Stabilizing the clear-water state in eutrophic ponds after biomanipulation: submerged vegetation versus fish recolonization

Biomanipulation through fish removal is a tool commonly used to restore a clear-water state in lakes. Biomanipulation of ponds is, however, less well documented, although their importance for biodiversity conservation and public amenities is undisputed. In ponds, a more complete fish removal can be carried out as compared to lakes and therefore a stronger response is expected. Fish recolonization can, however, potentially compromise the longer term success of biomanipulation. Therefore, we investigated the impact of fish recolonization on zooplankton, phytoplankton, and nutrients for several years after complete drawdown and fish removal in function of submerged vegetation cover in 12 peri-urban eutrophic ponds situated in Brussels (Belgium). Fish recolonization after biomanipulation had a considerable impact on zooplankton grazers, reducing their size and density substantially, independent of the extent of submerged vegetation cover. Only ponds with <30% cover of submerged vegetation shifted back to a turbid state after fish recolonization, coinciding with an increase in density of small cladocerans, rotifers, and cyclopoid copepods. In ponds with >30% submerged vegetation cover, macrophytes prevented an increase in phytoplankton growth despite the disappearance of large zooplankton grazers. Our results suggest that macrophytes, rather than by providing a refuge for zooplankton grazers, control phytoplankton through other associated mechanisms and confirm that the recovery of submerged macrophytes is essential for biomanipulation success. Although the longer term effect of biomanipulation is disputable, increased ecological quality could be maintained for several years, which is particularly interesting in an urban area where nutrient loading reduction is often not feasible.     

Biomanipulation of lake ecosystems: successful applications and expanding complexity in the underlying science

SUMMARY 1. To illustrate advances made in biomanipulation research during the last decade, seven main topics that emerged after the first biomanipulation conference in 1989 are discussed in relation to the papers included in this special issue and the general literature. 2. The substantially higher success rates of biomanipulations in shallow as opposed to stratified lakes can be attributed to several positive feedback mechanisms relating mainly to the recovery of submerged macrophytes. 3. The role of both nutrient loading and in‐lake concentrations in predicting the success of biomanipulations is emphasised and supported by empirically defined threshold values. Nutrient recycling by aquatic organisms (such as fish) can contribute to the bottom‐up effects on lake food webs, although the degree can vary greatly among lakes. 4. Ontogenetic niche shifts and size‐structured interactions particularly of fish populations add to the complexity of lake food webs and make scientifically sound predictions of biomanipulation success more difficult than was previously envisaged. 5. Consideration of appropriate temporal and spatial scales in biomanipulation research is crucial to understanding food web effects induced by changes in fish communities. This topic needs to be further developed. 6. An appropriate balance between piscivorous, planktivorous and benthivorous fishes is required for long‐lasting success of biomanipulations. Recommended proportions and absolute densities of piscivorous fish are currently based on data from only a few biomanipulation experiments and need to be corroborated by additional and quantitative assessments of energy flow through lake food webs. 7. Biomanipulation effects in stratified lakes can be sustained in the long term only by continued interventions. Alternate stable states of food web composition probably exist only in shallow lakes, but even here repeated interventions may be needed as long as nutrient inputs remain high. 8. Biomanipulation is increasingly used as a lake restoration technique by considering the needs of all lake users (sustainability approach). The combination of water quality management and fisheries management for piscivores with positive effects for both appears to be particularly promising. 9. Biomanipulation research has contributed substantially to progress in understanding complex lake food webs, which should in turn promote a higher success rate of future whole‐lake biomanipulations.     

