Marcel S. van den Berg

Macroinvertebrate communities in relation to submerged vegetation in two Chara-dominated lakes

Relationships between macroinvertebrates and the presence ofsubmerged vegetation were studied in two shallow eutrophiclakes inThe Netherlands, Lake Veluwemeer and Lake Wolderwijd. A shiftfromturbid water with sparse macrophyte cover (Potamogetonperfoliatus, Potamogeton pectinatus) towards clearwaterwith a dense cover of submerged vegetation (Chara spp.)hasbeen observed in the lakes over the past 10 years. RelativelylargeChara meadows (300–500 ha) have recently developed inbothlakes. The composition of macroinvertebrate fauna wasdetermined atsites varying in cover and dominant vegetation type bysamplingsediment and water during 1992 and 1994. Macrophyte biomass,sampling year and vegetation type were the major determinantsofmacroinvertebrate community composition. Valvatapiscinalis,Bithynia tentaculata, Gammarus tigrinus and Chironomussp.characterized the sites with high charophyte biomass, whereasPotamopyrgus antipodarum, Cladotanytarsus sp., Stictochironomus sp. dominated the samples with lowcharophytebiomass. Chara vegetation was different from Potamogeton sp. by showing lower densities of the midgelarvaeEinfeldia dissidens and Cricotopus gr. sylvestris.Seasonal variations in densities of the dominant molluscspecies(V. piscinalis, P. antipodarum) were closelyrelated tothe development of Chara biomass as well as toperiphytoncover on charophytes. Thus, changes of the light climate inbothlakes, which have led to an increase in colonization bysubmergedvegetation (particular Chara meadows), indirectly had alargeimpact on macroinvertebrate communities.

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.

Response of herbivorous water-birds to the return of Chara in Lake Veluwemeer, The Netherlands

Abstract Water-birds were almost absent from Lake Veluwemeer during the 1970s and 1980s, due to low food availability related to eutrophication. Several restoration measures have resulted in the recolonization of the lake by Chara spp. during the 1990s. Bird numbers in autumn and winter have increased along with charophyte biomass. Linear regression analysis was used to relate bird numbers to the abundance of their potential food sources Chara, pondweeds, filamentous macro-algae (FA) and zebra mussels (Dreissena polymorpha) within the depth range available to each species. Numbers of mute swan (Cygnus olor), pochard (Aythya ferina), tufted duck (A. fuligula), coot (Fulica atra) (R2≥0.98), gadwall (Anas strepera) (R2=0.78), Bewick’s swan (Cygnus columbianus) (R2=0.79), red-crested pochard (Netta rufina) (R2=0.76) and pintail (Anas acuta) (R2=0.38) showed significant correlations with these food sources. Chara biomass explained most of the variance in all species except for gadwall. Both bird numbers and their duration of stay were closely associated with the presence of Chara. Grazing pressure was low during spring and summer and Chara colonized the lake in spite of consumption. It is argued that birds did not slow down colonization of the lake by Chara during the 1990s, but may have accelerated succession from pondweeds Potamogeton spp. to charophytes. As more birds foraged on Chara compared to any other food resource, the preservation of large areas of Chara is of great importance to the water-bird community in Lake Veluwemeer.

Competition between Chara aspera and Potamogeton pectinatus as a function of temperature and light

Abstract Two experiments were carried out to study the interaction between Chara aspera Deth. ex Willd. and Potamogeton pectinatus L. The purpose of the first experiment was to assess the effect of temperature on the rate of emergence and the second was designed to study the effect of light on the competition during the established phase. P. pectinatus tubers sprouted after about 4 days at 16°C and 9 days at 10°C, which was sooner than either oospores or bulbils of C. aspera (about 15 days at 16°C and 27 days at 10°C). When the irradiance was 36 μ mol m −2 s −1 , the ash-free dry weight biomass of both species was about 80% lower than at 416 μ mol m −2 s −1 and the biomass was not affected by neighbouring plants. However, in the higher light treatment the presence of P. pectinatus suppressed the biomass of C. aspera by maximally 63%. The individual biomass of P. pectinatus at high initial plant densities was reduced by maximally 70% by other plants of the same species, but was not affected by C. aspera . It is concluded that P. pectinatus rather than C. aspera has an advantage in the phase of emergence during spring. Furthermore, under higher light conditions and high inorganic carbon concentrations P. pectinatus may be a better competitor for light than C. aspera , because of its canopy placement near the water surface.

Aquatic macrophytes: restore, eradicate or is there a compromise?

