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.     

Resuspension in a shallow eutrophic lake

The frequency and the importance of wind-induced resuspension were studied in the shallow, eutrophic Lake Arresø, Denmark (41 km², mean depth 3 m). During storm events in autumn 1988 lake water samples were collected every 2–8 hours by an automatic sampler at a mid-lake station. The concentration of suspended solids and Tot-P was found to increase markedly. During storms up to 2 cm of the superficial sediment was resuspended, and the concentration of resuspended solids in the water column rose to 140 mg l^–1. The resuspended particles had a relatively high settling velocity and on average, a relatively short residence time in the water column of 7 hours.A model which describes the concentration of resuspended solids as a function of wind velocity and of settling velocity of the resuspended particles is presented. Using additional wind velocity data from a nearby meteorological station, the model has been used to calculate the frequency of resuspension events and concentration of resuspended solids for the period from May to November 1988.These calculations show that resuspension occurred about 50% of the time. Average flux of suspended solids from the sediment to the water was 300 g m^–2 d^–1 and during 50% of the time lake water concentration of suspended solids was more than 32 mg l^–1. A relationship between concentration of suspended solids and Secchi-depth is presented. Because of resuspension, Secchi-depth in Lake Arresø is reduced to 0.5 m.Resuspension also had a marked effect on Tot-P concentration in the lake water, and P input to the lake water being totally dominated by resuspension events.     

Seasonal water quality of shallow and eutrophic Lake Pamvotis, Greece: implications for restoration

Lake Pamvotis is a moderately sized (22 km²) shallow (z _avg=4 m) lake with a polymictic stratification regime located in northwest Greece. The lake has undergone cultural eutrophication over the past 40 years and is currently eutrophic (annual averages of FRP=0.07 mg P l^-1, TP=0.11 mg P l^-1, NH₄ ⁺=0.25 mg N l^-1, NO₃ ^–=0.56 mg N l^-1). FRP and NH₄ ⁺ levels are correlated to external loading from streams during the winter and spring, and to internal loading during multi-day periods of summer stratification. Algal blooms occurred in summer (July–August green algae, August–September blue-green algae), autumn (October blue-green algae and diatoms), and winter (February diatoms), but not in the spring (March–June). The phytoplankton underwent brief periods of N- and P-limitation, though persistent low transparency (secchi depth of 60–80 cm) also suggests periods of light limitation. Rotifers counts were highest from mid-summer to early autumn whereas copepods were high in the spring and cladocerans were low in the summer. Removal of industrial and sewage point sources a decade ago resulted in a decrease in FRP. A phosphorus mass balance identified further reductions in external loading from the predominately agricultural catchment will decrease FRP levels further. The commercial fishery and lake hatchery also provides opportunities to control algal biomass through biomanipulation measures.     

Dynamics of phytoplankton detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

A study was made of the mortality and aerobic decomposition of light- and phosphorus-limited cultures of Oscillatoria limnetica, a dominant phytoplankton species in shallow, eutrophic Lake Loosdrecht (The Netherlands). When placed in the dark at 20 °C, most cells died and lysed within twelve days. The labile organic matter was completely decomposed within three weeks. Absorbance spectra indicated that blue green algae may contributed significantly to the refractory dissolved substances in the lake. Refractory particulate matter constituted from 7 to 24% of the biomass of O. limnetica, depending on the growth rate before incubation in the dark. The decomposition rate of this fraction was 0.005 d^–1. On a basis of a steady-state model of the dynamics of phytoplankton detritus, the areal organic dry weight concentration of the detritus in the lake is ca. 60 g m^–2. This means the quantities of detritus in the seston and epipelon are about equal.     

Biomanipulation,new perspectives for restoration of shallow,eutrophic lakes in The Netherlands

Eutrophication of Dutch lakes has led to massive algal growth, disappearance of most of the macrophytes and disturbance of the food chain. The pike population has fallen sharply and bream developed very strongly, in the absence of this predator. Eutrophication problems are primarily being tackled by reducing nutrient loading. Restoration of water quality, however, seems to be impeded by the present structure of the food chain,i.e. the large bream stock. Biomanipulation, especially fish stock control with the aim of reducing the bream stock and increasing that of predatory fish, can accelerate the process of restoration. For the further development of biomanipulation it is very important that water authorities and managers of fish stocks agree on a common strategy.     

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.

