., Tockner, K., Venohr, M. (2016). Changing river temperatures in northern Germany. Hydrological Processes 30, 3084-3096,

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Changing river temperatures in northern Germany

Climate change is one of the main drivers of river warming worldwide. However, the response of river temperature to climate change differs with the hydrology and landscape properties, making it difficult to generalize the strength and the direction, of river temperature trends across large spatial scales and various river types. Additionally, there is a lack of long-term and large-scale trend studies in Europe as well as globally. In this study, we investigated the long-term (25years; 132 sites) and the short-term (10years; 475 sites) river temperature trends, patterns and underlying drivers within the period 1985-2010 in seven river basins of Germany. The majority of the sites underwent significant river warming during 1985-2010 (mean warming trend: 0.03°Cyear-1, SE=0.003), with a faster warming observed during individual decades (1985-1995 and 2000-2010) within this period. Seasonal analyses showed that, while rivers warmed in all seasons, the fastest warming had occurred during summer. Among all the considered hydro-climatological variables, air temperature change, which is a response to climate forcing, was the main driver of river temperature change because it had the strongest correlation with river temperature, irrespective of the period. Hydrological variables, such as average flow and baseflow, had a considerable influence on river temperature variability rather than on the overall trend direction. However, decreasing flow probably assisted in a faster river temperature increase in summer and in rivers in NE basins (such as the Elbe basin). The North Atlantic Oscillation Index had a greater significant influence on the winter river temperature variability than on the overall variability. Landscape and basin variables, such as altitude, ecoregion and catchment area, induced spatially variable river temperature trends via affecting the thermal sensitivity of rivers, with the rivers in large catchments and in lowland areas being most sensitive.

Baattrup-Pedersen, A., Göthe, E., Riis, T., O'Hare, M. T. (2016). Functional trait composition of aquatic plants can serve to disentangle multiple interacting stressors in lowland streams. Science of the Total Environment 543, 230-238,

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Functional trait composition of aquatic plants can serve to disentangle multiple interacting stressors in lowland streams

Historically, close attention has been paid to negative impacts associated with nutrient loads to streams and rivers, but today hydromorphological alterations are considered increasingly implicated when lowland streams do not achieve good ecological status. Here, we explore if trait-abundance patterns of aquatic plants change along gradients in hydromorphological degradation and eutrophication in lowland stream sites located in Denmark. Specifically, we hypothesised that: i) changes in trait-abundance patterns occur along gradients in hydromorphological degradation and ii) trait-abundance patterns can serve to disentangle effects of eutrophication and hydromorphological degradation in lowland streams reflecting that the mechanisms behind changes differ. We used monitoring data from a total of 147 stream reaches with combined data on aquatic plant species abundance, catchment land use, hydromorphological alterations (i.e. planform, cross section, weed cutting) and water chemistry parameters. Traits related to life form, dispersal, reproduction and survival together with ecological preference values for nutrients and light (Ellenberg N and L) were allocated to 41 species representing 79% of the total species pool. We found clear evidence that habitat degradation (hydromorphological alterations and eutrophication) mediated selective changes in the trait-abundance patterns of the plant community. Specific traits could distinguish hydromorphological degradation (free-floating, surface; anchored floating leaves; anchored heterophylly) from eutrophication (free-floating, submerged; leaf area). We provide a conceptual framework for interpretation of how eutrophication and hydromorphological degradation interact and how this is reflected in trait-abundance patterns in aquatic plant communities in lowland streams. Our findings support the merit of trait-based approaches in biomonitoring as they shed light on mechanisms controlling structural changes under environmental stress. The ability to disentangle several stressors is particularly important in lowland stream environments where several stressors act in concert since the impact of the most important stressor can be targeted first, which is essential to improve the ecological status.

Boll, T., Levi, E. E., Bezirci, G., Özuluğ, M., Tavsanoğlu, Ü. N., Çakıroğlu, A. I., Özcan, S., Brucet, S., Jeppesen, E., Beklioğlu, M. (2016). Fish assemblage and diversity in lakes of western and central Turkey - role of geo-climatic and environmental gradients, Hydrobiologia 771, 31-44, http:/"

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Fish assemblage and diversity in lakes of western and central Turkey: role of geo-climatic and other environmental variables

We conducted a fish survey in 40 lakes in western and central Turkey. Fifty species (one to eleven per lake) were recorded, including eighteen endemic and seven alien species. We investigated which local geo-climatic and other environmental variables shaped the fish assemblages. Altitude and temperature turned out to be the most important factors for total species richness as well as richness of omnivorous and zooplanktivorous species and the Shannon–Wiener diversity index, with more species and higher diversity occurring in the warmer lowland lakes. Altitude may affect the fish assemblage directly through dispersal limitation or indirectly by creating a gradient in temperature with which it was strongly correlated. Cyprinidae was the most species-rich and widespread family. Atherinidae, Gobiidae, and Mugilidae (families of marine origin) were mainly found in the lowland regions, while Salmonidae exclusively appeared in the high-altitude lakes. The presence of widely distributed translocated native and alien species revealed a large human impact on the fish assemblages, potentially threatening the rich endemic fish fauna in lakes in this region.

Bondar-Kunze, E., Maier, S., Schönauer, D., Bahl, N., Hein, T. (2016). Antagonistic and synergistic effects on a stream periphyton community under the influence of pulsed flow velocity increase and nutrient enrichment. Science of the Total Environment 573, 594-602,

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Antagonistic and synergistic effects on a stream periphyton community under the influence of pulsed flow velocity increase and nutrient enrichment

Aquatic ecosystems are generally affected by multiple stressors, and therefore, analysing single stressor responses is not appropriate to evaluate the whole range of effects on these ecosystems. We assessed the interaction effects of two strong stressors (higher flow velocity due to e.g. daily hydropeaking) and nutrient enrichment for an oligotrophic stream periphyton community. As periphyton has a rapid reproduction rate and very short life cycles, it can therefore be expected to reflect short-term impacts and sudden changes/disturbances in the environment. We measured biomass development, algal group distribution and photosynthesis efficiency during a time period of 33 days in an experimental flume setting in Lunz am See (Austria). We conducted the experiment with two treatments (no hydropeaking and hydropeaking) and three nutrient enrichments (nitrate, phosphate and nitrate + phosphate enrichment) and control (no nutrient addition). The results showed a significant lower biomass development in the hydropeaking treatment (HP), compared to the no-hydropeaking treatment (NHP) in a later successional stage (day 33). Nutrient subsidy effects were not observed, because the biomass development (chlorophyll-a) of periphyton was highly diminished through the pulsed flow velocity increase. Also a negative synergistic interaction (more negative than predicted additively) was observed. Our study confirmed for periphyton communities that for different algal groups and functional guilds the same multiple stressor combination can be detrimental for one species group (e.g. chlorophyta) while beneficial for another (e.g. diatoms). We conclude for multiple stressor studies to consider the successional stage and community composition, when estimating the interaction effects of these stressors.

Bowes, M. J., Loewenthal, M., Read, D. S., Hutchins, M. G., Prudhomme, C., Armstrong, L. K., Harman, S. A., Wickham, H. D., Gozzard E., Carvalho, L. (2016). Identifying multiple stressor controls on phytoplankton dynamics in the River Thames (UK) using high-frequency water quality data. Science of the Total Environment 569-570, 1489-1499,

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Identifying multiple stressor controls on phytoplankton dynamics in the River Thames (UK) using high-frequency water quality data

River phytoplankton blooms can pose a serious risk to water quality and the structure and function of aquatic ecosystems. Developing a greater understanding of the physical and chemical controls on the timing, magnitude and duration of blooms is essential for the effective management of phytoplankton development. Five years of weekly water quality monitoring data along the River Thames, southern England were combined with hourly chlorophyll concentration (a proxy for phytoplankton biomass), flow, temperature and daily sunlight data from the mid-Thames. Weekly chlorophyll data was of insufficient temporal resolution to identify the causes of short term variations in phytoplankton biomass. However, hourly chlorophyll data enabled identification of thresholds in water temperature (between 9 and 19°C) and flow (<30m(3)s(-1)) that explained the development of phytoplankton populations. Analysis showed that periods of high phytoplankton biomass and growth rate only occurred when these flow and temperature conditions were within these thresholds, and coincided with periods of long sunshine duration, indicating multiple stressor controls. Nutrient concentrations appeared to have no impact on the timing or magnitude of phytoplankton bloom development, but severe depletion of dissolved phosphorus and silicon during periods of high phytoplankton biomass may have contributed to some bloom collapses through nutrient limitation. This study indicates that for nutrient enriched rivers such as the Thames, manipulating residence time (through removing impoundments) and light/temperature (by increasing riparian tree shading) may offer more realistic solutions than reducing phosphorus concentrations for controlling excessive phytoplankton biomass.

Branco, P., Santos, J. M., Amaral, S., Romao, F., Pinheiro, A. N., Ferreira, M. T. (2016). Potamodromous fish movements under multiple stressors: Connectivity reduction and oxygen depletion. Science of the Total Environment 572, 520-525,

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Potamodromous fish movements under multiple stressors: Connectivity reduction and oxygen depletion

Rivers are impacted by multiple stressors that can interact to create synergistic, additive or antagonistic effects, but experimental studies on fish encompassing more than one stressor are seldom found. Thus, there is the need to study stressors through multifactorial approaches that analyse the impact of fish exposure to multiple stressors and evaluate fish sensitivity to stressor combinations. Some of the most common impacts to Mediterranean rivers are of two natures: i) water abstraction and ii) diffuse pollution. Therefore, the present study aims at studying the responses of potamodromous fish facing combinations of: 1) a primary stressor (two levels of connectivity reduction due to water scarcity), and 2) a secondary stressor (three levels of oxygen depletion due to increase organic load - of anthropogenic nature). Schools of five wild fish from a cyprinid species (Luciobarbus bocagei) were placed in a flume, equipped with see-through sidewalls to allow for behavioural analysis, and subjected to different combinations of the stressors. Results show that at the unconnected level the primary stressor (lack of connectivity) overrode the effect of the secondary stressor (oxygen depletion), but when connectivity existed oxygen depletion caused a reduction of fish movements with decreasing oxygen concentrations. This multifactorial study contributes to improved prediction of fish responses upon actual or projected pressure scenarios.

Bruno, D., Gutiérrez-Cánovas, T., Sánchez-Fernández, D., Velasco, J., Nilsson, C. (2016). Impacts of environmental filters on funcational redundancy in riparian vegetation. Journal of Applied Ecology 53, 846-855,

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Impacts of environmental filters on funcational redundancy in riparian vegetation

Understanding and predicting ecosystem responses to multiple environmental pressures is a long-standing interest in ecology and environmental management. However, few studies have examined how the functional features of freshwater biological communities vary along multiple gradients of environmental stress. Furthermore, modelling these functional features for a whole river network constitutes a strong potential basis to improve ecosystem management. 2.We explored how functional redundancy of biological communities (FR, a functional feature related to the stability, resistance and resilience of ecosystems) responds to single and multiple environmental filters. We compared these responses with those of functional richness, evenness and divergence. We used riparian vegetation of a Mediterranean basin, and three of the main environmental filters affecting freshwater communities in such regions, i.e. drought, flow regulation and agricultural intensity, thus considering the potential effect of natural environmental variability. We also assessed the predictability of FR and estimated it for the entire river network. 3.We found that all functional measures decreased with increasing environmental filter intensity. However, FR was more sensitive to single and multiple environmental filters compared to other functional measures. The best-fitting model explained 59% of the FR variability and included agriculture, drought and flow regulation and the pairwise interactions of agriculture with drought and flow regulation. The parameters of the FR models differed from null model expectations reflecting a non-random decline along stress gradients. 4.Synthesis and applications. We found non-random detrimental effects along environmental filters for riparian functional redundancy (FR, the most sensitive functional index), meaning that increased stress could jeopardize stability, resistance and resilience of these systems. In general, agriculture caused the greatest impact on FR and functional diversity measures, being the most important stressor for riparian functionality in the study area. Temporary streams flowing through an agricultural, regulated basin had reduced values of FR, whereas the free-flowing medium-sized, perennial water courses flowing through unaltered sub-basins displayed higher values of FR and potentially greater stability against human impacts. All these findings along with the predicted basin-wide variation of FR can assist environmental managers in improving monitoring and ecosystem management. This article is protected by copyright. All rights reserved.

Bruno, D., Gutiérrez-Cánovas, T., Velasco, J., Sánchez-Fernández, D. (2016). Functional redundancy as a tool for bioassessment: A test using riparian vegetation. Science of the Total Environment 566-567, 1269-1276,

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Functional redundancy as a tool for bioassessment: A test using riparian vegetation

There is an urgent need to track how natural systems are responding to global change in order to better guide management efforts. Traditionally, taxonomically based metrics have been used as indicators of ecosystem integrity and conservation status. However, functional approaches offer promising advantages that can improve bioassessment performance. In this study, we aim to test the applicability of functional redundancy (FR), a functional feature related to the stability, resistance and resilience of ecosystems, as a tool for bioassessment, looking at woody riparian communities in particular. We used linear mixed-effect models to investigate the response of FR and other traditional biomonitoring indices to natural (drought duration) and anthropogenic stress gradients (flow regulation and agriculture) in a Mediterranean basin. Such indices include species richness, a taxonomic index, and the Riparian Quality Index, which is an index of ecological status. Then, we explored the ability of FR and the other indices to discriminate between different intensities of human alteration. FR showed higher explanatory capacity in response to multiple stressors, although we found significant negative relationships between all the biological indices (taxonomic, functional and ecological quality) and stress gradients. In addition, FR was the most accurate index to discriminate among different categories of human alteration in both perennial and intermittent river reaches, which allowed us to set threshold values to identify undisturbed (reference condition), moderately disturbed and highly disturbed reaches in the two types of river. Using these thresholds and the best-fitting model, we generated a map of human impact on the functional redundancy of riparian communities for all the stretches of the river network. Our results demonstrate that FR presents clear advantages over traditional methods, which suggests that it should be part of the biomonitoring toolbox used for environmental management so as to obtain better predictions of ecosystem response to environmental changes.

Bucak T., Beklioglu M., Cakiroglu A.I., Erdogan S., Levi E. (2016). Metadata of the Lake Beysehir and Its Catchment, Turkey. Freshwater Metadata Journal 11, 1-8,

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Metadata of the Lake Beysehir and Its Catchment, Turkey

This database contains climatic, hydrologic, water quality and biological information for the Lake Beysehir catchment, Turkey. The dataset includes meteorological data (precipitation, air temperature, wind speed, solar radiation, relative humidity), discharges for the main inflows and lake outflow, lake water level, water chemistry data for inflows and lake. In addition, lake biological data (phytoplankton, zooplankton, fish and macrophyte) is avaiable. Data was compiled during the METU-DPT-TEAB project, EU-FP7 REFRESH project and EU-FP7 MARS project.

Bussi, G., Whitehead, P. G., Bowes, M. J., Read, D. S., Prudhomme, C., Dadson, S. J. (2016). Impacts of climate change, land-use change and phosphorus reduction on phytoplankton in the River Thames (UK). Science of the Total Environment 572, 1507-1519,

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Impacts of climate change, land-use change and phosphorus reduction on phytoplankton in the River Thames (UK)

Potential increases of phytoplankton concentrations in river systems due to global warming and changing climate could pose a serious threat to the anthropogenic use of surface waters. Nevertheless, the extent of the effect of climatic alterations on phytoplankton concentrations in river systems has not yet been analysed in detail. In this study, we assess the impact of a change in precipitation and temperature on river phytoplankton concentration by means of a physically-based model. A scenario-neutral methodology has been employed to evaluate the effects of climate alterations on flow, phosphorus concentration and phytoplankton concentration of the River Thames (southern England). In particular, five groups of phytoplankton are considered, representing a range of size classes and pigment phenotypes, under three different land-use/land-management scenarios to assess their impact on phytoplankton population levels. The model results are evaluated within the framework of future climate projections, using the UK Climate Projections 09 (UKCP09) for the 2030s. The results of the model demonstrate that an increase in average phytoplankton concentration due to climate change is highly likely to occur, with the magnitude varying depending on the location along the River Thames. Cyanobacteria show significant increases under future climate change and land use change. An expansion of intensive agriculture accentuates the growth in phytoplankton, especially in the upper reaches of the River Thames. However, an optimal phosphorus removal mitigation strategy, which combines reduction of fertiliser application and phosphorus removal from wastewater, can help to reduce this increase in phytoplankton concentration, and in some cases, compensate for the effect of rising temperature.

Cavalli, G., Baattrup-Pedersen, A., Riis, T. (2016). Nutrient availability and nutrient use efficiency in plants growing in the transition zone between land and water. Plant Biology 18, 301-306,

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Nutrient availability and nutrient use efficiency in plants growing in the transition zone between land and water

The transition zone between terrestrial and freshwater habitats is highly dynamic, with large variability in environmental characteristics. Here, we investigate how these characteristics influence the nutritional status and performance of plant life forms inhabiting this zone. Specifically, we hypothesised that: (i) tissue nutrient content differs among submerged, amphibious and terrestrial species, with higher content in submerged species; and (ii) PNUE gradually increases from submerged over amphibious to terrestrial species, reflecting differences in the availability of N and P relative to inorganic C across the land-water ecotone. We found that tissue nutrient content was generally higher in submerged species and C:N and C:P ratios indicated that content was limiting for growth for ca. 20% of plant individuals, particularly those belonging to amphibious and terrestrial species groups. As predicted, the PNUE increased from submerged over amphibious to terrestrial species. We suggest that this pattern reflects that amphibious and terrestrial species allocate proportionally more nutrients into processes of importance for photosynthesis at saturating CO2 availability, i.e. enzymes involved in substrate regeneration, compared to submerged species that are acclimated to lower availability of CO2 in the aquatic environment. Our results indicate that enhanced nutrient loading may affect relative abundance of the three species groups in the land-water ecotone of stream ecosystems. Thus, species of amphibious and terrestrial species groups are likely to benefit more from enhanced nutrient availability in terms of faster growth compared to aquatic species, and that this can be detrimental to aquatic species growing in the land-water ecotone, e.g. Ranunculus and Callitriche. © 2015 German Botanical Society and The Royal Botanical Society of the Netherlands.

Çakiroğlu, A.I., Levi, E. E., Tavşanoğlu, N., Bezirci,G., Erdoğan, Ş., Filiz, N., Andersen, T. J., Davidson, T. A., Jeppesen, E., Beklioğlu, M. (2016). Inferring past environmental change of three Turkish shallow lakes from community changes of sub-fossil cladocera. Hydrobiologia 778, 295-312,

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Inferring past environmental changes in three Turkish lakes from sub-fossil Cladocera

Cladocerans are increasingly used in palaeolimnological studies as their community composition is sensitive to both anthropogenic and natural forces in lakes. We present the results of a palaeolimnological investigation of three Turkish shallow lakes located in cold dry steppe and semi-dry Mediterranean climatic regions. The aim was to elucidate historical changes in environmental conditions by analysing sub-fossil cladocerans in 210Pb-dated sediment cores. Sub-fossil cladoceran remains from the surface sediment of 40 Turkish lakes were analysed to examine the environmental factors that most correlated with variation in the cladoceran assemblage. Redundancy analysis showed that salinity, macrophyte abundance, fish density, depth and total phosphorus were the most correlated with change in cladoceran assemblage composition with eigenvalues for the first and the second axes being λ 1 = 0.312 and λ 2 = 0.061, respectively. Sedimentary cladoceran assemblages from three cores were placed passively within the framework of the surface sediment ordination. The results reveal a prevalent impact of salinity, fish abundance and water level changes from the past to present. Thus, using cladoceran-based inferences, we traced key environmental changes related to variation in climate change, restoration and water level regulation over the last century.

Chen, J., Cao, T., Zhang, X., Xi, Y., Ni, L., Jeppesen, E. (2016). Differential photosynthetic and morphological adaptations to low light affect depth distribution of two submersed macrophytes in lakes. Scientific Reports 6 (E34028), 1-9,

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Differential photosynthetic and morphological adaptations to low light affect depth distribution of two submersed macrophytes in lakes

To evaluate the relative importance of photosynthetic versus morphological adaptations of submersed macrophytes to low light intensity in lakes, rapid light curves (RLCs), morphological parameters, relative growth rate (RGR), clonal reproduction and abundance of two submersed macrophytes (Potamogeton maackianus and Vallisneria natans) were examined under 2.8%, 7.1%, 17.1% and 39.5% ambient light in a field and outdoor experimental study. The plants increased their initial slope of RLCs (α) and decreased their minimum saturating irradiance (Ek) and maximum relative electron transport rate (ETRm) of RLCs under low light stress, but V. natans was more sensitive in RLCs than P. maackianus. Accordingly, the RGR, plant height and abundance of P. maackianus were higher in the high light regimes (shallow water) but lower in the low light regimes than those of V. natans. At the 2.8% ambient light, V. natans produced ramets and thus fulfilled its population expansion, in contrast to P. maackianus. The results revealed that P. maackianus as a canopy-former mainly elongated its shoot length towards the water surface to compensate for the low light conditions, however, it became limited in severe low light stress conditions. V. natans as a rosette adapted to low light stress mainly through photosynthetic adjustments and superior to severely low light than shoot elongation.

Coppens J., Hejzlar, J., Šorf, M., Jeppesen, E., Papastergiadou, E., Stefanidis, K., Erdoğan, Ş., Scharfenberger, U., Mahdy, A., Nõges, P., Tuvikene, A., Baho, D. L., Trigalj, C., Olsen, S., Beklioğlu, M. (2016). The influence of nutrient loading, climate and water depth on nitrogen and phosphorus loss in shallow lakes: a pan-European mesocosm experiment. Hydrobiologia 778, 10-32,

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The influence of nutrient loading, climate and water depth on nitrogen and phosphorus loss in shallow lakes: a pan-European mesocosm experiment

Losses of phosphorus (P) and nitrogen (N) have important influences on in-lake concentrations and nutrient loading to downstream ecosystems. We performed a series of mesocosm experiments along a latitudinal gradient from Sweden to Greece to investigate the factors influencing N and P loss under different climatic conditions. In six countries, a standardised mesocosm experiment with two water depths and two nutrient levels was conducted concurrently between May and November 2011. Our results showed external nutrient loading to be of key importance for N and P loss in all countries. Almost all dissolved inorganic nitrogen (DIN) and soluble reactive phosphorus (SRP) were lost or taken up in biomass in all mesocosms. We found no consistent effect of temperature on DIN and SRP loss but a significant, though weak, negative effect of temperature on total nitrogen (TN) and total phosphorus (TP) loss in the deeper mesocosms, probably related to higher organic N and P accumulation in the water in the warmer countries. In shallow mesocosms, a positive trend in TN and TP loss with increasing temperature was observed, most likely related to macrophyte growth.