Phytoplankton biomass in Lake Xolotlán (Managua): Its seasonal and horizontal distribution

Some well-documented studies on restoring eutrophic lake systems in The Netherlands by fish stock management have been evaluated with the emphasis on the role of macrophytes. Furthermore, the factors determining the light climate for submerged macrophytes in a large shallow eutrophic lake (Lake Veluwe) have been assessed and the potential success of biomanipulation in large scale projects is discussed. Today relatively little attention has been paid to macrophyte management although the importance of macrophytes in lake restoration has been recognized regularly. The biomanipulation strategy was successful in small scale projects. In a large scale project, however, wind-induced resuspension may largely determine the underwater light climate through attenuation by the water column and periphytic layer. Therefore, restoration of relatively large waterbodies by fish stock management only is expected not to lead to any noteworthy improvement of the light climate for submerged macrophytes. Additional measures aimed at reducing wind-induced resuspension of sediment particles and reestablishing of the macrophyte stands are required for successful biomanipulation strategies. Water quality managers should pay more attention to macrophyte stands in biomanipulation projects because macrophytes enhance a more stable and diverse ecosystem. Restoration objectives and the methods of their achievement must be carefully planned since an abundant submerged macrophyte vegetation may have undesirable effects as well.     

Trophic interactions in a shallow lake following a reduction in nutrient loading: a long-term study

After the diversion of a nutrient-rich inflow, the eutrophic lake, Alderfen Broad, initially showed reduced total phosphorus concentrations and phytoplankton populations, clear water and the establishment of submerged macrophytes. Internal P loading then increased, perhaps stimulated by the senescence of submerged macrophytes and exacerbated by the lack of flushing. Cyanophytes appeared in the summer of two years. As a consequence of poor recruitment of roach (Rutilus rutilus (L.)), the chief zooplanktivore, and a summerkill of the fish population, populations of large-bodied Cladocera (Daphnia hyalina/ longispina and ultimately D. magna) developed. In the long-term, these may have limited the further development of phytoplankton populations and clear water and submerged macrophytes returned. During this latter period, internal P release has remained high (> 380 µg l^-1), thereby indicating the scope for biomanipulation even in eutrophic conditions. However, isolation of the lake has led to a decrease in water level (which through increased temperatures and lowered dissolved oxygen levels was probably responsible for the fish deaths) and further concentration of internal P load. Sediment is now being removed to reestablish greater water depth.     

Top-down control of phytoplankton: the role of time scale, lake depth and trophic state

SUMMARY 1. One of the most controversial issues in biomanipulation research relates to the conditions required for top-down control to cascade down from piscivorous fish to phytoplankton. Numerous experiments have demonstrated that Phytoplankton biomass Top-Down Control (PTDC) occurs under the following conditions: (i) in short-term experiments, (ii) shallow lakes with macrophytes, and (iii) deep lakes of slightly eutrophic or mesotrophic state. Other experiments indicate that PTDC is unlikely in (iv) eutrophic or hypertrophic deep lakes unless severe light limitation occurs, and (v) all lakes characterised by extreme nutrient limitation (oligo to ultraoligotrophic lakes).
2. Key factors responsible for PTDC under conditions (i) to (iii) are time scales preventing the development of slow-growing inedible phytoplankton (i), shallow depth allowing macrophytes to become dominant primary producers (ii), and biomanipulation-induced reduction of phosphorus (P) availability for phytoplankton (iii).
3. Under conditions (iv) and (v), biomanipulation-induced reduction of P-availability might also occur but is insufficient to alter the epilimnetic P-content enough to initiate effective bottom-up control (P-limitation) of phytoplankton. In these cases, P-loading is much too high (iv) or P-content in the lake much too low (v) to initiate or enhance P-limitation of phytoplankton by a biomanipulation-induced reduction of P-availability. However, PTDC may exceptionally result under condition (iv) if high mixing depth and/or light attenuation cause severe light limitation of phytoplankton.
4. Recognition of the five different conditions reconciles previous seemingly contradictory results from biomanipulation experiments and provides a sound basis for successful application of biomanipulation as a tool for water management.     

Is reduction of the benthivorous fish an important cause of high transparency following biomanipulation in shallow lakes?

