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.     

Characterization of bacterial communities in four freshwater lakes differing in nutrient load and food web structure

The phylogenetic composition of bacterioplankton communities in the water column of four shallow eutrophic lakes was analyzed by partially sequencing cloned 16S rRNA genes and by PCR-DGGE analysis. The four lakes differed in nutrient load and food web structure: two were in a clearwater state and had dense stands of submerged macrophytes, while two others were in a turbid state characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had very high nutrient levels (total phosphorus > 100 microg/l), while the other lakes were less nutrient rich (total phosphorus < 100 microg/l). Cluster analysis, multidimensional scaling and ANOSIM (analysis of similarity) were used to investigate differences among the bacterial community composition in the four lakes. Our results show that each lake has its own distinct bacterioplankton community. The samples of lake Blankaart differed substantially from those of the other lakes; this pattern was consistent throughout the year of study. The bacterioplankton community composition in lake Blankaart seems to be less diverse and less stable than in the other three lakes. Clone library results reveal that Actinobacteria strongly dominated the bacterial community in lake Blankaart. The relative abundance of Betaproteobacteria was low, whereas this group was dominant in the other three lakes. Turbid lakes had a higher representation of Cyanobacteria, while clearwater lakes were characterized by more representatives of the Bacteroidetes. Correlating our DGGE data with environmental parameters, using the BIOENV procedure, suggests that differences are partly related to the equilibrium state of the lake.     

Enhanced Input of Terrestrial Particulate Organic Matter Reduces the Resilience of the Clear-Water State of Shallow Lakes: A Model Study

The amount of terrestrial particulate organic matter (t-POM) entering lakes is predicted to increase as a result of climate change. This may especially alter the structure and functioning of ecosystems in small, shallow lakes which can rapidly shift from a clear-water, macrophyte-dominated into a turbid, phytoplankton-dominated state. We used the integrative ecosystem model PCLake to predict how rising t-POM inputs affect the resilience of the clear-water state. PCLake links a pelagic and benthic food chain with abiotic components by a number of direct and indirect effects. We focused on three pathways (zoobenthos, zooplankton, light availability) by which elevated t-POM inputs (with and without additional nutrients) may modify the critical nutrient loading thresholds at which a clear-water lake becomes turbid and vice versa. Our model results show that (1) increased zoobenthos biomass due to the enhanced food availability results in more benthivorous fish which reduce light availability due to bioturbation, (2) zooplankton biomass does not change, but suspended t-POM reduces the consumption of autochthonous particulate organic matter which increases the turbidity, and (3) the suspended t-POM reduces the light availability for submerged macrophytes. Therefore, light availability is the key process that is indirectly or directly changed by t-POM input. This strikingly resembles the deteriorating effect of terrestrial dissolved organic matter on the light climate of lakes. In all scenarios, the resilience of the clear-water state is reduced thus making the turbid state more likely at a given nutrient loading. Therefore, our study suggests that rising t-POM input can add to the effects of climate warming making reductions in nutrient loadings even more urgent.     

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.     

Effects of elevated turbidity on shallow lake fish communities

Synopsis We compared the fish communities of two shallow lakes in the lower Waikato River basin, North Island, New Zealand, to determine the effects of elevated suspended solids (SS) and collapse of submerged macrophytes. Lake Waahi was turbid (20–40 g m^-3 SS) and devoid of submerged macrophytes whereas Lake Whangape was clearer (5 g m^-3 SS) and dominated by submerged macrophytes. The lakes had similar fish species richness and had nine major species in common; representing eight families including Anguillidae, Retropinnidae, Galaxiidae, Eleotridae, Mugilidae, Ictaluridae, Poeciliidae, and Cyprinidae (two species). The only major fish that was absent from Lake Waahi was a lacustrine form of the common smelt, Retropinna retropinna, which disappeared after the lake became turbid in the late 1970s. CPUE, condition, and size of most species in Lake Waahi were similar to, or greater than, those in Lake Whangape. Lake Whangape clearly exceeded Lake Waahi only for CPUE of two species. Within Lake Whangape two species displayed significantly greater condition, and one species greater size, in a turbid arm of the lake than in the main basin. Apart from lacustrine Retropinna retropinna, the fish in these lakes appear well adapted to cope with, or to avoid, the direct toxic effects of suspended and settleable solids on sensitive early developmental stages. In Lake Waahi loss of cover and food provided by submerged macrophytes appears to have been compensated for by increased turbidity and an associated increase in the biomass of the mysid, Tenagomysis chiltoni (a major prey item).     

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.

     

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.     

Does roach behaviour differ between shallow lakes of different environmental state?

