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A considerable number of lakes in Sweden have high phosphorus internal loading from the sediments which cause cyanobacterial blooms every summer. Due to potential risks with such blooms for human health, drinking water supply, and ecosystem services, measures need to be taken to control the phosphorus content. Measures to control the phosphorus input from the surrounding land has been in focus. However, the measures have not been sufficient. This is because phosphorus deposited at the bottom of the lakes for many years are finally starting to leak to the water phase when the decomposition of sediments leads to anoxic conditions. In order to determine effective and efficient lake restoration measures, methods for lake restoration decision support by a multi-criteria analysis and the application of a decision analysis are developed. The multi-criteria analysis includes the determination of costs, longevity, and efficacy of six common lake restoration measures to reduce internal phosphorous loads in two lakes selected as a case study. The results show that aluminum treatment combines a highest efficacy with a high-cost efficiency being thus the optimal identified measure. The method involves adding an aluminum solution to the lakes’ sediment, which binds phosphorus, preventing it to be released to the water column. The multi-criteria model is integrated to a decision analytical model. The decision analytical model is used to identify the monetary socio-economic and environmental boundaries for the implementation of the optimal lake restoration measure.

In freshwater habitats, invasive species and the increase of cyanobacterial blooms have been identified as a major cause of biodiversity loss. The invasive cyanobacteria Raphidiopsis raciborskii a toxin-producing and bloom-forming species affecting local biodiversity and ecosystem services is currently expanding its range across Europe. We used species distribution models (SDMs) and regional bioclimatic environmental variables, such as temperature and precipitation, to identify suitable areas for the colonization and survival of R. raciborskii, with special focus on the geographic extent of potential habitats in Northern Europe. SDMs predictions uncovered areas of high occurrence probability of R. raciborskii in locations where it has not been recorded yet, e.g. some areas in Central and Northern Europe. In the southeastern part of Sweden, areas of suitable climate for R. raciborskii corresponded with lakes of high concentrations of total phosphorus, increasing the risk of the species to thrive. To our knowledge, this is the first attempt to predict areas at high risk of R. raciborskii colonization in Europe. The results from this study suggest several areas across Europe that would need monitoring programs to determine if the species is present or not, to be able to prevent its potential colonization and population growth. Regarding an undesirable microorganism like R. raciborskii, authorities may need to start information campaigns to avoid or minimize the spread.

Phytoplankton is a functionally important taxonomic group commonly used in the assessment of lakes. Using data from minimally disturbed lakes (n = 151) and relatively easily obtained environmental variables, we modeled the probability of taxon occurrence and taxon-specific biomass (mg L−1) of 183 phytoplankton taxa using variables characterizing spatial context, ecosystem size and land cover. Data from lakes putatively affected by one or more pressures (n = 162) were used to evaluate responses to anthropogenic stress. Longitude and latitude were the two best predictors of phytoplankton biomass, followed by the percent of catchment area classified as coniferous forest area and altitude. Model accuracy was highest for relatively predominant taxa, and taxa with relatively wide distributions did not improve model accuracy. Assemblage similarity (1-BC index) and the phytoplankton trophic index (PTI) calculated using predicted and observed phytoplankton assemblages were used to evaluate model performance. Significant changes in phytoplankton assemblages were associated with agricultural and forestry pressures. Encouragingly the PTI index responded as predicted to agricultural land use but not forestry, implying that PTI was a robust indicator of agricultural effects and that forestry effects on assemblage composition were not related to increased nutrients. Combined, our findings showed that phytoplankton assemblages can be reliably predicted using a few, often readily available, environmental variables, and that predictions of assemblage composition and taxon-specific biomass and biological metrics based on phytoplankton taxon-specific biomass are reliable indicators of anthropogenic pressures, supporting their use in lake management.

Despite their key role in biogeochemical processes, particularly the methane cycle, archaea are widely underrepresented in molecular surveys because of their lower abundance compared to bacteria and eukaryotes. Here, we use parallel high‐resolution small subunit rRNA gene sequencing to explore archaeal diversity in 109 Swedish lakes and correlate archaeal community assembly mechanisms to large‐scale latitudinal, climatic (nemoral to arctic), and nutrient (oligotrophic to eutrophic) gradients. Sequencing with universal primers showed the contribution of archaea was on average 0.8% but increased up to 1.5% of the three domains in forest lakes. Archaea‐specific sequencing revealed that freshwater archaeal diversity could be partly explained by lake variables associated with nutrient status. Combined with deterministic co‐occurrence patterns this finding suggests that ecological drift is overridden by environmental sorting, as well as other deterministic processes such as biogeographic and evolutionary history, leading to lake‐specific archaeal biodiversity. Acetoclastic, hydrogenotrophic and methylotrophic methanogens as well as ammonia‐oxidizing archaea were frequently detected across the lakes. Archaea‐specific sequencing also revealed representatives of Woesearchaeota and other phyla of the DPANN superphylum. This study adds to our understanding of the ecological range of key archaea in freshwaters and links these taxa to hypotheses about processes governing biogeochemical cycles in lakes. peerReviewed