Experimental reduction of the fish stock in two shallow lakes in The Netherlands shows that such a biomanipulation can lead to a substantial increase in transparency, which is caused not only by a decrease in algal biomass, but also by a decrease in resuspended sediment and detritus. A model was developed to describe transparency in relation to chlorophyll-a and inorganic, suspended solids (resuspended sediment). With the use of this model it is shown that more than 50% of the turbidity in these shallow lakes before biomanipulation was determined by the sediment resuspension, mainly caused by benthivorous fish. Another analysis reveals that the concentration of inorganic suspended solids and the biomass of benthivorous fish are positively correlated, and that even in the absence of algae a benthivorous fish biomass of 600 kg ha⁻¹ can reduce the Secchi depth to 0.4 m in shallow lakes. In addition, it is argued that algal biomass is also indirectly reduced by removal of benthivorous fish. Reduction of benthivorous fish is necessary to get macrophytes and macrophytes seem to be necessary to keep the algal biomass low in nutrient-rich shallow lakes. It is concluded that the impact of benthivorous fish on the turbidity can be large, especially in shallow lakes.     

Major changes in CO<Subscript>2</Subscript> efflux when shallow lakes shift from a turbid to a clear water state

Lakes can be sources or sinks of carbon, depending on local conditions. Recent studies have shown that the CO₂ efflux increases when lakes recover from eutrophication, mainly as a result of a reduction in phytoplankton biomass, leading to less uptake of CO₂ by producers. We hypothesised that lake restoration by removal of coarse fish (biomanipulation) or invasion of mussels would have a similar effect. We studied 14–22 year time series of five temperate Danish lakes and found profound effects on the calculated CO₂ efflux of major shifts in ecosystem structure. In two lakes, where limited colonisation of submerged macrophytes occurred after biomanipulation or invasion of zebra mussels (Dreissena polymorpha), the efflux increased significantly with decreasing phytoplankton chlorophyll a. In three lakes with major interannual variation in macrophyte abundance, the efflux declined with increasing macrophyte abundance in two of the lakes, while no relation to macrophytes or chlorophyll a was found in the third lake, likely due to high groundwater input to this lake. We conclude that clearing water through invasive mussels or lake restoration by biomanipulation may increase the CO₂ efflux from lakes. However, if submerged macrophytes establish and form dense beds, the CO₂ efflux may decline again.     

The role of solid waste materials as habitats for macroinvertebrates in a lowland dam reservoir

Eight hypereutrophic phytoplankton dominated ponds from the Brussels Capital Region (Belgium) were biomanipulated (emptied with fish removal) to restore their ecological quality and reduce the risk of cyanobacterial bloom formation. Continuous monitoring of the ponds before and after the biomanipulation allowed the effects of the management intervention on different compartments of pond ecosystems (phytoplankton, zooplankton, submerged vegetation and nutrients) to be assessed. Fish removal resulted in a drastic reduction in phytoplankton biomass and a shift to the clear-water state in seven out of eight biomanipulated ponds. The reduction in phytoplankton biomass was associated with a marked increase in density and size of large cladocerans in six ponds and a restoration of submerged macrophytes in five ponds. The phytoplankton biomass in the ponds with extensive stands of submerged macrophytes was less affected by planktivorous fish recolonisation of some of the ponds later in the summer. The two non-vegetated ponds as well as one pond with sparse submerged vegetation showed a marked increase in phytoplankton biomass associated with the appearance of fish. Phytoplankton biomass increase coincided with the decrease in large Cladocera density and size. One pond lacking submerged macrophytes could maintain very low phytoplankton biomass owing to large Cladocera grazing alone. The results of this study confirmed the importance of large zooplankton grazing and revegetation with submerged macrophytes for the maintenance of the clear-water state and restoration success in hypereutrophic ponds. They also showed that large Cladocera size is more important than their number for efficient phytoplankton control and when cladocerans are large enough, they can considerably restrain phytoplankton growth, including bloom-forming cyanobacteria, even when submerged vegetation is not restored. The positive result of fish removal in seven out of eight biomanipulated ponds clearly indicated that such management intervention can be used, at least, for the short-term restoration of ecological water quality and prevention of noxious cyanobacterial bloom formation. The negative result of biomanipulation in one pond seems to be related to the pollution by sewage water. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