     

Estimation of reference conditions for phytoplankton in a naturally eutrophic shallow lake

Classification of waters using biological quality elements and determination of the degree of deviation from reference levels is a key issue in the Water Framework Directive of EU. Lakes in reference conditions with sufficient biological data are available for several boreal lake types with the exception of naturally eutrophic lakes. An empirical approach is one alternative for estimating the reference conditions of such lakes. We used the water transparency of the naturally eutrophic Lake Tuusulanjärvi recorded in August in the early 1910s to estimate reference values for phytoplankton biomass and chlorophyll a concentrations. Three phytoplankton samples during August 2000–2001 corresponded to the estimated reference values for total biomass (<5.6 mg l^?1) and chlorophyll a (<28 μg l^?1), as did the simultaneous Secchi depths. The phytoplankton assemblage in these samples with 24 eutrophy indicators (17% of the total taxa number) corresponded in general the species list from the early 1900s, which as such could be regarded as reference assemblage. Furthermore, in August 2000, 3 years after intensive fish removal a prominent decrease in cyanobacterial biomass and toxin concentration was observed. The costs of the measures and studies in Lake Tuusulanjärvi during 1989–2003 have been approximately 2.5 million euros.     

Engineering and biological approaches to the restoration from eutrophication of shallow lakes in which aquatic plant communities are important components

Engineering approaches (nutrient removal, sediment pumping, hypolimnion oxygenation, alum treatments) may be most appropriate to deep lakes where the aim of restoration from eutrophication is simply to reduce the production and crop of one component, the phytoplankton. They do not always give the desired results because the nutrient loading may only be reduced to a limited extent. There are additional problems in shallow lakes where change of state between community dominance (aquatic plants versus plankton) is wanted. Each community has powerful buffering mechanisms and biomanipulation may be essential to switch one state to another even with considerable nutrient reduction. For the phytoplankton-dominated community the buffers include the advantages of early growth, lower diffusion pathways for CO₂, overhead shading, and an absence of large cladoceran grazers. This later is because open-water shallow environments provide no refuges against predation for the large Cladocera which are both the most efficient grazers and the most favoured prey for fish. Restoration of aquatic plants may then require provision of refuges for the grazers. Different sorts of refuge are discussed using case studies of Hoveton Great Broad and Cockshoot Broad in the Norfolk Broadland.     

Indirect effects of fish community structure on submerged vegetation in shallow,eutrophic lakes: an alternative mechanism

The loss of submerged macrophytes during eutrophication of shallow lakes is a commonly observed phenomenon. The proximate reason for this decline is a reduction of available light due to increasing phytoplankton and/or epiphyton biomass. Here we argue that the ultimate cause for the transition from a macrophyte-dominated state to a phytoplankton-dominated state is a change in fish community structure. A catastrophic disturbance event (e.g. winterkill) acting selectively on piscivores, cascades down food chains, eventually reducing macrophyte growth through shading by epiphyton, an effect that is reinforced by increasing phytoplankton biomass. The transition back from the phytoplankton to the macrophyte state depends on an increase in piscivore standing stock and a reduction of planktivores. A conceptual model of these mechanisms is presented and supported by literature data and preliminary observations from a field experiment.     

A comparison of shallow Danish and Canadian lakes and implications of climate change

1. Winter temperatures differ markedly on the Canadian prairies compared with Denmark. Between 1 January 1998 and 31 December 2002, average weekly and monthly temperatures did not drop below 0 °C in the vicinity of Silkeborg, Denmark. Over this same time, weekly average temperatures near Calgary, Alberta, Canada, often dropped below −10 °C for 3–5 weeks and the average monthly temperature was below 0 °C for 2–4 months. Accordingly, winter ice conditions in shallow lakes in Canada and Denmark differed considerably. 2. To assess the implications of winter climate for lake biotic structure and function we compared a number of variables that describe the chemistry and biology of shallow Canadian and Danish lakes that had been chosen to have similar morphometries. 3. The Danish lakes had a fourfold higher ratio of chlorophyll‐a: total phosphorus (TP). Zooplankton : phytoplankton carbon was related to TP and fish abundance in Danish lakes but not in Canadian lakes. There was no significant difference in the ratio log total zooplankton biomass : log TP and the Canadian lakes had a significantly higher proportion of cladocerans that were Daphnia. These differences correspond well with the fact that the Danish lakes have more abundant and diverse fish communities than the Canadian lakes. 4. Our results suggest that severe Canadian winters lead to anoxia under ice and more depauperate fish communities, and stronger zooplankton control on phytoplankton in shallow prairie lakes compared with shallow Danish lakes. If climate change leads to warmer winters and a shorter duration of ice cover, we predict that shallow Canadian prairie lakes will experience increased survivorship of planktivores and stronger control of zooplankton. This, in turn, might decrease zooplankton control on phytoplankton, leading to ‘greener’ lakes on the Canadian prairies.     