The partial pressure of CO2 (pCO(2)) in lake water, and thus CO2 emissions from lakes are controlled by hydrologic inorganic carbon inputs into lakes, and in-lake carbon transformation (mainly organic carbon mineralization and CO2 uptake by primary producers). In boreal lakes, CO2 uptake by phytoplankton is often considered to be of minor importance. At present, however, it is not known in which and how many boreal lakes phytoplankton CO2 uptake has a sizeable influence on the lake water pCO(2). Using water physico-chemical and phytoplankton data from 126 widely spread Swedish lakes from 1992 to 2012, we found that pCO(2) was negatively related to phytoplankton carbon in lakes in which the phytoplankton share in TOC (C-phyto:TOC ratio) exceeded 5%. Total phosphorus concentration (TP) was the strongest predictor of spatial variation in the C-phyto:TOC ratio, where C-phyto:TOC ratios>5% occurred in lakes with TP>30 mu gl(-1). These lakes were located in the hemi-boreal zone of central and southern Sweden. We conclude that during summer, phytoplankton CO2 uptake can reduce the pCO(2) not only in warm eutrophic lakes, but also in relatively nutrient poor hemi-boreal lakes.

Important drivers of gross primary production (GPP) and ecosystem respiration (ER) in lakes are temperature, nutrients, and light availability, which are predicted to be affected by climate change. Little is known about how these three factors jointly influence shallow lakes metabolism and metabolic status as net heterotrophic or autotrophic. We conducted a pan-European standardized mesocosm experiment covering a temperature gradi- ent from Sweden to Greece to test the differential temperature sensitivity of GPP and ER at two nutrient levels (mesotrophic or eutrophic) crossed with two water levels (1 m and 2 m) to simulate different light regimes. The findings from our experiment were compared with predictions made according the metabolic theory of ecology (MTE). GPP and ER were significantly higher in eutrophic mesocosms than in mesotrophic ones, and in shallow mesocosms compared to deep ones, while nutrient status and depth did not interact. The estimated temperature gains for ER of ~ 0.62 eV were comparable with those predicted by MTE. Temperature sensitivity for GPP was slightly higher than expected ~ 0.54 eV, but when corrected for daylight length, it was more consistent with predictions from MTE ~ 0.31 eV. The threshold temperature for the switch from autotrophy to heterotrophy was lower under mesotrophic (~ 11 C) than eutrophic conditions (~ 20 C). Therefore, despite a lack of signifi- cant temperature-treatment interactions in driving metabolism, the mesocosm’s nutrient level proved to be cru- cial for how much warming a system can tolerate before it switches from net autotrophy to net heterotrophy. We thank the technical staff at the various experimental sites for their support. We thank Alena S. Gesell, Deniz Özkundakci, Jan-Hendrik Schlei- mer, Silke Schmidt, Torsten Seltmann, and Tom Shatwell for their helpful discussions during the preparation of this manuscript. We thank Anne Mette Poulsen, Adam Wilkins, and Michael Thayne for their valuable edit- ing of the manuscript. We are also grateful to reviewers for their valuable comments on the manuscript. This study was supported by FP-7 REFRESH (Adaptive strategies to Mitigate the Impacts of Climate Change on European Freshwater Ecosystems, Contract No. 244121) and the MARS project (Managing Aquatic ecosystems and water Resources under multi- ple Stress), funded under the 7th EU Framework Programme, Theme 6 (Environment including Climate Change), Contract No. 603378 (http:// www.mars-project.eu), TUBITAK- ÇAYDAG (projects No. 105Y332 and 110Y125), the Middle East Technical University (METU)-BAP program of Turkey. EJ was further supported by AU Centre for Water Technology and A_lÇ was also supported by TUBITAK (project 296 Nos. 105Y332 and 110Y125). We thank the technical staff at the various experimental sites for their support. We thank Alena S. Gesell, Deniz Özkundakci, Jan-Hendrik Schlei- mer, Silke Schmidt, Torsten Seltmann, and Tom Shatwell for their helpful discussions during the preparation of this manuscript. We thank Anne Mette Poulsen, Adam Wilkins, and Michael Thayne for their valuable edit- ing of the manuscript. We are also grateful to reviewers for their valuable comments on the manuscript. This study was supported by FP-7 REFRESH (Adaptive strategies to Mitigate the Impacts of Climate Change on European Freshwater Ecosystems, Contract No. 244121) and the MARS project (Managing Aquatic ecosystems and water Resources under multi- ple Stress), funded under the 7th EU Framework Programme, Theme 6 (Environment including Climate Change), Contract No. 603378 (http:// www.mars-project.eu), TUBITAK- ÇAYDAG (projects No. 105Y332 and 110Y125), the Middle East Technical University (METU)-BAP program of Turkey. EJ was further supported by AU Centre for Water Technology and A_lÇ was also supported by TUBITAK (project 296 Nos. 105Y332 and 110Y125).