Biomanipulation and its feasibility for water quality management in shallow eutrophic water bodies in The Netherlands

Biomanipulation as a tool for lake restoration is discussed mainly using literature data. It is based on the exploitation of the interactions both within and between the trophic levels in an aquatic ecosystem. Important among the interactions are: competition for light and nutrients between aquatic macrophytes and phytoplankton and among different phytoplankton species; grazing by planktonic and benthic filter feeders; and size-selective predation by fish. In several case studies biomanipulation has proved to be successful in restorating mildly eutrophic small waterbodies. However, for long-term stability of the restored ecosystems supplementary measures like reducing the external nutrient loadings are needed. The feasibility of the different biomanipulation measures to improve the water quality in shallow Dutch lakes is discussed. Preliminary results on biomanipulation experiments in enclosures withOscillatoria agardhii and the benthic filter feederDreissena polymorpha are given.     

Fish manipulation as a lake restoration tool in shallow,eutrophic, temperate lakes 2: threshold levels,long-term stability and conclusions

In order to evaluate short-term and long-term effects of fish manipulation in shallow, eutrophic lakes, empirical studies on relationships between lake water concentration of total phosphorus (P) and the occurrence of phytoplankton, submerged macrophytes and fish in Danish lakes are combined with results from three whole-lake fish manipulation experiments. After removal of less than 80 per cent of the planktivorous fish stock a short-term trophic cascade was obtained in the nutrient regimes, where large cyanobacteria were not strongly dominant and persistent. In shallow Danish lakes cyanobacteria were the most often dominating phytoplankton class in the P-range between 200 and 1 000μg P l⁻¹. Long-term effects are suggested to be closely related to the ability of the lake to establish a permanent and wide distribution of submerged macrophytes and to create self-perpetuating increases in the ratio of piscivorous to planktivorous fish. The maximum depth at which submerged macrophytes occurred, decreased exponentially with increasing P concentration. Submerged macrophytes were absent in lakes>10 ha and with P levels above 250–300μg P l⁻¹, but still abundant in some lakes<3 ha at 650μg P l⁻¹. Lakes with high cover of submerged macrophytes showed higher transparencies than lakes with low cover aboveca. 50μg P l⁻¹. These results support the alternative stable state hypothesis (clear or turbid water stages). Planktivorous fish>10 cm numerically contributed more than 80 per cent of the total planktivorous and piscivorous fish (>10 cm) in the pelagical of lakes with concentrations above 100μg P l⁻¹. Below this threshold level the proportion of planktivores decreased markedly toca. 50 per cent at 22μg P l⁻¹. The extent of the shift in depth colonization of submerged macrophytes and fish stock composition in the three whole-lake fish manipulations follows closely the predictions from the relationships derived from the empirical study. We conclude that a long-term effect of a reduction in the density of planktivorous fish can be expected only when the external phosphorus loading is reduced to below 0.5–2.0 g m⁻² y⁻¹. This loading is equivalent to an in-lake summer concentration below 80–150μg P l⁻¹. Furthermore, fish manipulation as a restoration tool seems most efficient in shallow lakes.     

Influence of submerged vegetation and fish abundance on water clarity in peri-urban eutrophic ponds