Promoting indigenous submerged macrophytes is considered an important measure to restore shallow lakes. On the other hand, dense stands of aquatic vegetation often cause nuisance for boating, swimming and by obstruction of water flow. Consequently, the interests of recreational users may conflict with nature conservation. At first sight, aiming lake management at intermediate vegetation biomass seems a good solution for this controversy. However, as can be shown with a simple economical model, such a compromise may not always be an optimal solution in terms of total welfare across all users of a lake. Here, we show that a management strategy aimed at moderate plant biomass can also be unfeasible for ecological reasons. This is shown by applying two very different models: a minimal logistic model of plant growth which is easy to understand, and the relatively complex and realistic simulation model Charisma which, among other things, includes the seasonal cycle and a detailed description of the dynamics of light availability for macrophytes. The qualitative results of both models were remarkably similar. If the ecosystem has no alternative stable states, it is possible to set vegetation to any desired sustainable level using an appropriate harvesting strategy. However, if an intermediate vegetation biomass is to be realized the costs of harvesting are high, because the yield is predicted to be maximal in that case. If the ecosystem has alternative stable states, harvesting becomes risky because the vegetation may collapse entirely below a certain, in practice unknown, biomass. Also, even moderate harvesting may reduce the resilience of the vegetated state, making the ecosystem more vulnerable to adverse conditions such as unfavorable weather conditions or bird grazing.

Defining chlorophyll-a reference conditions in European lakes.

The concept of “reference conditions” describes the benchmark against which current conditions are compared when assessing the status of water bodies. In this paper we focus on the establishment of reference conditions for European lakes according to a phytoplankton biomass indicator—the concentration of chlorophyll-a. A mostly spatial approach (selection of existing lakes with no or minor human impact) was used to set the reference conditions for chlorophyll-a values, supplemented by historical data, paleolimnological investigations and modelling. The work resulted in definition of reference conditions and the boundary between “high” and “good” status for 15 main lake types and five ecoregions of Europe: Alpine, Atlantic, Central/Baltic, Mediterranean, and Northern. Additionally, empirical models were developed for estimating site-specific reference chlorophyll-a concentrations from a set of potential predictor variables. The results were recently formulated into the EU legislation, marking the first attempt in international water policy to move from chemical quality standards to ecological quality targets.

Propagule bank buildup of Chara aspera and its significance for colonization of a shallow lake

Several aspects of the propagule bank dynamics including germination, burial and storage characteristics, and the relationship between the distribution of the propagule bank and cover of an expanding Chara vegetation (dominated by C. asperaDeth. ex Willd.) was studied in the shallow lake Veluwemeer. The density of oospores in the sediment was positively correlated with the number of years that Chara was present at that particular site. After six years of Chara presence, at least 1.7 × 106 oospores m−2 had accumulated in the sediment. Oospores in sediment were evenly distributed in the 15 cm top layer, which was very similar to the foraging depth of Bewicks Swans (Cygnus columbianus bewickii Yarr). Burial in the sediment may be an important mechanism by which oospores are stored. On the other hand, at shallow areas about 50% of the biomass was consumed enhancing the potential dispersal of oospores by water birds. In a laboratory experiment, 100% of the tested bulbils of C. aspera emerged and thus may be important for short time survival of established vegetation. In contrast, oospore germination varied between 1 and 15% depending on light level and burial depth. The low germination and the high accumulation of oospores suggest that oospores are adapted to long time survival in a dormant state. Charophytes colonized Veluwemeer step by step in the course of about nine years. Not all the sites with suitable light conditions were colonized at the same speed. C. aspera established a dense vegetation only at sites with high oospores densities (> c. 1 × 104 m−2). The results indicate that the high density of oospores needed for attaining complete vegetation cover may be an important limiting factor during colonization.

Defining ecologically relevant water quality targets for lakes in Europe

Summary 1. The implementation of the Water Framework Directive requires EU member states to establish and harmonize ecological status class boundaries for biological quality elements. In this paper, we describe an approach for defining ecological class boundaries that delineates shifts in lake ecosystem functioning and, therefore, provides ecologically meaningful targets for water policy in Europe. 2. We collected an extensive data set of 810 lake-years from nine Central European countries, and we used phytoplankton chlorophyll a, a metric widely used to measure the impact of eutrophication in lakes. Our approach establishes chlorophyll a target values in relation to three significant ecological effects of eutrophication: the decline of aquatic macrophytes, the dominance of potentially harmful cyanobacteria and the major functional switch from a clear water to a turbid state. 3. Ranges of threshold chlorophyll a concentrations are given for the two most common lake types in lowland Central Europe: for moderately deep lakes (mean depth 3–15 m), the greatest ecological shifts occur in the range 10–12 l gL � 1 chlorophyll a, and for shallow lakes (<3 m mean depth), in the range 21–23 l gL � 1 chlorophyll a. 4. Synthesis and applications. Our study provides class boundaries for determining the ecological status of lakes, which have robust ecological consequences for lake functioning and which, therefore, provide strong and objective targets for sustainable water management in Europe. The results have been endorsed by all participant member states and adopted in the European Commission legislation, marking the first attempt in international water policy to move from physico-chemical quality standards to harmonized ecologically based quality targets.

Macrophyte assessment in European lakes: Diverse approaches but convergent views of ‘good’ ecological status

Graphical abstract