International audience; Assessment of ecological status for the European Water Framework Directive (WFD) is based on "Biological Quality Elements" (BQEs), namely phytoplankton, benthic flora, benthic invertebrates and fish. Morphological identification of these organisms is a time-consuming and expensive procedure. Here, we assess the options for complementing and, perhaps, replacing morphological identification with procedures using eDNA, metabarcoding or similar approaches. We rate the applicability of DNA-based identification for the individual BQEs and water categories (rivers, lakes, transitional and coastal waters) against eleven criteria, summarised under the headlines representativeness (for example suitability of current sampling methods for DNA-based identification, errors from DNA-based species detection), sensitivity (for example capability to detect sensitive taxa, unassigned reads), precision of DNA-based identification (knowledge about uncertainty), comparability with conventional approaches (for example sensitivity of metrics to differences in DNA-based identification), cost effectiveness and environmental impact. Overall, suitability of DNA-based identification is particularly high for fish, as eDNA is a well-suited sampling approach which can replace expensive and potentially harmful methods such as gill-netting, trawling or electrofishing. Furthermore, there are attempts to replace absolute by relative abundance in metric calculations. For invertebrates and phytobenthos, the main challenges include the modification of indices and completing barcode libraries. For phytoplankton, the barcode libraries are even more problematic, due to the high taxonomic diversity in plankton samples. If current assessment concepts are kept, DNA-based identification is least appropriate for macrophytes (rivers, lakes) and angiosperms/macroalgae (transitional and coastal waters), which are surveyed rather than sampled. We discuss general implications of implementing DNA-based identification into standard ecological assessment, in particular considering any adaptations to the WFD that may be required to facilitate the transition to molecular data.

Freshwater ecosystems are important for global biodiversity and provide essential ecosystem services. There is consensus in the scientific literature that freshwater ecosystems are vulnerable to the impacts of environmental change, which may trigger irreversible regime shifts upon which biodiversity and ecosystem services may be lost. There are profound uncertainties regarding the management and assessment of the vulnerability of freshwater ecosystems to environmental change. Quantitative approaches are needed to reduce this uncertainty. We describe available statistical and modeling approaches along with case studies that demonstrate how resilience theory can be applied to aid decision-making in natural resources management. We highlight especially how long-term monitoring efforts combined with ecological theory can provide a novel nexus between ecological impact assessment and management, and the quantification of systemic vulnerability and thus the resilience of ecosystems to environmental change.

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.

Research focusing on biodiversity responses to the interactions of ecosystem size and anthropogenic stressors are based mainly on correlative gradient studies, and may therefore confound size-stress relationships due to spatial context and differences in local habitat features across ecosystems. We investigated how local factors related to anthropogenic stressors (e.g., eutrophication) interact with ecosystem size to influence species diversity. In this study, constructed lake mesocosms (with two contrasting volumes: 1020 (shallow mesocosms) and 2150 (deep mesocosms) litres) were used to simulate ecosystems of different size and manipulated nutrient levels to simulate mesotrophic and hypertrophic conditions. Using a factorial design, we assessed how the interaction between ecosystem size and nutrients influences phytoplankton diversity. We assessed community metrics (richness, diversity, evenness and total biovolumes) and multivariate community structure over a growing season (May to early November 2011). Different community structures were found between deep and shallow mescosoms with nutrient enrichment: Cyanobacteria dominated in the deep and Charophyta in the shallow mesocosms. In contrast, phytoplankton communities were more similar to each other in the low nutrient treatments; only Chlorophyta had generally a higher biovolume in the shallow compared to the deep mesocosms. These results suggest that ecosystem size is not only a determinant of species diversity, but that it can mediate the influence of anthropogenic effects on biodiversity. Such interactions increase the uncertainty of global change outcomes, and should therefore not be ignored in risk/impact assessment and management.