Coupling of phytoplankton and detritus in a shallow,eutrophic lake (Lake Loosdrecht,The Netherlands)

An oscillating steady state is described of phytoplankton, dominated by Prochlorothrix hollandica and Oscillatoria limnetica, and sestonic detritus in shallow, eutrophic Lake Loosdrecht (The Netherlands). A steady-state model for the coupling of the phytoplankton and detritus is discussed in relation to field and experimental data on phytoplankton growth and decomposition. According to model predictions, the phytoplankton to detritus ratio decreases hyperbolically at increasing phytoplankton growth rate and is independent of a lake's trophic state. The seston in L. Loosdrecht contains more detritus than phytoplankton as will apply to many other lakes. The model provides a basis for estimating the loss rate of the detritus, including decomposition, sedimentation and hydraulic loss. In a shallow lake like L. Loosdrecht detritus will continue to influence the water quality for years.     

Fisheries management as an additional lake restoration measure: biomanipulation scaling-up problems

Restoration of the highly eutrophic Reeuwijk lakes (ca. 700 ha) started in 1986 by reducing the external phosphorus loading. As an additional measure to improve the quality of the lake water, the structure of the fish population in Lake Klein Vogelenzang (18 ha) was altered in 1989 by the removal of ca. 100 kg ha^–1 bream from the lake in April and December. This constituted about 50% of the total bream biomass in the lake.The fish-stock reduction in April, 1989, was initially followed by high phosphorus concentrations, probably the result of considerable phosphorus release from the sediments. The resulting heavy algal blooms that occurred reduced the transparency to very low values. During the summer the zooplankton population increased markedly in numbers coinciding with reductions in total suspended matter including (blue-green) algae. A great improvement in Secchi-disc transparency was observed and by the end of December, 1989, the bottom of the lake (1.5–2.0 m) was visible. After heavy storms in January and February 1990, transparency dropped to < 1 m as a result of resuspension of high concentrations of suspended matter from the bottom sediments. Although transparency over the rest of 1990 was higher than in 1988, i.e. the year preceding the removal of fish (biomanipulation), it was lower than expected, based on the results of 1989. The study shows that technical and biological factors can cause serious management problems for the implementation of biomanipulation in larger water bodies.     

Effects of different fish species and biomass on plankton interactions in a shallow lake

Thirty-six enclosures, surface area 4 m², were placed in Little Mere, a shallow fertile lake in Cheshire, U.K. The effects of different fish species (common carp, common bream, tench and roach) of zooplanktivorous size, and their biomass (0, 200 and 700 kg ha^–1) on water chemistry, zooplankton and phytoplankton communities were investigated. Fish biomass had a strong effect on mean zooplankton size and abundance. When fish biomass rose, larger zooplankters were replaced by more numerous smaller zooplankters. Consequently phytoplankton abundance rose in the presence of higher densities of zooplanktivorous fish, as zooplankton grazing was reduced. Fish species were also significant in determining zooplankton community size structure. In enclosures with bream there were significantly greater densities of small zooplankters than in enclosures stocked with either carp, tench and, in part, roach. When carp or roach were present, the phytoplankton had a greater abundance of Cyanophyta than when bream or tench were present. Whilst top-down effects of fish predation controlled the size partitioning of the zooplankton community, this, in turn apparently controlled the bottom-up regeneration of nutrients for the phytoplankton community. At the zooplankton–phytoplankton interface, both top-down and bottom-up processes were entwined in a reciprocal feedback mechanism with the extent and direction of that relationship altered by changes in fish species. This has consequences for the way that top-down and bottom-up processes are generalised.     

