Ecological threshold responses in European lakes and their applicability for the Water Framework Directive (WFD) implementation: synthesis of lakes results from the REBECCA project

The objective of this synthesis is to present the key messages and draw the main conclusions from the work on lakes in the REBECCA project, pointing out their links to theoretical ecology and their applicability for the WFD implementation. Type-specific results were obtained from analyses of large pan-European datasets for phytoplankton, macrophytes, macroinvertebrates and fish, and indicators and relationships showing the impact of eutrophication or acidification on these biological elements were constructed. The thresholds identified in many of the response curves are well suited for setting ecological status class boundaries and can be applied in the intercalibration of classification systems. Good indicators for phytoplankton (chrysophytes, cyanobacteria) and macrophytes (isoetids and charaphytes) responses to eutrophication were identified, and the level of eutrophication pressure needed to reach the thresholds for these indicators was quantified. Several existing metrics developed for macrophytes had low comparability and need further harmonisation to be useful for intercalibration of classification systems. For macroinvertebrates, a number of metrics developed for rivers turned out to be less useful to describe lake responses to eutrophication and acidification, whereas other species based indicators were more promising. All the biological elements showed different responses in different lake types according to alkalinity and humic substances, and also partly according to depth. Better harmonisation of monitoring methods is needed to achieve better precision in the dose–response curves. Future research should include impacts of hydromorphological pressures and climate change, as well as predictions of timelags involved in responses to reduction of pressures.     

Biological assessment of European lakes: ecological rationale and human impacts

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Water‐level fluctuations regulate the structure and functioning of natural lakes

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A hitchhiker's guide to European lake ecological assessment and intercalibration

The Water Framework Directive is the first international legislation to require European countries to establish comparable ecological assessment schemes for their freshwaters. A key element in harmonising quality classification within and between Europe's river basins is an “Intercalibration” exercise, stipulated by the WFD, to ensure that the good status boundaries in all of the biological assessment methods correspond to similar levels of anthropogenic pressure. In this article, we provide a comprehensive overview of this international comparison, focusing on the assessment schemes developed for freshwater lakes. Out of 82 lake ecological assessment methods reported for the comparison, 62 were successfully intercalibrated and included in the EC Decision on intercalibration, with a high proportion of phytoplankton (18), macrophyte (17) and benthic fauna (13) assessment methods. All the lake assessment methods are reviewed in this article, including the results of intercalibration. Furthermore, the current gaps and way forward to reach consistent management objectives for European lakes are discussed.     

Assessment of European streams with diatoms, macrophytes, macroinvertebrates and fish: a comparative metric-based analysis of organism response to stress

1. Periphytic diatoms, macrophytes, benthic macroinvertebrates and fish were sampled with standard methods in 185 streams in nine European countries to compare their response to degradation. Streams were classified into two main stream type groups (i.e. lowland, mountain streams); in addition, the lowland streams were grouped into four more specific stream types. 2. Principal components analysis with altogether 43 environmental parameters was used to construct complex stressor gradients for physical–chemical, hydromorphological and land use data. About 30 metrics were calculated for each sample and organism group. Metric responses to different stress types were analysed by Spearman Rank Correlation. 3. All four organism groups showed significant response to eutrophication/organic pollution gradients. Generally, diatom metrics were most strongly correlated to eutrophication gradients (85% and 89% of the diatom metrics tested correlated significantly in mountain and lowland streams, respectively), followed by invertebrate metrics (91% and 59%). 4. Responses of the four organism groups to other gradients were less strong; all organism groups responded to varying degrees to land use changes, hydromorphological degradation on the microhabitat scale and general degradation gradients, while the response to hydromorphological gradients on the reach scale was mainly limited to benthic macroinvertebrates (50% and 44% of the metrics tested correlated significantly in mountain and lowland streams, respectively) and fish (29% and 47%). 5. Fish and macrophyte metrics generally showed a poor response to degradation gradients in mountain streams and a strong response in lowland streams. 6. General recommendations on European bioassessment of streams were derived from the results.     

A phytoplankton trophic index to assess the status of lakes for the Water Framework Directive

Despite improvements in wastewater treatment systems, the impact of anthropogenic nutrient sources remains a key issue for the management of European lakes. The Water Framework Directive (WFD) 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 derived from summer phytoplankton data summarised by genus from 1,795 lakes, covering 20 European countries. We show that it is significantly related to total phosphorus concentrations, but that it is also 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 other national phytoplankton assessment systems and illustrate for a single European (intercalibration) lake type how it was used to intercalibrate WFD boundaries from different countries.     

Strength and uncertainty of phytoplankton metrics for assessing eutrophication impacts in lakes

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.     

