Marit Mjelde

Classifying aquatic macrophytes as indicators of eutrophication in European lakes

Aquatic macrophytes are one of the biological quality elements in the Water Framework Directive (WFD) for which status assessments must be defined. We tested two methods to classify macrophyte species and their response to eutrophication pressure: one based on percentiles of occurrence along a phosphorous gradient and another based on trophic ranking of species using Canonical Correspondence Analyses in the ranking procedure. The methods were tested at Europe-wide, regional and national scale as well as by alkalinity category, using 1,147 lakes from 12 European states. The grouping of species as sensitive, tolerant or indifferent to eutrophication was evaluated for some taxa, such as the sensitive Chara spp. and the large isoetids, by analysing the (non-linear) response curve along a phosphorous gradient. These thresholds revealed in these response curves can be used to set boundaries among different ecological status classes. In total 48 taxa out of 114 taxa were classified identically regardless of dataset or classification method. These taxa can be considered the most consistent and reliable indicators of sensitivity or tolerance to eutrophication at European scale. Although the general response of well known indicator species seems to hold, there are many species that were evaluated differently according to the database selection and classification methods. This hampers a Europe-wide comparison of classified species lists as used for the status assessment within the WFD implementation process.

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.

Ecological status assessment of European lakes: a comparison of metrics for phytoplankton, macrophytes, benthic invertebrates and fish

Data on phytoplankton, macrophytes, benthic invertebrates and fish from more than 2000 lakes in 22 European countries were used to develop and test metrics for assessing the ecological status of European lakes as required by the Water Framework Directive. The strongest and most sensitive of the 11 metrics responding to eutrophication pressure were phytoplankton chlorophyll a, a taxonomic composition trophic index and a functional traits index, the macrophyte intercalibration taxonomic composition metric and a Nordic lake fish index. Intermediate response was found for a cyanobacterial bloom intensity index (Cyano), the Ellenberg macrophyte index and a multimetric index for benthic invertebrates. The latter also responded to hydromorphological pressure. The metrics provide information on primary and secondary impacts of eutrophication in the pelagic and the littoral zone of lakes. Several of these metrics were used as common metrics in the intercalibration of national assessment systems or have been incorporated directly into the national systems. New biological metrics have been developed to assess hydromorphological pressures, based on aquatic macrophyte responses to water level fluctuations, and on macroinvertebrate responses to morphological modifications of lake shorelines. These metrics thus enable the quantification of biological impacts of hydromorphological pressures in lakes.

Maximum growing depth of submerged macrophytes in European lakes

Submerged macrophytes are important elements for the structure and functioning of lake ecosystems. In this study, we used chemical and maximum colonisation depth (C_max) data from 12 European countries in order to investigate how suitable C_max may describe the impact by eutrophication. The analyses include data from 757 lakes and 919 lake years covering oligotrophic to eutrophic lakes. Overall, C_max was closely related to Secchi depth (R2xa0=xa00.58) and less closely to chlorophyll a (R2xa0=xa00.31), TP (R2xa0=xa00.31) and total nitrogen, TN (R2xa0=xa00.24). The low coefficients of determination between C_max and nutrient concentrations suggest that other response factors than nutrient-phytoplankton-light conditions are important for C_max and that it will be difficult to establish strong relationships between external nutrient loading and C_max. Yearly monitoring for 13–16xa0years in eight Danish lakes showed considerable year-to-year variability in C_max, which for the individual lakes only related weakly to changes in Secchi depth. The use of C_max as an eutrophication indicator is especially relevant in not very shallow lakes (maximum depth >4–5xa0m), not too turbid lakes (C_max >1xa0m) and not very humic lakes (colour <60xa0mg Pt/l).

A water level drawdown index for aquatic macrophytes in Nordic lakes

Many northern lakes are regulated to enhance hydropower production and flood protection. This bears hydromorphological pressures which are important factors causing lowered ecological status. Water level fluctuation triggers erosion on the shoreline and, depending on fluctuation range, also affects species composition or disappearance of sensitive aquatic macrophytes. We developed a water level-drawdown index (WIc) for Nordic lakes using macrophyte data from 73 lakes with varying water level fluctuation in Finland, Norway and Sweden. The index is based on the ratio between sensitive and tolerant macrophyte species. The sensitive and tolerant species are identified based on a percentile approach, analysing the presence or absence of species along the winter drawdown range. The index correlates well with winter drawdown in Finnish and Norwegian lakes with strongest correlations with winter drawdown in storage lakes (lakes regulated for hydroelectric power and with a considerable winter drawdown). The WIc-index is applicable in low alkalinity, oligotrophic and ice-covered lakes, and is suggested to be a useful tool to identify and designate heavily modified water bodies in Nordic lakes according to the European Water Framework Directive.

