Influence of plant habitats on denitrification in lowland agricultural streams.

The aim of this study was to assess potential differences in denitrification in contrasting stream habitats in agricultural lowland streams located in Denmark. The study focused on three types of habitats i) vegetated habitats with emergent plants, ii) vegetated habitats with submerged plants, iii) bare sediments. Denitrification rates were measured in situ using denitrification chambers and nitrogen isotope pairing technique three times during a growing season. Denitrification rates across all habitats and samplings were 73 ± 116 µmol N m-2 h-1 (mean ± sd) with greater denitrification rates in vegetated habitats compared to bare sediments. Habitats with emergent plants had significantly higher denitrification rates than habitats with submerged plants. The habitats exhibited differences in oxygen and carbon availability probably connected to differences in flow velocity and physical effect of the vegetation (if present) which likely acted as a trap for finer organic-rich particles. Placing these results in the context of stream and river restoration highlights the potential of in-stream vegetation to mitigate nitrogen pollution, especially by restoring plant habitats in degraded and channelized streams to sustain vegetation promoting higher denitrification rates.

Effects of increased flooding on riparian vegetation: Field experiments simulating climate change along five European lowland streams.

In many parts of the world, the magnitude and frequency of cold-season precipitation are expected to increase in the near future. This will result in an increased magnitude and duration of winter and spring flooding by rain-fed streams and rivers. Such climate-driven increases in flooding are likely to affect riparian plant communities, but future vegetation changes are hard to predict due to current lack of data. To fill this knowledge gap, we experimentally modified the hydrology of five streams across three countries in north-western Europe during late winter/early spring over a period of 3 years. We assessed the responses in riparian plant species richness, biomass, plant-available nitrogen and phosphorus and seed deposition to increased flooding depth (+18 cm on average at the lowest positions along the riparian gradient) and prolonged flooding duration (6 weeks on average). After 3 years of increased flooding, there was an overall decline in riparian species richness, while riparian plant biomass increased. Extractable soil nitrogen and phosphorus also increased and are likely to have contributed to the increased biomass. Increased flooding resulted in the arrival of more seeds of additional species to the riparian zone, thereby potentially facilitating the shifts in riparian plant species composition we observed. The results of our concerted experimental effort demonstrate that changes in stream riparian plant communities can occur rapidly following increased winter flooding, leading to strong reductions in plant species diversity.

Trait Characteristics Determine Pyrethroid Sensitivity in Nonstandard Test Species of Freshwater Macroinvertebrates: A Reality Check.

We exposed 34 species of stream macroinvertebrates, representing 29 families, to a 90 min pulse of the pyrethroid λ-cyhalothrin. For 28 of these species, no pyrethroid ecotoxicity data exist. We recorded mortality rates 6 days post-exposure, and the behavioral response to pyrethroid exposure was recorded using automated video tracking. Most arthropod species showed mortality responses to the exposure concentrations (0.01-10 µg L(-1)), whereas nonarthropod species remained unaffected. LC50 varied by at least a factor of 1000 among arthropod species, even within the same family. This variation could not be predicted using ecotoxicity data from closely related species, nor using species-specific indicator values from traditional ecological quality indices. Moreover, LC50 was not significantly correlated to effect thresholds for behavioral responses. Importantly, however, the measured surface area-weight ratio and the preference for coarse substrates significantly influenced the LC50 for arthropod species, with the combination of small individuals and strong preference for coarse substrates indicating higher pyrethroid sensitivity. Our study highlights that existing pesticide ecotoxicity data should be extrapolated to untested species with caution and that actual body size (not maximum potential body size, as is usually available in traits databases) and habitat preference are central parameters determining species sensitivities to pyrethroids.

Riparian plant community responses to increased flooding: a meta-analysis.