Development of a fish-based index to assess the eutrophication status of European lakes

The use of the CEN (European Committee for Standardization) standard method for sampling fish in lakes using multi-mesh gillnets allowed the collection of fish assemblages of 445 European lakes in 12 countries. The lakes were additionally characterised by environmental drivers and eutrophication proxies. Following a site-specific approach including a validation procedure, a fish index including two abundance metrics (catch per unit effort expressed as fish number and biomass) and one functional metric of composition (abundance of omnivorous fish) was developed. Correlated with the proxy of eutrophication, this index discriminates between heavily and moderately impacted lakes. Additional analyses on a subset of data from Nordic lakes revealed a stronger correlation between the new fish index and the pressure data. Despite an uneven geographical distribution of the lakes and certain shortcomings in the environmental and pressure data, the fish index proved to be useful for ecological status assessment of lakes applying standardised protocols and thus supports the development of national lake fish assessment tools in line with the European Water Framework Directive.     

Effects of macrophyte growth forms on invertebrate communities in saline lakes of the Wyoming High Plains

In saline lakes, areal cover and both species and structural diversity of macrophytes often decline as salinity increases. To assess effects of the loss of certain macrophyte growth forms, we characterized benthic and epiphytic invertebrates in three growth forms (thin-stemmed emergents, erect aquatics, and low macroalgae) in oligosaline lakes (0.8–4.2 mS cm⁻¹) of the Wyoming High Plains, USA. We also measured the biomass and taxonomic composition of epiphytic and benthic invertebrates in two erect aquatics with very similar structure that are found in both oligosaline (Potamogeton pectinatus) and mesosaline (9.3–23.5 mS cm⁻¹) (Ruppia maritima) lakes. Although total biomass of epiphytic invertebrates varied among oligosaline lakes, the relative distribution of biomass among growth forms was similar. For epiphytic invertebrates, biomass per unit area of lake was lowest in emergents and equivalent in erect aquatics and low macroalgae; biomass per unit volume of macrophyte habitat was greatest in low macroalgae. For benthic invertebrates, biomass was less beneath low macroalgae than other growth forms. Taxonomic composition did not differ appreciably between growth forms for either benthic or epiphytic invertebrates, except that epiphytic gastropods were more abundant in erect aquatics. Total biomass of epiphytic and benthic invertebrates for the same growth form (erect aquatic) did not differ between oligosaline (Potamogeton pectinatus) and mesosaline (Ruppia maritima) lakes, but taxonomic composition did change. In the oligosaline to mesosaline range, direct toxic effects of salinity appeared important for some major taxa such as gastropods and amphipods. However, indirect effects of salinity, such as loss of macrophyte cover and typically higher nutrient levels at greater salinities, probably have larger impacts on total invertebrate biomass lake-wide.     

A new procedure for comparing class boundaries of biological assessment methods: A case study from the Danube Basin

For the implementation of the European Water Framework Directive class boundaries of biological assessment methods need to be intercalibrated. The most commonly applied intercalibration approach of national river assessment methods in Europe requires data on near-natural reference sites; however, these data are generally scarce. We developed an alternative approach based on sites impacted by similar levels of disturbance and tested it with national assessment methods based on diatoms and benthic invertebrates from countries in the Danube River Basin. Using environmental variables we screened for sampling sites of at least good environmental status. Relations between different assessment methods were established by common metrics, and we standardized these metrics by “biological benchmarks” obtained from the screened datasets. This approach allows for intercalibration even if near-natural reference sites are absent; relatively few and easily available data are required.     

Quantifying uncertainties in biologically-based water quality assessment: A pan-European analysis of lake phytoplankton community metrics

Lake phytoplankton are adopted world-wide as a sensitive indicator of water quality. European environmental legislation, the EU Water Framework Directive (WFD), formalises this, requiring the use of phytoplankton to assess the ecological status of lakes and coastal waters. Here we provide a rigorous assessment of a number of proposed phytoplankton metrics for assessing the ecological quality of European lakes, specifically in response to nutrient enrichment, or eutrophication, the most widespread pressure affecting lakes. To be useful indicators, metrics must have a small measurement error relative to the eutrophication signal we want them to represent among lakes of different nutrient status. An understanding of variability in metric scores among different locations around a lake, or due to sampling and analytical variability can also identify how best this measurement error is minimised.To quantify metric variability, we analyse data from a multi-scale field campaign of 32 European lakes, resolving the extent to which seven phytoplankton metrics (including chlorophyll a, the most widely used metric of lake quality) vary among lakes, among sampling locations within a lake and through sample replication and processing. We also relate these metrics to environmental variables, including total phosphorus concentration as an indicator of eutrophication.For all seven metrics, 65–96% of the variance in metric scores was among lakes, much higher than variability occurring due to sampling/sample processing. Using multi-model inference, there was strong support for relationships between among-lake variation in three metrics and differences in total phosphorus concentrations. Three of the metrics were also related to mean lake depth. Variability among locations within a lake was minimal (<4%), with sub-samples and analysts accounting for much of the within-lake metric variance. This indicates that a single sampling location is representative and suggests that sub-sample replication and standardisation of analyst procedures should result in increased precision of ecological assessments based upon these metrics.For three phytoplankton metrics being used in the WFD: chlorophyll a concentration, the Phytoplankton Trophic Index (PTI) and cyanobacterial biovolume, >85% of the variance in metric scores was among-lakes and total phosphorus concentration was well supported as a predictor of this variation. Based upon this study, we can recommend that these three proposed metrics can be considered sufficiently robust for the ecological status assessment of European lakes in WFD monitoring schemes.     