Coppens J., Özen, A., Çakıroğlu, I., Tavşanoğlu, Ü. N., Yozgatlıgil, C., Jeppesen, E., Beklioğlu, M. (2016). Impact of alternating wet and dry periods on long-term seasonal phosphorus and nitrogen budgets of two shallow Mediterranean lakes. Science of the Total Environment 563–564, 456–467,

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Impact of alternating wet and dry periods on long-term seasonal phosphorus and nitrogen budgets of two shallow Mediterranean lakes

The water balance, with large seasonal and annual water level fluctuations, has a critical influence on the nitrogen and phosphorus dynamics of shallow lakes in the semi-arid climate zone. We constructed seasonal water and nutrient budgets for two connected shallow lakes, Lakes Mogan and Eymir, located in Central Anatolia, Turkey. The study period covered 20 years with alternations between dry and wet years as well as restoration efforts including sewage effluent diversion and biomanipulations in Lake Eymir. Both lakes experienced a 1–2 m water level drop during a drought period and a subsequent increase during the wet period, with seasonal water level fluctuations of 0.60 to 0.70 m. During wet years with high water levels, small seasonal differences were observed with a nutrient peak in spring caused by external loading and nutrient loss via retention during summer. During years with low water levels, nutrient concentrations increased due to internal and external loading, exacerbated by evaporative water loss. In Lake Eymir, a shift to eutrophic conditions with turbid water occurred under low water level conditions and consequent internal loading of P from the sediment, causing high nutrient concentrations in summer. Our results indicate a threat of lakes drying out in the semi-arid climate zone if evaporation increases and precipitation decreases as anticipated from the global climate change predictions. In addition, our results show the influence of the water balance on the eutrophication of shallow lakes in the Mediterranean climate zone and highlight the ultimate consequences for lake management.

Cremona, F., Laas, A., Arvola, L., Pierson, D., Nõges, P., Nõges, T. (2016). Numerical exploration of the planktonic to benthic primary production ratios in lakes of the Baltic Sea catchment. Ecosystems 19(8), 1386-1400,

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Numerical exploration of the planktonic to benthic primary production ratios in lakes of the Baltic Sea catchment

Autotrophic structure refers to the partitioning of whole-ecosystem primary production between benthic and planktonic primary producers. Autotrophic structure remains poorly understood especially because of the paucity of estimates regarding benthic primary production. We used a conceptual model for numerically exploring the autotrophic structure of 13 hemiboreal lakes situated in the Baltic Sea catchment. We also used diel variations in primary production profiles to graphically evaluate levels of light and/or nutrient limitation in lakes. The input morphometric data, light extinction coefficients and dissolved carbon parameters were mostly obtained from in situ measurements. Results revealed that cross- and within-lake autotrophic structure varied greatly: one lake was clearly dominated by benthic production, and three lakes by phytoplankton production. In the rest, phytoplankton production was generally dominant but switch to benthic dominance was possible. The modelled primary production profiles varied according to lake water clarity and bathymetry. Our results clearly indicate that the relative contribution of benthic primary production to whole-lake primary production should be taken into account in studies about hemiboreal and boreal lakes.

Cremona, F., Laas, A., Nõges, P., Nõges, T. (2016). An estimation of diel metabolic rates of eight limnological archetypes from Estonia using high-frequency measurements. Inland Waters 6 (3), 352-363,

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An estimation of diel metabolic rates of eight limnological archetypes from Estonia using high-frequency measurements

We employed a Bayesian model to assess the metabolic state of 8 Estonian lakes representing the 8 lake types according to the European Union Water Framework Directive. We hypothesized that long-term averages of light-related variables would be better predictors of lake metabolism than nutrient-related variables. Model input parameters were in situ high-frequency measurements of dissolved oxygen, temperature, and irradiance. Model simulations were conducted for several (5–12) diel cycles for each lake during the summer season. Accounting for uncertainty, the results from the Bayesian model revealed that 2 lakes were autotrophic for the duration of the experiment, 1 was heterotrophic, and 5 were balanced or had an ambiguous metabolic state. Cross-comparison with a traditional bookkeeping model showed that the majority of lakes were in metabolic balance. A strong coupling between primary production and respiration was observed, with the share of autochthonous primary production respired by consumers increasing with light extinction and nutrient-related variables. Unlike gross primary production, community respiration was strongly related to light extinction, dissolved organic carbon (DOC) and total phosphorus. These findings suggest that a drastic decrease in light-limited primary production along the DOC gradient counter-balanced nutrient supply in the darker lakes and thus blurred the relationship between primary production and nutrients. Thus, contrary to our hypothesis, both light and nutrient-related variables seemed to be good predictors of lake respiration and its coupling to lake primary production.

Feld, C. K. (2016). Zustandsverbesserung in kleinen Schritten—Das gute ökologische Potenzial in landwirtschaftlich geprägten Gewässern. Korrespondenz Wasserwirtschaft 9, 606–612,


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Improvement of Conditions in Small Steps – The Good Ecological Potential in Agriculturally Characterized Bodies of Water

The large majority of surface waters in Germany fail to meet the good ecological condition required by the Water Framework Directive. Above all, the flowing waters are encumbered through the combination of diffuse material loads as well as through a hydrological and morphological transformation. Both loading factors are closely connected with agriculture. The achievement of the good ecological potential in agriculturally characterized bodies of water is, however, possible with the implementation of hydromorphological key measures. The central key measure is the establishment of riparian woodlands. These have a positive effect on the habitat availability and diversity, temper climate effects and reduce the input of nutrients, fine sediments and pesticides from agricultural areas. With the current practice in the implementation of buffer strips the establishment of riparian woodlands is, however, almost impossible. It is essential to increase both the acceptance of this key decision and also its implementation through suitable promotional instruments.

Feld, C. K., Segurado, P., Gutiérrez-Cánovas, C. (2016). Analysing the impact of multiple stressors in aquatic biomonitoring data: A 'cookbook' with applications in R. Science of the Total Environment 573, 1320-1339,


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Analysing the impact of multiple stressors in aquatic biomonitoring data: A ‘cookbook’ with applications in R

Multiple stressors threaten biodiversity and ecosystem integrity, imposing new challenges to ecosystem management and restoration. Ecosystem managers are required to address and mitigate the impact of multiple stressors, yet the knowledge required to disentangle multiple-stressor effects is still incomplete. Experimental studies have advanced the understanding of single and combined stressor effects, but there is a lack of a robust analytical framework, to address the impact of multiple stressors based on monitoring data. Since 2000, the monitoring of Europe's waters has resulted in a vast amount of biological and environmental (stressor) data of about 120,000 water bodies. For many reasons, this data is rarely exploited in the multiple-stressor context, probably because of its rather heterogeneous nature: stressors vary and are mixed with broad-scale proxies of environmental stress (e.g. land cover), missing values and zero-inflated data limit the application of statistical methods and biological indicators are often aggregated (e.g. taxon richness) and do not respond stressor-specific. Here, we present a ‘cookbook’ to analyse the biological response to multiple stressors using data from biomonitoring schemes. Our ‘cookbook’ includes guidance for the analytical process and the interpretation of results. The ‘cookbook’ is accompanied by scripts, which allow the user to run a stepwise analysis based on his/her own data in R, an open-source language and environment for statistical computing and graphics. Using simulated and real data, we show that the recommended procedure is capable of identifying stressor hierarchy (importance) and interaction in large datasets. We recommend a minimum number of 150 independent observations and a minimum stressor gradient length of 75% (of the most relevant stressor's gradient in nature), to be able to reliably rank the stressor's importance, detect relevant interactions and estimate their standardised effect size. We conclude with a brief discussion of the advantages and limitations of this protocol.

Gieswein, A. (2016). Metadata describing the biological and environmental database of the Ruhr catchment (Germany). Freshwater Metadata Journal 14, 1-7,

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Metadata describing the biological and environmental database of the Ruhr catchment (Germany)

The database of the EU project MARS contains point data of macroinvertebrates, fish and macrophytes including hydromorphological, physico-chemical and land use information. Most biotic and physico-chemical data have been collected as part of an extensive national monitoring survey. Land use infromation (ATKIS land cover data) was GIS-based generated for fixed buffer strips with different widths and lengths.

González-Bergonzoni, I., Jeppesen, E., Vidal, N., Teixeira-de Mello, F., Goyenola, G., López-Rodríguez, A., Meerhoff, M. (2016). Potential drivers of seasonal shifts in fish omnivory in a subtropical stream. Hydrobiologia 768, 183-196,

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Potential drivers of seasonal shifts in fish omnivory in a subtropical stream

The trophic structure of fish assemblages often varies seasonally, following the changes in food availability and supposedly water temperature. To unveil potential drivers of trophic shifts, we studied changes in fish trophic structure at both whole-assemblage and species levels at contrasting food availability and water temperatures in a subtropical stream. We analysed the diet of the most abundant omnivorous species (Bryconamericus iheringii) monthly along the year, searching for relationships with environmental variables changing seasonally (i.e. temperature and water level) and with fish reproductive stage. We ran a single-species food choice field experiment with fixed animal and vegetal food availability in contrasting seasons to test food availability as driver of diet shifts. At the assemblage level, we found a higher consumption of vegetal during summer, reflecting the increased proportion of vegetal in the diet of omnivores, which dominated the assemblage. At the species level, the enhanced vegetal consumption was related to increases in temperature and reduction in water level. Moreover, fish selected for vegetal during summer and for animal food in winter under experimental conditions. Our findings support the role of temperature driving food web dynamics by increasing fish herbivory towards warmer scenarios, with potential strong implications for whole-assemblage trophic structure.

Grizzetti, B., Lanzanova, D., Liquete, C., Reynaud, A., Cardoso, A. C. (2016). Assessing water ecosystem services for water resource management. Environmental Science & Policy, 61, 194-203,

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Assessing water ecosystem services for water resource management

Ecosystem service concepts can offer a valuable approach for linking human and nature, and arguments for the conservation and restoration of natural ecosystems. Despite an increasing interest in the topic, the application of these concepts for water resource management has been hampered by the lack of practical definitions and methodologies. In this study we review and analyse the current literature and propose an approach for assessing and valuing ecosystem services in the context of water management. In particular, to study the link between multiple pressures, ecological status and delivery of ecosystem services in aquatic ecosystems under different scenarios of measures or future changes. This is of interest for the development of River Basin Management Plans under the EU Water Framework Directive. We provide a list of proxies/indicators of natural capacity, actual flow and social benefit for the biophysical assessment of the ecosystem services. We advocate the use of indicators of sustainability, combining information on capacity and flow of services. We also suggest methods for economic valuation of aquatic ecosystem for each service and spatial scale of application. We argue that biophysical assessment and economic valuation should be conducted jointly to account for the different values of ecosystem services (ecologic, social and economic) and to strengthen the recognition of human dependency on nature. The proposed approach can be used for assessing the benefits of conservation and restoration of aquatic ecosystems in the implementation of the EU water policy.

Gu, J., Jin, H, He, H., Ning, X., Yu, J., Tan, B., Jeppesen, E., Li, K. (2016). Effects of small-sized crucian carp Carassius auratus on the growth of submerged macrophytes: implications for shallow lake restoration. Ecological Engineering 95, 567-573,

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Effects of small-sized crucian carp (Carassius carassius) on the growth of submerged macrophytes: Implications for shallow lake restoration

Reestablishment of submerged macrophytes is considered important when restoring shallow eutrophic lakes. To improve water clarity and consequently the growth conditions of macrophytes, removal of plankti-benthivorous fish has been used. In subtropical shallow lakes, however, rapid recruitment of small fish, especially benthivores during restoration, may hamper early reestablishment of submerged macrophytes. Crucian carp (Carassius carassius) and Vallisneria natans are common species dominated in subtropical shallow lakes. To investigate the effect of small benthivorous fish on the growth of Vallisneria natans, a 28-day outdoor controlled experiment was undertaken in 12 mesocosms with three densities of Carassius carassius − low (10 g m −2), high (40 g m −2) and no fish (all in four replicates). The results showed that the fish significantly increased chlorophyll-a concentrations and periphyton biomass in both fish treatments, most significantly at high density for Chl a. This concurs with an increase in nutrient concentrations , likely mediated by fish sediment disturbance and excretion, and a reduction of zooplankton biomass (less algal grazing). Increased concentrations of inorganic suspended solids with increasing fish density further enhanced turbidity, causing shading of the macrophytes. Accordingly, the relative growth rate, ramet number and root/leaf ratio of V. natans decreased significantly at low and high fish density compared with the controls, but the effects did not depend on fish density. However, mean leaf length rose with increasing fish density, likely to allow the plants to obtain more light. Overall, our results show that not only large-bodied carp, as demonstrated frequently, but also small-sized crucian carp posed a constraint on submerged macrophyte reestablishment, and we conclude that crucian carp may hamper restoration efforts in subtropical shallow lakes. Restoration by biomanipulation should therefore target also small-sized crucian carp.

Gu, J., Xu, Z., Ning, X., Jin, H., He, H., Yu, J., Jeppesen E., Li, K. (2016). Response of eelgrass (Vallisneria natans) to increasing nitrogen loading is depending on sediment nutrient characteristics. Water 8, 563,

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Response of eelgrass (Vallisneria natans) to increasing nitrogen loading is depending on sediment nutrient characteristics

High nitrogen (N) loading may contribute to recession of submerged macrophytes in shallow lakes; yet, its influences vary depending on environmental conditions. In August 2013, we conducted a 28-day factorial-designed field mesocosm experiment in Lake Taihu at the Taihu Laboratory for Lake Ecosystem Research (TLLER) to examine the effects of high N loading on the growth of Vallisneria natans in systems with contrasting sediment types. We ran the experiments with two levels of nutrient loading—present-day external nutrient loading (average P: 5 µg·L −1 ·day −1 , N: 130 µg·L −1 ·day −1) and P: 5 µg·L −1 ·day −1 , and with three times higher N loading (N: 390 µg·L −1 ·day −1) and used sediment with two contrasting nutrient levels. V. natans growth decreased significantly with increasing N loading, the effect being dependent, however, on the nutrient status of the sediment. In low nutrient sediment, relative growth rates, leaf biomass and root biomass decreased by 11.9%, 18.2% and 23.3%, respectively, at high rather than low N loading, while the decline was larger (44.0%, 32.7% and 41.8%, respectively) when using high nutrient sediment. The larger effect in the nutrient-rich sediment may reflect an observed higher shading of phytoplankton and excess nutrient accumulation in plant tissue, though potential toxic effects of the high-nutrient sediment may also have contributed. Our study confirms the occurrence of a negative effect of increasing N loading on submerged plant growth in shallow nutrient-enriched lakes and further shows that this effect is augmented when the plants grow in nutrient-rich sediment. External N control may, therefore, help to protect or restore submerged macrophytes, especially when the sediment is enriched with nutrients and organic matter.

Gutierrez, M.F., Devercelli, M., Brucet, S., Lauridsen, T. L., Søndergaard M., Jeppesen, E. (2016). Is recovery of large-bodied zooplankton after nutrient loading reduction hampered by climate warming? A long-term study of shallow hypertrophic Lake Søbygaard, Denmark. Water 8(8), 341,

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Nutrient Loading Reduction Hampered by Climate Warming? A Long-Term Study of Shallow Hypertrophic Lake Søbygaard, Denmark

Nutrient fluctuations and climate warming can synergistically affect trophic dynamics in lakes, resulting in enhanced symptoms of eutrophication, thereby potentially counteracting restoration measures. We performed a long-term study (23 years) of zooplankton in Danish Lake Søbygaard, which is in recovery after nutrient loading reduction, but now faces the effects of climate warming. We hypothesized that the recovery of large-bodied zooplankton after nutrient loading reduction would be hampered by climate warming through indirect effects on fish size structure. We found a shift in macrozooplankton from initial dominance of Daphnia spp. towards Bosmina spp. as well as a decline in the body size of copepods and an increase in the abundance of nauplii. These changes coincided with the increase in small sized fish as a result of rising water temperature. Despite a reduction in body size, the total biomass of cladocerans increased coinciding with a diminished fish catch per unit effort (CPUE), and likely then an overall reduction in the predation on zooplankton. A cascading effect to phytoplankton was evidenced by enhanced zooplankton:phytoplankton and cladoceran:phytoplankton ratios and a decrease in Chl-a:TP and Chl-a:TN ratios. Our results indicate that climate warming, through changes in the size structure of fish community, has major effects on zooplankton size structure. In Lake Søbygaard, the decline in zooplankton size did not prevent, but modulated, the positive cascading effect on phytoplankton through an expected diminished fish CPUE related to nutrient loading reduction.

Haande, S., Moe, S. J., Couture, R. M., (2016). Phytoplankton and other monitoring data from Lake Vansjø. Freshwater Metadata Journal 15, 1-8,

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Phytoplankton and other monitoring data from Lake Vansjø

This report gives an overview of biological and other environmental monitoring data from Lake Vansjø in the Morsa river basin district, South-East Norway. The lake is impacted by eutrophication and has been subject to several nutrient abatement measures. The dataset comprises phytoplankon data at species level from 2005-2015, and water chemistry data from the period 1980-2015. The dataset is available online from NIVA's web portal AquaMonitor and will be updated regularly with new monitoring data.

Hu, F., Bolding, K., Bruggeman, J., Jeppesen, E., Flindt, M. R., Mooij, W, M., Janse, J., Janssen, A., van Gerven, L., Trolle, D. (2016). FABM-PCLake – linking aquatic ecology with hydrodynamics. Geoscientific Model Development 9, 2271-2278, 10.5194/gmd-9-2271-2016

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FABM-PCLake – linking aquatic ecology with hydrodynamics

This study presents FABM-PCLake, a complete redesign of the PCLake aquatic ecosystem model, which we implemented into the Framework for Aquatic Biogeochemical Models (FABM). In contrast to the original model, which was designed for temperate, fully mixed freshwater lakes, the new FABM-PCLake represents an integrated aquatic ecosystem model that enables simulations of hydrodynamics and biogeochemical processes for zero dimensional, one-dimensional as well as three-dimensional heterogeneous environments. FABM-PCLake describes interactions between multiple trophic levels, including piscivorous, zooplanktivorous and benthivorous fish, zooplankton, zoobenthos, three groups of phytoplankton and rooted macrophytes. The model also accounts for oxygen dynamics and nutrient cycling for nitrogen, phosphorus and silicon, both within the pelagic and benthic domains. FABM-PCLake includes a two-way communication between the biogeochemical processes and the physics, where some biogeochemical state variables (e.g., phytoplankton) influence light attenuation and thereby the spatial and temporal distributions of light and heat. At the same time, the physical environment, including water currents, light and temperature influence a wide range of biogeochemical processes. The model enables studies on ecosystem dynamics in physically heterogeneous environments (e.g., stratifying water bodies, and water bodies with horizontal gradient in physical and biogeochemical properties), and through FABM also enables data assimilation and multi-model ensemble simulations. Examples of relevant model applications include climate change impact studies and environmental impact assessment scenarios for lakes and reservoirs worldwide.

Hu, E., He, H., Su, Y., Jeppesen, E., Liu, Z. (2016). The uses of multi-carbon sources by zooplankton in an oligotrophic lake from Tibetan Plateau. Water 8, 565,

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The uses of multi-carbon sources by zooplankton in an oligotrophic lake from Tibetan Plateau

We applied natural abundance stable isotope δ 13 C and radiocarbon ∆ 14 C analyses to investigate trophic linkages between zooplankton and their potential food sources (phytoplankton, submersed plants, and allochthonous organic carbon) in Lake Nam Co, one of the largest oligosaline and oligotrophic lakes in the Tibetan Plateau, in southwest China. The δ 13 C and ∆ 14 C levels of the calanoid copepod Arctodiaptomus altissimus pectinatus indicate that it uses different carbon sources. Thus, based on a two-isotope mixing model, our results suggested that recently synthesized but 14 C-depleted primary producers (phytoplankton and submersed plants) were the most important sources of carbon, together contributing 92.2% of the zooplankton biomass. Allochthonous organic carbon and dissolved organic carbon constituted 4.7% and 3.1% of the carbon in the diet of zooplankton, respectively. Our findings from Lake Nam Co suggest that the carbon in the food webs of lakes located in a glaciated environment originates from various sources of different ages.

Idrizaj, A., Laas, A., Anijalg, U., Nõges, P. (2016). Horizontal differences in ecosystem metabolism of a large shallow lake. Journal of Hydrology 535, 93–100,

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Horizontal differences in ecosystem metabolism of a large shallow lake

The causes of horizontal differences in metabolic activities between lake zones are still poorly understood. We carried out a two-year study of lake metabolism in two contrasting parts of a large shallow lake using the open-water technique based on high-frequency measurements of dissolved oxygen concentrations. We expected that the more sheltered and macrophyte-rich southern part of the lake receiving a high hydraulic load from the main inflow will exhibit equal or higher rate of metabolic processes compared to the open pelagic zone, and higher temporal variability, including anomalous metabolic estimates such as negative gross primary production (GPP) or community respiration (CR) due to rapid water exchange. Our results showed that anomalous metabolic estimates occurred at both stations with a similar frequency and were related rather to certain wind directions, which likely contributed to stronger water exchange between the littoral and pelagic zones. Periods of auto- and heterotrophy (daily mean NEP> or <0) had a 50:50 distribution at the Central Station while the proportions were 30:70 at the Southern Station. High areal GPP estimated in our study exceeding nearly twice the long-term average 14C primary production, showed the advantages of the free-water technique in integrating the metabolism of all communities, a large part of which has remained undetected by the traditional bottle or chamber incubation techniques.

Jeppesen, E., Trolle, D., Davidson, T. A., Bjerring, R., Søndergaard, M., Johansson, L. S., Lauridsen, T. L. (2016). Major changes in CO2 efflux when shallow lakes shift from turbid to a clear water state. Hydrobiologia 778, 33-44,

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Major changes in CO2 efflux when shallow lakes shift from a turbid to a clear water state

Lakes can be sources or sinks of carbon, depending on local conditions. Recent studies have shown that the CO2 efflux increases when lakes recover from eutrophication, mainly as a result of a reduction in phytoplankton biomass, leading to less uptake of CO2 by producers. We hypothesised that lake restoration by removal of coarse fish (biomanipulation) or invasion of mussels would have a similar effect. We studied 14–22 year time series of five temperate Danish lakes and found profound effects on the calculated CO2 efflux of major shifts in ecosystem structure. In two lakes, where limited colonisation of submerged macrophytes occurred after biomanipulation or invasion of zebra mussels (Dreissena polymorpha), the efflux increased significantly with decreasing phytoplankton chlorophyll a. In three lakes with major interannual variation in macrophyte abundance, the efflux declined with increasing macrophyte abundance in two of the lakes, while no relation to macrophytes or chlorophyll a was found in the third lake, likely due to high groundwater input to this lake. We conclude that clearing water through invasive mussels or lake restoration by biomanipulation may increase the CO2 efflux from lakes. However, if submerged macrophytes establish and form dense beds, the CO2 efflux may decline again.