Clear and Turbid Water in Shallow Norwegian Lakes Related to Submerged Vegetation

Timms and Moss (1984) suggested that fertile shallow lakes may have alternative stable states, a clearwater state with dense vegetation and a turbid water state dominated by phytoplankton and with little submerged and floating-leaved vegetation. This phenomenon has also been observed and discussed by several other authors (e.g., Irvine et al., 1990; Jeppesen et al., 1990; van Donk et al., 1990; Blindow et al., 1993; Scheffer et al., 1993). According to the model of Scheifer (1989, 1990), the main controlling factor for the two alternative states is the turbidity of water regulating the vertical light penetration. When the nutrient level increases, phytoplankton growth is often stimulated, which in turn increases the turbidity. This leads to increased light attenuation and a reduction of the maximum growth depth of submerged vegetation. Increased nutrient concentrations will therefore reduce the bottom area covered by submerged vegetation until it is virtually absent. In shallow lakes with most bottom areas at similar depths, the change from high plant cover to plantless bottoms may be abrupt. Uncovered sediments in shallow lakes are much more vulnerable to resuspen-sion and more easily give rise to turbid water during periods of wind and wave action than in lakes with plant-covered sediments. Benthivorous fish may be favored under these circumstances and add to the turbidity by foraging on the sediment. A more-or-less stable turbid state may be the result.

Macrophyte responses to water level fluctuation in Fennoscandinavian Lakes — Applying a common index

Hydromorphological pressure caused by artificial water level regulation is one ofthe major factors influencing the distribution of aquatic vegetation in Fennoscandinavian Iakes. More than 350 lakes in Finland and 900 lakes in Norway are regulated or were artificially created (MARTIUNEN et al. 2006). Aquatic macrophytes are key indicators ofhydromorphological changes in lakes due to their sensitivity to water level fluctuation as shown in several Nordic studies (R0RSLETI 1989, HELLSTEN 2001, PARTANEN & HELLSTEN 2005, KETO et al. 2006). Aquatic macrophyte data are very often collected by diverse sets of field methodology; the term macrophyte is also unclear, ranging from a very strict definition ofhydrophytes to a wide definition that includes helophytes and even some shore plants. Therefore, full scientific comparison of different national datasets is difficult although implementation of the European Water Framework Directive demands relatively uniform datasets for a common intercalibration exercise. The aim of our study was to compare Finnish and Norwegian macrophyte data from regulated and non-regulated lakes to develop a common index for estimation of the ecological status of regulated lakes.

Potential conflicts between environmental legislation and conservation exemplified by aquatic macrophytes

It is important that legislation on water quality issues of freshwaters is not in conflict with nature conservation purposes. So far, it is however unknown how the assessment of ecological status according to for example the Water Framework Directive (WFD) of the European Community relates to the status of lakes according to the Habitat Directive (HD) or to national environmental objectives including, e.g., the protection of important wetland areas and red-listed species. We used lake macrophyte classification schemes of Norway, Sweden, and Finland and a total of 1,014 lakes to evaluate the possible conflict between these directives and national legislation. The classification schemes represent mainly trophic indices penalizing lakes with elevated phosphorous concentrations. In general, high ecological status according to the WFD did not mean high number of red-listed species or high status according to the HD or other national environmental objectives. In Sweden 78%, in Norway 47%, and in Finland 29% of lakes with red-listed species were classified as lakes of moderate or worse ecological status based on the macrophyte classification scheme. These lakes thus did not fulfill the demands of the WFD. Restoration of surface water toward fulfilling the demands requires in practice a reduction of the trophic status. This might potentially result in for example the loss of red-listed species. To avoid such potential conflicts, we primarily suggest revising the national quality assessment systems toward implicitly incorporating nature conservation aspects, e.g., the number of red-listed species in a multi-metric assessment system.

Quantifying interspecific spatial overlap in aquatic macrophyte communities

Levins’s asymmetrical α index quantifies between species overlap over resources more realistically than similar-purpose single-value indices. The associated community-wide