A future higher risk of severe flooding of streams and rivers has been projected to change riparian plant community composition and species richness, but the extent and direction of the expected change remain uncertain. We conducted a meta-analysis to synthesize globally available experimental evidence and assess the effects of increased flooding on (1) riparian adult plant and seedling survival, (2) riparian plant biomass and (3) riparian plant species composition and richness. We evaluated which plant traits are of key importance for the response of riparian plant species to flooding. We identified and analysed 53 papers from ISI Web of Knowledge which presented quantitative experimental results on flooding treatments and corresponding control situations. Our meta-analysis demonstrated how longer duration of flooding, greater depth of flooding and, particularly, their combination reduce seedling survival of most riparian species. Plant height above water level, ability to elongate shoots and plasticity in root porosity were decisive for adult plant survival and growth during longer periods of flooding. Both 'quiescence' and 'escape' proved to be successful strategies promoting riparian plant survival, which was reflected in the wide variation in survival (full range between 0 and 100%) under fully submerged conditions, while plants that protrude above the water level (>20 cm) almost all survive. Our survey confirmed that the projected increase in the duration and depth of flooding periods is sufficient to result in species shifts. These shifts may lead to increased or decreased riparian species richness depending on the nutrient, climatic and hydrological status of the catchment. Species richness was generally reduced at flooded sites in nutrient-rich catchments and sites that previously experienced relatively stable hydrographs (e.g. rain-fed lowland streams). Species richness usually increased at sites in desert and semi-arid climate regions (e.g. intermittent streams).

Monitoring fish communities in wadeable lowland streams: comparing the efficiency of electrofishing methods at contrasting fish assemblages.

Electrofishing is considered a reliable tool to assess the assemblages and biodiversity of fish in wadeable streams. The most widely used electrofishing techniques (point [P], single-pass [S-P], and multiple-pass [M-P]) vary as to the effort needed for sample collection, and this may potentially influence the degree of accuracy. Moreover, little is known about the comparability of the methods and their specific performance in streams with different fish assemblages. The aim of this investigation was to validate (using M-P sampling as reference) the use of P and S-P electrofishing techniques to accurately assess the richness, density and size distribution of fishes in small streams at both regional and global scale independently of fish assemblages and geographical region. We sampled 50-m-long reaches in a total of 33 lowland stream reaches that were located in different climatic and biogeographical regions (Uruguay and Denmark) and hosted different fish assemblages. Subtropical fish communities exhibited higher richness (Uy: 12-32, Dk: 1-9) and densities (Uy: 1.3-5.2, DK: 0.1-4.9 in. m(-2)) than temperate streams. We applied both "global models" using the entire database (33 sites) and "local models" including the same number of sites but using the climatic region as a model variable. Regression analyses revealed that the P, S-P and M-P methods all provided an adequate picture of the species composition and size distribution, and transfer equations for comparison between methods are thus not required. Conversely, richness was better predicted by S-P and by P techniques for regional and global models, respectively. Transfer equations obtained for abundance revealed that the P and S-P models can accurately transform catch data into M-P estimations. The transfer equations provided here may have great relevance as they allow relatively reliable comparisons to be made between data obtained by different techniques. We also show that less intensive sampling techniques may be equally useful for monitoring purposes as those requiring more intensive efforts (and costs). We encourage validation of our developed transfer equations on data from other regions of the world.

Photosynthetic performance of submerged macrophytes from lowland stream and lake habitats with contrasting CO2 availability.

· We examine the photosynthetic response of submerged plants from streams and lakes with contrasting free-CO2 and nitrogen (N) availability. We hypothesized that: the photosynthetic capacity of stream plants is higher because of higher N availability; the photosynthetic N-use efficiency (PNUE) is also higher because stream plants are acclimated to higher free-CO2; and PNUE is lower in aquatic compared to terrestrial plants. · We tested these hypotheses by measuring tissue-N, photosynthetic capacity and inorganic C extraction capacity in plants collected from streams and lakes and by comparing the PNUE of aquatic plants with previously published PNUE of terrestrial plants. · We found that the organic N content was consistently higher in stream (3.8-6.3% w/w) than in lake plants (1.2-4.3% w/w). The photosynthetic capacity correlated positively with tissue-N. The relationships were similar for stream and lake plants, indicating that N allocation patterns were similar despite variability in free-CO2 between the two habitats. · The slope of the relationship between photosynthetic capacity and tissue-N was lower than found for terrestrial plants, whereas the compensatory N content for photosynthesis was similar. This suggests that PNUE is lower in aquatic plants, perhaps reflecting that the selection pressure for a high C fixation rate per unit N is reduced as a result of low inorganic C availability in the aquatic environment.