Jo, H., Ventura, M., Vidal, N., Gim J.-S., Buchaca, T., Barmuta, L. A., Jeppesen, E., Joo, G.-J. (2016). Discovering hidden biodiversity: the use of complementary monitoring of fish diet based on DNA barcoding in freshwater ecosystems. Ecology and Evolution 6, 219–232,

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Discovering hidden biodiversity: The use of complementary monitoring of fish diet based on DNA barcoding in freshwater ecosystems

Ecological monitoring contributes to the understanding of complex ecosystem functions. The diets of fish reflect the surrounding environment and habitatsand may, therefore, act as useful integrating indicators of environmental status. It is, however, often difficult to visually identify items in gut contents to species level due to digestion of soft-bodied prey beyond visual recognition, but new tools rendering this possible are now becoming available. We used a molecular approach to determine the species identities of consumed diet items of an introduced generalist feeder, brown trout (Salmo trutta), in 10 Tasmanian lakes and compared the results with those obtained from visual quantification of stomach contents. We obtained 44 unique taxa (OTUs) belonging to five phyla, including seven classes, using the barcode of life approach from cytochrome oxidase I (COI). Compared with visual quantification, DNA analysis showed greater accuracy, yielding a 1.4-fold higher number of OTUs. Rarefaction curve analysis showed saturation of visually inspected taxa, while the curves from the DNA barcode did not saturate. The OTUs with the highest proportions of haplotypes were the families of terrestrial insects Formicidae, Chrysomelidae, and Torbidae and the freshwater Chironomidae. Haplotype occurrence per lake was negatively correlated with lake depth and transparency. Nearly all haplotypes were only found in one fish gut from a single lake. Our results indicate that DNA barcoding of fish diets is a useful and complementary method for discovering hidden biodiversity.

Jovanovic, B., Bezirci, G., Cagan, A. S., Coppens, J., Levi, E. E., Oluz, Z., Tuncel, E., Duran, H., Beklioglu, M. (2016). Food web effects of titanium dioxide nanoparticles in an outdoor freshwater mesocosm experiment. Nanotoxicology 10(7), 902-912,

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Food web effects of titanium dioxide nanoparticles in an outdoor freshwater mesocosm experiment

Over the course of 78 days, 9 outdoor mesocosms, each with 1350 L capacity, were situated on a pontoon platform in the middle of a lake and exposed to 0 µg L−1 TiO2, 25 µg L−1 TiO2, or 250 µg L−1 TiO2 nanoparticles in the form of E171 TiO2 human food additive five times a week. Mesocosms were inoculated with sediment, phytoplankton, zooplankton, macroinvertebrates, macrophytes, and fish before exposure, ensuring a complete food web. Physicochemical parameters of the water, nutrient concentrations, and biomass of the taxa were monitored. Concentrations of 25 µg L−1 TiO2 and 250 µg L−1 TiO2 caused a reduction in available soluble reactive phosphorus in the mesocosms by 15% and 23%, respectively, but not in the amount of total phosphorous. The biomass of Rotifera was significantly reduced by 32% and 57% in the TiO2 25 µg L−1 and TiO2 250 µg L−1 treatments, respectively, when compared to the control; however, the biomass of the other monitored groups—Cladocera, Copepoda, phytoplankton, macrophytes, chironomids, and fish—remained unaffected. In conclusion, environmentally relevant concentrations of TiO2 nanoparticles may negatively affect certain parameters and taxa of the freshwater lentic aquatic ecosystem. However, these negative effects are not significant enough to affect the overall function of the ecosystem, as there were no cascade effects leading to a major change in its trophic state or primary production.

Kangur, K., Ginter, K., Kangur, P., Kangur, A., Nõges, P., Laas, A. (2016). Changes in water temperature and chemistry preceding a massive kill of bottom-dwelling fish: An analysis of high–frequency buoy data of shallow Lake Võrtsjärv (Estonia). Inland Waters 6(4), 535-542,

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Changes in water temperature and chemistry preceding a massive kill of bottom-dwelling fish: An analysis of high–frequency buoy data of shallow Lake Võrtsjärv (Estonia)

Although massive fish kills are wide-spread and can be economically devastating, there is little information on exact causal mechanisms of fish kills in nature. In large shallow Lake Vortsjarv, sporadic fish kills have been registered mainly in cold winters, yet in 2013, an unexpected fish kill occurred beginning mid-June. At the time of the fish kill, an investigation was conducted to determine species composition, number, and sizes of dead fish along the lake shore. To determine possible causes of the fish kill, we analysed the dynamics of key physical and chemical parameters of lake water, including diurnal fluctuations of water temperature (WT), pH, dissolved oxygen (DO), ammonium ion concentrations (NH4-N), and the development of water stratification, during the growing season of 2013 using high-frequency water quality monitoring buoy and monthly manual monitoring data. Environmental data between 2010 and 2012 were used as a reference because no fish kill occurred. The results suggest that the fish kill was induced by a combination of successive and co-occurring extreme water parameters such as high WT (up to 24.5 °C), pH (up to 9.2), and NH4-N (up to 0.13 mg L⁻¹), short-term stratification, and low DO concentration in the bottom water (0.49 mg L-1, saturation 5.4%) induced by quick warming of this shallow lake after a long ice-covered period and leading to a likely ammonia poisoning and hypoxia. The main target species was the bottom-dwelling ruffe (Gymnocephalus cernuus), indicating that the summer kill started at the bottom of the lake. The event highlights the significance of short-term disturbances on fish populations, which can be detected only using high-frequency monitoring data.

Kramer, L., Kuijper, M. (2016). Metadata of the Regge & Dinkel catchment for the MARS project. Freshwater Metadata Journal 19, 1-8,

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Metadata of the Regge & Dinkel catchment for the MARS project

This data set contains an overview of biological and environmental data from the Regge and Dinkel catchment, the Netherlands. The main factors that impact this catchment are hydrological alteration, groundwater abstraction and drainage, point source and diffuse nutrient loading. The data has been collected between 2000 and 2012 by the waterboard Vechtstromen and contains macroinvertebrate, macrophyte, fish, and physico-chemical parameters. The dataset is available upon request from the waterboard Vechtstromen.

Kristensen, P. B., Riis, T., Dylmer, H. E., Kristensen, E. A., Meerhoff, M., Olesen, B., Teixeira-de Mello, F., Baattrup-Pedersen, A., Cavalli, G., Jeppesen, E. (2016). Baseline identification in stable isotope studies of temperate lotic systems and implications for calculated trophic positions. Freshwater Science 35, 909-921,

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Baseline identification in stable-isotope studiesof temperate lotic systems and implications for calculated trophic positions

Stable-isotope analysis is widely used in aquatic ecosystem studies to evaluate trophic structure and resource dynamics. Because δ15N values vary in freshwater systems, e.g., reflecting variations in land use, suitable baseline indicators must be specified. Few investigators have identified specific baseline organisms based on thorough and methodical screening. We screened for baseline organisms in temperate lotic waters based on 4 criteria: 1) baseline organisms should be easy to collect, 2) within-site variation in δ15N levels should be low, 3) δ15N should reflect land use, and 4) trophic position (TP) of consumers calculated from the baseline should be independent of system-specific δ15N variability as long as no systematic change in food consumption occurred. We investigated individual taxa and bulked groups representing different feeding modes as baselines. We found that Simuliidae, a sestonic filter feeder, fulfilled all criteria. Furthermore, TP estimates of 2 common fishes that were based on the Simuliidae or grouped filterers as baselines were the only estimates in our study that were independent of landuse changes. In addition, the diet of these fishes did not change across land use as based on stable-isotope mixing-model analysis. Simuliidae also had the lowest within-site variation, i.e., the lowest trophic level range, probably a result of uniform feeding behavior. Therefore, Simuliidae and grouped filterers could be suitable baseline indicators in future studies. We recommend minimizing δ15N variability in and among systems because the precise, complex choice, timing, or proportions of food sources consumed cannot be mimicked. We also promote combining TP estimation and mixing-model analyses as a strong tool in studies of stream food webs.

Kuha, J., Arvola, L., Hanson, P.C., Huotari, J., Huttula, T., Juntunen, J., Järvinen, M., Kallio, K., Ketola, M., Kuoppamäki, K., Lepistö, A., Lohila, A., Paavola, R., Vuorenmaa, J., Winslow, L., Karjalainen, J. (2016). Response of boreal lakes to episodic weather events. Inland Waters 6, 523-534,

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Response of boreal lakes to episodic weather events

Weather-induced episodic mixing events in lake ecosystems are often unpredictable, and their impacts are therefore poorly known. The impacts can be short-lived, including changes in water temperature and stratification, but long-lasting effects on the lake’s biology may also occur. In this study we used automated water quality monitoring (AWQM) data from 8 boreal lakes to examine how the episodic weather-induced mixing events influenced thermal structure, hypolimnetic dissolved oxygen (DO), fluorometric chlorophyll estimates (Chl-a), and lake metabolism and how these events varied in frequency and magnitude in lakes with different characteristics. Rise in wind speed alone had an effect on the lakes with the weakest thermal stability, but a decrease in air temperature together with strong wind induced mixing events in all lakes. The return period of these mixing events varied widely (from 20 to 92 d) and was dependent on the magnitude of change in weather. In lakes with strong stability, thermal structure and hypolimnetic DO concentration were only slightly affected. Weather-induced mixing in the upper water column diluted the surface water Chl-a repeatedly, whereas seasonal maximum occurred in late summer on each lake. Although Finnish lakes have been characterized with stable stratification during summer, we observed many substantial mixing events of relatively short return periods relevant to both chemical and biological properties of the lakes.

Laas, A., Cremona, F., Meinson, P., Rõõm, E.-I., Nõges, T., Nõges, P. (2016). Summer depth distribution profiles of dissolved CO2 and O2 in shallow temperate lakes reveal trophic state and lake type specific differences. Science of the Total Environment 566–567, 63–75,

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Summer depth distribution profiles of dissolved CO2 and O2 in shallow temperate lakes reveal trophic state and lake type specific differences

Knowledge about dissolved oxygen (DO) and carbon dioxide (CO2) distribution in lakes has increased considerably over the last decades. However, studies about high resolution dynamics of dissolved CO2 in different types of lakes over daily or weekly time scales are still very scarce. We measured summertime vertical DO and CO2 profiles at sub-hourly intervals during one week in eight Estonian lakes representing different lake types according to European Water Framework Directive. The lakes showed considerable differences in thermal stratification and vertical distribution of dissolved oxygen and CO2 as well as different diurnal dynamics over the measurement period. We observed a continuous CO2 supersaturation in the upper mixed layer of the alkalitrophic (calcareous groundwater-fed) lake and the dark soft-water lake showing them as CO2 emitting "chimneys" although with different underlying mechanisms. In three lake types strong undersaturation with CO2 occurred in the surface layer characterising them as CO2 sinks for the measurement period while in another three types the surface layer CO2 was mostly in equilibrium with the atmosphere. Factor analysis showed that DO% in the surface layer and the strength of its relationship with CO2% were positively related to alkalinity and negatively to trophic state and DOC gradients, whereas deeper lakes were characterised by higher surface concentration but smaller spatial and temporal variability of CO2. Multiple regression analysis revealed lake area, maximum depth and the light attenuation coefficient as variables affecting the largest number of gas regime indicators. We conclude that the trophic status of lakes in combination with type specific features such as morphometry, alkalinity and colour (DOC) determines the distribution and dynamics of dissolved CO2 and DO, which therefore may indicate functional differences in carbon cycling among lakes.

Levi, E. E., Bezirci, G., Çakıroğlu, A. İ., Turner, S., Bennion, H., Kernan, M., Jeppesen, E., Beklioğlu, M. (2016). Multi-proxy palaeoecological responses to water-level fluctuations in three shallow Turkish lakes. Palaeogeography, Palaeoclimatology, Palaeoecology 449, 553-566,

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Multi-proxy palaeoecological responses to water-level fluctuations in three shallow Turkish lakes

Natural or human-induced water-level fluctuations influence the structure and function of shallow lakes, especially in semi-arid to arid climate regions. In order to reliably interpret the effect of water-level changes from sedimentary remains in the absence of historical data, it is crucial to understand the variation in sedimentary proxies in relation to water level measurements. Here, we took advantage of existing water surface elevation data on three large shallow lakes in Turkey to elucidate the impact of lake-level changes on benthic-pelagic primary production over the last 50–100 years. Sub-fossil cladocerans, diatoms, plant remains and pigments were investigated as biological variables; X-ray fluorescence (XRF) and loss on ignition (LOI) analyses were conducted as geochemical-physical variables on a set of 210Pb and 137Cs dated cores. Dating of the cores were robust, with the exception of uncertainties in Lake Marmara littoral core due to low unsupported 210Pb activities and high counting errors. Results indicated that Lake Marmara was dominated by benthic species throughout the sediment record, while Lakes Beyşehir and Uluabat shifted from a littoral-dominated system to one with increased pelagic species abundance. In all cores there was a stronger response to longer-term (decadal) and pronounced water-level changes than to short-term (annual-biennial) and subtle changes. It was also noted that degree of alteration in proxies differed between lakes, through time and among pelagic-littoral areas, likely emphasising differences in depositional environments and/or resolution of sampling and effects of other stressors such as eutrophication. Our results highlight lake-specific changes associated with water-level fluctuations, difficulties of conducting studies at required resolution in lakes with rather mixed sediment records and complexity of palaeolimnological studies covering recent periods where multiple drivers are in force. They further emphasise the need to include instrumental records when interpreting effects of recent water-level changes from sediment core data in large shallow lakes.

Levi, P. S., Starnawski, P., Poulsen, B., Baattrup-Pedersen, A., Schramm, A., Riis, T. (2016). Microbial diversity varies with habitat complexity and biofilm function in macrophyte-rich streams. Oikos 126(3), 398-409,

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Microbial diversity varies with habitat complexity and biofilm function in macrophyte-rich streams

Biofilms in streams play an integral role in ecosystem processes and function yet few studies have investigated the broad diversity of these complex prokaryotic and eukaryotic microbial communities. Physical habitat characteristics can affect the composition and abundance of microorganisms in these biofilms by creating microhabitats. Here we describe the prokaryotic and eukaryotic microbial diversity of biofilms in sand and macrophyte habitats (i.e. epipsammon and epiphyton, respectively) in five macrophyte-rich streams in Jutland, Denmark. The macrophyte species varied in growth morphology, C:N stoichiometry, and preferred stream habitat, providing a range in environmental conditions for the epiphyton. Among all habitats and streams, the prokaryotic communities were dominated by common phyla, including Alphaproteobacteria, Bacteriodetes, and Gammaproteobacteria, while the eukaryotic communities were dominated by Stramenopiles (i.e. diatoms). For both the prokaryotes and eukaryotes, the epipsammon were consistently the most diverse communities and the epiphytic communities were generally similar among the four macrophyte species. However, the communities on the least complex macrophyte, Sparganium emersum, had the lowest richness and evenness and fewest unique OTUs, whereas the macrophyte with the most morphological complexity, Callitriche spp., had the highest number of unique OTUs. In general, the microbial taxa were ubiquitously distributed across the relatively homogeneous Danish landscape as determined by measuring the similarity among communities (i.e. Sørensen similarity index). Furthermore, we found significant correlations between microbial diversity (i.e. Chao1 rarefied richness and Pielou's evenness) and biofilm structure and function (i.e. C:N ratio and ammonium uptake efficiency, respectively); communities with higher richness and evenness had higher C:N ratios and lower uptake efficiency. In addition to describing the prokaryotic and eukaryotic community composition in stream biofilms, our study indicates that 1) physical habitat characteristics influence microbial diversity and 2) the variation in microbial diversity may dictate the structural and functional characteristics of stream biofilm communities.

Liquete, C., Cid, N., Lanzanova, D., Grizzetti, B., Reynaud, A. (2016). Perspectives on the link between ecosystem services and biodiversity: The assessment of the nursery function. Ecological Indicators 63, 249-257,

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Perspectives on the link between ecosystem services and biodiversity: The assessment of the nursery function

The relationship between biodiversity and each ecosystem service or bundle of ecosystem services (e.g. win−win, win−lose or win−neutral) is an active field of research that requires structured and consistent information. The application of that research for conservation and decision-making can be hampered by the ambiguity found in the definition of the nursery function under the ecosystem service perspective. In this paper, we review how the role of nursery habitats is included in the ecosystem services literature, covering conceptual, biophysical and economic reflections. The role of ecosystems as nurseries is mostly analyzed in coastal environments. The main observation is that there is no consensus on the consideration of the nursery function as a service (e.g. which species or habitats) or on how to assess it (e.g. which indicators or valuation methods). After that review, we analyze three different interpretations given to the nursery function, namely the ecological, conservationist and economic point of view; and we distinguish between different types of assessment that may consider the nursery function. We conclude that the nursery function can be considered an ecosystem service on its own right when it is linked to a concrete human benefit and not when it is represented with indicators of general biodiversity or ecosystem condition. Thus, the analysis of the delivery of ecosystem services should be differentiated from the analysis of ecological integrity. Only with this distinction science may be able to quantify the link between biodiversity and ecosystem services and policy may be effective in halting biodiversity loss. Similar considerations could apply for other biodiversity constituents that may be treated as ecosystem services.

Lorenz, S., Leszinski, M., Graeber, D. (2016). Meander reconnection method determines restoration success for macroinvertebrate communities in a German lowland river. International Review of Hydrobiology 101, 123-131,

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Meander reconnection method determines restoration success for macroinvertebrate communities in a German lowland river

Re-meandering of degraded rivers is a frequently implemented measure in river restoration. A simple solution is reconnection of old meanders; however, its success likely depends on the reconnection method. We conducted a field study to analyze the benefits of a fullyreconnected (fully opened meander, blocked main channel) and a partially reconnected meander (opened downstream, pipe bypass from main channel upstream, still open main channel) for macroinvertebrate communities in a German lowland river. Immediately upon reconnection of the two meanders, habitat diversity, and macroinvertebrates were recorded for three years with sampling in spring and in summer each year. The results of a principal response curve analysis show that the macroinvertebrate community in the fully reconnected meander reflected main channel reference conditions after 1.5 years. The macroinvertebrate community composition was not improved relative to in-stream reference conditions within the partially reconnected meander, which could be attributed to the almost complete lack of flow changes that resulted in missing improvements of substrate diversity. Our results show that the meander reconnection method must sufficiently affect the basic hydromorphological requirements to achieve reference macroinvertebrate community composition. Measures including hydromorphological conditions are therefore recommended for employment in environmental management.

Malve, O., Hjerppe, T., Tattari, S., Vaisanen, S., Huttunen, I., Kotamaki, N., Kallio, K., Taskinen, A., Kauppila, P. (2016). Participatory operations model for cost-efficient monitoring and modeling of river basins – A systematic approach. Science of the Total Environment 540, 79-89,

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Participatory operations model for cost-efficient monitoring and modeling of river basins – A systematic approach

The worldwide economic downturn and the climate change in the beginning of 21st century have stressed the need for cost efficient and systematic operations model for the monitoring and management of surface waters. However, these processes are still all too fragmented and incapable to respond these challenges. For example in Finland, the estimation of the costs and benefits of planned management measures is insufficient. On this account, we present a new operations model to streamline these processes and to ensure the lucid decision making and the coherent implementation which facilitate the participation of public and all the involved stakeholders. The model was demonstrated in the real world management of a lake. The benefits, pitfalls and development needs were identified. After the demonstration, the operations model was put into operation and has been actively used in several other management projects throughout Finland.

Meinson, P., Idrizaj, A., Nõges, P., Nõges, T., Laas, A. (2016). Continuous and high-frequency measurements in limnology: History, applications and future challenges. Environmental Reviews 24(1), 52-62,

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Continuous and high-frequency measurements in limnology: History, applications and future challenges

Over the past 15 years, an increasing number of studies in limnology have been using data from high-frequency measurements (HFM).This new technology offers scientists a chance to investigate lakes at time scales that were not possible earlier and in places where regular sampling would be complicated or even dangerous. This has allowed capturing the effects of episodic or extreme events, such as typhoons on lakes. In the present paper we review the various fields of limnology such as monitoring, studying highly dynamic processes, lake metabolism studies, and budget calculations where HFM has been applied, and which have benefitted most from the application. Our meta-analysis showed that more than half of the high-frequency studies from lakes were made in North-America and Europe. The main field of application has been lake ecology (monitoring, lake metabolism) followed by physical limnology. Water temperature and dissolved oxygen have been the most universal and commonly measured parameters and we review the various study purposes for which these measurements have been used. Although a considerable challenge forthe future, our review highlights that broadening the spatial scale of HFM would substantially broaden the applicability of these data across a spectrum of different fields.

Moe, S. J., Haande, S., Couture, R. M. (2016). Climate change, cyanobacteria and ecological status of lakes: A Bayesian network approach. Ecological Modelling 337, 330-347,

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Climate change, cyanobacteria and ecological status of lakes: A Bayesian network approach

Eutrophication of lakes and the risk of harmful cyanobacterial blooms due is a major challenge for management of aquatic ecosystems, and climate change is expected to reinforce these problems. Modelling of aquatic ecosystems has been widely used to predict effects of altered land use and climate change on water quality, assessed by chemistry and phytoplankton biomass. However, the European Water Framework Directive requires more advanced biological indicators for the assessment of ecological status of water bodies, such as the amount of cyanobacteria. We applied a Bayesian network (BN) modelling approach to link future scenarios of climate change and land-use management to ecological status, incorporating cyanobacteria biomass as one of the indicators. The case study is Lake Vansjø in Norway, which has a history of eutrophication and cyanobacterial blooms. The objective was (i) to assess the combined effect of changes in land use and climate on the ecological status of a lake and (ii) to assess the suitability of the BN modelling approach for this purpose. The BN was able to model effects of climate change and management on ecological status of a lake, by combining scenarios, process-based model output, monitoring data and the national lake assessment system. The results showed that the benefits of better land-use management were partly counteracted by future warming under these scenarios. Most importantly, the BN demonstrated the importance of including more biological indicators in the modelling of lake status: namely, that inclusion of cyanobacteria biomass can lower the ecological status compared to assessment by phytoplankton biomass alone. Thus, the BN approach can be a useful supplement to process-based models for water resource management.

Nõges, P., Argillier, C., Borja, Á., Garmendia J. M., Hanganu, J., Kodeš, V., Pletterbauer, F., Sagouis, A., Birk S. (2016). Quantified biotic and abiotic responses to multiple stress in freshwater, marine and ground waters. Science of the Total Environment 540, 43-52,

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Quantified biotic and abiotic responses to multiple stress in freshwater, marine and ground waters

We reviewed 219 papers and built an inventory of 532 items of ecological evidence on multiple stressor impacts in rivers, lakes, transitional and coastal waters, as well as groundwaters. Our review revealed that, despite the existence of a huge conceptual knowledge base in aquatic ecology, few studies actually provide quantitative evidence on multi-stress effects. Nutrient stress was involved in 71% to 98% of multi-stress situations in the three types of surface water environments, and in 42% of those in groundwaters. However, their impact manifested differently along the groundwater-river-lake-transitional-coastal continuum, mainly determined by the different hydro-morphological features of these ecosystems. The reviewed papers addressed two-stressor combinations most frequently (42%), corresponding with the actual status-quo of pressures acting on European surface waters as reported by the Member States in the WISE WFD Database (EEA, 2015). Across all biological groups analysed, higher explanatory power of the stress-effect models was discernible for lakes under multi-stressor compared to single stressor conditions, but generally lower for coastal and transitional waters. Across all aquatic environments, the explanatory power of stress-effect models for fish increased when multi-stressor conditions were taken into account in the analysis, qualifying this organism group as a useful indicator of multi-stress effects. In contrast, the explanatory power of models using benthic flora decreased under conditions of multiple stress.