Despite the presence of high nutrient concentrations, most ponds located around Brussels (Belgium) show a considerable variation in turbidity. The importance of submerged macrophytes in maintaining the clear-water state requires identification of the main factors determining macrophyte abundance and diversity in ponds and small lakes. In this study, the inter-relationships between submerged macrophyte cover, fish abundance and turbidity were investigated in 13 eutrophic peri-urban ponds. Along a turbidity gradient, vegetation switched from dominance by Stoneworts (Chara and Nitella spp.) in the clearest ponds, to dominance by Potamogeton pectinatus in ponds with a slightly lower water transparency. Despite the presence of both P. pectinatus and Stoneworts in each of the vegetated ponds, only one became dominant. Only a very low abundance (around 20%) of submerged vegetation was found in ponds of intermediate turbidity, while macrophytes were absent in turbid ponds. Multi- and univariate analysis showed a marked difference in chemical, physical and biological properties between ponds deliberately used for fish stocking and ponds that were not. Macrophyte cover was significantly negatively correlated with turbidity and plankti-benthivorous fish abundance. No such correlation was observed with piscivorous fish abundance, except for pike that were associated with a charophyte vegetation in the study ponds. The strong relationship found between fish abundance and turbidity, its negative effect on submerged vegetation cover, and the importance of submerged vegetation in controlling phytoplankton abundance, should be taken into account when selecting ponds for fish stocking. It also suggests that the study ponds have a good potential for ecological quality restoration by biomanipulation.     

Why biomanipulation can be effective in peaty lakes

The effects of fish stock reduction (biomanipulation) was studied in an 85 ha shallow peaty turbid lake. The lake cleared in a 4-week period in April–May 2004, which demonstrated that biomanipulation can be effective in peaty lakes. We demonstrated that it is possible to reduce the fish stock to <25 kg ha⁻¹ benthivorous fish and <15 kg ha⁻¹ planktivorous fish, sufficiently low to switch the lake from a turbid to a clear state. Knowledge of lake morphology, fish stock, fish behaviour, and a variety of fishing methods was necessary to achieve this goal. It is expected that continuation of fisheries to remove young of the year planktivorous species is needed for several years, until macrophytes provide sufficient cover for zooplankton and can compete with phytoplankton. Cladocerans developed strongly after fish removal. The clearing of the lake coincided with a sudden decrease of filamentous cyanobacteria and suspended detritus, and a strong increase of Bosmina. We assume that Bosmina was able to reduce filamentous prokaryotes and detritus. After the disappearance of the cyanobacteria, Bosmina disappeared too. After the clearing of the lake Daphnia dominated in zooplankton and apparently was able to keep phytoplankton levels low. In our case, wind resuspension did not prevent biomanipulation from being successful. No correlation between windspeed and turbidity was found, neither in an 85 ha nor in a 230 ha shallow peaty lake. Regression analysis showed that on average 50% of the amount of suspended detritus can be explained by resuspension by fish and 50% by phytoplankton decomposition. The main goal of this biomanipulation experiment, clear water and increased submerged plant cover in a shallow peaty lake, was reached.     

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.     

Identification of total phosphate,submerged vegetation cover and zooplankton size thresholds for success of biomanipulation in peri-urban eutrophic ponds

In small shallow lakes and ponds, the clear-water state can generally be maintained at higher nutrient concentrations compared to larger shallow lakes. The main objective of this study was to identify thresholds for total phosphorus (TP), submerged vegetation cover and zooplankton size that determine biomanipulation success in peri-urban eutrophic ponds. Additionally, the relationship between transparency and TP is discussed with regard to similar relationships and thresholds reported for shallow lakes. Using classification trees, a threshold TP concentration of 0.300 mg P L^?1 was determined below which a clear-water state was generally maintained after biomanipulation. When the average TP concentration was >0.300 mg P L^?1, the stability of the clear-water state largely depended on the presence of sufficiently large zooplankton (>0.87 mm) or a submerged vegetation cover of >82% at some point during the year. This threshold TP concentration is considerably higher than the threshold of 0.1 mg L^?1 which is generally suggested for longer-term success of biomanipulation in shallow lakes. Such threshold nutrient concentration is important when restoring ecological quality in eutrophic small lakes and ponds. Extended follow-up of biomanipulation success in eutrophic ponds could provide more insight into the feasibility of these thresholds on the longer term.     

Biomanipulation additional to nutrient control for restoration of shallow lakes in The Netherlands

Eutrophication control is one of the major issues in the environmental policy in The Netherlands. As a result of international action programmes the average phosphorus loading of freshwater systems should decrease by 50% between 1985 and 1995. However, in many cases the restoration of water quality requires additional measures. Recovery is hampered by the structure and functioning of the present food-chain.