Boreal lakes undergo broad-scale environmental change over time, but biodiversity responses to these changes, particularly at macroecological scales, are not well known. We studied long-term trends (1992-2009) of environmental variables and assessed α, β, and γ diversity responses of phytoplankton and littoral invertebrates to these changes. Diversity was assessed based on taxon richness ("richness") and the exponentiated Shannon entropy ("diversity"). Almost all environmental variables underwent significant monotonic change over time, indicating mainly decreasing acidification, water clarity and nutrient concentrations in the lakes. These variables explained about 54 and 38% of variance in regression models of invertebrates and phytoplankton, respectively. Despite this, most diversity-related variables fluctuated around a long-term mean. Only α and γ richness and diversity of invertebrates increased monotonically through time, and these patterns correlated significantly with local and regional abundances. Results suggest that biodiversity in boreal lakes is currently stable, with no evidence of regional biotic homogenization or local diversity loss. Results also show that richness trends between phytoplankton and invertebrates were widely uncorrelated, and the same was found for diversity trends. Also, within each taxonomic group, temporal patterns of richness and diversity were largely uncorrelated with each other. From an applied perspective, this suggest that long-term trends of biodiversity in boreal lakes at a macroecological scale cannot be accurately assessed without multiple lines of evidence, i.e. through the use of multiple taxa and diversity-related variables in the analyses.

It is frequently observed that the local relative abundances of aquatic microbial taxa are correlated with their average relative abundance at the regional scale, which results in the composition of different communities being more similar than expected by chance or invariant. The degree to which communities within a region match the regional average community is variable and likely depends on several different mechanisms that control the process of microbial community assembly. Here, we show that environmental variables were associated with the community specific degree of regional invariance in 9 of 10 datasets of microbial communities in aquatic systems, being the main set of variables explaining differences in regional invariance in 5 of them. This indicates that variation in local environmental conditions across a region reduces the degree of regional invariance amongst communities. Spatial distances between communities were not related to the degrees of regional invariance, but in 7 of the datasets, regional invariance differed among different parts of the regions, particularly for phytoplankton communities. This suggests an influence of spatial or historical processes on the community specific degree of regional invariance. We conclude that both local environmental conditions and spatial/historical processes cause between-site differences in the degree of invariance between local and regional abundances in aquatic microbial metacommunities. We argue that studies of regional invariance can be an important complement to other statistical methods due to its propensity to detect variation in stochastic processes along gradients.

A central goal in ecology is to grasp the mechanisms that underlie and maintain biodiversity and patterns in its spatial distribution can provide clues about those mechanisms. Here, we investigated what might determine the bacterioplankton richness (BR) in lakes by means of 454 pyrosequencing of the 16S rRNA gene. We further provide a BR estimate based upon a sampling depth and accuracy, which, to our knowledge, are unsurpassed for freshwater bacterioplankton communities. Our examination of 22 669 sequences per lake showed that freshwater BR in fourteen nutrient-poor lakes was positively influenced by nutrient availability. Our study is, thus, consistent with the finding that the supply of available nutrients is a major driver of species richness; a pattern that may well be universally valid to the world of both micro- and macro-organisms. We, furthermore, observed that BR increased with elevated landscape position, most likely as a consequence of differences in nutrient availability. Finally, BR decreased with increasing lake and catchment area that is negative species–area relationships (SARs) were recorded; a finding that re-opens the debate about whether positive SARs can indeed be found in the microbial world and whether positive SARs can in fact be pronounced as one of the few ‘laws' in ecology.

The turnover and distribution of energy and nutrients in food webs is influenced by consumer stoichiometry. Although the stoichiometry of heterotrophs is generally considered to vary only little, t ...

The complex nature of ecological systems limits the unambiguous determination of mechanisms that drive resilience to natural disturbance or anthropogenic stress. Using eight-year time series data from boreal lakes with and without bloom formation of an invasive alga (Gonyostomum semen, Raphidophyceae), we studied resilience of phytoplankton communities in relation to recurring bloom impacts. We first characterized phytoplankton community dynamics in both lake types using univariate metrics of community structure (evenness, species richness, biovolume and Simpson diversity). All metrics, except species richness, were substantially altered and showed an inherent stronger variability in bloom lakes relative to reference lakes. We assessed resilience mechanisms using a multivariate time series modelling technique. The models captured clear successional dynamics of the phytoplankton communities in all lakes, whereby different groups of species were substituted sequentially over the ice-free period. The models also identified that G. semen impacts in bloom lakes were only manifested within a single species group, not across species groups, highlighting the rapid renewal of the phytoplankton communities upon bloom collapse. These results provide empirical support of the cross-scale resilience model. Cross-scale resilience could provide an explanation for the paradox that similar species richnesses are seen in bloom-forming lakes and reference lakes despite the clear difference between the community features of the two different sets of lakes investigated.