Nõges, P., Cremona, F., Laas, A., Martma, T., Rõõm, E.-I., Toming, K., Viik, M., Vilbaste, S., Nõges, T. (2016). Role of a productive lake in carbon sequestration within a calcareous catchment. Science of the Total Environment 550, 225–230,

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Role of a productive lake in carbon sequestration within a calcareous catchment

For a long time, lakes were considered unimportant in the global carbon (C) cycle because of their small total area compared to the ocean. Over the last two decades, a number of studies have highlighted the important role of lakes in both sequestering atmospheric C and modifying the C flux from the catchment by degassing CO2 and methane and burying calcite and organic matter in the sediment. Based on a full C mass balance, high frequency measurements of lake metabolism and stable isotope analysis of a large shallow eutrophic lake in Estonia, we assess the role alkaline lakes play in augmenting the strength of terrestrial carbonate weathering as a temporary CO2 sink. We show that a large part of organic C buried in the sediments in this type of lakes originates from the catchment although a direct uptake from the atmosphere during periods of intensive phytoplankton growth in eutrophic conditions contributes to the carbon sink.

Nõges, T., Järvalt, A., Haberman, J., Zingel, P., Nõges, P. (2016). Is fish able to regulate filamentous blue-green dominated phytoplankton? Hydrobiologia 780(1), 59-69,

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Is fish able to regulate filamentous blue-green dominated phytoplankton? 

Efficient zooplankton grazing is a prerequisite for establishing a cascading food web control over phytoplankton in a lake. We studied if the top-down impact of fish could reach phytoplankton in a lake where the grazing pressure of small-sized zooplankton on filamentous phytoplankton is considered weak. We analysed >30 years of data on plankton, fish catches, hydrochemistry, hydrology, and meteorology from Võrtsjärv, a large and shallow eutrophic lake in Estonia with intensive commercial fisheries. The lake’s unregulated water level has been considered the strongest factor affecting the ecosystem through modifying sediment resuspension, internal loading of nutrients, and underwater light conditions and spawning conditions for fish. We found a negative relationship between phytoplankton biomass and pikeperch biomass indicating a potential top-down cascading effect in the food web. Top-down control of phytoplankton by zooplankton was reflected in a negative relationship between phyto- and zooplankton biomasses. A decrease of the individual weight of crustacean zooplankton with increasing biomass of small fish suggested top-down control of zooplankton by planktivorous fish. In contrast, we could not demonstrate a direct linkage between piscivorous fish and small fish. The top-down food web impact of piscivores, however, was manifested at zooplankton level in a positive correlation of pikeperch biomass with the biomass of dominating cladoceran species Bosmina coregoni and the individual weight of copepods. At high biomasses of small fish, ciliate domination over metazooplankton increased and thus enhanced the strength of the microbial food web. According to our results, fishery management measures that increase small plankti- and benthivorous fish biomass have to be avoided as they have a cascading negative effect on the ecosystem health.

Rankinen, K., Butterfield, D., Faneca Sanchez, M., Grizzetti, B., Whitehead, P., Pitkanen, T., Uusi-Kamppa, J., Leckie, H. (2016). The INCA-Pathogens model: An application to the Loimijoki River basin in Finland. Science of the Total Environment 572, 1611-1621,

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The INCA-Pathogens model: An application to the Loimijoki River basin in Finland

Good hygienic quality of surface waters is essential for drinking water production, irrigation of crops and recreation. Predictions of how and when microbes are transported by rivers are needed to protect downstream water users. In this study we tested the new process-based INCA-Pathogens model in the agricultural Loimijoki River basin (3138km(2)) in Finland, and we quantified ecosystem services of water purification and water provisioning for drinking and recreation purposes under different scenarios. INCA is a catchment scale process based model to calculate pollutant transfer from terrestrial environment and point sources to the catchment outlet. A clear gradient was observed in the numbers of faecal coliforms along the River Loimijoki. The highest bacterial counts were detected in the middle part of the main stream immediately after small industries and municipal sewage treatment plants. In terms of model performance, the INCA-Pathogen model was able to produce faecal coliform counts and seasonality both in the low pollution level sampling points and in the high pollution level sampling points. The model was sensitive to the parameters defining light decay in river water and in soil compartment, as well as to the amount of faecal coliforms in the manure spread on the fields. The modeling results showed that the number of faecal coliforms repeatedly exceeded 1000 bacteria 100ml(-1). Moreover, results lead to the following conclusions: 1) Climate change does not cause a major threat to hygienic water quality as higher precipitation increases runoff and causes diluting effect in the river, 2) Intensification of agriculture is not a threat as long as animal density remains relatively low and environmental legislation is followed, 3) More intensive agriculture without environmental legislation causes a threat especially in tributaries with high field percentage and animal density, and 4) Hygienic water quality in the River Loimijoki can best be improved by improving sewage treatment. We conclude that this catchment scale model is a useful tool for addressing catchment management and water treatment planning issues.

Rääpysjärvi, J., Hämäläinen, H., Aroviita, J. (2016). Macrophytes in boreal streams: Characterizing and predicting nativeoccurrence and abundance to assess human impact. Ecological Indicators 64, 309-318,

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Macrophytes in boreal streams: Characterizing and predicting native occurrence and abundance to assess human impact

Macrophytes are a structurally and functionally essential element of stream ecosystems and therefore indispensable in assessment, protection and restoration of streams. Modelling based on continuous environmental gradients offers a potential approach to predict natural variability of communities and thereby improve detection of anthropogenic community change. Using data from minimally disturbed streams, we described natural macrophyte assemblages in pool and riffle habitats separately and in combination, and explored their variation across large scale environmental gradients. Specifically, we developed RIVPACS-type models to predict the presence and abundance of macrophyte taxa at stream sites in the absence of human influence and, used data from impacted streams to explore the responses of three biotic indices to anthropogenic stress. The indices used, taxonomic completeness (O/E-taxa), a measure of compositional dissimilarity (BC-index) and an index taking into account the abundance of species (AB-index), are based on predicted and observed macrophyte communities. We found that size of the catchment area, altitude, latitude and percentage of lakes in the catchment were the large scale environmental variables that best predicted the natural variation of assemblages. The RIVPACS approach substantially improved both the precision and accuracy to predict the natural communities and the sensitivity to human disturbance. O/E-taxa performed best in relation to the null model decreasing the variation by 20% in pools, 29% in riffles and 32% in combined data. In general, models based on the riffle assemblages performed better than models based on pool assemblages, but including both habitats and predicting abundances instead of only presence/absence yielded the greatest accuracy and sensitivity. Our results support the use of multivariate modelling techniques in predicting reference condition to assess status of stream macrophyte communities.

Sagouis, A., Jabot, F., Argillier, C. (2016). Taxonomic versus functional diversity metrics: How do fish communities respond to anthropogenic stressors in reservoirs? Ecology of Freshwater Fish 26(4), 621-635,

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Taxonomic versus functional diversity metrics: How do fish communities respond to anthropogenic stressors in reservoirs?

Biological indicators are frequently used to assess the effects of anthropogenic stressors on freshwater ecosystems. The structure of fish communities and their response to stressors have been commonly described by taxonomic richness, diversity and evenness. More recently, functional structure of communities has also been suggested to be a reliable indicator of disturbance. This article aimed at testing whether taxonomic and functional diversity metrics can provide comparable or complementary information on the response of fish communities to eutrophication and abundance of non-native species in reservoirs. Comparison of the responses of taxonomic and functional diversities to biogeography, habitat and stressors was made in 112 French reservoirs. Widely observed effects of biogeographic and habitat variables on taxonomic and functional diversities were identified. Taxonomic and functional richness metrics notably increased with lake area and temperature respectively. Taxonomic diversity metrics did not respond to any stressor, while all functional diversity metrics were found to be impacted by non-native species. Eutrophication was further found to decrease the impact of non-native species on two functional diversity metrics: evenness and divergence. Our study therefore reveals that functional metrics are more sensitive than taxonomic metrics to anthropogenic stressors in the studied reservoirs. Still, the multiple linear regressions tested had overall low explanatory power. Further refinements will thus be needed to use this type of metrics in an impact assessment scheme.

Schinegger, R., Palta, M., Segurado, P., Schmutz, S. (2016). Untangling the effects of multiple human stressors and their impacts on fish assemblages in European running waters. Sciene of the Total Environment, 573, 1079-1088,

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Untangling the effects of multiple human stressors and their impacts on fish assemblages in European running waters

This work addresses human stressors and their impacts on fish assemblages at pan-European scale by analysing single and multiple stressors and their interactions. Based on an extensive dataset with 3105 fish sampling sites, patterns of stressors, their combination and nature of interactions, i.e. synergistic, antagonistic and additive were investigated. Geographical distribution and patterns of seven human stressor variables, belonging to four stressor groups (hydrological-, morphological-, water quality- and connectivity stressors), were examined, considering both single and multiple stressor combinations. To quantify the stressors' ecological impact, a set of 22 fish metrics for various fish assemblage types (headwaters, medium gradient rivers, lowland rivers and Mediterranean streams) was analysed by comparing their observed and expected response to different stressors, both acting individually and in combination. Overall, investigated fish sampling sites are affected by 15 different stressor combinations, including 4 stressors acting individually and 11 combinations of two or more stressors; up to 4 stressor groups per fish sampling site occur. Stressor-response analysis shows divergent results among different stressor categories, even though a general trend of decreasing ecological integrity with increasing stressor quantity can be observed. Fish metrics based on density of species ‘intolerant to water quality degradation’ and ‘intolerant to oxygen depletion” responded best to single and multiple stressors and their interactions. Interactions of stressors were additive (40%), synergistic (30%) or antagonistic (30%), emphasizing the importance to consider interactions in multi-stressor analyses. While antagonistic effects are only observed in headwaters and medium-gradient rivers, synergistic effects increase from headwaters over medium gradient rivers and Mediterranean streams to large lowland rivers. The knowledge gained in this work provides a basis for advanced investigations in European river basins and helps prioritizing further restoration and management actions.

Schülting, L., Feld, C. K., Graf, W. (2016). Effects of hydro- and thermopeaking on benthic macroinvertebrate drift. Science of the Total Environment 61, 194-203,

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Effects of hydro- and thermopeaking on benthic macroinvertebrate drift

The operation of storage hydropower plants is commonly linked to frequent fluctuations in discharge and water level (hydropeaking) of downstream river stretches and is often accompanied by cooling or warming of the water body downstream (cold or warm thermopeaking, respectively). The objective of this study is to assess the single and combined effects of hydropeaking and cold thermopeaking on the drift of selected aquatic macroinvertebrates in experimental flumes. The study specifically aims to (1) investigate the macroinvertebrate drift induced by hydropeaking, (2) identify taxon-specific drift patterns following combined hydropeaking and cold thermopeaking and (3) quantify diurnal drift differences under both impact types. Overall, hydropeaking induced significantly higher drift rates of most macroinvertebrate taxa. Combined hydropeaking and cold thermopeaking, however, revealed reduced total drift rates, however with strong taxon-specific response patterns. Hydropeaking during night led to significantly higher drift rates than during daytime, while in combination with thermopeaking the same trend was observable, although insignificant. Taxon-specific analysis revealed lower drift rates following hydropeaking for rheophilic and interstitial taxa (e.g. Leuctra sp., Hydropsyche sp.), whereas many limnophilic taxa adapted to low current showed markedly increased drift (e.g. Lepidostoma hirtum and Leptoceridae). In line with previous studies, our results confirm a significant loss of limnophilic macroinvertebrate taxa following hydraulic stress. The mitigating effect of cold thermopeaking might be explained by behavioural patterns, but requires further investigation to clarify if macroinvertebrates actively avoid drift and intrude into the interstitial, when cold water is discharged. Our results imply that river restoration projects must address the hydrological regime and, if necessary need to include suitable management schemes for hydropower plants. Besides operative management measures, the construction of reservoirs to buffer hydropeaks or the diversion of hydropeaks into larger water bodies could mitigate hydropeaking effects and foster biological recovery including limnophilic taxa.

Segurado, P., Branco, P., Jauch, E., Neves, R., Ferreira, M. T. (2016). Sensitivity of river fishes to climate change: the role of hydrological stressors on habitat range shifts. Science of the Total Environment 562, 435–445,

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Sensitivity of river fishes to climate change: the role of hydrological stressors on habitat range shifts

Climate change will predictably change hydrological patterns and processes at the catchment scale, with impacts on habitat conditions for fish. The main goal of this study is to assess how shifts in fish habitat favourability under climate change scenarios are affected by hydrological stressors. The interplay between climate and hydrological stressors has important implications in river management under climate change because management actions to control hydrological parameters are more feasible than controlling climate. This study was carried out in the Tamega catchment of the Douro basin. A set of hydrological stressor variables were generated through a process-based modelling based on current climate data (2008–2014) and also considering a high-end future climate change scenario. The resulting parameters, along with climatic and site-descriptor variables were used as explanatory variables in empirical habitat models for nine fish species using boosted regression trees. Models were calibrated for the whole Douro basin using 254 fish sampling sites and predictions under future climate change scenarios were made for the Tamega catchment. Results show that models using climatic variables but not hydrological stressors produce more stringent predictions of future favourability, predicting more distribution contractions or stronger range shifts. The use of hydrological stressors strongly influences projections of habitat favourability shifts; the integration of these stressors in the models thinned shifts in range due to climate change. Hydrological stressors were retained in the models for most species and had a high importance, demonstrating that it is important to integrate hydrology in studies of impacts of climate change on freshwater fishes. This is a relevant result because it means that management actions to control hydrological parameters in rivers will have an impact on the effects of climate change and may potentially be helpful to mitigate its negative effects on fish populations and assemblages.

Søndergaard, M., Larsen, S. E., Johansson, L. S., Lauridsen T. L., Jeppesen, E. (2016). Ecological classification of lakes: Uncertainty and the influence of year-to-year variability. Ecological Indicators 61, 248- 257,

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Ecological classification of lakes: Uncertainty and the influence of year-to-year variability

Regular monitoring of lakes is important to determine their ecological state and development and of key significance when deciding whether action should be taken to improve their quality, for instance by reducing the external loading of nutrients. Imprecise or inadequate knowledge of the ecological state increases the risk of misclassification and of wrong management decisions. Based on Danish lake data, we aimed to determine temporal variations, in particular natural year-to-year differences, and to describe the uncertainty in assessing the ecological state of lakes. We analysed environmental data from ca. 350 Danish lakes (1100 lake years), including three case studies, with long-term data series (up to 24 years), with no significant changes in external nutrient loading. We used summer means of selected water chemical variables, phytoplankton and submerged macrophytes as indicators of ecological state and found considerable variations in all indicators, which could not be ascribed alone to meteorological variation. In shallow lakes, chlorophyll a concentrations exhibited large year-to-year variations, especially at TP ranging between 0.05 and 0.15 mg L −1 where the lakes may shift between a macrophyte-and a phytoplankton-dominated state. For example, chlorophyll a varied by a factor 5–10 between years and was particularly low when submerged macrophyte coverage exceeded 20% compared with lakes without macrophytes. Use of a multimetric index including four phytoplankton indicators reduced the coefficient of variation. Generally, the 95% confidence interval of ecological classification was approximately 50% lower when the assessment of ecological state was based on 4–5 years' measurements than if based on only one year's measurements. Knowledge and awareness of the uncertainty of indicators used in ecological classification are highly relevant for lake managers and policy makers when defining efficient monitoring and restoration strategies.

Stefanidis, K., Panagopoulos, Y., Mimikou, M. (2016). Impact assessment of agricultural driven stressors on benthic macroinvertebrates using simulated data. Science of the Total Environment 540, 32-42,

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Impact assessment of agricultural driven stressors on benthic macroinvertebrates using simulated data

Agricultural land use poses a significant threat to the ecological integrity of rivers in Europe. Particularly in the Mediterranean, water abstraction and nutrient application are anthropogenic pressures that have a significant impact on aquatic habitats and biodiversity. In this article, we assessed the effects of agricultural management practices on benthic macroinvertebrates in a large river basin of central Greece using simulated data based on the application of SWAT (Soil Water Assessment Tool) model. Physicochemical and hydrological output variables of the model were used as predictors of the ASPT (Average Score Per Taxon) metric based on a correlated component regression analysis (CCR) built on empirical data. The estimation of ASPT was performed for the wet and dry seasons within a 20-year period for a total of 47 subbasins under the baseline conditions and after implementing three management scenarios that reduced: a) irrigation water applied to crops by 30%, b) chemical fertilization applied to crops by 30% and c) both irrigation and fertilization by 30%. The results revealed that application of the reduced irrigation resulted to a slight increase of the simulated dissolved inorganic nitrogen concentration (DIN), which in turn decreased the mean ASPT in 21 of the 47 subbasins implying a negative effect on the macroinvertebrate communities. On the contrary, the reduction of fertilization as well as the combined scenario decreased both the simulated DIN and phosphate concentration causing an increase of the mean ASPT for a total of 40 of the 47 subbasins. Based on these results, we suggest that the best management option is a combined practice of deficit irrigation and fertilization reduction since it improved water quality, increased ASPT values and saved a considerable amount of water. Overall, this work demonstrates a simple methodology that can efficiently assess the effects of agricultural management practices on biotic indicators.

Stefanidis, K., Panagopoulos, Y., Psomas, A., Mimikou, M. (2016). Assessment of the natural flow regime in a Mediterranean river impacted from irrigated agriculture. Science of the Total Environment 573, 1492-1502,

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Assessment of the natural flow regime in a Mediterranean river impacted from irrigated agriculture

Over the last few decades, the natural flow regime of most rivers has been significantly altered influencing the ecological integrity and functioning of river ecosystems. Especially in the Mediterranean region, irrigated agriculture is considered one of the most important drivers of hydro-morphological modifications in river systems. In this study we employ the Indicators of Hydrologic Alteration (IHA) methodology for the Pinios River and its tributaries, located in a Mediterranean catchment in central Greece, with the purpose to assess the natural flow regime under a simulated no-agriculture scenario and compare with the current situation. The work is based on the use of the SWAT (Soil Water Assessment Tool) model for the simulation of long time series of daily stream flows, which were analyzed under the actual conditions (baseline), and the hypothetical scenario. The key characteristics of the flow regime projected under each model run were assessed through the implementation of the IHA methodology that utilizes a number of indicators to characterize the intra- and inter-annual variability in the hydrologic conditions. The results of this study revealed that without agricultural activities in the catchment, annual and monthly flows would increase, with significant alterations in the flow characteristics of the winter months, and much smaller in summer. However, the analysis showed that the frequency of droughts and low flow summer events would be smaller. The article provides a comprehensive and easy-to-implement methodology that can facilitate the impact assessment of agricultural human activities on river flow variability under the typical Mediterranean conditions, allowing experimentation on setting river flow thresholds required for a good ecological status within the context of the European Water Framework Directive.

Taipale, S. J., Vuorio, K., Strandberg, U., Kahilainen, K. K., Jarvinen, M., Hiltunen, M., Peltomaa, E., Kankaala, P. (2016). Lake eutrophication and brownification downgrade availability and transfer of essential fatty acids for human consumption. Environment International 96, 156-166,

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Lake eutrophication and brownification downgrade availability and transfer of essential fatty acids for human consumption

Fish are an important source of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) for birds, mammals and humans. In aquatic food webs, these highly unsaturated fatty acids (HUFA) are essential for many physiological processes and mainly synthetized by distinct phytoplankton taxa. Consumers at different trophic levels obtain essential fatty acids from their diet because they cannot produce these sufficiently de novo. Here, we evaluated how the increase in phosphorus concentration (eutrophication) or terrestrial organic matter inputs (brownification) change EPA and DHA content in the phytoplankton. Then, we evaluated whether these changes can be seen in the EPA and DHA content of piscivorous European perch (Perca fluviatilis), which is a widely distributed species and commonly consumed by humans. Data from 713 lakes showed statistically significant differences in the abundance of EPA- and DHA-synthesizing phytoplankton as well as in the concentrations and content of these essential fatty acids among oligo-mesotrophic, eutrophic and dystrophic lakes. The EPA and DHA content of phytoplankton biomass (mg HUFA g− 1) was significantly lower in the eutrophic lakes than in the oligo-mesotrophic or dystrophic lakes. We found a strong significant correlation between the DHA content in the muscle of piscivorous perch and phytoplankton DHA content (r = 0.85) as well with the contribution of DHA-synthesizing phytoplankton taxa (r = 0.83). Among all DHA-synthesizing phytoplankton this correlation was the strongest with the dinoflagellates (r = 0.74) and chrysophytes (r = 0.70). Accordingly, the EPA + DHA content of perch muscle decreased with increasing total phosphorus (r2 = 0.80) and dissolved organic carbon concentration (r2 = 0.83) in the lakes. Our results suggest that although eutrophication generally increase biomass production across different trophic levels, the high proportion of low-quality primary producers reduce EPA and DHA content in the food web up to predatory fish. Ultimately, it seems that lake eutrophication and brownification decrease the nutritional quality of fish for human consumers.

Teichert, N., Borja, A., Chust, G., Uriate, A., Lepage, M. (2016). Restoring fish ecological quality in estuaries: Implication of interactive and cumulative effects among anthropogenic stressors. Science of the Total Environment 542, 383-393,

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Restoring fish ecological quality in estuaries: Implication of interactive and cumulative effects among anthropogenic stressors

Estuaries are subjected to multiple anthropogenic stressors, which have additive, antagonistic or synergistic effects. Current challenges include the use of large databases of biological monitoring surveys (e.g. the European Water Framework Directive) to help environmental managers prioritizing restoration measures. This study investigated the impact of nine stressor categories on the fish ecological status derived from 90 estuaries of the North East Atlantic countries. We used a random forest model to: 1) detect the dominant stressors and their non-linear effects; 2) evaluate the ecological benefits expected from reducing pressure from stressors; and 3) investigate the interactions among stressors. Results showed that largest restoration benefits were expected when mitigating water pollution and oxygen depletion. Non-additive effects represented half of pairwise interactions among stressors, and antagonisms were the most common. Dredged sediments, flow changes and oxygen depletion were predominantly implicated in non-additive interactions, whereas the remainder stressors often showed additive impacts. The prevalence of interactive impacts reflects a complex scenario for estuaries management; hence, we proposed a step-by-step restoration scheme focusing on the mitigation of stressors providing the maximum of restoration benefits under a multi-stress context.