European river typologies fail to capture diatom, fish, and macrophyte community composition.

Typology systems are frequently used in applied and fundamental ecology and are relevant for environmental monitoring and conservation. They aggregate ecosystems into discrete types based on biotic and abiotic variables, assuming that ecosystems of the same type are more alike than ecosystems of different types with regard to a specific property of interest. We evaluated whether this assumption is met by the Broad River Types (BRT), a recently proposed European river typology system, that classifies river segments based on abiotic variables, when it is used to group biological communities. We compiled data on the community composition of diatoms, fishes, and aquatic macrophytes throughout Europe and evaluated whether the composition is more similar in site groups with the same river type than in site groups of different river types using analysis of similarities, classification strength, typical species analysis, and the area under zeta diversity decline curves. We compared the performance of the BRT with those of four region-based typology systems, namely, Illies Freshwater Ecoregions, the Biogeographic Regions, the Freshwater Ecoregions of the World, and the Environmental Zones, as well as spatial autocorrelation (SA) classifications. All typology systems received low scores from most evaluation methods, relative to predefined thresholds and the SA classifications. The BRT often scored lowest of all typology systems. Within each typology system, community composition overlapped considerably between site groups defined by the types of the systems. The overlap tended to be the lowest for fishes and between Illies Freshwater Ecoregions. In conclusion, we found that existing broad-scale river typology systems fail to delineate site groups with distinct and compositionally homogeneous communities of diatoms, fishes, and macrophytes. A way to improve the fit between typology systems and biological communities might be to combine segment-based and region-based typology systems to simultaneously account for local environmental variation and historical distribution patterns, thus potentially improving the utility of broad-scale typology systems for freshwater biota.

Diversity and distribution of riparian plant communities in relation to stream size and eutrophication.

The present study was conducted in 47 different riparian areas distributed throughout Denmark to investigate diversity and distributional patterns of plant communities along a lowland stream size gradient (first to fifth order). The investigated areas were representative for Danish riparian areas not in use for agricultural production. We investigated plant community richness along a stream size gradient and the influence of eutrophication on the abundance of different plant communities. Vegetation analyses were performed in transects placed perpendicular to the stream channel, with a total of 1798 plots analyzed. Overall, we found a positive relationship between stream mean depth as a measure of stream size and the number of plant community types identified in the riparian areas. We also found that the abundance of the identified communities was positively correlated with their nutrient preference and negatively correlated with their moisture preference. The abundance of alkaline fens and Molinia meadows (protected community types) in riparian areas decreased with increasing size of the stream, whereas the abundance of humid meadows and wet herb fringes increased with increasing size of the stream. Based on our findings, we recommend that wide buffer zones be established along streams with protected habitat types in the associated riparian areas to reduce the direct impact from agriculture. Furthermore, we recommend that wide buffer zones be established along middle-sized and large streams because several community types may develop.

Phosphorus load to surface water from bank erosion in a Danish lowland river basin.

Phosphorus loss from bank erosion was studied in the catchment of River Odense, a lowland Danish river basin, with the aim of testing the hypothesis of whether stream banks act as major diffuse phosphorus (P) sources at catchment scale. Furthermore, the study aimed at analyzing the impact of different factors influencing bank erosion and P loss such as stream order, anthropogenic disturbances, width of uncultivated buffer strips, and the vegetation of buffer strips. A random stratified procedure in geographical information system (GIS) was used to select two replicate stream reaches covering different stream orders, channelized vs. naturally meandering channels, width of uncultivated buffer strips (≤ 2 m and ≥ 10 m), and buffer strips with different vegetation types. Thirty-six 100-m stream reaches with 180 bank plots and a total of 3000 erosion pins were established in autumn 2006, and readings were conducted during a 3-yr period (2006-2009). The results show that neither stream size nor stream disturbance measured as channelization of channel or the width of uncultivated buffer strip had any significant ( < 0.05) influence on bank erosion and P losses during each of the 3 yr studied. In buffer strips with natural trees bank erosion was significantly ( < 0.05) lower than in buffer strips dominated by grass and herbs. Gross and net P input from bank erosion amounted to 13.8 to 16.5 and 2.4 to 6.3 t P, respectively, in the River Odense catchment during the three study years. The net P input from bank erosion equaled 17 to 29% of the annual total P export and 21 to 62% of the annual export of P from diffuse sources from the River Odense catchment. Most of the exported total P was found to be bioavailable (71.7%) based on a P speciation of monthly suspended sediment samples collected at the outlet of the river basin. The results found in this study have a great importance for managers working with P mitigation and modeling at catchment scale.

