The importance of drawdown and sediment removal for the restoration of the eutrophied shallow Lake Kraenepoel (Belgium)

Lake Kraenepoel (Belgium) is a shallow lake (22 ha), divided in two basins since 1957 by a shallow dike. The lake was used for fish farming until World War II and was drawn down about every 5 years to harvest fish. Despite its dense historical carp population, it had clear water and a rich Littorelletea vegetation. During the course of the 20th century, the lake became eutrophic and the Littorelletea vegetation degraded. The northern basin, which was still drawn down about every decade after 1957, retained its clear water and had a dense submerged macrophyte vegetation. The southern basin, which was never drawn down after 1957 and which received direct surface water inputs, had become a turbid shallow lake with phytoplankton blooms in summer. In 2000, efforts were taken to restore the lake: the entire lake was drawn down, the fish community was biomanipulated, nutrient-rich surface water inputs were diverted from the southern basin and sediments were removed (only in the northern basin). Fish biomanipulation and sediment removal were successful in the northern basin, as nutrient levels declined and the Littorelletea vegetation recovered. In the southern basin, sediment analyses indicated that drawdown resulted in sediments with a lower water and organic matter content and water column turbidity decreased after the drawdown. But pH in the southern basin declined to <4, probably because sulphides in the sediment were oxidized during drawdown and sediment desiccation. In contrast, desiccated sediments were removed from the northern basin and pH did not decline below 6 after restoration. In spite of the still high dissolved nutrient concentrations, phytoplankton biomass declined significantly in the southern basin, probably due to acidification. However, no Littorelletea species colonised the lake bottom in the southern basin. Thus, lake drawdown may be a useful management technique to promote clear water conditions in shallow lakes. However, acidification due to sulphide oxidation may be an undesirable outcome and should be considered in drawdown and sediment desiccation manipulations.     

OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Invasive Crayfish Threaten the Development of Submerged Macrophytes in Lake Restoration

Submerged macrophytes enhance water transparency and aquatic biodiversity in shallow water ecosystems. Therefore, the return of submerged macrophytes is the target of many lake restoration projects. However, at present, north-western European aquatic ecosystems are increasingly invaded by omnivorous exotic crayfish. We hypothesize that invasive crayfish pose a novel constraint on the regeneration of submerged macrophytes in restored lakes and may jeopardize restoration efforts. We experimentally investigated whether the invasive crayfish (Procambarus clarkii Girard) affects submerged macrophyte development in a Dutch peat lake where these crayfish are expanding rapidly. Seemingly favourable abiotic conditions for macrophyte growth existed in two 0.5 ha lake enclosures, which provided shelter and reduced turbidity, and in one lake enclosure iron was added to reduce internal nutrient loading, but macrophytes did not emerge. We transplanted three submerged macrophyte species in a full factorial exclosure experiment, where we separated the effect of crayfish from large vertebrates using different mesh sizes combined with a caging treatment stocked with crayfish only. The three transplanted macrophytes grew rapidly when protected from grazing in both lake enclosures, demonstrating that abiotic conditions for growth were suitable. Crayfish strongly reduced biomass and survival of all three macrophyte species while waterfowl and fish had no additive effects. Gut contents showed that crayfish were mostly carnivorous, but also consumed macrophytes. We show that P. clarkii strongly inhibit macrophyte development once favourable abiotic conditions for macrophyte growth are restored. Therefore, expansion of invasive crayfish poses a novel threat to the restoration of shallow water bodies in north-western Europe. Prevention of introduction and spread of crayfish is urgent, as management of invasive crayfish populations is very difficult.     

Top-down control of phytoplankton in a shallow hypertrophic lake: Little Mere (England)

Top-down control of phytoplankton by zooplankton is possible through reductions in density of zooplanktivorous fish. Little Mere is a shallow lake where the effects of sewage effluent caused such a reduction. This allowed the large-bodied cladoceran, Daphnia magna Straus, to develop huge populations, preventing potentially large algal crops from developing.Subsequent diversion of the effluent is anticipated to lead to recovery of the fish community, reduced numbers of large-bodied grazers, and increased phytoplankton biomass. Whether the aquatic plant community, present in Little Mere, is resilient to such changes may depend upon whether cyanophytes are favoured, or not.     