Performance of a new phytoplankton composition metric along a eutrophication gradient in Nordic lakes

A new phytoplankton metric is presented, which is developed from a large dataset of Norwegian lakes (>2,000 samples from >400 lakes). In contrast to previous metrics, this index is not built on selected ‘indicative’ taxa, but uses all available taxonomic information at genus and species level. Taxa optima with respect to lake trophic status (derived from total phosphorus concentrations) are used to calculate a phytoplankton trophic index (TI) for each sample. Analysis of the TI shows that phytoplankton communities exhibit highly non-linear responses to eutrophication in Norwegian lakes. Reference lakes are characterized by very similar TIs despite having considerable variation in total phosphorus and chlorophyll a concentrations. TI exhibits a non-linear distribution along the eutrophication gradient which separates unimpacted from impacted sites in the study area. We further show that TI exhibits smaller seasonal variations than chlorophyll a, making it a more reliable indicator for lake monitoring. Implications for its applicability within the WFD are discussed.     

Landscape-scale Variation in Taxonomic Diversity in Four Groups of Aquatic Organisms: The Influence of Physical, Chemical, and Biological Properties

We evaluated several factors influencing the taxonomic richness of macrophytes, benthic invertebrates, snails, and fish in a series of northern Wisconsin lakes. We chose the study lakes to decouple the potential effects of ionic strength of lake water and stream connection, two factors that are usually highly correlated and therefore have been confounded in previous studies. In addition, our study lakes covered a wide range in a variety of characteristics, including residential development, abundance of exotic species, nutrient concentrations, predator abundance, and lake size. Species richness within each of the four taxonomic groups was significantly positively related to ionic strength (as measured by specific conductance); we also found secondary associations with other variables, depending on the specific group of organisms. The relationship between richness and lake area was dependent on the specific conductance of the lake and the vagility of the organisms; less vagile groups of organisms showed stronger and steeper species–area relationships in low-conductivity lakes. Further, after variance owing to specific conductance was removed, the presence of stream connections was positively related to species richness for fish, snails, and macrophytes as well as familial richness in benthic invertebrates. Our results indicate that lakes with relatively more groundwater input have lower extinction rates for all four groups of taxa and that lakes with stream inlets and outlets have enhanced immigration rates for fish, snails, benthic invertebrate families, and macrophytes. These findings link processes of immigration and extinction of four groups of organisms of varying vagility to landscape-level hydrologic characteristics related to the glacial history of the region.     

First steps in the Central-Baltic intercalibration exercise on lake macrophytes: where do we start?

The Water Framework Directive (WFD 2000) defines macrophytes as one of the biological groups required for the ecological assessment of European surface waters. Several indices for macrophyte assessment have been proposed or are currently in use by different European states. As a first step towards performing an intercalibration of these indices a common dataset was developed. This dataset contains abundance data on 789 macrophyte species from 316 different lake sites in ten European countries. Various common species and genera were identified as indicators of reference and impacted conditions within the dataset. Cluster analysis of macrophyte data, supported by non-metric multidimensional scaling, indicated that clusters formed were more reflective of their source country rather than lake type. This might be caused by differences in regional climate, biogeography, monitoring techniques, or a combination of these factors. A total of six national indices were applied to assign quality classes to the lakes. However, this produced results that often differed by one or two quality classes for the same site. We foresee that a more precise intercalibration exercise is necessary, and it should be based on more detailed data considering both seasonality and the latitudinal differences within the area covered.     