Prediction of stream fish assemblages from land use characteristics: implications for cost-effective design of monitoring programmes.

Increasing human impact on stream ecosystems has resulted in a growing need for tools helping managers to develop conservations strategies, and environmental monitoring is crucial for this development. This paper describes the development of models predicting the presence of fish assemblages in lowland streams using solely cost-effective GIS-derived land use variables. Three hundred thirty-five stream sites were separated into two groups based on size. Within each group, fish abundance data and cluster analysis were used to determine the composition of fish assemblages. The occurrence of assemblages was predicted using a dataset containing land use variables at three spatial scales (50 m riparian corridor, 500 m riparian corridor and the entire catchment) supplemented by a dataset on in-stream variables. The overall classification success varied between 66.1-81.1% and was only marginally better when using in-stream variables than when applying only GIS variables. Also, the prediction power of a model combining GIS and in-stream variables was only slightly better than prediction based solely on GIS variables. The possibility of obtaining precise predictions without using costly in-stream variables offers great potential in the design of monitoring programmes as the distribution of monitoring sites along a gradient in ecological quality can be done at a low cost.

Alkalinity and diatom assemblages in lowland streams: How to separate alkalinity from inorganic phosphorus in ecological assessments?

Benthic algae are widely used as ecological indicators of the ecological status of streams because they are widely distributed, they show high species diversity and they respond rapidly to human pressures in particular eutrophication and organic pollution. Recent findings have highlighted that in addition to human pressures, alkalinity may also play a role for community composition as bicarbonate becomes an increasingly important carbon source for photosynthesis when alkalinity increases. With this study, we aimed to elucidate how alkalinity influences the distribution of diatoms in Danish lowland streams, and to explore ifdiatom assemblage patterns can be affected by alkalinity in a way that interferes with the ecological assessment using diatom-based indices. We found that alkalinity affect the benthic algae community in lowland streams and that different species of diatoms were associated with different levels of alkalinity, a finding that might indicate dissimilarities in the efficiency of their HCO3- use. Nitzschia intermedia, Synedra acus, Nitzschia recta, Diatoma tenue, and Nitzschia linearis were associated with high alkalinity, whereas Synedra rumpens, Fragilaria vaucheriae, Psammothidium bioretii, and Gomphonema parvulum were associated with low alkalinity in streams with very low levels of phosphate. We also found that the Danish indicator for ecological status in streams (a combination of two Austrian indices, the Saprobic Index (SID) and the Trophic Index (TID) may exceed levels acceptable for good ecological status in moderate to high alkaline streams despite low phosphate levels. These findings highlight the need for the development of a diagnostic method to disentangle the effects of alkalinity from eutrophication and, additionally, that we need more insight into the autecology of species to interpret ecological assessments to be able to guide management efforts.

Periphyton biomass and life-form responses to a gradient of discharge in contrasting light and nutrients scenarios in experimental lowland streams.

Climate-induced changes in precipitation and land-use intensification affect the discharge of streams worldwide, which, together with eutrophication and loss of riparian canopy, can affect periphyton biomass and composition, and therefore, ultimately the stream functioning. We investigated the responses of periphyton biomass and life-forms (i.e., high profile, low profile and motile) to these changes applying an experimental approach by modulating nutrients (nutrient diffusion substrates enriched with 0.5 M NH4NO3 + 0.031 M KH2PO4 and without nutrient enrichment) and light availability (50% shade and full light) along a gradient in discharge ranging from 0.46 to 3.89 L/s (0.7 to 6.5 cm/s) in twelve large-sized (12- m long) outdoor flumes resembling lowland streams. We also analysed the potential effects of other environmental variables including macroinvertebrates on the responses of periphyton to discharge, nutrients, and light. Light and nutrient availability drastically affected periphyton biomass and composition responses to discharge. Periphyton biomass decreased with increasing discharge when shaded but this did not happen when exposed to full light. Under full light conditions, nutrient enrichment mediated an increase in the periphyton biomass with increasing discharge, possibly reflecting an increased metabolism, but this did not happen under non-enriched conditions. Enrichment further affected the compositional responses of periphyton to discharge, with an increase in the biomass of motile, fast-growing, small-sized flagellated at low discharge conditions, and mitigating a loss of high profile periphyton under higher discharges. Light did not affect periphyton composition, and the abundance or feeding-group composition of the macroinvertebrates did not affect biomass or composition of the periphyton either. Our results suggest that nutrient enrichment and light play an important synergistic role in the responses of the periphyton biomass and composition to discharge and emphasize the relevance of riparian canopy conservation and eutrophication control to avoid periphyton growth under increased discharge scenarios in small lowland streams.