Fish induced macrophyte loss in shallow lakes: top–down and bottom–up processes in mesocosm experiments

SUMMARY 1. Macrophyte loss from Sites of Special Scientific Interest in England has become widespread over the last 20 years. One reason for this may be changing trends in angling, a multimillion pound industry that has an enormous impact on aquatic ecosystems. Stocking with cyprinid fish is a common angling management practice but the particular fish species and distribution of their biomass may be crucial to the ecosystem. 2. Carp (Cyprinus carpio), roach (Rutilus rutilus), bream (Abramis brama) and tench (Tinca tinca) at biomasses ranging from 0 to 800 kg ha^?1 and at various sizes were placed into experimental mesocosms in Little Mere, a shallow, fertile lake in Cheshire, U.K. The effects these treatments had on the aquatic ecosystem were studied over two summers. Specifically the effects of the treatments on macrophyte growth, benthic and macrophytic macro‐invertebrate populations, water chemistry, epiphyton production and plankton survival were investigated. 3. Carp had a greater detrimental effect on the macrophytes than bream, tench and in particular roach. A biomass of fish > 200 kg ha^?1 adversely affected the extent of macrophyte growth. 4. The decline in macrophyte growth was most likely as a result of increased epiphyton growth that probably reduced the amount of light and carbon dioxide available to the plant. There were no observed direct fish impacts on macrophytes. 5. The chemical data suggested that inorganic nitrogen levels were low and it is possible that release of nitrogen, from fish excreta, followed by immediate uptake, could have been a major factor stimulating epiphyton growth and subsequently macrophyte loss. Phosphorus concentrations increased even in the controls and substantial amounts were available. Phosphorus stimulation can therefore be discounted. Macrophyte‐associated macro‐invertebrates were positively correlated with epiphyton load but had no impact on the extent of epiphytic growth. Shading from disturbed sediment or phytoplankton was also unimportant.     

The rapid effects of a whole-lake reduction of coarse woody debris on fish and benthic macroinvertebrates

1. Ecosystems can enhance the biodiversity of adjacent ecosystems through subsidies of prey, nutrients and also habitat. For example, trees can fall into aquatic ecosystems and act as a subsidy that increases aquatic habitat heterogeneity. This habitat subsidy is vulnerable in lakes where anthropogenic development of shorelines coincides with a thinning of riparian forests and the removal of these dead trees (termed coarse woody debris: CWD). How the disruption of this subsidy affects lake ecosystems is not well understood.
2. We performed a whole ecosystem experiment on Little Rock Lake, a small (18 ha), undeveloped, and unfished lake in Vilas County, WI, U.S.A., that is divided into two similar-sized basins by a double poly-vinyl chloride curtain that prevents both fish and water exchange between basins. In 2002, we removed about 70% of the littoral CWD in the treatment basin, while the reference basin was left unaltered. We tested for changes in both fish and benthic macroinvertebrate community composition in the two years following the CWD reduction.
3. Yellow perch ( Perca flavescens ) was the most abundant fish species in the lake prior to our experiment, but declined to very low densities in the treatment basin after manipulation. We found no evidence of an effect on macroinvertebrates – the treatment basin's macroinvertebrate community composition, diversity and density did not change relative to the reference basin.
4. Our results indicate that different trophic groups may have differential responses to the loss of a habitat subsidy, even if anthropogenic effects on that subsidy are severe. In the case of Little Rock Lake, fish community responses were evident on a short-time scale, whereas the macroinvertebrate community did not rapidly change following CWD reduction.     

Relationship between epipelon,epiphyton and phytoplankton in two limnological phases in a shallow tropical reservoir with high Nymphaea coverage

Macrophytes and phytoplankton are recognized as having roles in determining alternative stable states in shallow lakes and reservoirs, while the role of periphyton has been poorly investigated. Temporal and spatial variation of phytoplankton, epipelon and epiphyton was examined in a shallow reservoir with high abundance of aquatic macrophytes. The relationships between algae communities and abiotic factors, macrophyte coverage and zooplankton density were also analyzed. Monthly sampling was performed in three zones of the depth gradient of the reservoir. Two phases of algal dominance were found: a phytoplankton phase and epipelon phase. The phase of phytoplankton dominance was characterized by high macrophyte coverage. Rotifera was the dominant zooplankton group in all the zones. Flagellate algae were dominant in phytoplankton, epipelon and epiphyton. Macrophyte coverage was found to be a predictor for algal biomass. Changes in biomass and species composition were associated with macrophyte cover variation, mainly the Nymphaea. In addition to the abiotic factors, the macrophyte coverage was a determining factor for changes to the algal community, contributing to the alternation between dominance phases of phytoplankton and epipelon. The macrophyte–phytoplankton–periphyton relationship needs to be further known in shallow reservoirs, especially the role of epipelon as an alternate stable state.