The feeding behaviour of the dominant fish species in Dutch lakes, bream and roach, tend to impose a homeostasis on the system, resisting restoration of water quality. In shallow lakes, biomanipulation, including drastic reduction of fish-stocks, may induce a shift from a stable ‘turbid-water state’ to a stable ‘clear-water state’.

To assess the possibilities of biomanipulation for the restoration of a particular lake, three questions are relevant: (1) is a drastic reduction of fish-stocks feasible?, (2) will a shift occur from ‘turbid to clear’ after the fish reduction? and (3) will the new situation of clear water be stable? This paper focuses attention on the last two questions. The increase in water clarity, following fish reduction, largely depends on the increase in the density of the Daphnia-population and the contribution of benthivorous fish to the resuspension of sediments. A ‘turbid to clear’ shift may be expected if the total biomass of planktivorous and benthivorous fish is reduced to levels<50 kg ha^?1. The stability of the achieved clear-water state largely depends on the development of submerged macrophytes in the lake and on the level of nutrient loading. It is tentatively concluded that a stable clear-water state may be expected at initial total-P concentrations<0.10 mg l^?1.

Because the water managers in The Netherlands have no fishing rights, they have to.co-operate with anglers and commercial fishermen to apply biomanipulation as a tool for water management.

     

Changes in phosphorus cycling in a shallow lake due to food web manipulations

SUMMARY. 1. Food web manipulation, by removal of planktivorous or benthivorous fish, is a promising method for reducing phytoplankton concentrations in shallow lakes. The part that nutrients may play in the success of such a measure is not well documented.
2. In this study, we analysed the flow of phosphorus through the food web of the shallow, eutrophic Lake Wolderwijd/Nuldernauw. Our studies occurred in the years 1981 (when a bloom of cyanobacteria occurred) and 1987 (no bloom); a hypothetical situation was also examined in which most of the bream are assumed to be removed.
3. The analysis shows that the success of biomanipulation is probably due not only to an increased grazing pressure on the phytoplankton, but also to a decreased availability of phosphorus. The reason for this is the removal of detrital phosphorus by increased sedimentation as a result of a predicted increase in growth of macrophytes after biomanipulation.     

Effects of an artificial protection structure on the sandy shore macrofaunal community: the special case of Lido di Dante (Northern Adriatic Sea)

Biomanipulation of eutropicated peaty lakes has rarely been successful; clear water with dense macrophyte stands fails to develop in most cases. It was unclear whether (1) high turbidity due to resuspension by benthivorous fish or wind is the major cause of low macrophyte density or whether (2) the establishment of submerged macrophyte stands is prevented by a lack of propagules, low cohesive strength of the lake sediment, high concentrations of phytotoxics, grazing by waterfowl and/or shading by periphyton growth. These hypotheses were tested in an experiment in a shallow peat lake in the Netherlands (Terra Nova). Removal of fish from a 0.5 ha experimental site resulted in clear water and the development of a dense (90% coverage) and species-rich (10 species) submerged vegetation. At a fish-stocked site and a control site the water remained turbid and dense macrophyte stands did not develop. The establishment of submerged macrophytes appeared not to be limited by a lack of propagules. Introduced plants grew poorly in turbid water, but very well in clear water. Exclosures showed that bird grazing reduced the plant biomass. In clear water grazing seemed to enhance the vegetation diversity. Periphyton development did not prevent plant growth in clear water. After the experiment, the fish stock was greatly reduced in the whole lake (85 ha), to test if (3) in a large lake, submerged macrophyte stands will not develop after biomanipulation. In the first season after fish reduction, transparency increased and species-rich submerged macrophyte stands developed, covering 60% of the shallow parts of the lake. Most of the species known to have occurred in the past re-established. The results indicate that high turbidity caused by benthivorous fish in combination with bird grazing were the major causes of the absence of submerged macrophyte stands in this lake. Abiotic conditions after the clearing of the lake were suitable for the growth of macrophytes. We infer that the restoration potential of submerged macrophyte stands in eutrophicated peaty lakes can be high, and results can be obtained quickly.     