Local physical habitat quality cloud the effect of predicted pesticide runoff from agricultural land in Danish streams.

The combination of intensive agricultural activities and the close connectivity between land and stream emphasise the potential risk of pesticide exposure in Danish streams. Benthic macroinvertebrates are applied in the assessment of stream ecological status, and some sensitive species have been shown to respond strongly to brief pulses of pesticide contamination. In this study we investigate the impact of agriculturally derived pesticides on stream macroinvertebrate communities in Denmark. As a measure of toxic pressure we apply the Runoff Potential. We investigated a total of 212 streams. These were grouped into distinct classes according to the magnitude of pesticide contamination in the period from 2003-2006. A total of 24 different macroinvertebrate indices were applied to detect effects of pesticide runoff (e.g. the SPEAR-index and the number of EPT taxa). We found high predicted pesticide runoff in 39% of the streams, but we found no significant effect of predicted pesticide exposure on stream macroinvertebrate indices. We, additionally, examined the influence of a series of environmental parameters ranging from site scale to catchment scale on the macroinvertebrate community. Relative proportions of gravel, sand and silt in bed sediments explained most of the variation in macroinvertebrate indices as well as the upstream riparian habitat quality. We suggest that the Runoff Potential model overestimate pesticide runoff contamination in Danish streams due the presence of buffer strips enforced by Danish legislation. When pesticide runoff contamination is low to moderate, poor physical properties (indirectly related to agricultural activity) are the main impediment for the ecological quality of Danish streams.

Microbial biofilm community dynamics in five lowland streams.

Stream biofilms are complex aggregates of diverse organism groups that play a vital role in global carbon and nitrogen cycles. Most of the current studies on stream biofilm focus on a limited number of organism groups (e.g., bacteria and algae), and few have included both prokaryote and eukaryote communities simultaneously. In this study, we incubated artificial substrates in five Danish lowland streams exhibiting different hydrological and physico-chemical conditions and explored the dynamics of community composition and diversity of the benthic biofilm, including both prokaryotes and eukaryotes. We found that few phyla in the prokaryote (Gammaproteobacteria and Bacteroidetes) and eukaryote (Cercozoa) communities accounted for over two-thirds of the total abundance at most of the sites. Both prokaryotic and eukaryotic diversity displayed the same temporal patterns, i.e., diversity peaked in July and January. We also found that hydrological and physico-chemical variables significantly explained the variation in the community composition at phylum level for both prokaryotes and eukaryotes. However, a large proportion of variation remained unexplained, which can be ascribed to important but unmeasured variables like light intensity and biological factors such as trophic and non-trophic interactions as revealed by network analysis. Therefore, we suggest that use of a multitrophic level perspective is needed to study biofilm i.e., the "microbial jungles", where high occurrences of trophic and non-trophic interactions are expected.

Danish wetlands remained poor with plant species 17-years after restoration.