1. We tested whether increasing atmospheric nitrogen (N) deposition along a north-south gradient intensifies epilithic phosphorus (P) limitation in oligotrophic Swedish lakes from the north to the ...

OA hybrid Although the Water Framework Directive specifies that macrophytes and phytobenthos should be used for the ecological assessment of lakes and rivers, practice varies widely throughout the EU. Most countries have separate methods for macrophytes and phytobenthos in rivers; however, the situation is very different for lakes. Here, 16 countries do not have dedicated phytobenthos methods, some include filamentous algae within macrophyte survey methods whilst others use diatoms as proxies for phytobenthos. The most widely-cited justification for not having a dedicated phytobenthos method is redundancy, i.e. that macrophyte and phytoplankton assessments alone are sufficient to detect nutrient impacts. Evidence from those European Union Member States that have dedicated phytobenthos methods supports this for high level overviews of lake condition and classification; however, there are a number of situations where phytobenthos may contribute valuable information for the management of lakes.

Species–area relationships (SAR) are fundamental in the understanding of biodiversity patterns and of critical importance for predicting species extinction risk worldwide. Despite the enormous attention given to SAR in the form of many individual analyses, little attempt has been made to synthesize these studies. We conducted a quantitative meta-analysis of 794 SAR, comprising a wide span of organisms, habitats and locations. We identified factors reflecting both pattern-based and dynamic approaches to SAR and tested whether these factors leave significant imprints on the slope and strength of SAR. Our analysis revealed that SAR are significantly affected by variables characterizing the sampling scheme, the spatial scale, and the types of organisms or habitats involved. We found that steeper SAR are generated at lower latitudes and by larger organisms. SAR varied significantly between nested and independent sampling schemes and between major ecosystem types, but not generally between the terrestrial and the aquatic realm. Both the fit and the slope of the SAR were scale-dependent. We conclude that factors dynamically regulating species richness at different spatial scales strongly affect the shape of SAR. We highlight important consequences of this systematic variation in SAR for ecological theory, conservation management and extinction risk predictions.

A gradient of dissolved organic carbon (DOC) was added to enclosures in a high Arctic lake (Svalbard, 79° N). The aim was to simulate the effect of increased concentrations of DOC that will be a predicted effect of increased temperature and precipitation. The study aimed to provide information on the overall effects of such increased levels of DOC on the pelagic food-web, as well as the increased attenuation of UV-radiation (UV-R) caused by increased DOC. The biomass development of phytoplankton, heterotrophic bacteria and ciliates from 15. July (shortly after ice-off) to 8. August revealed a consistent pattern across all enclosures. Initial phytoplankton biomass decreased from maxima around 600μg Cl -1 towards 50μg Cl -1 by the end of July. Similarly, ciliate biomass decreased from ∼100 to 5 μg Cl -1 , while heterotrophic bacteria decreased from initially 280μg Cl -1 to biomasses near 100μg Cl -1 . Over the same period, zooplankton biomass (almost a monoculture of Daphnia tenebrosa) increased from

1. Field data from five unproductive Swedish lakes were used to investigate the occurrence of mixotrophic flagellates in relation to bacterioplankton, autotrophic phytoplankton, heterotrophic flage ...

1. Biomass and production of picophytoplankton, phytoplankton and heterotrophic bacterioplankton were measured in seven lakes, exhibiting a broad range in water colour because of humic substances. ...

Factors controlling pelagic and sediment heterotrophic bacterial numbers and production in lakes.

1. The biomass and production of picophytoplankton, large phytoplankton and heterotrophic bacterioplankton were measured in humic Lake Ortrasket, northern Sweden during four consecutive summers. 2. ...

1. Two small humic lakes in northern Sweden with concentrations of dissolved organic carbon (DOC) between 15 and 20 mg L-1 were fertilized with inorganic phosphorus (P) and inorganic nitrogen (N), ...

The aim of this study was to investigate the interactions between the cyanobacteria (Microcystis aeruginosa) and the potential grazer (Ochromonas sp.) with regard to colony formation. Two kinds of treatments were carried out: (i) In the dialyse experiment Microcystis aeruginosa and Ochromonas sp. were physically separated by a dialyse tubing. (ii) In the contact experiment interactions between Microcystis aeruginosa, heterotrophic bacteria and Ochromonas sp. in different concentrations were investigated. In one treatment where the predator Ochromonas sp. came in direct contact with Microcystis, aggregates were formed.