Thomas, S. M., Griffiths, S. W., Ormerod, S. (2016). Beyond cool: Adapting upland streams for climate change using riparian woodlands. Global Change Biology 22, 310-324,

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Beyond cool: Adapting upland streams for climate change using riparian woodlands

Managed adaptation could reduce the risks of climate change to the world’s ecosystems, but there have been surprisingly few practical evaluations of the options available. For example, riparian woodland is advocated widely as shade to reduce warming in temperate streams, but few studies have considered collateral effects on species composition or ecosystem functions. Here, we use cross sectional analyses at two scales (region and within streams) to investigate whether four types of riparian management, including those proposed to reduce potential climate change impacts, might also affect the composition, functional character, dynamics and energetic resourcing of macroinvertebrates in upland Welsh streams (UK). Riparian land use across the region had only small effects on invertebrate taxonomic composition, while stable isotope data showed how energetic resources assimilated by macroinvertebrates in all functional guilds were split roughly 50:50 between terrestrial and aquatic origins irrespective of riparian management. Nevertheless, streams draining the most extensive deciduous woodland had the greatest stocks of coarse particulate matter (CPOM) and greater numbers of “shredding” detritivores. Stream-scale investigations showed that macroinvertebrate biomass in deciduous woodland streams was around twice that in moorland streams, and lowest of all in streams draining non-native conifers. The unexpected absence of contrasting terrestrial signals in the isotopic data implies that factors other than local land use affect the relative incorporation of allochthonous subsidies into riverine food webs. Nevertheless, our results reveal how planting deciduous riparian trees along temperate headwaters as an adaptation to climate change can modify macroinvertebrate function, increase biomass and potentially enhance resilience by increasing basal resources where cover is extensive (>60m riparian width). We advocate greater urgency in efforts to understand the ecosystem consequences of climate change adaptation in order to guide future actions.

Toming, K., Kutser, T., Tuvikene, L., Viik, M., Nõges, T. (2016). Dissolved organic carbon and its potential predictors in large shallow eutrophic temperate lake. Water Research 102, 32-40,

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Dissolved organic carbon and its potential predictors in large shallow eutrophic temperate lake

Understanding of the true role of lakes in the global carbon cycle requires reliable estimates of dissolved organic carbon (DOC) and there is a strong need to develop remote sensing methods for mapping lake carbon content at larger regional and global scales. Part of DOC is optically inactive. Therefore, lake DOC content cannot be mapped directly. The objectives of the current study were to estimate the relationships of DOC and other water and environmental variables in order to find the best proxy for remote sensing mapping of lake DOC. The Boosted Regression Trees approach was used to clarify in which relative proportions different water and environmental variables determine DOC. In a studied large and shallow eutrophic lake the concentrations of DOC and coloured dissolved organic matter (CDOM) were rather high while the seasonal and interannual variability of DOC concentrations was small. The relationships between DOC and other water and environmental variables varied seasonally and interannually and it was challenging to find proxies for describing seasonal cycle of DOC. Chlorophyll a (Chl a), total suspended matter and Secchi depth were correlated with DOC and therefore are possible proxies for remote sensing of seasonal changes of DOC in ice free period, while for long term interannual changes transparency-related variables are relevant as DOC proxies. CDOM did not appear to be a good predictor of the seasonality of DOC concentration in Lake Võrtsjärv since the CDOM–DOC coupling varied seasonally. However, combining the data from Võrtsjärv with the published data from six other eutrophic lakes in the world showed that CDOM was the most powerful predictor of DOC and can be used in remote sensing of DOC concentrations in eutrophic lakes.

Turunen, J., Muotka, T., Vuori, K-M., Karjalainen, S.M., Rääpysjärvi, J., Sutela, T., Aroviita, J. (2016). Disentangling the responses of boreal stream assemblages to low stressor levels of diffuse pollution and altered channel morphology. Science of the Total Environment 544, 954–962,

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Disentangling the responses of boreal stream assemblages to low stressor levels of diffuse pollution and altered channel morphology

Non-point diffuse pollution from land use and alteration of hydromorphology are among the most detrimental stressors to stream ecosystems. We explored the independent and interactive effects of morphological channel alteration (channelization for water transport of timber) and diffuse pollution on species richness and community structure of four organism groups in boreal streams: diatoms, macrophytes, macroinvertebrates, and fish. Furthermore, the effect of these stressors on stream condition was evaluated by Ecological Quality Ratios (EQR) from the national Water Framework Directive (WFD) assessment system. We grouped 91 study sites into four groups that were impacted by either diffuse pollution or hydromorphological alteration, by both stressors, or by neither one. Macroinvertebrate richness was reduced by diffuse pollution, whereas other biological groups were unaltered. Hydromorphological modification had no effect on taxon richness of any of the assemblages. Community structure of all groups was significantly affected by diffuse pollution but not by hydromorphology. Similarly, EQRs indicated negative response by diatoms, macroinvertebrates and fish to diffuse pollution, but not to hydromorphological alteration. Agricultural diffuse pollution thus affected species identities and abundances rather than taxonomic richness. Our results suggest that channelization of boreal streams for timber transport has not altered hydromorphological conditions sufficiently to have a strong impact on stream biota, whereas even moderate nutrient enrichment may be ecologically harmful. Controlling diffuse pollution and associated land use stressors should be prioritized over restoration of in-stream habitat structure to improve the ecological condition of boreal streams.

Ventelä, A.-M., Amsinck, S.L., Kauppila, T., Johansson, L. S., Jeppesen, E., Kirkkala, T., Søndergaard, M., Weckström, J., Sarvala, J. (2016). Ecosystem changes in large and shallow Lake Säkylän Pyhäjärvi, Finland, during the past 400 years - implications for management. Hydrobiologia 778, 273-294,

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Ecosystem change in the large and shallow Lake Säkylän Pyhäjärvi, Finland, during the past ~400 years: implications for management

Lake Säkylän Pyhäjärvi has been an important fishing site and drinking water source for the local population for centuries. The lake has undergone significant changes: (1) the water level was lowered in the 1600s and in the 1850s; (2) planktivorous coregonid fish were successfully introduced in the early 1900s; (3) nutrient input from intensified agriculture has increased since the 1950s and (4) the effects of the current variable climate on the lake and its catchment have become more evident since the 1990s. We determined the phases of oligotrophication, eutrophication and recovery and elucidated the ecosystem changes by combining palaeolimnological records with detailed neolimnological data. The sedimentary diatom and cladoceran assemblages first showed a relatively eutrophic period followed by oligotrophic periods, linked with the artificial changes in water level and consequent shifts in macrophyte abundance. The oligotrophic period in the early 1900s is thought to represent the target trophic state for the lake. After the 1950s, introduction of vendace resulted in higher planktivory reflected by an increased relative abundance of small-bodied pelagic cladocerans. Signs of eutrophication occurred due to increased nutrient load. During the last 10 years, signs of recovery have been recorded. A complex history such as that of Lake Pyhäjärvi illustrates the difficulties in selecting management targets, and the risk of setting false targets, for lakes based solely on monitoring data—both neolimnological and palaeolimnological approach are needed.

Verberk, W. C. E. P., Durance, I., Vaughan, I. P., Ormerod, S. J. (2016). Field and laboratory studies reveal interacting effects of stream oxygenation and warming on aquatic ectotherms. Global Change Biology 22, 1769–1778,

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Field and laboratory studies reveal interacting effects of stream oxygenation and warming on aquatic ectotherms

Aquatic ecological responses to climatic warming are complicated by interactions between thermal effects and other environmental stressors such as organic pollution and hypoxia. Laboratory experiments have demonstrated how oxygen limitation can set heat tolerance for some aquatic ectotherms, but only at unrealistic lethal temperatures and without field data to assess whether oxygen shortages might also underlie sub-lethal warming effects. Here we test whether oxygen availability affects both lethal and non-lethal impacts of warming on two widespread Eurasian mayflies, Ephemera danica Müller 1764 and Serratella ignita (Poda 1761). Mayfly nymphs are often a dominant component of the invertebrate assemblage in streams, and play a vital role in aquatic and riparian food webs. In the laboratory, lethal impacts of warming were assessed under three oxygen conditions. In the field, effects of oxygen availability on non-lethal impacts of warming were assessed from mayfly occurrence in 42,293 UK stream samples where water temperature and biochemical oxygen demand were measured. Oxygen limitation affected both lethal and sub-lethal impacts of warming in each species. Hypoxia lowered lethal limits by 5.5 (±2.13) °C and 8.2 (±0.62) °C for E. danica and S. ignita respectively. Field data confirmed the importance of oxygen limitation in warmer waters; poor oxygenation drastically reduced site occupancy, and reductions were especially pronounced under warm water conditions. Consequently, poor oxygenation lowered optimal stream temperatures for both species. The broad concordance shown here between laboratory results and extensive field data suggests that oxygen limitation not only limits survival at thermal extremes but also restricts species abundance in the field at temperatures well below upper lethal limits. Stream oxygenation could thus control the vulnerability of aquatic ectotherms to global warming. Improving water oxygenation and reducing pollution can provide key facets of climate change adaptation for running waters.

Vilbaste, S., Järvalt, A., Kalpus, K., Nõges, T., Pall, P., Piirsoo, K., Tuvikene, L. and Nõges, P. (2016). Ecosystem services of Lake Võrtsjärv under multiple stress: A case study. Hydrobiologia 780(1), 145-159,

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Ecosystem services of Lake Võrtsjärv under multiple stress: A case study

This study is the first attempt at the European scale to make an inventory of ecosystem services (ESS) of a large lake. We analysed a set of ESS indicators against the annual mean values of environmental parameters for 2006–2013. According to principal component analysis, the trophic state- and hydrology-related factors explained about 70% of the environmental variability of the lake and showed strong relationships with some ESS. Among the provisioning ESS, the annual eel catch and the total fish catch were positively related to different eutrophication indicators while the catches of pike, bream, and burbot depended rather on hydrological factors. Reed harvesting efficiency was related to the lake’s water level. The indicators of regulating, maintenance, and cultural ESS showed very high variability in different years, the latter depending on socio-economic conditions rather than environmental factors. We discovered numerous trade-offs between ESS benefitting from higher trophic state or regulated water level of the lake and the goals of good ecological status of the lake. Our analysis showed a clear need for rules prioritizing life supporting regulatory services against other ESS.

Vilmi, A., Karjalainen, S. M., Hellsten, S., Heino, J. (2016). Bioassessment in a metacommunity context: Are diatom communities structured solely by species sorting? Ecological Indicators 62, 86–94,

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Bioassessment in a metacommunity context: Are diatom communities structured solely by species sorting?

Aquatic ecosystems face a variety of anthropogenic pressures, urging the development of efficient biological indicators. In addition to local environmental conditions, the community structure of indicator organisms is affected by spatial processes, such as high and limited dispersal rates. Understanding the relative roles of environmental factors and spatial processes for ecological communities should thus be associated with bioassessment practices. We examined the main drivers, both environmental and spatial, influencing community structure and several indices derived from diatom communities. We sampled 81 stony littoral sites in a large boreal lake system (305 km 2), where relatively large gradients in water chemistry (35 variables measured) exist, but no dispersal limitation can be expected. Instead, high dispersal rates should interfere with species sorting. Our response variables, including commonly-used diatom indices, diversity indices and taxonomic distinctness indices, were better explained by pure effects of spatial variables and shared effects of spatial and environmental variables than by pure effects of environmental variables. Thus, high dispersal rates between sites are likely to interfere with environmental filtering and can result in clear spatial structures in index values used in bioassessment. Bioassessment should thus acknowledge the importance of spatial processes and not take it for granted that only local environmental conditions determine index values. Failure to consider high dispersal rates may lead to biased information about the state of freshwater ecosystems. The same idea should also be considered in systems with similarly highly-connected sets of bioassessment sites, such as marine coastal systems and stream networks.

Wiegleb, G., Gebler, D., de Weyer, K., Birk, S. (2016). Comparative test of ecological assessment methods of lowland streams based on long-term monitoring data of macrophytes. Science of the Total Environment 541, 1269–1281,

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Comparative test of ecological assessment methods of lowland streams based on long-term monitoring data of macrophytes

Ecological assessment of water courses is required by the European Water Framework Directive (WFD). Assessment by means of macrophytes is impeded by insufficient knowledge on the relations between assessment scores and the dynamics of environmental parameters. Data from a long-term observation of macrophyte dynamics over 21years in two lowland rivers were used for testing the performance of six widely used assessment methods. Six sample sites situated in two lowland streams were selected. Four sites were classified as of moderate habitat quality and two sites as of poor habitat quality in the context of WFD. Assessment methods generally showed a poor performance in recognizing the ecological status of the annual observations. Status was more often over- than underestimated. Performance of methods differed among individual rivers and among river zones. Assessment scores mostly showed a steady decline, even though all sites obviously remained in the same habitat quality class throughout the observation period. Variation of most environmental factors was largely unrelated to assessment scores. Fluctuations of assessment scores were partly related to single natural disturbance events such as high discharge. Increased shading by marginal trees was reflected negatively by most assessment scores. Assessment scores were highly correlated with species richness and total abundance. The best overall performance was shown by the North-Rhine Westphalian (NRW) method. In contrast to single metric methods it can be adapted to individual properties of a reach in a flexible way. Macrophyte assessment based on the pressure-impact framework did not lead to a satisfying result in our case study. Improvement of species assessment scores and inclusion of functional properties such as growth form may help to overcome the present difficulties.

Yu, J., Liu, Z., He, H., Zhen, W., Guan, B., Chen, F., Li, K., Zhong, P., Teixeira-de Mello, F., Jeppesen, E. (2016). Submerged macrophytes facilitate the omnivorous dominance of fish community in a subtropical shallow lake: Implication for shallow lake restoration. Hydrobiologia 775, 97-107,

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Submerged macrophytes facilitate dominance of omnivorous fish in a subtropical shallow lake: implications for lake restoration

Biomanipulation based on removal of coarse fish, piscivorous fish stocking and sometimes also planting of submerged macrophytes has been used to restore temperate eutrophic shallow lakes. However, in warmer lakes, omnivorous fish are more abundant and apparently less well controlled by the piscivores. We investigated the food web structure and energy pathways of fish in the restored part of subtropical Lake Wuli, China, using gut contents analysis (GCA) and the IsoSource model based on stable isotope analysis (SIA) data. We found that omnivores dominated the fish community in terms of numbers. GCA showed that cyclopoid copepods constituted the main food item for the planktivores, while all adult omnivorous fish fed mainly on macrophytes. The IsoSource SIA model supported these results. Furthermore, piscivores consumed shrimps rather than juvenile omnivores, and the SIA analysis revealed no trophic links between piscivores and adult omnivores or zooplanktivores. We conclude that macrophytes constituted an important food item for omnivores, potentially promoting population growth of omnivores as control by piscivores was weak. This may yield a high predation pressure on both zooplankton and on macrophytes, possibly preventing the establishment of a stable macrophyte state following restoration of eutrophic lakes unless the fish density is regularly controlled.

Yu, J., Liu, Z., Li, K., Chen, F., Guan, B., Hu, Y., Zhong, P., Tang, Y., Zhao, X., He, H., Zeng, H., Jeppesen, E. (2016). Restoration of Shallow Lakes in Subtropical and Tropical China: Response of Nutrients and Water Clarity to Biomanipulation by Fish Removal and Submerged Plant Transplantation. Water 8(10), 438,

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Restoration of Shallow Lakes in Subtropical and Tropical China: Response of Nutrients and Water Clarity to Biomanipulation by Fish Removal and Submerged Plant Transplantation

Fish removal has been used to restore temperate lakes, and positive effects on ecological state and water clarity have frequently been recorded in many lakes. Recently, a supplementary measure, transplantation of submerged macrophytes after fish removal, has been applied to restore warm Chinese shallow lakes in order to compensate for the expected lack of increasing grazing control of phytoplankton after the biomanipulation. These measures have successfully shifted turbid warm lakes to a clear water state, but little is known about the responses to restoration of key physico-chemical variables. We analyzed the seasonal variation in nutrient concentrations in two subtropical and one tropical biomanipulated shallow Chinese lakes subjected to restoration. In all three lakes, a marked decline occurred in the concentrations of lake total nitrogen (TN), total phosphorus (TP), total suspended solids (TSS), and chlorophyll a (Chl a), while the transparency (SD:WD ratio, Secchi depth to water depth ratio) increased. A clear water state was established, lasting so far for 7 to 23 months, and TN, TP, Chl a, and TSS levels in the three restored lakes decreased to, on average, 49%, 58%, 41%, and 18% of the level prior to restoration and/or the level in a reference lake, respectively, while the annual mean SD:WD ratio exhibited a 1.5–4 fold increase. In conclusion, lake restoration by transplantation of submerged macrophytes after fish removal had major positive effects on the physico-chemical variables in our study lakes. However, continuous control of omnivorous and herbivorous fish biomass is recommended as the fish typically present in warm, shallow lakes to some extent feed on submerged macrophytes, when available.

Yu, J., Zhen, W., Guan, B., Zhong, P., Jeppesen, E., Liu, Z. (2016). Dominance of Myriophyllum spicatum in submerged macrophyte communities associated with grass carp. Knowledge and Management of Aquatic Ecosystems 417, 24,

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Dominance of Myriophyllum spicatum in submerged macrophyte communities associated with grass carp

Re-establishment of macrophyte communities is a key to restore eutrophic shallow lakes. The species com- position of the plant communities may change rapidly during the recovery period. A basin in subtropical Lake Qinhu (China) was restored by biomanipulation including fish removal followed by planting of submerged macrophytes in 2011. In September-December 2011, dominance of Vallisneria spinulosa and Ceratophyllum demersum shifted to dom- inance of Myriophyllum spicatum. Meanwhile, the CPUE (catch per unit effort) showed that the number and biomass of grass carp (Ctenopharyngodon idella) demonstrated a marked increase. Stable isotope analyses revealed that grass carp preferred V. spinulosa and C. demersum to M. spicatum. We propose that grazing by grass carp was responsible for the observed shift in dominance towards M. spicatum but as alternative explanations are possible, further tests by controlled experiments are needed to draw firm conclusions.

Zamora-Marín, J. M., Gutiérrez-Cánovas, C., Abellán, P., Millán, A. (2016). The role of protected areas in representing aquatic biodiversity: A test using α, β and γ diversity of water beetles from the Segura River Basin (SE Spain). Limnetica 35(1), 179-192,

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The role of protected areas in representing aquatic biodiversity: A test using α, β and γ diversity of water beetles from the Segura River Basin (SE Spain)

The role of protected areas in representing aquatic biodiversity: a test using α, β and γ diversity of water beetles from the Segura River Basin (SE Spain) Networks of protected areas represent one of the main strategies to reduce the rapid loss of biodiversity. However, most of these protected areas have been designed by considering only charismatic groups of vertebrates and plants, most linked to terrestrial environments. Thus, little is known about how well protected areas perform in representing aquatic biodiversity. This study analyses the suitability of national and European protected area networks (Natural Protected Areas and Natura 2000) in representing such biodiversity. For this purpose, we studied the different components of diversity (α, β and γ) using water beetles from the Segura River Basin as surrogates of overall macroinvertebrate biodiversity. Our results revealed no significant differences in α-diversity between protected and non-protected areas. Similarly, we did not find significant differences in β-diversity components (species replacement and nestedness, i.e., differences in among-site richness without species replacement) between protected and non-protected areas. The species replacement contributed more than nestedness to explain overall β-diversity changes. Finally, we found that the γ-diversity component was significantly higher in both protected areas, when compared to an equivalent number of randomly selected locations. Thus, the protected area networks from the Segura River Basin currently seem to have gaps in embracing the different aquatic biodiversity components. These results for river management and biodiversity conservation are discussed, providing some guidelines for future research.

Zhang, Z., Cao, Y., Jeppesen, E., Li, W. (2016). The response of Vallisneria spinulosa (Hydrocharitaceae) and plankton to pulse addition of inorganic nitrogen with different loading patterns. Hydrobiologia 767, 175–184,

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The response of Vallisneria spinulosa (Hydrocharitaceae) and plankton to pulse addition of inorganic nitrogen with different loading patterns

The global climate change may lead to more extreme climate events such as severe flooding creating excessive pulse-loading of nutrients, includ- ing nitrogen (N), to freshwaters. We conducted a 3-month mesocosm study to investigate the responses of phytoplankton, zooplankton and Vallisneria spin- ulosa to different N loading patterns using weekly and monthly additions of in total 14 g N m - 2 month - 1 during the first 2 months. The monthly additions led to higher phytoplankton chlorophyll a and total phyto- plankton biomass than at ambient conditions as well as lower leaf biomass and a smaller ramet number of V. spinulosa . Moreover, the biomass of cyanobacteria was higher during summer (August) in the monthly treatments than those with weekly or no additions. However, the biomass of plankton and macrophytes did not differ among the N treatments at the end of the experiment, 1 month after the termination of N addition. We conclude that by stimulating the growth of phytoplankton (cyanobacteria) and reducing the growth of submerged macrophytes, short-term extreme N loading may have significant effects on shallow nutrient-rich lakes and that the lakes may show fast recovery if they are not close to the threshold of a regime shift from a clear to a turbid state. .

Zhang, X., Liu, Z., Jeppesen, E., Taylor, W. D., Rudstam, L. G. (2016). Effects of benthic-feeding common carp and filter-feeding silver carp on benthic-pelagic coupling: Implications for shallow lake management. Ecological Engineering 88, 256–264,

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Effects of benthic-feeding common carp and filter-feeding silver carp on benthic-pelagic coupling: Implications for shallow lake management

Benthic–pelagic coupling is a key factor in the dynamics of shallow lakes. A 12-week mesocosm exper-iment was set up to test the hypotheses that benthic-feeding common carp (Cyprinus carpio) reduce the growth of benthic algae and promote eutrophication and that filter-feeding silver carp (Hypoph-thalmichthys molitrix) stimulate benthic algae growth and promote the establishment of a clear-water state. Compared to the controls, the common carp treatment had higher concentrations of total nitrogen (TN) and total phosphorus (TP) in the water column, higher biomass of pelagic algae (measured as chloro-phyll a), higher total suspended solids (TSS) concentrations, lower light intensity, and lower biomass of benthic algae at the sediment surface. Silver carp did not change the chlorophyll a of pelagic algae relative to the controls, but they did decrease the biomass of benthic algae and increase TP and TSS. A microcosm experiment using 32P radiotracer was conducted to examine effects of the two carp species on the release of sediment phosphorus (P). The P release to the water column was higher with common carp present than without common carp. This was not the case in the silver carp experiments. Our findings show that both common carp and silver carp deteriorate water quality by increasing TP and TSS concentrations and decreasing the biomass of benthic algae at the sediment surface. Common carp had a larger negative effect on water quality than silver carp, perhaps because only common carp enhanced P release from the sediment. The implications for lake management are that removal of both common carp and silver carp from shallow lakes may enhance the growth of benthic algae and help promote the establishment of a clear-water state.

Zhao, S., Yin, L., Chang, F., Olsen, Søndergaard, M., Jeppesen, E., Li, W. (2016). Response of Vallisneria spinulosa (Hydrocharitaceae) to contrasting nitrogen loadings in controlled lake mesocosms. Hydrobiologia 766, 215–223,

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Response of Vallisneria spinulosa (Hydrocharitaceae) to contrasting nitrogen loadings in controlled lake mesocosms

The role of nitrogen (N) in the shift from a macrophyte-dominated state to a phytoplankton-dominated one at high N concentrations in shallow lakes is still debated. To elucidate possible toxic and ecological effects of high N on macrophyte growth, we conducted a short-term (40 day) study of a eutrophication-tolerant macrophyte, Vallisneria spinulosa (Hydrocharitaceae), incubated in pots in a mesocosm system subjected to different N concentrations (1, 3, and 5 mg l-1). Plant leaf and root length as well as growth rate decreased significantly with increased N concentrations, but most N-and P-related physiological parameters, including the soluble protein content, nitrate reductase activity, acid phosphatase activity, and tissue N and P contents, did not differ significantly among the N treatments. Only the alkaline phosphatase activity differed, being lower at high nitrogen loading, likely due to P limitation. Epiphyton and phytoplankton biomasses increased significantly with increasing N loading. Our results including a large number of physiological tests of the macrophytes, therefore, provide supporting evidence that the loss of submerged macrophytes, like V. spinulosa, seen at high N loading in shallow lakes, can be attributed to competition with phytoplankton and epiphyton rather than to toxic effects.