The diel activity levels and spatial distribution of roach Rutilus rutilus differed markedly between two shallow lakes of different environmental state. The movements of roach (12–25 cm L _T), with surgically implanted mini‐radio transmitters, were monitored regularly during several 48 h tracking sessions in a clearwater and in a turbid lake. In both lakes, the roach in general were most active during dawn and dusk and least active during the night. Activity level in midsummer was lowest around noon in the clear lake and high around noon in the turbid lake. In summer, roach in the clear lake stayed passively in a restricted area of water lilies during the day and moved into the central part of the lake during the night. In the turbid lake, roach were dispersed all over the lake during the day and moved close to the shoreline at night. Predator : prey fish ratios did not differ in the two lakes, however the observed behaviour of roach in the clearwater lake may be explained by a larger predation pressure from fish and birds, both being favoured in the clear water.     

Migratory benthic fishes may induce regime shifts in a tropical floodplain pond

1. Alternative states are a widely recorded phenomenon in shallow lakes, which may shift between turbid‐ and clear‐water conditions. Here, we investigate whether such shifts in a tropical floodplain pond may be related to the effect of the flood pulse regime on the community structures of fish and macrophytes. 2. Using a long‐term data set, we demonstrate how benthic fish migration together with colonisation by submerged plants affected the transition from a turbid to a macrophyte‐dominated state in a floodplain pond without top‐down control. 3. In our study, the turbid state occurred mostly during low water phases and was largely characterised by high values for the biomass of benthic fish, chlorophyll‐a and total phosphorous. 4. During the period of rising water levels, the migration of benthic fish out of the pond occurs simultaneously with the establishment of submerged plants, while water turbidity decreases along with phytoplankton and nutrient concentrations, inducing a clear‐water phase. However, when submerged plants are absent and fish migration is low, a transient state is generated. 5. We suggest that, in contrast to temperate ponds and shallow lakes, where the main driving mechanisms establishing alternative states are related to cascading effects via the food chain, in tropical ponds and shallow lakes it is resuspension of sediments by benthic fish that plays the most significant role in establishing alternative states. However, the effect of the flood pulse regime plays an important role in the temporal dynamics of fish community structure by controlling benthic fish migration.     

Can artificial plant beds be used to enhance macroinvertebrate food resources for perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis> L.) during the initial phase of lake restoration by cyprinid removal?

Restoration of shallow lakes to a clear-water state, often characterized by high submerged macrophyte cover and a high proportion of piscivores such as perch, Perca fluviatilis L., frequently involves removal of a large proportion of the zoobenthivorous fish, such as bream, Abramis brama L., and roach, Rutilus rutilus L. (i.e. biomanipulation). However, establishment of submerged macrophytes is often delayed following fish removal. This is unfortunate because plant beds typically host high densities of the macroinvertebrates constituting the diet of small perch and thus help perch to go through the bottleneck from feeding on macroinvertebrates to feeding on fish. Establishment of artificial plant beds may be a useful tool to enhance macroinvertebrate population growth and thus food resources for small perch until the natural plants have established. To investigate this restoration option, we studied during two growing seasons (June–October) the composition and abundance of the macroinvertebrate community in artificial plant beds installed in shallow Lake Væng (Denmark) comprising the initial phase of a biomanipulation effort by fish removal. Lake areas with artificial plant beds exhibited substantially higher macroinvertebrate densities than the lake bottom. This suggests that artificial plant beds may be used as feeding grounds for small perch, similarly to the well-known refuge effect for zooplankton against fish predation. In this way, artificial plant beds could help maintain a clear-water state during the transient period when natural submerged vegetation is not yet established in the lake.     

Relationship between bacterial community composition and bottom-up versus top-down variables in four eutrophic shallow lakes

Bacterial community composition was monitored in four shallow eutrophic lakes during one year using denaturing gradient gel electrophoresis (DGGE) of PCR-amplified prokaryotic rDNA genes. Of the four lakes investigated, two were of the clearwater type and had dense stands of submerged macrophytes while two others were of the turbid type characterized by the occurrence of phytoplankton blooms. One turbid and one clearwater lake had high nutrient levels (total phosphorus, >100 micro g liter(-1)) while the other lakes had relatively low nutrient levels (total phosphorus, <100 micro g liter(-1)). For each lake, seasonal changes in the bacterial community were related to bottom-up (resources) and top-down (grazers) variables by using canonical correspondence analysis (CCA). Using an artificial model dataset to which potential sources of error associated with the use of relative band intensities in DGGE analysis were added, we found that preferential amplification of certain rDNA genes over others does not obscure the relationship between bacterial community composition and explanatory variables. Besides, using this artificial dataset as well as our own data, we found a better correlation between bacterial community composition and explanatory variables by using relative band intensities compared to using presence/absence data. While bacterial community composition was related to phytoplankton biomass in the high-nutrient lakes no such relation was found in the low-nutrient lakes, where the bacterial community is probably dependent on other organic matter sources. We used variation partitioning to evaluate top-down regulation of bacterial community composition after bottom-up regulation has been accounted for. Using this approach, we found no evidence for top-down regulation of bacterial community composition in the turbid lakes, while grazing by ciliates and daphnids (Daphnia and Ceriodaphnia) was significantly related to changes in the bacterial community in the clearwater lakes. Our results suggest that in eutrophic shallow lakes, seasonality of bacterial community structure is dependent on the dominant substrate source as well as on the food web structure.     