The influence of water colour and effective light climate on mixotrophic phytoflagellates in three small Swedish dystrophic lakes

Humic lakes with high inputs of allochthonous dissolved organic carbon have a pelagic food chain that, to a large extent, is based on bacterioplankton energy mobilization from allochthonous organic carbon compounds. This is in contrast to clear lakes in which total pelagic production is based mainly on phytoplankton photosynthesis. The energy economy in humic lakes may be less efficient than in clear lakes, because it is likely that one more link is included in the food chain. Lake data from Scandinavia and North America demonstrate that shifts between food chains based on heterotrophic production and food chains based on primary production can take place at moderate increases or decreases in the concentration of dissolved organic carbon from allochthonous sources. Large variations in the loading of allochthonous organic carbon (e.g., due to climatic variations) may have considerable effects on the biostructure and productivity of lakes.

Climate change is expected to profoundly affect both temperature and net precipitation, with implications for lake water level. We describe the design of a harmonized, simultaneous, cross- European mesocosm experiment to elucidate the effects of climate change on community structure, functioning, and metabolism in shallow lakes at low and high nutrient levels with contrasting depths (1 and 2 m). We used cylindrical (D = 1.2 m) tanks that were either 1.2 or 2.2 m high, each having a 10-cm sediment layer. We inoculated the mesocosms with a mixed sample of sediment and plankton from lakes with contrasting nutrient concentrations and added macrophytes and planktivorous fish. Sediment was pre-equilibrated to the required experimental nutrient concentration. During the experiment the water level decreased with increasing temperature (up to 90 cm in the Mediterranean mesocosms) while conductivity increased. The average chlorophyll a concentration increased with temperature in the deep mesocosms but was more variable in the shallow mesocosms. Macrophyte

Liming has been used extensively in Scandinavia and elsewhere since the 1970s to counteract the negative effects of acidification. Communities in limed lakes usually return to acidified conditions once liming is discontinued, suggesting that liming is unlikely to shift acidified lakes to a state equivalent to pre-acidification conditions that requires no further management intervention. While this suggests a low resilience of limed lakes, attributes that confer resilience have not been assessed, limiting our understanding of the efficiency of costly management programs. In this study, we assessed community metrics (diversity, richness, evenness, biovolume), multivariate community structure and the relative resilience of phytoplankton in limed, acidified and circum-neutral lakes from 1997 to 2009, using multivariate time series modeling. We identified dominant temporal frequencies in the data, allowing us to track community change at distinct temporal scales. We assessed two attributes of relative resilience (cross-scale and within-scale structure) of the phytoplankton communities, based on the fluctuation frequency patterns identified. We also assessed species with stochastic temporal dynamics. Liming increased phytoplankton diversity and richness; however, multivariate community structure differed in limed relative to acidified and circum-neutral lakes. Cross-scale and within-scale attributes of resilience were similar across all lakes studied but the contribution of those species exhibiting stochastic dynamics was higher in the acidified and limed compared to circum-neutral lakes. From a resilience perspective, our results suggest that limed lakes comprise a particular condition of an acidified lake state. This explains why liming does not move acidified lakes out of a “degraded” basin of attraction. In addition, our study demonstrates the potential of time series modeling to assess the efficiency of restoration and management outcomes through quantification of the attributes contributing to resilience in ecosystems.

24 páginas, 2 figuras, 1 tabla., The protection, preservation and restoration of aquatic ecosystems and their functions are of global importance. For European states it became legally binding mainly through the EUWater Framework Directive (WFD). In order to assess the ecological status of a given water body, aquatic biodiversity data are obtained and compared to a reference water body. The quantified mismatch obtained determines the extent of potential management actions. The current approach to biodiversity assessment is based on morpho-taxonomy. This approach has many drawbacks such as being time consuming, limited in temporal and spatial resolution, and error-prone due to the varying individual taxonomic expertise of the analysts. Novel genomic tools can overcome many of the aforementioned problems and could complement or even replace traditional bioassessment. Yet, a plethora of approaches are independently developed in different institutions, thereby hampering any concerted routine application. The goal of this Action is to nucleate a group of researchers across disciplines with the task to identify gold-standard genomic tools and novel ecogenomic indices for routine application in biodiversity assessments of European fresh- and marine water bodies. Furthermore, DNAqua-Net will provide a platform for training of the next generation of European researchers preparing them for the new technologies. Jointly with water managers, politicians, and other stakeholders, the group will develop a conceptual framework for the standard application of eco-genomic tools as part of legally binding assessments.