     

Fish community assemblages changed but biomass remained similar after lake restoration by biomanipulation in a Chinese tropical eutrophic lake

While the effects of lake restoration by fish manipulation are well-studied in the temperate zone, comparatively little information is available on this issue from tropical lakes. It may be expected that fish removal leads to faster recovery of the fish stock here than in temperate lakes due to more frequent and earlier reproduction, which may, in turn, delay positive effects of the restoration. We studied the community composition, feeding type and abundance of fish in three basins of a tropical shallow lake: one unrestored basin (UR) and two basins restored by fish manipulation and transplantation of submerged macrophytes. While omni-benthivorous fish dominated the biomass in the restored basins 3 and 5 years after restoration, planktivores were most abundant in the UR, although total fish biomass remained similar. One-way analyses of similarities based on fish species presence/absence, abundance and biomass data revealed significant differences in fish community composition among the restored basins and UR, and redundancy analyses further indicated that submerged macrophytes were a key driver behind this difference. Our results indicate that active implantation of submerged macrophytes to stabilise the fish community is a tool to consider when planning lake restoration by biomanipulation in the tropics.     

新疆开都河流域水生植物多样性及其影响因素

河流是一个连续的、流动的、独特而完整的系统,研究河流生态系统中水生植物的多样性分布格局及其影响因素对河流生态学研究具有重要意义。本文通过野外调查,研究了新疆开都河流域水生植物多样性、主要水生植物群落特征及与环境因子之间的关系,并利用水分-能量动态假说和栖息地异质性假说对该流域水生植物物种多样性的地理格局进行解释。结果表明: 开都河流域共有水生植物71种,隶属于24科39属;聚类分析可将开都河流域水生植物群落划分为10个主要群落类型,其中芦苇群落物种丰富度最高,狭叶香蒲群落和金鱼藻群落物种丰富度最低;流域水生植物群落Shannon指数与pH呈显著负相关,Simpson指数与pH、经度呈显著负相关,与海拔呈显著正相关;流域水生植物群落类型主要受海拔、水深及水温的影响;流域水生植物物种多样性随经纬度无明显变化规律。水分-能量动态假说和栖息地异质性假说共解释开都河流域水生植物多样性格局变量的31.4%,表明这两个假说对于该流域水生植物多样性格局的解释力并不高。     

Influences of the lowland river Spree on phytoplankton dynamics in the flow-through Lake Müggelsee (Germany)

The influences of imports of nutrients and planktonic algae from the River Spree on the dynamics of phytoplankton were examined in the shallow, eutrophic Müggelsee, which has a retention time of only 42 days. Phytoplankton biomass and nutrient concentrations were measured in both the lake and its inflow from 1980–1990. On a long-term average, mean biomass as well as vitality of most dominant phytoplankton populations in the lake were not significantly different from those in the river. Nevertheless, during distinct periods the external rates of biomass change of single lake populations (due to dilution or enrichment) were as high as the lake internal ones. The import of inocula populations from the river probably induced the formation of the typical community structure in the lake. Growth and decay of phytoplankton populations in the river strongly influenced the load of dissolved nutrients and thus indirectly the dynamics of planktonic algae in the downstream lake. For example, intensive assimilation of phosphorus by riverine algae in spring intensified the P-shortage and supported possible P-limitation of algal growth in the lake at that time. In years with high vernal biomass of centric diatoms in the river, and thus diminished import of dissolved silicon, the growth of diatoms was suppressed and that of cyanobacteria was favoured in the lake during summer.     