For more than two decades, wetland restoration has been successfully applied in Denmark as a tool to protect watercourses from elevated nutrient inputs from agriculture, but little is known about how the flora and fauna respond to restoration. The main objective of this study was therefore to: (1) examine plant community characteristics in 10 wetland sites in the River Odense Kratholm catchment, restored between 2001 and 2011 by re-meandering the stream and disconnecting the tile drains, and (2) explore whether the effects of restoration on plant community characteristics change with the age of the restoration. Specifically, we hypothesised that plant community composition, species richness and diversity would improve with the age of the restoration and eventually approach the state of natural wetland vegetation. We found that the prevailing plant communities could be characterised as humid grasslands, moist fallow fields and improved grasslands, whereas the abundance of natural wetland plant communities (e.g., rich fens, fen-sedge beds and humid grasslands) was lower in both the recently restored as well as in older restored wetlands. Additionally, species richness and diversity did not seem to improve with the age of the restoration. We suggest that the continued high nutrient input at the restored sites in combination with restricted dispersal of wetland plant species may hamper the recovery of natural plant communities and that the sites therefore may stay botanically poor for many decades.

Small-sized omnivorous fish induce stronger effects on food webs than warming and eutrophication in experimental shallow lakes.

Warming, eutrophication, and increased omnivory by small-sized fish are global change processes that induce major effects on the food web structure and primary producers of shallow lakes. Despite the key relevance of phytoplankton and periphyton in freshwaters, the combined and potential synergistic effects of fish omnivory, warming and eutrophication, especially on periphyton, remains little addressed, particularly for subtropical shallow lakes. We experimentally tested the food web effects on phytoplankton and periphyton induced by small visually feeding omnivorous fish (Rhodeus ocellatus), high nutrient enrichment and warming (+4.5 °C) in thirty-two 1000 L-mesocosms simulating littoral conditions of subtropical shallow lakes. We aimed at analysing the mechanisms and responses of periphyton and phytoplankton to these experimental factors. All mesocosms included the submerged macrophytes Vallisneria denseserrulata and Potamogeton lucens and artificial plants at 50% plant volume inhabited, plankton and macroinvertebrates. Small-sized visually feeding omnivorous fish enhanced phytoplankton dominance and periphyton loss. These changes coincided with a decrease in zooplankton biomass and a diversity loss of both zooplankton and macroinvertebrates as well as an increase in snail abundance. Fish presence led to a collapse of cladocerans, thereby releasing the grazing pressure on phytoplankton, and predator and collector macroinvertebrates were replaced with small snails (Radix peregra < 0.5 cm) resulting in enhanced grazing on periphyton. Eutrophication reinforced the fish effects, while warming had weak or no effects. Our results indicate that omnivory by small-sized visually feeding fish may induce stronger effects on the food webs of shallow lakes, towards phytoplankton-dominated states, than the combined effect of nutrient enrichment and warming under the present experimental conditions.

Short-period hydrological regimes override physico-chemical variables in shaping stream diatom traits, biomass and biofilm community functions.

Despite increasing interest in hydrological effects on riverine ecosystems, few studies have documented the impact of hydrology on biofilm community functions, and those existing have typically focused on annual-based hydrological indices. In this study, we conducted monthly samplings during a year in five lowland streams with different flow regimes and investigated the impacts of hydrological conditions and physico-chemical variables on the trait composition of diatoms growing on artificial substrates, biomass (chlorophyll a and ash free dry weight), and biofilm community functions (biochemical processes, i.e., biofilm metabolism and nutrient uptake rates measured in the laboratory). Instead of the commonly used annual-based hydrological indices, we calculated indices for shorter periods (14 and ~28 days) of the hydrological regimes. Results of species-based variation partitioning showed that short-period hydrological indices (10.10 ± 7.18%) contributed more to explain species distribution than physico-chemical variables (5.90 ± 3.83%), indicating the dominant role of hydrology in structuring the diatom community. Specifically, we found different response patterns for different guilds and size classes to the hydrological and physico-chemical variables, and our results demonstrated that species tolerating high disturbance may be more appropriate as indicators of environmental disturbance than low-tolerant species. We also found dominant effects of short-period hydrological events on biomass and biofilm community functions. Despite an overall negative effect of high flow events and flow variations on biomass and biofilm community functions, positive effects on function-biomass ratios were also observed, indicating that the effects of flow regimes on biofilm are complex. In conclusion, our study highlights the importance of including short-period hydrological conditions in studies on environmental factors shaping benthic algae. Based on our results, we recommend use of short-period hydrological conditions when investigating the effects of flow regime on biofilm community composition and functions.

Effects of low flow and co-occurring stressors on structural and functional characteristics of the benthic biofilm in small streams.