14 páginas, 1 figura, 4 tablas., Phytoplankton constitutes a diverse array of short-lived organisms which derive their nutrients from the water column of lakes. These features make this community the most direct and earliest indicator of the impacts of changing nutrient conditions on lake ecosystems. It also makes them particularly suitable for measuring the success of restoration measures following reductions in nutrient loads. This paper integrates a large volume of work on a number of measures, or metrics, developed for using phytoplankton to assess the ecological status of European lakes, as required for the Water Framework Directive. It assesses the indicator strength of these metrics, specifically in relation to representing the impacts of eutrophication. It also examines how these measures vary naturally at different locations within a lake, as well as between lakes, and how much variability is associated with different replicate samples, different months within a year and between years. On the basis of this analysis, three of the strongest metrics (chlorophyll-a, phytoplankton trophic index (PTI), and cyanobacterial biovolume) are recommended for use as robust measures for assessing the ecological quality of lakes in relation to nutrient-enrichment pressures and a minimum recommended sampling frequency is provided for these three metrics., This paper is a result of the project WISER (Water bodies in Europe: Integrative Systems to assess Ecological status and Recovery) funded by the European Union under the 7th Framework Programme, Theme 6 (Environment including Climate Change) (contract No. 226273).

The recognition and discrimination of phytoplankton species is one of the foundations of freshwater biodiversity research and environmental monitoring. This step is frequently a bottleneck in the analytical chain from sampling to data analysis and subsequent environmental status evaluation. Here we present phytoplankton diversity data from 49 lakes including three seasonal surveys assessed by next generation sequencing (NGS) of 16S ribosomal RNA chloroplast and cyanobacterial gene amplicons and also compare part of these datasets with identification based on morphology. Direct comparison of NGS to microscopic data from three time-series showed that NGS was able to capture the seasonality in phytoplankton succession as observed by microscopy. Still, the PCR-based approach was only semi-quantitative, and detailed NGS and microscopy taxa lists had only low taxonomic correspondence. This is probably due to, both, methodological constraints and current discrepancies in taxonomic frameworks. Discrepancies included Euglenophyta and Heterokonta that were scarce in the NGS but frequently detected by microscopy and Cyanobacteria that were in general more abundant and classified with high resolution by NGS. A deep-branching taxonomically unclassified cluster was frequently detected by NGS but could not be linked to any group identified by microscopy. NGS derived phytoplankton composition differed significantly among lakes with different trophic status, showing that our approach can resolve phytoplankton communities at a level relevant for ecosystem management. The high reproducibility and potential for standardization and parallelization makes our NGS approach an excellent candidate for simultaneous monitoring of prokaryotic and eukaryotic phytoplankton in inland waters. peerReviewed

Boreal lakes undergo broad-scale environmental change over time, but biodiversity responses to these changes, particularly at macroecological scales, are not well known. We studied long-term trends (1992-2009) of environmental variables and assessed α, β, and γ diversity responses of phytoplankton and littoral invertebrates to these changes. Diversity was assessed based on taxon richness ("richness") and the exponentiated Shannon entropy ("diversity"). Almost all environmental variables underwent significant monotonic change over time, indicating mainly decreasing acidification, water clarity and nutrient concentrations in the lakes. These variables explained about 54 and 38% of variance in regression models of invertebrates and phytoplankton, respectively. Despite this, most diversity-related variables fluctuated around a long-term mean. Only α and γ richness and diversity of invertebrates increased monotonically through time, and these patterns correlated significantly with local and regional abundances. Results suggest that biodiversity in boreal lakes is currently stable, with no evidence of regional biotic homogenization or local diversity loss. Results also show that richness trends between phytoplankton and invertebrates were widely uncorrelated, and the same was found for diversity trends. Also, within each taxonomic group, temporal patterns of richness and diversity were largely uncorrelated with each other. From an applied perspective, this suggest that long-term trends of biodiversity in boreal lakes at a macroecological scale cannot be accurately assessed without multiple lines of evidence, i.e. through the use of multiple taxa and diversity-related variables in the analyses.