Zhou, Y., Jeppesen, E., Li, L., Zhang, Y., Zhang, X., Li, X. (2016). Impacts of Three Gorges Reservoir on the sedimentation regimes in the downstream-linked two largest Chinese freshwater lakes. Scientific Reports 6 (E35396),

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Impacts of Three Gorges Reservoir on the sedimentation regimes in the downstream-linked two largest Chinese freshwater lakes

We studied the impacts of Three Gorges Reservoir (TGR) on the sedimentation regimes in the downstream-linked two largest Chinese freshwater lakes, Lake Dongting and Lake Poyang. Our results indicate that up to 1.73 × 10⁹ t sediment was retained in TGR from June 2003 to December 2014. This resulted in a 145.9 × 10⁶ t yr⁻¹ decline in the suspended sediment load at Zhicheng and a 16.8 × 10⁶ t yr⁻¹ lower sediment flow from Yangtze River to Lake Dongting, which partially explains the 13.4 × 10⁶ t yr⁻¹ lower sedimentation in Lake Dongting during the post-TGR period. Furthermore, TGR resulted in a 0.5 ± 0.3 m reduction of the multi-year mean water level at the Lake Poyang outlet Hukou, accelerating the suspended sediment export discharge from the lake. The reduced sedimentation in Lake Poyang during the post-TGR period was estimated to 6.3 × 10⁶ t yr⁻¹. We estimate that a monthly mean concentration of sediment flow from TGR below 0.60 kg m⁻³ will lead to erosion in Lake Dongting and Lake Poyang. Better regulation of TGR may extend the life expectancy of the two vanishing large lakes.

Zhou, Y., Jeppesen, E., Zhang, Y., Shi, K., Liu, X., Zhu, G. (2016). Dissolved organic matter fluorescence at wavelength 275/342 nm as a key indicator for detection of point-source contamination in a large Chinese drinking water lake. Chemosphere 144, 503-509,

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Dissolved organic matter fluorescence at wavelength 275/342 nm as a key indicator for detection of point-source contamination in a large Chinese drinking water lake

Surface drinking water sources have been threatened globally and there have been few attempts to detect point-source contamination in these waters using chromophoric dissolved organic matter (CDOM) fl uorescence. To determine the optimal wavelength derived from CDOM fl uorescence as an indicator of point-source contamination in drinking waters, a combination of fi eld campaigns in Lake Qiandao and a laboratory wastewater addition experiment was used. Parallel factor (PARAFAC) analysis identi fi ed six components, including three humic-like, two tryptophan-like, and one tyrosine-like component. All metrics showed strong correlation with wastewater addition ( r 2 > 0.90, p < 0.0001). Both the fi eld campaigns and the laboratory contamination experiment revealed that CDOM fl uorescence at 275/ 342 nm was the most responsive wavelength to the point-source contamination in the lake. Our results suggest that pollutants in Lake Qiandao had the highest concentrations in the river mouths of upstream in fl ow tributaries and the single wavelength at 275/342 nm may be adapted for online or in situ fl uo- rescence measurements as an early warning of contamination events. This study demonstrates the po- tential utility of CDOM fl uorescence to monitor water quality in surface drinking water sources.

Zhou, Y., Zhang, Y., Jeppesen, E., Murphy, K. R., Shi, K., Liu, M., Liu, X., Zhu, G. (2016). Inflow rate-driven changes in the composition and dynamics of chromophoric dissolved organic matter in a large drinking water lake. Water Research 100, 211-221,

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Inflow rate-driven changes in the composition and dynamics of chromophoric dissolved organic matter in a large drinking water lake

Drinking water lakes are threatened globally and therefore in need of protection. To date, few studies have been carried out to investigate how the composition and dynamics of chromophoric dissolved organic matter (CDOM) in drinking water lakes are influenced by inflow rate. Such CDOM can lead to unpleasant taste and odor of the water and produce undesirable disinfection byproducts during drinking water treatment. We studied the drinking water Lake Qiandao, China, and found that the concentrations of suspended particulate matter (SPM) in the lake increased significantly with inflow rate (p < 0.001). Similarly, close relationships between inflow rate and the CDOM absorption coefficient at 350 nm a(350) and with terrestrial humic-like fluorescence C3 and a negative relationship between inflow rate and the first principal component (PC1) scores, which, in turn, were negatively related to the concentrations and relative molecular size of CDOM (p < 0.001), i.e. the concentration and molecular size of CDOM entering the lake increased proportionately with inflow rate. Furthermore, stable isotopes (δD and δ18O) were depleted in the upstream river mouth relative to downstream remaining lake regions, substantiating that riverine CDOM entering the lake was probably driven by inflow rate. This was further underpinned by remarkably higher mean chlorophyll-a and in situ measured terrestrial CDOM fluorescence (365/480 nm) and apparent oxygen utilization (AOU), and notably lower mean PC1 and CDOM spectral slope (S275-295) recorded in the upstream river mouth than in the downstream main lake area. Strong negative correlations between inflow rate and a(250):a(365), S275-295, and the spectral slope ratio (SR) implied that CDOM input to the lake in rainy period was dominated by larger organic molecules with a more humic-like character. Rainy period, especially rainstorm events, therefore poses a risk to drinking water safety and requires higher removal efficiency of CDOM during drinking water treatment processes.

Zhou, Y., Zhou, J., Jeppesen, E., Zhang, Y., Qin, B., Shi, K., Tang, X., Han, X. (2016). Will enhanced turbulence in inland waters result in elevated production of autochthonous dissolved organic matter? Science of the Total Environment 543, 405–415,

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Will enhanced turbulence in inland waters result in elevated production of autochthonous dissolved organic matter?

Biological activity in lakes is strongly influenced by hydrodynamic conditions, not least turbulence intensity; which increases the encounter rate between plankter and nutrient patches. To investigate whether enhanced turbulence in shallow and eutrophic lakes may result in elevated biological production of autochthonous chromophoric dissolved organic matter (CDOM), a combination of field campaigns and mesocosm experiments was used. Parallel factor analysis identified seven components: four protein-like, one microbial humic-like and two terrestrial humic-like components. During our field campaigns, elevated production of autochthonous CDOM was recorded in open water with higher wind speed and wave height than in inner bays, implying that elevated turbulence resulted in increased production of autochthonous CDOM. Confirming the field campaign results, in the mesocosm experiment enhanced turbulence resulted in a remarkably higher microbial humic-like C1 and tryptophan-like C3 (p<0.01), indicating that higher turbulence may have elevated the production of autochthonous CDOM. This is consistent with the significantly higher mean concentrations of chlorophyll-a (Chl-a) and dissolved organic carbon (DOC) and the enhanced phytoplanktonic alkaline phosphatase activity (PAPA) recorded in the experimental turbulence groups than in the control group (p<0.05). The C:N ratio (from 3.34 to 25.72 with a mean of 13.13±4.08) for the mesocosm CDOM samples further suggested their probable autochthonous origin. Our results have implications for the understanding of CDOM cycling in shallow aquatic ecosystems influenced by wind-induced waves, in which the enhanced turbulence associated with extreme weather conditions may be further stimulated by the predicted global climate change.

Zingel, P., Agasild, H., Karus, K., Kangro, K., Tammert, H., Tonno, I., Feldmann, T., Noges, T. (2016). The influence of zooplankton enrichment on the microbial loop in a shallow, eutrophic lake. European Journal of Protistology 52, 22-35,

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The influence of zooplankton enrichment on the microbial loop in a shallow, eutrophic lake

With increasing primary productivity, ciliates may become the most important members of the microbial loop and form a central linkage in the transformation of microbial production to upper trophic levels. How metazooplankters, especially copepods, regulate ciliate community structure in shallow eutrophic waters is not completely clear. We carried out mesocosm experiments with different cyclopoid copepod enrichments in a shallow eutrophic lake to examine the responses of ciliate community structure and abundance to changes in cyclopoid copepod biomass and to detect any cascading effects on bacterioplankton and edible phytoplankton. Our results indicate that an increase in copepod zooplankton biomass favours the development of small-sized bacterivorous ciliates. This effect is unleashed by the decline of predaceous ciliate abundance, which would otherwise graze effectively on the small-sized ciliates. The inverse relationship between crustacean zooplankton and large predaceous ciliates is an important feature adjusting not only the structure of the ciliate community but also the energy transfer between meta- and protozooplankton. Still we could not detect any cascading effects on bacterio- or phytoplankton that would be caused by the structural changes in the ciliate community.


Borja, A., Garmendia, J. M. (2015). Nervion catchment (Spain). Freshwater Metadata Journal 4, 1-14,

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Nervion catchment (Spain)

This database comes from different sources (monitoring programmes) aiming to create an information system on thestate of water environment in the Basque Country. This information includes environmental (water, sediment) andbiological (benthic macroinvertebrates, fish, phytoplankton, macroalgae, macrophytes) elements obtained from samplingstations placed in rivers, estuarine and coastal zones.

Cao, Y., Neif, E. M., Li, W., Coppens, J., Filiz, N., Lauridsen, T. L., Davidson, T. A., Søndergaard, M., Jeppesen, E. (2015). Heat wave effects on biomass and vegetative growth of macrophytes after long-term adaptation to different temperatures: a mesocosm study. Climate Research 66, 265-274,

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Heat wave effects on biomass and vegetative growth of macrophytes after long-term adaptation to different temperatures: a mesocosm study

Elevated temperatures and extreme climatic events, such as heat waves, can negatively affect submerged macrophytes. Here, we investigated how submerged macrophytes adapted to three different temperatures: 1) ambient, 2) ca. +3 oC and 3) ca. +4.5 oC responded to a heat wave. After ten years of adaptation, the shoots of two species of submerged macrophytes, Elodea canadensis and Potamogeton crispus, were collected from each of the three temperature treatments and transferred to the two heated treatments for one month. Thereafter, the two heated treatments were exposed to a one-month heat wave with an additional 5 oC temperature increase. For P. crispus, total biomass did not differ among the plants adapted to the different temperatures or between the two heated treatments for the whole duration of the experiment. Plants adapted to the highest temperatures, however, produced a larger number of smaller turions before the heat wave and allocated less biomass to elongation before and after the heat wave. As to E. canadensis, the plants adapted to higher temperatures had higher total biomass before and during the heat wave and allocated more biomass to roots and leaves during the heat wave. Most indicators (e.g. length and biomass) of the macrophyte performance measured during the experiment did not differ between the two heated treatments. In summary, after the ten-year adaptation to higher temperatures, the submerged macrophytes showed adaptive changes in growth and asexual reproduction and responded in a complex way to the heat wave depending on species, growth status and adaptation temperature.

Davidson, T. A., Audet, J., Svenning, J.-C., Lauridsen, T. L., Søndergaard, M., Landkildehus, F., Larsen, S. E., Jeppesen, E. (2015). Eutrophication effects on greenhouse gas fluxes from shallow-lake mesocosms override those of climate warming. Global Change Biology 21, 4449–4463,

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Eutrophication effects on greenhouse gas fluxes from shallow-lake mesocosms override those of climate warming

Fresh waters make a disproportionately large contribution to greenhouse gas (GHG) emissions, with shallow lakes being particular hot spots. Given their global prevalence, how GHG fluxes from shallow lakes are altered by climate change may have profound implications for the global carbon cycle. Empirical evidence for the temperature dependence of the processes controlling GHG production in natural systems is largely based on the correlation between seasonal temperature variation and seasonal change in GHG fluxes. However, ecosystem-level GHG fluxes could be influenced by factors, which while varying seasonally with temperature are actually either indirectly related (e.g. primary producer biomass) or largely unrelated to temperature, for instance nutrient loading. Here, we present results from the longest running shallow-lake mesocosm experiment which demonstrate that nutrient concentrations override temperature as a control of both the total and individual GHG flux. Furthermore, testing for temperature treatment effects at low and high nutrient levels separately showed only one, rather weak, positive effect of temperature (CH4 flux at high nutrients). In contrast, at low nutrients, the CO2 efflux was lower in the elevated temperature treatments, with no significant effect on CH4 or N2O fluxes. Further analysis identified possible indirect effects of temperature treatment. For example, at low nutrient levels, increased macrophyte abundance was associated with significantly reduced fluxes of both CH4 and CO2 for both total annual flux and monthly observation data. As macrophyte abundance was positively related to temperature treatment, this suggests the possibility of indirect temperature effects, via macrophyte abundance, on CH4 and CO2 flux. These findings indicate that fluxes of GHGs from shallow lakes may be controlled more by factors indirectly related to temperature, in this case nutrient concentration and the abundance of primary producers. Thus, at ecosystem scale, response to climate change may not follow predictions based on the temperature dependence of metabolic processes.

Hanganu, J., Navodaru, I., Tudor, I. M., Ibram, O., Doroftei, M., Nastase, A. (2015). Introduction of the Danube Delta Database. Freshwater Metadata Journal 8, 1-11,

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Introduction of the Danube Delta Database

A description of biological and ecological data of the Danube delta lakes and channels is presented. The biologicalindicators refer to aquatic macrophytes, fish, zoo-plankton, and macro-invertebrates. Environmental data includephysio-chemical data as well as hydrological parameters.

Havens, K. E., Pinto-Coelho, R. M., Beklioğlu, M., Christoffersen, K. S., Jeppsen, E., Lauridsen, T. L., Mazumder A., Méthot, G., Pinel Alloul, B., Tavşanoğlu, Ü. N., Erdoğan, Ş., Vijverberg, J. (2015). Temperature effects on body size of feshwater crustacean zooplanktion from Greenland to the tropics. Hydrobiologia 743, 27-35,

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Temperature effects on body size of feshwater crustacean zooplanktion from Greenland to the tropics

The body size of zooplankton has many substantive effects on the function of aquatic food webs. A variety of factors may affect size, and earlier studies indicate that water temperature may be a particularly important variable. Here we tested the hypothesis that the body size of cladocerans, cala-noids, and cyclopoids declines with increasing water temperature, a response documented in an earlier study that considered only cladoceran zooplankton. We tested the hypothesis by comparing body size data that were available from prior studies of lakes ranging from 6 to 74 o latitude and encompassing a temperature range of 2–30°C. Cladoceran body size declined with temperature, in a marginally significant manner (P = 0.10). For cyclopoids, the decline was more significant (P = 0.05). In both cases, there was considerably more variation around the regression lines than previously observed; suggesting that other variables such as fish predation played a role in affecting size. Calanoid body size was unrelated to temperature. In contrast with cladocerans and cyclo-poids, perhaps calanoid body size is not metabolically constrained by temperature or is differently affected by changes in fish predation occurring with increasing temperature. The unexpected result for calanoids requires further investigation.

He, H., Zhu, X., Song, X., Jeppesen, E., Liu, Z. (2015). Phytoplankton response to winter warming modified by large-bodied zooplankton: an experimental microcosm study. Journal of Limnology 74(3), online,

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Phytoplankton response to winter warming modified by large-bodied zooplankton: an experimental microcosm study

While several field investigations have demonstrated significant effects of cool season (winter or spring) warming on phytoplankton development, the role played by large-bodied zooplankton grazers for the responses of phytoplankton to winter warming is ambiguous. We conducted an outdoor experiment to compare the effect of winter warming (heating by 3°C) in combination with presence and absence of Daphnia grazing (D. similis) on phytoplankton standing crops and community structure under eutrophic conditions. When Daphnia were absent, warming was associated with significant increases in phytoplankton biomass and cyanobacterial dominance. In contrast, when Daphnia were present, warming effects on phytoplankton dynamics were offset by warming-enhanced grazing, resulting in no significant change in biomass or taxonomic dominance. These results emphasize that large-bodied zooplankton like Daphnia spp. may play an important role in modulating the interactions between climate warming and phytoplankton dynamics in nutrient rich lake ecosystems.

Hering, D., Carvalho, L., Argillier, C., Beklioglu, M., Borja, A., Cardoso, A. C., Duel, H., Ferreira, T., Globevnik, L., Hanganu, J., Hellsten, S., Jeppesen, E., Kodeš, V., Lyche Solheim, A., Nõges, T., Ormerod, S., Panagopoulos, Y., Schmutz, S. Venohr, M., Birk, S. (2015). Managing aquatic ecosystems and water resources under multiple stress - an introduction to the MARS project. Science of the Total Environment 503-504, 10-21,

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Managing aquatic ecosystems and water resources under multiple stress — An introduction to the MARS project

Water resources globally are affected by a complex mixture of stressors resulting from a range of drivers, including urban and agricultural land use, hydropower generation and climate change. Understanding how stressors interfere and impact upon ecological status and ecosystem services is essential for developing effective River Basin Management Plans and shaping future environmental policy. This paper details the nature of these problems for Europe's water resources and the need to find solutions at a range of spatial scales. In terms of the latter, we describe the aims and approaches of the EU-funded project MARS (Managing Aquatic ecosystems and water Resources under multiple Stress) and the conceptual and analytical framework that it is adopting to provide this knowledge, understanding and tools needed to address multiple stressors. MARS is operating at three scales: At the water body scale, the mechanistic understanding of stressor interactions and their impact upon water resources, ecological status and ecosystem services will be examined through multi-factorial experiments and the analysis of long time-series. At the river basin scale, modelling and empirical approaches will be adopted to characterise relationships between multiple stressors and ecological responses, functions, services and water resources. The effects of future land use and mitigation scenarios in 16 European river basins will be assessed. At the European scale, large-scale spatial analysis will be carried out to identify the relationships amongst stress intensity, ecological status and service provision, with a special focus on large transboundary rivers, lakes and fish. The project will support managers and policy makers in the practical implementation of the Water Framework Directive (WFD), of related legislation and of the Blueprint to Safeguard Europe's Water Resources by advising the 3rd River Basin Management Planning cycle, the revision of the WFD and by developing new tools for diagnosing and predicting multiple stressors.

Jeppesen, E., Brucet, S., Naselli-Flores, L., Papastergiadou, E., Stefanidis, K., Nõges, T., Nõges, P., Attayde, J. L., Zohary, T., Coppens, J., Bucak, T., Fernandes Menezes, R., Sousa Freitas, F. R., Kernan, M., Søndergaard, M., Beklioğlu, M. (2015). Ecological impacts of global warming and water abstraction on lakes and reservoirs due to changes in water level and related changes in salinity. Hydrobiologia 750, 201-227,

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Ecological impacts of global warming and water abstraction on lakes and reservoirs due to changes in water level and related changes in salinitys

According to the Intergovernmental Panel on Climate Change report released in September 2014, unprecedented changes in temperature and precipitation patterns have been recorded globally in recent decades and further change is predicted to occur in the near future, mainly as the result of human activity. In particular, projections show that the Mediterranean climate zone will be markedly affected with significant implications for lake water levels and salinity. This may be exacerbated by increased demands for irrigation water. Based on long-term data from seven lakes and reservoirs covering a geographical gradient of 52° of latitudes and a literature review, we discuss how changes in water level and salinity related to climate change and water abstraction affect the ecosystem structure, function, biodiversity and ecological state of lakes and reservoirs. We discuss mitigation measures to counteract the negative effects on ecological status that are likely to result from changes in climate and water abstraction practices. Finally, we highlight research required to improve knowledge of the impacts of anthropogenically induced changes on lake water level and consequent changes in salinity.

Kattel, G., Gell, P., Perga, M.-E., Jeppesen, E., Grundell, R., Weller S., Zawadzki, A., Barry, L. (2015). Tracking a century of change in trophic structure and dynamics in a floodplain wetland: Integrating palaeoecological and palaeoisotopic evidence. Freshwater Biology 60, 711–723 ,

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Tracking a century of change in trophic structure and dynamics in a floodplain wetland: integrating palaeoecological and palaeoisotopic evidence

The palaeoecological assessment, and the use of stable isotopes of carbon in subfossils of herbivores and omnivores, represents a novel approach to understand transitions in past food-web structure and the dynamics of lake ecosystems in response to natural perturbations and human impacts. Combined with records of subfossil assemblages of cladocerans and chironomids, it may be possible to decipher whether changes are attributable to external forces or internally derived system shifts.A sediment record taken from the shallow (2.3 m depth) Kings Billabong in the River Murray floodplain (Australia) was analysed to explore changes in trophic dynamics over the past century.The palaeoecological assessment revealed that littoral assemblages of cladocerans and benthic diatoms were gradually replaced by planktonic (planktonic and facultative planktonic) assemblages after river regulation in the 1920s.The stable isotopic composition of carbon (δ13C), derived from chironomid head capsules, was relatively constant down-core, ranging between −26.1‰ and −24.0‰, and coincided largely with the δ13C of bulk sediment samples (−25.6‰ to −22.0‰). The δ13C values of pelagic (Daphnia) and ubiquitous (Bosmina, Alona) cladocerans, however, varied markedly, with that for Daphnia between −29.8‰ (10–20 cm) and −23.2‰ (60–70 cm), and for ubiquitous cladocerans, between −29.4‰ (20–30 cm) and −24.5‰ (80–70 cm).The temporal changes in the δ13C values of cladocerans also suggest a gradual transition from a macrophyte-dominated state to a phytoplankton-dominated state after river regulation and further indicate changes in the horizontal migration behaviour of Daphnia depending on macrophyte abundance and predation risk.Our study demonstrates the potential of reconstructing, more precisely, the trophic dynamics of large river floodplain lakes and their ecological resilience by combining subfossil analyses with stable isotope analyses of selected subfossil groups.

Kotamaki, N., Patynen, A., Taskinen, A., Huttula, T., Malve, O. (2015). Statistical Dimensioning of Nutrient Loading Reduction: LLR Assessment Tool for Lake Managers. Environmental Management 56(2), 480-491,

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Statistical Dimensioning of Nutrient Loading Reduction: LLR Assessment Tool for Lake Managers

Implementation of the EU Water Framework Directive (WFD) has set a great challenge on river basin management planning. Assessing the water quality of lakes and coastal waters as well as setting the accepted nutrient loading levels requires appropriate decision supporting tools and models. Uncertainty that is inevitably related to the assessment results and rises from several sources calls for more precise quantification and consideration. In this study, we present a modeling tool, called lake load response (LLR), which can be used for statistical dimensioning of the nutrient loading reduction. LLR calculates the reduction that is needed to achieve good ecological status in a lake in terms of total nutrients and chlorophyll a (chl-a) concentration. We show that by combining an empirical nutrient retention model with a hierarchical chl-a model, the national lake monitoring data can be used more efficiently for predictions to a single lake. To estimate the uncertainties, we separate the residual variability and the parameter uncertainty of the modeling results with the probabilistic Bayesian modeling framework. LLR has been developed to answer the urgent need for fast and simple assessment methods, especially when implementing WFD at such an extensive scale as in Finland. With a case study for an eutrophic Finnish lake, we demonstrate how the model can be utilized to set the target loadings and to see how the uncertainties are quantified and how they are accumulating within the modeling chain.