A model study on the role of wetland zones in lake eutrophication and restoration

Shallow lakes respond in different ways to changes in nutrient loading (nitrogen, phosphorus). These lakes may be in two different states: turbid, dominated by phytoplankton, and clear, dominated by submerged macrophytes. Both states are self-stabilizing; a shift from turbid to clear occurs at much lower nutrient loading than a shift in the opposite direction. These critical loading levels vary among lakes and are dependent on morphological, biological, and lake management factors. This paper focuses on the role of wetland zones. Several processes are important: transport and settling of suspended solids, denitrification, nutrient uptake by marsh vegetation (increasing nutrient retention), and improvement of habitat conditions for predatory fish. A conceptual model of a lake with surrounding reed marsh was made, including these relations. The lake-part of this model consists of an existing lake model named PCLake. The relative area of lake and marsh can be varied. Model calculations revealed that nutrient concentrations are lowered by the presence of a marsh area, and that the critical loading level for a shift to clear water is increased. This happens only if the mixing rate of the lake and marsh water is adequate. In general, the relative marsh area should be quite large in order to have a substantial effect. Export of nutrients can be enhanced by harvesting of reed vegetation. Optimal predatory fish stock contributes to water quality improvement, but only if combined with favourable loading and physical conditions. Within limits, the presence of a wetland zone around lakes may thus increase the ability of lakes to cope with nutrients and enhance restoration. Validation of the conclusions in real lakes is recommended, a task hampered by the fact that, in the Netherlands, many wetland zones have disappeared in the past.     

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.     

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.     

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.     

Biomanipulation-induced reduction of sediment phosphorus release in a tropical shallow lake

Biomanipulation via fish regulation combined with submerged plant introduction is an effective measure to restore eutrophic shallow lakes. Improved water quality and clarity promote growth of benthic algae, which with submerged plants may limit sediment phosphorus (P) release, thereby reinforce lake recovery. Our study sought to evaluate the effect of such a biomanipulation on water quality, benthic algal development and sediment P release in a shallow, tropical lake by (1) comparing porewater and lake water quality, light intensity and benthic algal development in restored and unrestored sections; (2) conducting a ³²P radiotracer experiment to track P release from sediment cores sampled from both sections. The biomanipulation led to lower total P, total dissolved P, and soluble reactive P concentrations in lake water, lower phytoplankton biomass, and increased light intensity at sediment surface, stimulating benthic algal development. Moreover, sediment ³²P release was lower in the restored than unrestored section. Concurrently, dissolved oxygen levels in upper layers of the sediment cores were higher in the restored section. Our study indicates that the biomanipulation improved water quality and enhanced growth of benthic algae, thereby reducing sediment P release, which may be one of the main mechanisms to create successful restoration.     

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.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

Shallow lakes theory revisited: various alternative regimes driven by climate,nutrients, depth and lake size

Shallow lakes have become the archetypical example of ecosystems with alternative stable states. However, since the early conception of that theory, the image of ecosystem stability has been elaborated for shallow lakes far beyond the simple original model. After discussing how spatial heterogeneity and fluctuation of environmental conditions may affect the stability of lakes, we review work demonstrating that the critical nutrient level for lakes to become turbid is higher for smaller lakes, and seems likely to be affected by climatic change too. We then show how the image of just two contrasting states has been elaborated. Different groups of primary producers may dominate shallow lakes, and such states dominated by a particular group may often represent alternative stable states. In tropical lakes, or small stagnant temperate waters, free-floating plants may represent an alternative stable state. Temperate shallow lakes may be dominated alternatively by charophytes, submerged angiosperms, green algae or cyanobacteria. The change of the lake communities along a gradient of eutrophication may therefore be seen as a continuum in which gradual species replacements are interrupted at critical points by more dramatic shifts to a contrasting alternative regime dominated by different species. The originally identified shift between a clear and a turbid state remains one of the more dramatic examples, but is surely not the only discontinuity that can be observed in the response of these ecosystems to environmental change.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.