A fish-based index for the assessment of the ecological quality of temperate lakes

A fish – based index for the assessment of the ecological quality of natural temperate lakes was developed, in accordance to the requirements of the Water Framework Directive (WFD) 2000/60/EC. As a case study, 11 natural lakes located at northern and western Greece were selected. Fish surveys were conducted during mid summer to mid autumn in 2010, 2011 and 2012 using Nordic gillnets and electrofishing. Environmental parameters and anthropogenic pressures were assessed for each lake. Fish species richness, abundance, trophic, reproductive and habitat functional guilds were used for extracting a set of 107 metrics, meeting the requirements of the WFD. All metrics were initially tested as candidates for the index development. A stepwise linear regression of each metric against environmental parameters (lake area, altitude, maximum depth, alkalinity) and anthropogenic pressures (drainage area covered by non-natural land uses – NNLC, water total phosphorus concentrations – TP, Lake Habitat Modification Score – LHMS) was initially conducted for ensuring pressure-response relationships. Reference conditions for each lake were estimated by the hindcasting procedure and the ecological quality for each lake was expressed as the ecological quality ratio (EQR) by a value ranging from 0 (poor quality) to 1 (excellent quality). Two fish fauna metrics, the relative numerical abundance of introduced species (Introduced_a) and the relative biomass of omnivorous species (OMNI_b) were finally extracted as the most significant, responding to LHMS and TP, respectively. The final index was expressed as the mean values of the EQRs of these two metrics. The multimetric fish index presented herein could serve as a tool for assessing the ecological quality of natural lakes at broad geographical scale and generally, in the Mediterranean temperate lakes with similar hydromorphological characteristics.     

Using aquatic macrophyte community indices to define the ecological status of European lakes

Defining the overall ecological status of lakes according to the Water Framework Directive (WFD) is to be partially based on the species composition of the aquatic macrophyte community. We tested three assessment methods to define the ecological status of the macrophyte community in response to a eutrophication pressure as reflected by total phosphorus concentrations in lake water. An absolute species richness, a trophic index (TI) and a lake trophic ranking (LTR) method were tested at Europe-wide, regional and national scales as well as by alkalinity category, using data from 1,147 lakes from 12 European states. Total phosphorus data were used to represent the trophic status of individual samples and were plotted against the calculated TI and LTR values. Additionally, the LTR method was tested in some individual lakes with a relatively long time series of monitoring data. The TI correlated well with total P in the Northern European lake types, whereas the relationship in the Central European lake types was less clear. The relationship between total P and light extinction is often very good in the Northern European lake types compared to the Central European lake types. This can be one of the reasons for a better agreement between the indices and eutrophication pressure in the Northern European lake types. The response of individual lakes to changes in the abiotic environment was sometimes represented incorrectly by the indices used, which is a cause of concern for the use of single indices in status assessments in practice.     

Zooplankton as indicators in lakes: a scientific-based plea for including zooplankton in the ecological quality assessment of lakes according to the European Water Framework Directive (WFD)

With the implementation of the EU Water Framework Directive (WFD), the member states have to classify the ecological status of surface waters following standardised procedures. It was a matter of some surprise to lake ecologists that zooplankton were not included as a biological quality element (BQE) despite their being considered to be an important and integrated component of the pelagic food web. To the best of our knowledge, the decision of omitting zooplankton is not wise, and it has resulted in the withdrawal of zooplankton from many so-far-solid monitoring programmes. Using examples from particularly Danish, Estonian, and the UK lakes, we show that zooplankton (sampled from the water and the sediment) have a strong indicator value, which cannot be covered by sampling fish and phytoplankton without a very comprehensive and costly effort. When selecting the right metrics, zooplankton are cost-efficient indicators of the trophic state and ecological quality of lakes. Moreover, they are important indicators of the success/failure of measures taken to bring the lakes to at least good ecological status. Therefore, we strongly recommend the EU to include zooplankton as a central BQE in the WFD assessments, and undertake similar regional calibration exercises to obtain relevant and robust metrics also for zooplankton as is being done at present in the cases of fish, phytoplankton, macrophytes and benthic invertebrates.     

Global ecological impacts of invasive species in aquatic ecosystems

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta‐analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far‐reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems.     

Assessing river ecological quality using benthic macroinvertebrates in the Hindu Kush-Himalayan region

We developed a system for the assessment of ecological condition for rivers in the lower mountains and lowlands of the Hindu Kush-Himalayan region (Pakistan, India, Nepal, Bhutan, and Bangladesh). We used benthic invertebrates collected from 198 rivers, located in five different ecoregions and covering degradation gradients; samples were taken twice (pre-monsoon and post-monsoon) applying a multi-habitat sampling procedure. Out of 38 environmental parameters, we constructed complex principal component analysis (PCA) gradients, separately for the stressors organic pollution, eutrophication, floodplain land use, and hydromorphological degradation. Correlation analysis between invertebrate metrics and environmental parameters revealed those biological metrics that are most responsive to river deterioration. Redundant metrics were deleted, and the most robust metrics were selected. The range of the index values under reference conditions was defined, and a five-class river quality system was generated.     

Fish diversity in European lakes: geographical factors dominate over anthropogenic pressures

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