Despite improvements in waste water treatment systems, the impact of anthropogenic nutrient sources remains a key issue for the management of European lakes. The Water Framework Directive provides a mechanism through which progress can be made on this issue. The Directive requires a classification of the ecological status of phytoplankton which includes an assessment of taxonomic composition. In this paper we present a composition metric, the Plankton Trophic Index, that was developed in the WISER EU FP7 project and demonstrate how it has been used to compare national phytoplankton classification systems in Northern and Central Europe. The metric was deerived from sumaer phytoplankton data from 1795 lakes, summarised by genus covering 20 European countries. We show that it is significantly related to total phosphorus concentrations, but that it is alos sensitive to alkalinity, lake size and climatic variables. Through the use of country specific reference values for the index we demonstrate that it is significantly related to othrt national phytoplankton assessment systems and illustrate for a sinngle lake type how it was used to intercalibrate Water Framework Directive boundaries for different countries., JRC.H.1-Water Resources

Microbial ecology has focused much on causes of between-site variation in community composition. By analysing five data-sets each of aquatic bacteria and phytoplankton, we demonstrated that microbial communities show a large degree of similarity in community composition and that abundant taxa were widespread, a typical pattern for many metazoan metacommunities. The regional abundance of taxa explained on average 85 and 41% of variation in detection frequency and 58 and 31% of variation in local abundances for bacteria and phytoplankton, respectively. However, regional abundance explained less variation in local abundances with increasing environmental variation between sites within data-sets. These findings indicate that the studies of microbial assemblages need to consider similarities between communities to better understand the processes underlying the assembly of microbial communities. Finally, we propose that the degree of regional invariance can be linked to the evolution of microbes and the variation in ecosystem functions performed by microbial communities.

Species-area relationships (SAR) are fundamental in the understanding of biodiversity patterns and of critical importance for predicting species extinction risk worldwide. Despite the enormous attention given to SAR in the form of many individual analyses, little attempt has been made to synthesize these studies. We conducted a quantitative meta-analysis of 794 SAR, comprising a wide span of organisms, habitats and locations. We identified factors reflecting both pattern-based and dynamic approaches to SAR and tested whether these factors leave significant imprints on the slope and strength of SAR. Our analysis revealed that SAR are significantly affected by variables characterizing the sampling scheme, the spatial scale, and the types of organisms or habitats involved. We found that steeper SAR are generated at lower latitudes and by larger organisms. SAR varied significantly between nested and independent sampling schemes and between major ecosystem types, but not generally between the terrestrial and the aquatic realm. Both the fit and the slope of the SAR were scale-dependent. We conclude that factors dynamically regulating species richness at different spatial scales strongly affect the shape of SAR. We highlight important consequences of this systematic variation in SAR for ecological theory, conservation management and extinction risk predictions.

An oligotrophic clearwater lake, initially characterized by a pronounced dominance of autotrophic phytoplankton and mostly by one species, the green alga Botryococcus, was subject to additions of dissolved organic carbon in the form of white sugar (sucrose) during two consecutive years. The hypothesis tested was that it is organic carbon per se, and not other possible effects of humic substances, that determines the differences in structure of the planktonic ecosystem between humic and clearwater lakes. The additions of DOC resulted in a significant increase in bacterial biomass and a decrease in the biomass of autotrophic phytoplankton. The biomass of mixotrophic and heterotrophic flagellates instead increased significantly, whereas no effects were found to propagate to higher trophic levels. As a result of the changes among biota, total planktonic biomass also decreased to a level typical of nearby humic lakes. We suggest that it is the carbon component of humic material and its utilization by bacterioplankton that determines the structure and function of the pelagic food web in humic lakes.

A laboratory experiment was performed to test whether differences in nutrient and energy demands between picophytoplankton and heterotrophic bacteria can explain the apparent inverse biomass relationship between these organisms in lakes along gradients of organic carbon and nutrients. Growth rates and final yield of cells were analyzed in crossed gradients of glucose and phosphate. Concentrations of phosphate (10, 25, and 60 microg P L(-1)) and glucose (0, 0.3, and 3 mg C L(-1)) were used in all possible combinations giving 9 different treatments. Heterotrophic bacteria had higher maximum growth rates in all treatments and became larger than picophytoplankton in many treatments. The variance in abundance of heterotrophic bacteria between treatments could almost completely be explained by the combined effects of glucose and P. In treatments where carbon limitation slowed down the growth of heterotrophic bacteria, picophytoplankton became abundant and these organisms showed a positive response to P in combination with a negative response to glucose. The negative effect of glucose on picophytoplankton is suggested to be indirect and caused by competition with bacteria that are favored by organic C. The results suggest that competition for phosphate between phytoplankton and bacteria is not size-dependent, that heterotrophic bacteria are superior competitors for P, and that organic C produced by picophytoplankton was of minor importance for heterotrophic bacteria.