Wave exposure related growth of epiphyton: implications for the distribution of submerged macrophytes in eutrophic lakes

The distribution of submerged macrophytes in eutrophic lakes has been found to be skewed towards sites with intermediate exposure to waves. Low submerged macrophyte biomass at exposed sites has been explained by, for instance, physical damage from waves. The aim of this study was to investigate if lower biomass at sheltered sites compared to sites with intermediate exposure to waves can be caused by competition from epiphyton.Investigations were performed in eutrophic lakes in southern Sweden. Samples of submerged macrophytes and epiphytic algae on the macrophytes were taken along a wave exposure gradient. The amount of epiphyton (AFDW) per macrophyte biomass decreased with increased exposure. Biomass of submerged macrophytes, on the other hand, increased with increased exposure until a relatively abrupt disappearance of submerged vegetation occurred at high exposures. Production of epiphytic algae was monitored on artificial substrates from June to September at a sheltered and an exposed site in three lakes. It was higher at sheltered sites compared with exposed sites.We suggest that epiphytic algae may be an important factor in limiting the distribution of submerged macrophytes at sheltered sites in eutrophic lakes.     

Changes in benthic macroinvertebrate abundance and lake isotope (C,N) signals following biomanipulation: an 18-year study in shallow Lake Vaeng,Denmark

Change in the abundance of benthic macroinvertebrates and the stable isotope composition (C, N) of benthic invertebrates and zooplankton in Lake Vaeng, Denmark, was investigated over an 18-year period following biomanipulation (removal of cyprinids). During the first nine years after biomanipulation, the lake was clear and submerged macrophytes were abundant; after this period, a shift occurred to low plant abundance and high turbidity. Two years after the biomanipulation, total density of benthic macroinvertebrates reached a maximum of 17042 (±2335 SE) individuals m⁻² and the density was overall higher when the lake was in a clear state. Redundancy analysis (RDA) suggested macrophyte abundance and total nitrogen (TN) concentration were the dominant structuring forces on the benthic macroinvertebrate assemblage. Stable isotope analysis revealed that δ¹³C of macroinvertebrates and zooplankton was markedly higher in years with high submerged macrophyte abundance than in years without macrophytes, most likely reflecting elevated δ¹³C of phytoplankton and periphyton mediated by a macrophyte-induced lowering of lake water CO₂ concentrations. We conclude that the strong relationship between macrophyte coverage and δ¹³C of macroinvertebrates and cladocerans may be useful in paleoecological studies of past changes in the dynamics of shallow lakes, as change in macrophyte abundance may be tracked by the δ¹³C of invertebrate remains in the sediment.     

Spatial distribution of four freshwater fish species in different types of artificial European water bodies

Spatial distribution of young-of-the-year (YOY) and older roach, rudd, perch and ruffe was compared in two artificial lakes with macrophytes present and absent, and a valley reservoir, using gillnets. Almost all species of interest and both age categories preferred benthic habitats. The depth distribution in benthic habitats was relatively consistent across water bodies with the highest fish densities found in the shallowest depths. In the macrophyte-rich lake, YOY roach and perch utilize the 3–6 m benthic layer the most, whereas the fish preferred the 0–3 m benthic layer in the macrophyte-poor lake and reservoir. No differences were found in the depth distribution in pelagic habitats sampled by pelagic gillnets for YOY fish between the water bodies. Older fish usually utilized the surface water layer. Macrophytes influenced the depth distribution of YOY fish in benthic habitats, where their density maximum shifted deeper in the macrophyte-rich lake when fewer macrophytes were present in the shallowest benthic depth. In lakes, YOY fish utilized a wider depth spectrum due to the deeper thermocline when compared to the reservoir. Oxygen and temperature stratification are the main factors influencing fish distribution, whereas macrophyte presence particularly influences the depth distribution of YOY fish in benthic habitats.