Levi, P. S., Riis, T., Alnøe, A. B., Peipoch, M., Maetzke, K., Bruus, C., Baattrup-Pedersen, A. (2015). Macrophyte Complexity Controls Nutrient Uptake in Lowland Streams. Ecosystems 18, 914–931,

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Macrophyte Complexity Controls Nutrient Uptake in Lowland Streams

Macrophytes act as ecosystem engineers in lowland stream ecosystems, enhancing habitat complexity and physical structure. Studies have demonstrated that macrophyte abundance and growth form can dictate the degree to which physical and biological stream characteristics are altered. However, few studies have investigated the influence of macrophytes and their species-specific variation in morphological complexity on functional processes, such as nutrient uptake. We injected 15N-labeled ammonium (15N-NH4 +) into four macrophyte-rich lowland streams in Denmark to quantify the uptake of NH4 + by macrophytes, epiphytic biofilms, benthic biofilms, and suspended particulate organic matter in the water column. Overall, macrophytes and their epiphytic biofilms accounted for 71-98% of the reach-weighted uptake across the study streams. While macrophytes had the highest rates of NH4 + uptake among the compartments we measured, the epiphytic biofilms had the highest uptake efficiency, ranging from 0.06 to 0.6 mg N mg N biomass −1 d−1. Among all compartments, the uptake efficiency was inversely related to the carbon-to-nitrogen ratio. Macrophyte complexity, expressed as leaf perimeter-to-area ratio (P:A), varied among the five species found in the study streams. The uptake rates by macrophyte species with high leaf P:A were, on average, an order of magnitude higher than the rates for species with simple leaf morphology (430 vs. 49 mg N m−2 d−1). In summary, our results indicate that macrophytes regulate stream function both via direct uptake of NH4 + from the water column and by providing a substrate for epiphytic biofilms. Furthermore, the effect of leaf architecture on nutrient uptake rates provides evidence that physical complexity can enhance ecosystem function.

Teichert, N., Uriarte, A., Chust, G., Borja, A., Lepage, M. (2015). How detecting interaction effects among pressures can assist environmental managers to develop efficient strategies of restoration for estuaries? ECSA 55th Symposium, 06.-09. September 2015, London.

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How detecting interaction effects among pressures can assist environmental managers to develop efficient strategies of restoration for estuaries?

European estuaries are subjected to a large spectrum of anthropogenic pressures, which are known to have additive, antagonistic, synergistic and multiplicative impacts on the functioning of ecological systems. Understanding the pressure-impact relationships and the interactions effects among pressures are essential requirements to develop accurate strategy of restoration. In this context, a key component is to identify priority actions and sort them to maximize the expected ecological improvement at each step of restoration. This study investigated the impact of multiple pressures in fish communities for 91 European transitional waters, with the aim of assisting managers to develop efficient strategies of restoration. Twenty environmental indicators were selected to assess the intensity of 10 major pressures affecting estuaries. Random Forest regression method was used to predict the fish ecological status of systems, assessed by the Ecological Quality Ratio (EQR), according to the combined effects of indicators. The relative importance of indicators and their effects on the partial ecological response was evaluated, highlighting gradual nonlinear relationships and thresholds shifts. The expected improvements of EQR were investigated for both individual and combined measures of restoration using the predictive performances of the random forest. The effects of pairwise restoration events, i.e. additive or interactive, were defined by comparing the EQR improvement for separated (i.e. sum of individual predictions) and combined events of pressure restoration. Strict additive effects were highlighted for only 7 pairs of pressures, whereas the 38 other pairs showed significant interactions suggesting antagonistic and synergistic effects. Our results showed that the fish ecological status of estuaries cannot be summarized by the sum of the individual responses for each pressure. This observation should be considered to define the priority order of pressures in management restoration strategies.

Olsen, S., Jeppesen, E., Moss, B., Ozkan, K., Beklioglu, M., Feuchtmayr, H., González Sagrario, M., Li, W., Larsen, S. E., Søndergaard, M. (2015). Factors influencing nitrogen processing in lakes: An experimental approach. Freshwater Biology 60, 642-662,

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Factors influencing nitrogen processing in lakes: an experimental approach

  1. To help improve our understanding of the nitrogen cycle in lakes, particularly in the context of climate change, we analysed total nitrogen (TN) and nitrate (inline image-N) data from six mesocosm experiments (in Denmark, U.K., China and Turkey) covering different climatic regions. We assessed the effects of nitrogen (N) and phosphorus (P) loading, temperature, salinity and water level on N processing.
  2. Water column N loss (defined as the nitrogen processed in and lost from the water column in units of net amount processed per unit area and per unit of time, or in relative terms as the percentage loss of the total pool in 2 weeks) was particularly sensitive to external nutrient loading to the mesocosms. Mean water column TN loss at high N loading varied from 111 to 250 mg m−2 day−1 and increased with N loading. High P loading resulted in increased water column N loss, possibly because of increased uptake into plants and attached algae and sedimentation of the increased algal crop. High salinity generally decreased water column TN loss; on average, 10% more TN was in the water column at 12‰ salinity than at 2‰ salinity, while no significant effect of water level was found.
  3. Only weak relationships were observed between N processing and temperature, and mesocosms limited by P accumulated more nitrogen in their water columns than those with high P loadings. Our results suggest that N processing in lakes appears to be more sensitive to features of the catchment, such as hydrology and loading, than to climatic effects related to temperature, salinity and water level.

Ren, L., Jeppesen, E., He, D., Wang, J., Liboriussen, L., Xing, P., Wu, Q. L. (2015). pH influences the Importance of Niche-Related and Neutral Processes in Lacustrine Bacterioplankton Assembly. Applied and Environmental Microbiology,

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pH influences the Importance of Niche-Related and Neutral Processes in Lacustrine Bacterioplankton Assembly

pH is an important factor that shapes the structure of bacterial communities. However, we have very limited information about the patterns and processes by which overall bacterioplankton communities assemble across wide pH gradients in natural freshwater lakes. Here, we used pyrosequencing to analyze the bacterioplankton communities in 25 discrete freshwater lakes in Denmark with pH levels ranging from 3.8 to 8.8. We found that pH was the key factor in impacting lacustrine bacterioplankton community assembly. More acidic lakes imposed stronger environmental filtering, which decreased the richness and evenness of bacterioplankton operational taxonomic unit (OTU) and largely shifted community composition. Although environmental filtering was detected to be the most important determinant of bacterioplankton community assembly, the importance of neutral assembly processes must also be considered notably in acidic lakes, where the species (OTU) diversity was low. We observed that the strong effect of environmental filtering in more acidic lakes was weakened by the enhanced relative importance of neutral community assembly, and bacterioplankton communities tended to be less phylogenetically clustered in more acidic lakes. In summary, we propose that pH was a major environmental determinant in freshwater lakes, regulating the relative importance and interplay between niche-related and neutral processes and shaping the patterns of freshwater lake bacterioplankton biodiversity.

Schmidt-Kloiber, A., Hering, D. (2015). – An online tool that unifies, standardises and codifies more than 20,000 European freshwater organisms and their ecological preferences. Ecological Indicators 53, 271–282,

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Species’ ecological preferences are progressively important for understanding distribution patterns, for conserving biodiversity or for assessing and evaluating the status of freshwater ecosystems. Comprehensive databases compiling species traits are already established in the terrestrial realm, but widely missing in freshwater science. We established a database for European taxa of five aquatic organism groups by compiling information on taxonomy, ecology and distribution based on extensive literature surveys, which were performed by experts for the targeted organism groups. The database includes fishes (654 taxa/21 ecological preferences), macro-invertebrates (8586/40), macrophytes (1083/5), diatoms (8868/36) and phytoplankton (1976/4). It is available online with various options and tools for finding information and has currently over 800 users. The reviewed literature as well as examples given in this paper, highlight the importance of the general availability of knowledge on ecological preferences for various aspects in ecological assessment. is considered a service for basic research, applied scientists, water managers or other stakeholders. It serves as base for bioassessment and monitoring.

Segurado, P., Almeida, C., Santos, J. M., Neves, R., Ferreira, M. T. (2015). Biological and environmental database of Sorraia catchment (Portugal). Freshwater Metadata Journal 5, 1-10,

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Biological and environmental database of Sorraia catchment (Portugal)

The database presented here contains general and specific information for the Sorraia river basin in Portugal, compiled whithin the context of the FP7 MARS Project. The information is based on multiple datasets from multiple sources andcontains data on hydrology, climate, water quality, geomorphological pressures and several biotic elements, including fish, macroinvertebrates, macrophytes and diatoms. The main source of information is the Portuguese Environmental Agency(APA) from the Ministry of the Environment, Territory and Energy.

Šorf, M., Davidson, T. A., Brucet, S., Menezes, R. F., Søndergaard, M., Lauridsen, T. L., Landkildehus, F., Liboriussen, L., Jeppesen, E. (2015). Zooplankton response to climate warming: A mesocosm experiment at contrasting temperatures and eutrophication states. Hydrobiologia 742, 185-203,

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Zooplankton response to climate warming: a mesocosm experiment at contrasting temperatures and nutrient levels

Zooplankton community response to the combined effects of nutrients and fish (hereafter N-F) at contrasting temperatures was studied in a long-term experiment conducted in 24 shallow lake mesocosms with low and high nutrient levels. We found a positive effect of N-F on zooplankton biomass, chlorophyll-a and turbidity. In contrast, zooplankton species and size diversity decreased with added N-F, as did submerged macrophyte plant volume inhabited (PVI). The community composition of zooplankton in high N-F mesocosms was related to chlorophyll-a and turbidity and to macrophyte PVI in the low N ? F mesocosms. Macrophytes can protect zooplankton from fish predation. Compared to N ? F effects, temperature appeared to have little effect on the zooplankton community. Yet analysis of community heterogeneity among treatments indicated a significant temperature effect at high N-F levels. The results indicate an indirect temperature effect at high N-F levels that can be attributed to tempera-ture-dependent variation in fish density and/or chlorophyll-a concentration.

Tavşanoğlu, Ü. N., Brucet, S., Levi E. E., Bucak, T., Bezirci, G., Özen, A., Johansson, L. S., Jeppesen, E., Beklioğlu, M. (2015). Size-based diel migration of zooplankton in Mediterranean shallow lakes assessed from in situ experiments with artificial plants. Hydrobiologia 753(1), 47-59,

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Size-based diel migration of zooplankton in Mediterranean shallow lakes assessed from in situ experiments with artificial plants

In warm lakes, fish aggregate within macrophytes, thereby weakening the role of these as a daytime refuge for zooplankton and altering the zooplankton size structure, predation pressure, and water clarity. To elucidate the role of macrophytes as a refuge for zooplankton and their effect on zooplankton size distribution, we established three sets of strandardized artificial plant beds in 11 lakes in Turkey with contrasting fish predation risk and turbidity. Zooplankton were sampled within and outside of each plant beds during day and night. Fish, collected overnight in multimesh-sized gillnets, were abundant both inside and outside the artificial plant beds, impoverishing the usefulness of plants as a daytime refuge for particularly large-bodied zooplankton. Zooplankton size diversity was negatively related to fish abundance. Diel vertical migration was the frequent anti-predator avoidance behavior, but reverse migration was also observed when Chaoborus was present. In contrast to the small-bodied taxa, large- and medium-sized taxa showed intraspecific size-based migration (i.e., individuals of different sizes had different migration patterns). Predators influenced the size structure and diel movement of zooplankton, but the response changed with the size of zooplankton and water clarity.

Teixeira-de Mello, F., Meerhoff, M., González-Bergonzoni, I., Kristensen, E. A., Baattrup-Pedersen, A., Jeppesen, E. (2015). Influence of riparian forests on fish assemblages in temperate lowland streams. Environmental Biology of Fishes 99, 133-144,

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Influence of riparian forests on fish assemblages in temperate lowland streams

The characteristics of riparian vegetation along streams vary with natural and anthropogenic factors. Deforestation for agricultural purposes has consequences for the physical in-stream structure and function, such as the predominance of autotrophic or heterotrophic stream metabolism. Open canopy lowland streams are often dominated by macrophytes, with potential direct and indirect effects on the fish community. We tested for possible differences in the structure (relative abundance of species, mean body size, and density) and composition (species richness, species identity, and different trophic groups) of fish assemblages between open canopy streams (OCS) and riparian forest streams (RFS), including pool and riffle habitats, in temperate lowland Denmark. OCS reaches exhibited higher alpha and beta diversity and frequently hosted rare species. Almost 50 % of the recorded species appeared only in OCS. OCS also had smaller mean body size of fish and tended to have higher fish densities. The relative abundance of the different trophic groups did not differ between the two streams types, but the RFS had a higher abundance and occurrence frequency of intolerant salmonids. Our results suggest that modification of riparian habitats can affect richness patterns and that strong functional changes may occur as a consequence of forest clearance through changes in the relative importance of a keystone species, trout (Salmo trutta).

Trolle, D., Nielsen, A., Rolighed, J., Thodsen, H., Andersen, H. E., Karlsson, I. B., Refsgaard, J. C., Olesen, J. E., Bolding, K., Kronvang, B., Søndergaard, M., Jeppesen, E. (2015). Projecting the future ecological state of lakes in Denmark in a 6 degree warmin scenario. Climate Research 64, 55-72,

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Projecting the future ecological state of lakes in Denmark in a 6 degree warming scenario

Lakes are highly sensitive to climate change, and climate warming is known to induce eutrophication symptoms in temperate lakes. In Denmark, climate is projected to cause increased precipitation in winter and increased air temperatures throughout the year by the end of the 21st century. Looking further into the future, the warming trend is projected to continue and likely reach a 6°C increase around the 22nd century (relative to a baseline period of 1986-2005). In the present study, we evaluate the consequences of such extreme changes for temperate Danish lakes. We use a multifaceted modelling approach by combining an eco-hydrological model to estimate future water runoff and catchment nutrient exports with both mechanistic and empirical lake models, describing key biogeochemical indicators in lakes, in order to quantify the effects of future nutrient loads and air temperature on lake ecosystems. Our model projections for the future scenario suggest that annual water runoff will increase (46%), driving also increases in exports of nitrogen and phosphorus (13 and 64%, respectively). Both the mechanistic and empirical modelling approaches suggest that phytoplankton biomass will increase and that potentially toxin-producing cyanobacteria may become a dominant feature of the phytoplankton community from spring. Warming and increased nutrient loads also affect the food webs within the lakes in the direction of higher fish control of algae-grazing water fleas, further reinforcing eutrophication. To be able to mitigate these eutrophication effects, external nutrient loading to the lakes must be reduced considerably.

Weyhenmeyer, G. A., Kosten, S., Wallin, M. B., Tranvik, L. J., Jeppesen, E., Roland, F. (2015). Significant fraction of CO2 emissions from boreal lakes derived from hydrologic inorganic carbon inputs. Nature Geoscience 8, 933-936,

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Annual CO2 emissions from lakes and other inland waters into the atmosphere are estimated to almost entirely compensate the total annual carbon uptake by oceans1, 2, 3. CO2 supersaturation in lakes, which results in CO2 emissions, is frequently attributed to CO2 produced within the lake4, 5, 6, 7, 8. However, lateral inorganic carbon flux through watersheds can also be sizeable9, 10, 11. Here we calculated lake surface water CO2 concentrations and emissions using lake pH, alkalinity and temperature from a compilation of data from 5118 boreal lakes12. Autumn surface water CO2 concentrations and CO2 emissions from the 5118 lakes co-varied with lake internal autumn CO2 production. However, using a mass balance approach we found that CO2 emission in the majority of lakes was sustained by inorganic carbon loading from the catchment rather than by internal CO2 production. Small lakes with high dissolved organic carbon and phosphorus concentrations, shorter retention times and longer ice-free seasons had the highest CO2 concentrations. CO2 emissions from these small lakes was twice that of comparable lakes in colder regions, and similar to emissions from subtropical and tropical lakes. We conclude that changes in land use and climate that increase dissolved inorganic carbon may cause emission levels from boreal lakes to approach those of lakes in warmer regions.

Whitehead, P. G., Bussi, G., Bowes, M., Read, D., Dadson, S., Elliot, A. (2015). Dynamic Modelling of Multiple Phytoplankton Groups in Rivers with an Application to the Thames River System in the UK. Environmental Modelling and Software 74, 75-91,

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Dynamic Modelling of Multiple Phytoplankton Groups in Rivers with an Application to the Thames River System in the UK

A process-based phytoplankton model developed to simulate the movement and growth of phytoplankton in river systems is presented in this paper. The model is based on mass-balance, and takes into account water temperature, light, self-shading, dissolved phosphorus and silicon concentrations. It was implemented in five reaches of the River Thames (UK), and the results compared to a novel dataset of cytometric data which includes concentrations of chlorophytes, diatoms, cyanobacteria and picoalgae. A Multi-Objective General Sensitivity Analysis was carried out in order to test the model robustness and to quantify the sensitivity to its parameters. The results show a good agreement between the simulations and the measured phytoplankton abundance. The most influential parameters were phytoplankton growth and death rates, while phosphorus concentration showed little influence, due to the high concentration of phosphorus in the Thames. The model is an important step forward towards understanding and predicting algal dynamics in river systems.

Yu, Q., Wang, H.-J., Li, Y., Shao, J.-C., Liang, X.-M., Jeppesen, E., Wang, H.-J. (2015). Effects of high nitrogen concentrations on the growth of submersed macrophytes at moderate phosphorus concentrations. Water Research 83, 385-395,

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Effects of high nitrogen concentrations on the growth of submersed macrophytes at moderate phosphorus concentrations

Eutrophication of lakes leading to loss of submersed macrophytes and higher turbidity is a worldwide phenomenon, attributed to excessive loading of phosphorus (P). However, recently, the role of nitrogen (N) for macrophyte recession has received increasing attention. Due to the close relationship between N and P loading, disentanglement of the specific effects of these two nutrients is often difficult, and some controversy still exists as to the effects of N. We studied the effects of N on submersed macrophytes represented by Vallisneria natans (Lour.) Hara in pots positioned at three depths (0.4 m, 0.8 m, and 1.2 m to form a gradient of underwater light conditions) in 10 large ponds having moderate concentrations of P (TP 0.03 ± 0.04 mg L_1) and five targeted concentrations of total nitrogen (TN) (0.5, 2, 10, 20, and 100 mg L_1), there were two ponds for each treatment. To study the potential shading effects of other primary producers, we also measured the biomass of phytoplankton (ChlaPhyt) and periphyton (ChlaPeri) expressed as chlorophyll a.We found that leaf length, leaf mass, and root length of macrophytes declined with increasing concentrations of TN and ammonium, while shoot number and root mass did not. All the measured growth indices of macrophytes declined significantly with ChlaPhyt, while none were significantly related to ChlaPeri. Neither ChlaPhyt nor ChlaPeri were, however, significantly negatively related to the various N concentrations. Our results indicate that shading by phytoplankton unrelated to the variation in N loading and perhaps toxic stress exerted by high nitrogen were responsible for the decline in macrophyte growth.

Zhang, X., Odgaard, R., Olesen, B., Lauridsen, T. L., Liboriussen, L., Søndergaard, M., Liu, Z., Jeppesen, E. (2015). Warming shows differential effects on late-season growth and competitive capacity of Elodea canadensis and Potamogeton crispus in shallow lakes. Inland Waters 5, 421-432,

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Warming shows differential effects on late-season growth and competitive capacity of Elodea canadensis and Potamogeton crispus in shallow lakes

Submerged macrophytes are likely to be affected by climate changes through changes in water temperatures and length of growing season. We conducted a lab experiment to examine the influence of a late-season temperature increase on growth, biomass allocation, and acclimation of 2 submerged macrophyte species, Elodea canadensis and Potamogeton crispus. We also ran competitive interaction experiments between the 2 species with mono- and mixed-species cultures in pots placed in outdoor heated mesocosms (5 years at ambient temperature and a higher temperature following the IPCC A2 scenario downscaled to local conditions but enhanced by 50%). In the lab, macrophytes collected in the 2 types of mesocosms were grown at ambient temperatures (12 °C in September and 8 °C in October) and 4 °C higher. Warming had an overall stronger effect on E. candensis than P. crispus, particularly within the low temperature range studied. Hence, the relative growth rate (RGR) of E. canadensis acclimated to ambient mesocosm conditions increased 6-fold from low (8 °C) to high (16 °C) temperature, whereas the RGR of P. crispus increased

Zhou, Y., Jeppesen, E., Zhang, Y., Niu, C., Shi, K., Liu, X., Zhu, G., Qin, B. (2015). Chromophoric dissolved organic matter of black waters in a highly eutrophic Chinese lake: Freshly produced from algal scums? Journal of Hazardous Materials 299, 222–230,

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Chromophoric dissolved organic matter of black waters in a highly eutrophic Chinese lake: Freshly produced from algal scums?

Field campaigns and an incubation experiment were conducted to evaluate the sources of chromophoric dissolved organic matter (CDOM) in black water spots in highly polluted regions of the Chinese Lake Taihu. A significant positive correlation (< 0.0001) was found between chlorophyll a (Chl-a) and the CDOM absorption coefficient a(350), indicating that algae degradation was likely the primary source of CDOM in black waters. This is supported by our field results that Chl-a, a(350) and the spectral slope ratio (SR) were significantly higher in the black water samples than in the regular samples (< 0.001). Our incubation experiment further substantiated the primary significance of biological CDOM source where a(350) increased with decreasing Chl-a concentrations. After seven days’ incubation, a 72.2% decrease and a 74.9% increase were recorded for Chl-a and a(350), respectively, relative to the initial values. Parallel factor analysis identified five fluorescent components. The maximal fluorescence intensity (Fmax) of tryptophan-like C1 and microbial humic-like C3 of black water samples was significantly higher than in the regular water samples (< 0.0005). This is consistent with incubation experiment results showing a rapid increase in Fmax of the two components, emphasizing the priority of the in situ biological CDOM source in black water spots.

Ziegler, C. R., Webb, J. A., Norton, S. B., Pullin, A. S., Melcher, A. H. (2015). Digital repository of associations between environmental variables: A new resource to facilitate knowledge synthesis. Ecological Indicators 53, 61-69,

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Digital repository of associations between environmental variables: A new resource to facilitate knowledge synthesis

Responsible care and management of Earth's resources requires scientific support, but the pool of underused research is growing rapidly. Environmental science research studies describe associations between variables (e.g. statistical relationships between stressors and responses). We propose open-access and online sharing of such associations. This concept differs from various efforts around the world to promote sharing of primary research data, but holds similar goals of improved use of existing knowledge. The initiative is made possible by recent developments in information technology and evolving online culture (e.g. crowdsourcing and citizen science). We have begun to connect existing projects that catalog and store associations, thereby moving toward a single virtual repository. Researchers and decision makers may share and re-use associations for myriad purposes, including: increasing efficiency and timeliness of systematic reviews, environmental assessments and meta-analyses, identifying knowledge gaps and research opportunities, providing evolved metrics of research impact, and demonstrating connections between research and environmental improvement.