Weedbeds and big bugs: the importance of scale in detecting the influence of nutrients and predation on macroinvertebrates in plant‐dominated shallow lakes

1. The scale of investigations influences the interpretation of results. Here, we investigate the influence of fish and nutrients on biotic communities in shallow lakes, using studies at two different scales: (i) within‐lake experimental manipulation and (ii) comparative, among‐lake relationships. 2. At both scales, fish predation had an overriding influence on macroinvertebrates; fish reduced macroinvertebrate biomass and altered community composition. Prey selection appeared to be size based. Fish influenced zooplankton abundance and light penetration through the water column also, but there was no indication that fish caused increased resuspension of sediment. 3. There were effects of nutrients at both scales, but these effects differed with the scale of the investigation. Nutrients increased phytoplankton and periphyton at the within‐lake scale, and were associated with increased periphyton at the among‐lake scale. No significant effect of nutrients on macroinvertebrates was observed at the within‐lake scale. However, at the among‐lake scale, nutrients positively influenced the biomass and density of macroinvertebrates, and ameliorated the effect of fish on macroinvertebrates. 4. Increased prey availability at higher nutrient concentrations would be expected to cause changes in the fish community. However, at the among‐lake scale, differences were not apparent in fish biomass among lakes with different nutrient conditions, suggesting that stochastic events influence the fish community in these small and relatively isolated shallow lakes. 5. The intensity of predation by fish significantly influences macroinvertebrate community structure of shallow lakes, but nutrients also play a role. The scale of investigation influences the ability to detect the influence of nutrients on the different components of shallow lake communities, particularly for longer lived organisms such as macroinvertebrates, where the response takes longer to manifest.     

The potential of artificial refugia for maintaining a community of large-bodied cladocera against fish predation in a shallow eutrophic lake

The Norfolk Broads are a series of shallow lakes which are highly eutrophic and typified by dense populations of phytoplankton and an absence of submerged aquatic plants. The zooplankton community is subject to intense predation pressure by young fish and is dominated by small-bodied organisms which have a low potential for reducing phytoplankton populations through grazing. Various designs and densities of artificial refugia for zooplankton against fish predation were established in Hoveton Great Broad in order to enhance populations of large-bodied Cladocera. Initially some of the refuges contained higher densities and larger individuals ofDaphnia andCeriodaphnia than the surrounding open water. However, towards the end of the first season and throughout the subsequent two years, population densities and size-structure were similar both within and outside the refuges, although there was still evidence of enhanced body-size ofDaphnia within the refuges compared with the open water. The provision of habitat structures designed as refugia from fish predation did not enhance large-bodied cladoceran populations enough to promote this restoration strategy as feasible for eutrophic and shallow lakes.     

Community resistance and change to nutrient enrichment and fish manipulation in a vegetated lake littoral

1. High biomass of macrophytes is considered important in the maintenance of a clear‐water state in shallow eutrophic lakes. Therefore, rehabilitation and protection of aquatic vegetation is crucial to the management of shallow lakes. 2. We conducted field mesocosm experiments in 1998 and 1999 to study community responses in the plant‐dominated littoral zone of a lake to nutrient enrichment at different fish densities. We aimed to find the threshold fish biomass for the different nutrient enrichment levels below which large herbivorous zooplankton escapes control by fish. The experiments took place in the littoral of Lake Vesijärvi in southern Finland and were part of a series of parallel studies carried out jointly at six sites across Europe. 3. In 1998, when macrophyte growth was poor, a clear‐water state with low phytoplankton biomass occurred only in unenriched mesocosms without fish or with low fish biomass (4 g fresh mass m^?2). Both nutrient enrichment and high fish biomass (20 g fresh mass m^?2) provoked a turbid water state with high planktonic and periphytic algal biomass. The zooplankton community was dominated by rotifers and failed to control the biomass of algae in nutrient enriched mesocosms. The littoral community thus had low buffer capacity against nutrient enrichment. 4. In 1999, macrophytes, especially free‐floating Lemna trisulca L., grew well and the zooplankton community was dominated by filter‐feeding cladocerans. The buffer capacity of the littoral community against nutrient enrichment was high; a clear‐water state with low phytoplankton biomass prevailed even under the highest nutrient enrichment. High grazing rates by cladocerans, together with reduced light penetration into the water caused by L. trisulca, were apparently the main mechanisms behind the low algal biomass. 5. Effects of fish manipulations were less pronounced than effects of nutrient enrichment. In 1999, clearance rates of cladocerans were similar in fish‐free and low‐fish treatments but decreased in the high‐fish treatment. This suggests that the threshold fish biomass was between the low‐ and high‐fish treatments. In 1998, such a threshold was found only between fish‐free and low‐fish treatments. 6. The pronounced difference in the observed responses to nutrient enrichment and fish additions in two successive years suggests that under similar nutrient conditions and fish feeding pressure either clear or turbid water may result depending on the initial community structure and on weather.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Is the island biogeography model a poor predictor of biodiversity patterns in shallow lakes?