Low flow and co-occurring stress is a more and more frequent phenomenon these years in small agricultural streams as a consequence of climate change. In the present study we explored short and longer term structural responses of the stream benthic algae community and biofilm metabolism to multiple stress in small streams applying a semi-experimental approach. We hypothesized that i) a reduction in flow in combination with secondary stress (nutrients and sediments) have immediate effects on the benthic algae community in terms of biomass (chlorophyll a, biovolume), taxonomic and trait (lifeform and size distribution) compositions as well as on metabolism (GPP and CR), and ii) that changes in the benthic algae community persist due to altered environmental settings but that functional redundancy among benthic algae species provides a high level of resilience in metabolism (GPP and CR). Overall, we found that stress imposed by nutrients was less pronounced than stress imposed by fine sediments under low flow, and that nutrient enrichment to some extent mitigated effects of fine sediments. Fine sediment deposition mediated a decline in the fraction of erect algae and/or algae with mucilage stalks but this did not happen under co-occurring stress from both sediments and nutrients. Additionally, fine sediment deposition mediated a decline in GPP of the biofilm, but again this did not happen under co-occurring stress from nutrients. We conclude that 1) the benthic algae community and biofilm metabolism displayed similar resilience to stress imposed by low flow and co-occurring stress from nutrients and sediments on a short and longer time scale and 2) as structure-function adaptations may occur at several trophic levels in the biofilm, more research is needed to explore mechanisms underlying mitigating effects of nutrients in response to sediment deposition under low flow.

Impacts of multiple stressors on freshwater biota across spatial scales and ecosystems.

Climate and land-use change drive a suite of stressors that shape ecosystems and interact to yield complex ecological responses (that is, additive, antagonistic and synergistic effects). We know little about the spatial scales relevant for the outcomes of such interactions and little about effect sizes. These knowledge gaps need to be filled to underpin future land management decisions or climate mitigation interventions for protecting and restoring freshwater ecosystems. This study combines data across scales from 33 mesocosm experiments with those from 14 river basins and 22 cross-basin studies in Europe, producing 174 combinations of paired-stressor effects on a biological response variable. Generalized linear models showed that only one of the two stressors had a significant effect in 39% of the analysed cases, 28% of the paired-stressor combinations resulted in additive effects and 33% resulted in interactive (antagonistic, synergistic, opposing or reversal) effects. For lakes, the frequencies of additive and interactive effects were similar for all spatial scales addressed, while for rivers these frequencies increased with scale. Nutrient enrichment was the overriding stressor for lakes, with effects generally exceeding those of secondary stressors. For rivers, the effects of nutrient enrichment were dependent on the specific stressor combination and biological response variable. These results vindicate the traditional focus of lake restoration and management on nutrient stress, while highlighting that river management requires more bespoke management solutions.

Can a priori defined reference criteria be used to select reference sites in Danish streams? Implications for implementing the Water Framework Directive.

An important step in the implementation of the Water Framework Directive is to define and characterize the natural status, designated as the reference condition (RC). Here we present the results of a type-specific screening for reference stream sites in Denmark using two different approaches. First, we performed a screening applying physicochemical, hydro-morphological and pressure criteria at the catchment, reach and site level of a total of 128 sites a priori selected by the regional water authorities as representing the best sites in Denmark. Second, we performed a GIS screening of all mapped streams in Denmark (26,000 km representing app. 90% of all Danish streams) using solely land use characteristics in the catchment area to target the search for larger stream sites to comply with the WFD requirements of type-specificity. Among the 128 sites we did not find any that fulfilled all criteria applied at the catchment, reach and site level using recommended RC threshold values and only three sites using threshold values that were less strict. Similarly very few km (<1%) of the GIS screened streams fulfilled catchment land use criteria, suggesting that the potential of identifying RC sites in Denmark is very limited. The lack of success in the screening process clearly demonstrates a need for alternative methods to establish RC for Danish streams. We propose a combined approach that includes the development of a guiding image for RC for all the stream biota needed to evaluate the ecological quality. This guiding image should be based on historical data, expert knowledge and investigations in streams situated in countries that are subjected to less intense land use and, at the same time, share both topographical and climatic similarities with Denmark, e.g. the Baltic countries.