Birk, S., Ecke, F. (2014). The potential of remote sensing in ecological status assessment of coloured lakes using aquatic plants. Ecological Indicators 46, 398-406,

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The potential of remote sensing in ecological status assessment of coloured lakes using aquatic plants

Field-based survey methods for aquatic vegetation have been identified as resource-demanding. Recent advances in remote sensing (RS) with sub-decimetre resolution allow for surveying aquatic vegetation at the species level. Coloured lakes, mainly due to high concentrations of humic substances, are globally widespread. However, high colour impedes the identification of submerged vegetation via remote sensing. Here, we evaluate the potential of using only emergent, floating and floating-leaved taxa that are detectable by high-resolution RS (RS-taxa) to assess the ecological status of lakes. In a dataset covering 72 Swedish low alkaline coloured lakes, we identified 31 RS-taxa. The power of RS-taxa assemblages to predict non-RS assemblages was analysed by a combination of ordination (Detrended Correspondence Analysis, DCA, and Redundancy Analysis) and multiple regression analysis. We compared the performance of a trophic metric score based on RS-taxa with that based on field data from all taxa along different environmental gradients. Forty percent of the variability of the first non-RS taxa DCA-axis was predicted by the DCA-results based on RS-taxa. Correlations of the trophic metric score and total nitrogen concentrations were equally strong for the dataset based on RS-taxa compared to the dataset based on all taxa. For total phosphorous concentrations, the correlation was stronger for the dataset based on all taxa, but for a complex water quality gradient (including sulphate, N-species, chlorophyll and percent cover of wetlands in the riparian buffer) the correlation was higher for the RS-taxa dataset. The significant linkage between the two community fractions (remotely-sensible and non-sensible) revealed considerable assemblage concordance, suggesting a notable potential of the use of remote sensing in lake macrophyte monitoring. The established trophic metric score seems most qualified for surveillance monitoring that, in combination with the eased efforts of data acquisition, detects long-term changes of the aquatic environment caused by shifts in climate, land use and (related) eutrophication.

Çakıroğlu, A.I., Tavşanoğlu, Ü. N., Davidson,T. A., Levi, E. E., Bucak, T., Özen, A., Akyıldız, G. K., Jeppesen, E., Beklioğlu, M.(2014). Relatedness between contemporary and surface sediment subfossil cladocera assemblages in Turkish shallow lakes. Paleolim 52, 367-383,

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Relatedness between contemporary and subfossil cladoceran assemblages in Turkish lakes

Cladocerans are valuable indicators of environmental change in lakes. Their fossils provide information on past changes in lake environments. However, few studies have quantitatively examined the relationships between contemporary and sub-fossil cladoceran assemblages and no investigations are available from Mediterranean lakes where salinity, eutrophication and top-down control of large-bodied cladocerans are known to be important. Here we compared contemporary Cladocera assemblages, sampled in summer, from both littoral and pelagic zones, with their sub-fossil remains from surface sediment samples from 40 Turkish, mainly shallow, lakes. A total of 20 and 27 taxa were recorded in the contemporary and surface sediment samples, respectively. Procrustes rotation was applied to both the principal components analysis (PCA) and redundancy analysis (RDA) ordinations in order to explore the relationship between the cladoceran community and the environmental variables. Procrustes rotation analysis based on PCA showed a significant accord between both littoral and combined pelagic–littoral contemporary and sedimentary assemblages. RDA ordinations indicated that a similar proportion of variance was explained by environmental variation for the contemporary and fossil Cladocera data. Total phosphorus and salinity were significant explanatory variables for the contemporary assemblage, whereas salinity emerged as the only significant variable for the sedimentary assemblage. The residuals from the Procrustes rotation identified a number of lakes with a high degree of dissimilarity between modern and sub-fossil assemblages. Analysis showed that high salinity, deep water and high macrophyte abundance were linked to a lower accord between contemporary and sedimentary assemblages. This low accord was, generally the result of poor representation of some salinity tolerant, pelagic and macrophyte-associated taxa in the contemporary samples. This study provides further confirmation that there is a robust relationship between samples of modern cladoceran assemblages and their sedimentary remains. Thus, sub-fossil cladoceran assemblages from sediment cores can be used with confidence to track long-term changes in this environmentally sensitive group and in Mediterranean lakes, subjected to large inter-annual variation in water level, salinity and nutrients.

Cao Y., Li, W., Jeppesen, E. (2014). The response of two submerged macrophyte and periphyton to elevated temperature at high nutrient level: a microcosm approach. Hydrobiologia 738, 49–59,

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The response of two submerged macrophytes and periphyton to elevated temperatures in the presence and absence of snails: a microcosm approach

Global warming may affect snail–periphyton–macrophyte relationships in lakes with implications also for water clarity. We conducted a 40-day aquaria experiment to elucidate the response of submerged macrophytes and periphyton on real and artificial plants to elevated temperatures (3°C) under eutrophic conditions, with and without snails present. With snails, the biomass and length of Vallisneria spinulosa leaves increased more at the high temperature, and at both temperatures growth was higher than in absence of snails. The biomass of periphyton on V. spinulosa as well as on artificial plants was higher at the highest temperature in the absence but not in the presence of snails. The biomass of Potamogeton crispus (in a decaying state) declined in all treatments and was not affected by temperature or snails. While total snail biomass did not differ between temperatures, lower abundance of adults (size >1 cm) was observed at the high temperatures. We conclude that the effect of elevated temperature on the snail–periphyton–macrophyte relationship in summer differs among macrophyte species in active growth or senescent species in subtropical lakes and that snails, when abundant, improve the chances of maintaining actively growing macrophytes under eutrophic conditions, and more so in a warmer future with potentially denser growth of periphyton.

Cremona, F., Kõiv, T., Kisand, V., Laas, A., Zingel, P., Agasild, H., Feldmann, T., Järvalt, A., Nõges, P., Nõges, T. (2014). From Bacteria to Piscivorous Fish: Estimates of Whole-Lake and Component-Specific Metabolism with an Ecosystem Approach. PLOS one,

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From Bacteria to Piscivorous Fish: Estimates of Whole- Lake and Component-Specific Metabolism with an Ecosystem Approach

The influence of functional group specific production and respiration patterns on a lake's metabolic balance remains poorly investigated to date compared to whole-system estimates of metabolism. We employed a summed component ecosystem approach for assessing lake-wide and functional group-specific metabolism (gross primary production (GPP) and respiration (R)) in shallow and eutrophic Lake Võrtsjärv in central Estonia during three years. Eleven functional groups were considered: piscivorous and benthivorous fish; phyto-, bacterio-, proto- and metazooplankton; benthic macroinvertebrates, bacteria and ciliates; macrophytes and their associated epiphytes. Metabolism of these groups was assessed by allometric equations coupled with daily records of temperature and hydrology of the lake and measurements of food web functional groups biomass. Results revealed that heterotrophy dominated most of the year, with a short autotrophic period observed in late spring. Most of the metabolism of the lake could be attributed to planktonic functional groups, with phytoplankton contributing the highest share (90% of GPP and 43% of R). A surge of protozooplankton and bacterioplankton populations forming the microbial loop caused the shift from auto- to heterotrophy in midsummer. Conversely, the benthic functional groups had overall a very small contribution to lake metabolism. We validated our ecosystem approach by comparing the GPP and R with those calculated from O2 measurements in the lake. Our findings are also in line with earlier productivity studies made with 14C or chlorophyll a (chl-a) based equations. Ideally, the ecosystem approach should be combined with diel O2 approach for investigating critical periods of metabolism shifts caused by dynamics in food-web processes.

Cremona, F., Laas, A., Nõges, P., Nõges, T. (2014). High-frequency data within a modeling framework: On the benefit of assessing uncertainties of lake metabolism. Ecological Modelling 294, 27-35,


High-frequency data within a modeling framework: On the benefit of assessing uncertainties of lake metabolism

We used a Bayesian metabolic model for assessing the gross primary production (GPP), ecosystem respiration (ER) and their uncertainties in lake Võrtsjärv, a large eutrophic lake in Estonia (North-eastern Europe). Diel cycle modeling was based on high-frequency (10-min) measurements of irradiance, water temperature and dissolved oxygen during most of the growing season (from May to August 2011). Posterior distribution of production and respiration was successfully simulated with the model and displayed with highly credible intervals (2.5 and 97.5 percentiles). Considering the mean GPP and ER values, the lake was autotrophic from May to June, at equilibrium in July, and heterotrophic in August. However, adding the uncertainty to metabolism estimates revealed that an ambiguous metabolic state (no clear monthly predominance of auto- or hetero-trophy) represented between 12 and 32% of the period. It is thus incautious to conclude about lake metabolic state in these conditions. A comparison with the existing classical model based on dissolved oxygen measurements showed that metabolic dynamics differed between the two approaches. Though the classical model recorded highest ecosystem productivity in midsummer, the Bayesian model predicted that productivity peaked earlier in the season and gradually declined as the irradiance dropped and the water temperature rose. Coupling between GPP and ER during the whole study period was very variable, resulting that, depending on the month, 50–100% of primary production was consumed in the lake. This coupling variability was caused by extensive diel fluctuation of irradiance-dependent production compared to relatively stable water temperature and respiration. The background respiration was high in spring and declined progressively in summer, reflecting lower inputs of allochthonous organic matter to the lake. With a wider use of high-frequency techniques for measuring lake ecological parameters, this kind of performant models that are able to assess lake productivity within small time steps and take into account the uncertainty, will be increasingly needed in the future.

Karus, K., Feldmann, T., Nõges, P., Zingel, P. (2014). Ciliate communities of a large shallow lake: Association with macrophyte beds. European Journal of Protistology 50, 382–394,

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Ciliate communities of a large shallow lake: Association with macrophyte beds

We investigated the influence of macrophyte composition on ciliate community structure in a large, shallow, eutrophic Lake Võrtsjärv. We hypothesized that macrophyte composition must have strong influence on the dispersal of ecologically different ciliate groups in a shallow lake and that more diverse macrophyte stands cause also a greater diversity in the ciliate community. In Võrtsjärv macrophyte distribution is spatially strongly polarized both in east–west and north–south directions in relation to abiotic factors. Phragmites australis and Myriophyllum spicatum were the most widespread species occurring in most parts of the lake. Correlation of environmental, macrophyte and planktonic ciliate variables confirmed the suggested spatial gradients. More diverse macrophyte stands supported a high species richness and abundance of epiplanktonic community but showed negative influence on the number and abundance of euplanktonic ciliate taxa. Opposite trends were found relative to the abundance of P. australis. Benthic ciliates showed a similar distribution pattern to euplanktonic taxa being most abundant in sites were the Shannon–Weaver index for macrophytes was low. Strong polarizing effect of the lake's vegetation on planktonic ciliate diversity was reflected in correlations of the number of ciliate taxa as well as the numbers of eu- and epiplanktonic taxa with geographic co-ordinates.

Lin, Q., Jiang, X., Han, B.-P., Jeppesen, E. (2014). Does stocking of filter-feeding fish for production have cascading effects on zooplankton and the ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient levels. Hydrobiologia 736, 115-125,

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Does stocking of filter-feeding fish for production have a cascading effect on zooplankton and ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient concentrations

Stocking of filter-feeding fish is a common tool used in Chinese reservoirs to increase fish production because of low natural recruitment. Whether such stocking has important negative effects on zooplankton with cascading effects on phytoplankton is debated. We compared the zooplankton communities in fourteen reservoirs with different nutrient concentrations and fish densities. Both chlorophyll a (Chla) and fish catch were positively related with total phosphorus (TP), whereas zooplankton biomass did not show a similar relationship with TP. Zooplankton seemed to be influenced by fish as high fish catches coincided with a low proportion of calanoids of the total copepod biomass, a high proportion of rotifers of the total zooplankton biomass, a low zooplankton:phytoplankton biomass ratio, and the absence of Daphnia irrespective of TP concentration. Both zooplankton biomass and most of the zooplankton:phytoplankton biomass ratios were among the lowest reported in the literature for the nutrient range studied. Furthermore, the Chla:TP ratio was higher than what is typically observed in temperate lakes. We conclude that top-down control of zooplankton is of key importance in reservoirs in South China where frequent stocking of filter-feeding fish seems to contribute to poor water quality in the form of higher algal biomass and reduced clarity.

Menezes, F. R., Borchsenius, F., Svenning, J-C., Davidson, T. A., Søndergaard, M., Lauridsen, L. T., Landkildehus, F., Jeppesen, E. (2014). Homogenization of fish assemblages in different lake depth strata at local and regional scales. Freshwater Biology 60, 745-757,

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Homogenization of fish assemblages in different lake depth strata at local and regional scales

Eutrophication alters the trophic dynamics in lakes and may result in homogenisation of biotic communities. How nutrient enrichment drives patterns of homogenisation of fish species composition at within-lake (local) and among-lake (regional) scales is, however, not well studied. To test for homogenisation in fish communities, we analysed number, biomass and individual mean body mass of the different fish species present in 53 Danish lakes with contrasting depths, surface area and eutrophication. A combination of uni- and multivariate techniques revealed that eutrophication homogenises fish community composition in the littoral zone of both shallow and deep lakes at within- and among-lake scales, a notable contrast being that community composition was not homogenised in the offshore in deep lakes. In addition, fish species richness and diversity converged with progressive eutrophication and mean lake depth in all lake zones. For deep lakes, surface area was positively related to increasing differences in fish species richness and diversity. Increased homogeneity of this key assemblage may have profound implications for ecosystems and their stability (such as decreased resilience to disturbance, reduced biological complexity and increased vulnerability to large-scale and stochastic environmental events). Thus, to fully understand the impacts of eutrophication on aquatic communities at both local and regional scales, the effects of nutrient enrichment on compositional heterogeneity should be considered.

Özen, A., Bucak, T., Tavşanoğlu, Ü. N., Çakıroğlu, A. İ, Levi, E. E., Coppens, J., Jeppsen, E., Beklioğlu, M. (2014). Water level and fish-mediated cascading effects on the microbial community in eutrophic warm shallow lakes: A mesocosm experiment. Hydrobiologia 740, 25-35,

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Water level and fish-mediated cascading effects on the microbial community in eutrophic warm shallow lakes: a mesocosm experiment

Information on the effects of water level changes on microbial planktonic communities in lakes is limited but vital for understanding ecosystem dynamics in Mediterranean lakes subjected to major intra-and inter-annual variations in water level. We performed an in situ mesocosm experiment in an eutrophic Turkish lake at two different depths crossed with presence/absence of fish in order to explore the effects of water level variations and the role of top-down regulation at contrasting depths. Strong effects of fish were found on zooplankton, weakening through the food chain to ciliates, HNF and bacterioplankton, whereas the effect of water level variations was overall modest. Presence of fish resulted in lower biomass of zooplankton and higher biomasses of phytoplankton, ciliates and total plankton. The cascading effects of fish were strongest in the shallow mesocosms as evidenced by a lower zooplankton contribution to total plankton biomass and lower zooplankton:ciliate and HNF:bacteria biomass ratios. Our results suggest that a lowering of the water level in warm shallow lakes will enhance the contribution of bacteria, HNF and ciliates to the plankton biomass, likely due to increased density of submerged macrophytes (less phytoplankton); this effect will, however, be less pronounced in the presence of fish.

Olsen, S., Chan, F., Li, W., Zhao, S., Søndergaard, M., Jeppsen, E. (2014). Strong impact of nitrogen loading on submerged macrophytes and algae: a long-term mesocosm experiment in a shallow Chinese lake. Freshwater Biology 60, 1525-1536,

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Strong impact of nitrogen loading on submerged macrophytes and algae: a long-term mesocosm experiment in a shallow Chinese lake

We studied the effect of N loading on macrophytes (dominated by Potamogeton lucens and Cabomba caroliniana), periphyton, filamentous algae and phytoplankton in mesocosms over 10 months (starting in October) in subtropical China (Wuhan). There were three N treatments: controls (CN) without nitrogen addition (mean TN = 1.9 mg L−1), low nitrogen (LN) addition (mean TN = 3.5 mg L−1) and high nitrogen (HN) addition (mean TN = 5.5 mg L−1). Total phosphorus (TP) concentration in the water column remained moderate (0.05–0.07 mg L−1) during the experiment in all treatments. Macrophyte abundance declined in the LN and HN treatments in the first 6 months, but not in controls, followed by a partial recovery in the LN treatments. They disappeared completely in the HN treatments the following summer. Periphyton (biofilm on plastic) and phytoplankton biomass remained unaffected during the first 6 months but increased over the summer by two or three times, compared with controls, in low and high nitrogen treatments, respectively. By contrast, the abundance of filamentous algae increased over winter but declined during the summer with no obvious relationship to the N treatments. There was no difference in the TN or nitrate concentrations or soluble protein, soluble sugar and Chl-a content of P. lucens leaves and stems with increasing N load.Macrophyte populations are partially resilient to abrupt increases in N loading at moderate TP concentrations, but, after prolonged exposure, a complete collapse occurs. Our results further indicate that macrophyte loss is exacerbated by shading by filamentous algae during the winter, and by phytoplankton and periphyton in the summer, while there was no indication of direct N toxicity.

Olsen, S., Jeppesen, E., Moss, B., Özkan, K., Beklioğlu, M., Feuchtmayr, H., González Sagrario, M., Wei, L., Larsen, S., Lauridsen, T. S., Søndergaard, M. (2014). Factors influencing nitrogen processing in lakes: An experimental approach. Freshwater Biology 60, 646-662,

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Factors influencing nitrogen processing in lakes: an experimental approach

1. To help improve our understanding of the nitrogen cycle in lakes, particularly in the context of climate change, we analysed total nitrogen (TN) and nitrate (NO À 3 -N) data from six mesocosm experiments (in Denmark, U.K., China and Turkey) covering different climatic regions. We assessed the effects of nitrogen (N) and phosphorus (P) loading, temperature, salinity and water level on N processing. 2. Water column N loss (defined as the nitrogen processed in and lost from the water column in units of net amount processed per unit area and per unit of time, or in relative terms as the percentage loss of the total pool in 2 weeks) was particularly sensitive to external nutrient loading to the mesocosms. Mean water column TN loss at high N loading varied from 111 to 250 mg m À2 day À1 and increased with N loading. High P loading resulted in increased water column N loss, possibly because of increased uptake into plants and attached algae and sedimentation of the increased algal crop. High salinity generally decreased water column TN loss; on average, 10% more TN was in the water column at 12‰ salinity than at 2‰ salinity, while no significant effect of water level was found. 3. Only weak relationships were observed between N processing and temperature, and mesocosms limited by P accumulated more nitrogen in their water columns than those with high P loadings. Our results suggest that N processing in lakes appears to be more sensitive to features of the catchment, such as hydrology and loading, than to climatic effects related to temperature, salinity and water level.

Panagopoulos, Y., Stefanidis, K., Mimikou, M. (2014). Description of an environmental database for the catchment of river Pinios, Thessaly in Greece. Freshwater Metadata Journal 3, 1-7,

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Description of an environmental database for the catchment of river Pinios, Thessaly in Greece

This database contains climatic, hydrologic, water quality and biological data for the river catchment of Pinios in Thessaly Greece, that were compiled during the European FP7 funded project MARS. The database is a compilation of datasets that contain mostly information on precipitation, air temperature, river discharge, river nutrient concentrations, macroinvertabrates and fish of river Pinios in Greece. The majority of the data listed in the database were obtained by the Ministries of Agriculture and Environment, the Public Power Corporation of Greece, and other published sources.

Sandin, L., Schmidt-Kloiber, A., Svenning, J-C, Jeppesen, E., Friberg, N. (2014). A trait based approach to assess climate change sensitivity of freshwater invertebrates across Swedish ecoregions. Current Zoology 60, 221-232,

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A trait-based approach to assess climate change sensitivity of freshwater invertebrates across Swedish ecoregions

Freshwater habitats and organisms are among the most threatened on Earth, and freshwater ecosystems have been subject to large biodiversity losses. We developed a Climate Change Sensitivity (CCS) indicator based on trait information for a selection of stream-and lake-dwelling Ephemeroptera, Plecoptera and Trichoptera taxa. We calculated the CCS scores based on ten species traits identified as sensitive to global climate change. We then assessed climate change sensitivity between the six main ecoregions of Sweden as well as the three Swedish regions based on Illies. This was done using biological data from 1,382 stream and lake sites where we compared large-scale (ecoregional) patterns in climate change sensitivity with potential future exposure of these ecosystems to increased temperatures using ensemble-modelled future changes in air temperature. Current (1961~1990) measured temperature and ensemble-modelled future (2100) temperature showed an increase from the northernmost towards the southern ecoregions, whereas the predicted temperature change increased from south to north. The CCS indicator scores were highest in the two northernmost boreal ecoregions where we also can expect the largest global climate change-induced increase in temperature, indicating an unfortunate congruence of exposure and sensitivity to climate change. These results are of vital importance when planning and implementing management and conservation strategies in freshwater ecosystems, e.g., to mitigate increased temperatures using riparian buffer strips. We conclude that traits information on taxa spe-cialization, e.g., in terms of feeding specialism or taxa having a preference for high altitudes as well as sensitivity to changes in temperature are important when assessing the risk from future global climate change to freshwater ecosystems

Schmutz, S., Bakken, T. H., Friedrich, T., Greimel, F., Harby, A., Jungwirth, M., Meichler, A., Unfer, G., Zeiringer, B. (2014). Response of fish communities to hydrological and morphological alterations hydropeaking rivers of Austria. Rivers and Research Applications 31, 919-930,

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Response of fish communities to hydrological and morphological alterations hydropeaking rivers of Austria. Rivers and Research Applications

Climate change asks for the reduction in the consumption of fossil-based fuels and an increased share of non-regulated renewable energy sources, such as solar and wind power. In order to back up a larger share of these intermittent sources, ‘battery services’ are needed, currently provided only in large scale by hydropower, leading to more rapid and frequent changes in flows (hydropeaking) in the downstream rivers. Increased knowledge about the ecosystem response to such operations and design of cost-effective measures is needed.We analysed the response of fish communities to hydropeaking (frequency, magnitude, ramping rate and timing) and the interaction with the habitat conditions in Austrian rivers. An index of biotic integrity (Fish Index Austria) was used to compare river sections with varying degrees of flow fluctuations under near-natural and channelized habitat conditions. The results showed that habitat conditions, peak frequency (number of peaks per year), ramping rate (water level variation) and interaction between habitat and ramping rate explained most of the variation of the Fish Index Austria. In addition, peaking during the night seems to harm fish more than peaking during the day. Fish communities in hyporhithral and epipotamal types of rivers are more affected by hydropeaking than those in metarhithral type of rivers. The results support the findings of other studies that fish stranding caused by ramping rates >15 cm h−1 are likely to be the main cause of fish community degradation when occurring more often than 20 times a year. While the ecological status degrades with increasing ramping rate in nature-like rivers, fish communities are heavily degraded in channelized rivers regardless of the ramping rate. The mitigation of hydropeaking, therefore, requires an integrative approach considering the combined effects of hydrological and morphological alterations on fish.