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Spatial distribution of phytoplankton in Lake Baikal, Spring 1991

• 1 Studies carried out in Lake Baikal in late spring (late May-early June) 1991 showed marked differences in the species composition and abundance of phytoplankton in different regions of the lake. The south and north basins were characterized by small forms of algae. The middle basin, Maloe More and the shallow waters of the Selenga had species with large cells including Aulacoseira islandica subsp. helvetica, Dinobryon cylindricum and D. divergens.
• 2 Areas of high biomass were correlated with shallow waters and river inputs. This was especially apparent in the region of the Selenga delta. The vertical distribution of phytoplankton indicated the non-synchronous start of the spring homothermy throughout the lake. Phytoplankton were concentrated in the upper 100 m layer with subsurface maxima resulting from the sinking of large algae.
• 3 The concentration of phytoplankton biomass in general at this time characterized the lake as moderately productive.

     

A 2-year experimental study on nutrient and predator influences on food web constituents in a shallow lake of north-west Spain

1. A 2‐year study was carried out on the roles of nutrients and fish in determining the plankton communities of a shallow lake in north‐west Spain. Outcomes were different each year depending on the initial conditions, especially of macrophyte biomass. In 1998 estimated initial ‘per cent water volume inhabited’ (PVI) by submerged macrophytes was about 35%. Phytoplankton biomass estimated as chlorophyll a was strongly controlled by fish, whereas effects of nutrient enrichment were not significant. In 1999 estimated PVI was 80%, no fish effect was observed on phytoplankton biomass, but nutrients had significant effects. Water temperatures were higher in 1998 than in 1999. 2. In the 1998 experiment, cladoceran populations were controlled by fish and cyanobacteria were the dominant phytoplankton group. There were no differences between effects of low (4 g fresh mass m^?2) and high (20 g fresh mass m^?2) fish density on total zooplankton biomass, but zooplankton biomass was higher in the absence of fish. With the high plant density in 1999, fish failed to control any group of the zooplankton community. 3. Total biovolume of phytoplankton strongly decreased with increased nutrient concentrations in 1998, although chlorophyll a concentrations did not significantly change. At higher nutrient concentrations, flagellate algae became more abundant with likely growth rates that could have overcompensated cladoceran feeding rates. This change in phytoplankton community composition may have been because of increases in the DIN : SRP ratio. Both chlorophyll a concentration and total phytoplankton biovolume increased significantly with nutrients in the 1999 experiment. 4. A strong decline of submerged macrophytes was observed in both years as nutrients increased, resulting in shading by periphyton. This shading effect could account for the plant decline despite lower water turbidity at the very high nutrient levels in 1998.     

Consequences of Fish Kills for Long-Term Trophic Structure in Shallow Lakes: Implications for Theory and Restoration

Fish kills are a common occurrence in shallow, eutrophic lakes, but their ecological consequences, especially in the long term, are poorly understood. We studied the decadal-scale response of two UK shallow lakes to fish kills using a palaeolimnological approach. Eutrophic and turbid Barningham Lake experienced two fish kills in the early 1950s and late 1970s with fish recovering after both events, whereas less eutrophic, macrophyte-dominated Wolterton Lake experienced one kill event in the early 1970s from which fish failed to recover. Our palaeo-data show fish-driven trophic cascade effects across all trophic levels (covering benthic and pelagic species) in both lakes regardless of pre-kill macrophyte coverage and trophic status. In turbid Barningham Lake, similar to long-term studies of biomanipulations in other eutrophic lakes, effects at the macrophyte level are shown to be temporary after the first kill (c. 20 years) and non-existent after the second kill. In plant-dominated Wolterton Lake, permanent fish disappearance failed to halt a long-term pattern of macrophyte community change (for example, loss of charophytes and over-wintering macrophyte species) symptomatic of eutrophication. Important implications for theory and restoration ecology arise from our study. Firstly, our data support ideas of slow eutrophication-driven change in shallow lakes where perturbations are not necessary prerequisites for macrophyte loss. Secondly, the study emphasises a key need for lake managers to reduce external nutrient loading if sustainable and long-term lake restoration is to be achieved. Our research highlights the enormous potential of multi-indicator palaeolimnology and alludes to an important need to consider potential fish kill signatures when interpreting results.     

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.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.