Evaluating stocking of YOY pike Esox lucius as a tool in the restoration of shallow lakes

1. Stocking of piscivores in shallow, eutrophicated lakes to reduce cyprinid densities is a common approach in lake restorations. Young‐of‐the‐year (YOY) pike Esox lucius are frequently used to reduce cyprinid densities, but their effectiveness is equivocal. This study uses a simple model to assess the efficacy of pike stocking as a lake restoration tool. 2. The model evaluates YOY pike consumption in relation to different survival and growth patterns, timing of stocking, degree of piscivory of pike as well as YOY cyprinid growth patterns and productivity. The parameters used in the model are obtained from field surveys in eight study lakes stocked with YOY pike as well as from the literature. 3. Our model showed that all parameters studied were important for predicting the effects of pike stocking on cyprinids. In particular, body size at stocking, cyprinid production and pike survival were good predictors of biomanipulation success. 4. However, our model showed that only under very specific circumstances will stocking of YOY pike result in a reduction of YOY cyprinid densities large enough to affect water quality. 5. The findings of this study question the efficacy of pike stocking as an appropriate and reliable tool for restoration programmes in shallow lakes, and indicate that when used efforts should be made to optimize the timing of stocking in relation to YOY cyprinid production and to increase the stocking body size of the pike.     

Phytoplankton biomass reduction after planktivorous fish reduction in a shallow,eutrophic lake: a combined effect of reduced internal P-loading and increased zooplankton grazing

Hydrobiologia - No recovery was recorded in the shallow and eutrophic Lake Væng, Denmark, after a sewage diversion in 1981, due to an internal phosphorus loading and a dominance of...     

Centennial‐scale changes to the aquatic vegetation structure of a shallow eutrophic lake and implications for restoration

1. We investigate long‐term (>200 years) changes to the composition and spatial structure of macrophyte communities in a shallow, eutrophic lake (Barton Broad, eastern England) and consider the implications for lake restoration. 2. Historical macrophyte data were assembled from a variety of sources: existing plant databases, museum herbaria, journal articles, old photographs and eyewitness accounts. Additionally, two types of sediment core sample were analysed for plant macro‐remains and pollen; bulk basal samples from multiple core sites analysed to provide information on ‘pre‐disturbance’ macrophyte communities and two whole cores analysed to determine historical change. 3. Prior to the late 1800s, macrophyte communities were diverse and included a multilayered mosaic of short‐stature submerged taxa and taller submerged and floating‐leaved species. With the progression of eutrophication after around 1900, the former community was displaced by the latter. Diversity was maintained, however, since an encroaching Schoenoplectus–nymphaeid swamp generated extensive patches of low‐energy habitat affording refugia for several macrophytes otherwise unable to withstand the hydraulic forces associated with open water conditions. When this swamp vegetation disappeared in the 1950s, many of the ‘dependent’ aquatic macrophytes also declined leaving behind a sparse, species‐poor community (as today) resilient to both eutrophication and turbulent open waters. 4. The combination of historical and palaeolimnological data sources offers considerable benefits for reconstructing past changes to the aquatic vegetation of lakes and for setting restoration goals. In this respect, our study suggests that successful restoration might often be better judged by reinstatement of the characteristic structure of plant communities than the fine detail of species lists; when nutrients are low and the structure is right, the right species will follow.     

Seasonal changes in sediment and water chemistry of a subtropical shallow eutrophic lake

A field study was conducted (May 1981 to June 1982) to develop a data-base on seasonal changes of water and sediment chemistry of Lake Monroe (4 000 ha surface and ca. 2 m deep) located in central Florida, USA. This shallow eutrophic lake is a part of the St. Johns River. Quantitative samples of lake water and sediments were collected on a monthly basis from 16 stations and analyzed for various physico-chemical parameters. Relatively high levels of dissolved solids (mean electrical conductivity (EC) = 1832 µS cm¹) prevailed in the lake water, and seasonal changes in EC were probably associated with hydrologic flushing from external sources, such as incoming water from upstream as well as precipitation. Average monthly levels of total N and P during the study period were 1.82 and 0.21 mg l^–1, respectively. Nutrient concentrations in the water did not show any strong seasonal trends. Organic matter content of lake sediments ranged from 1 to 182 g C kg^–1 of dry sediment, reflecting considerable spatial variability. All nutrient elements in the sediments showed highly significant (P < 0.01) correlations with sediment organic C, though little or no significant relationship appeared at any sampling period between water and sediment chemistry of the lake. Temporal trends in water and sediment chemical parameters may have been concealed by periodic hydrologic flushing of the St. Johns River into Lake Monroe.Florida Agricultural Experiment Stations Journal Series No. 7836.     

Restoration and recovery of shallow eutrophic lake ecosystems in The Netherlands: epilogue

During the symposium Restoration and recovery of shallow lake ecosystems in The Netherlands studies on restoration of eutrophic lakes were addressed and discussed. Many Dutch shallow lakes have received high external loadings of phosphorus through supply water that is influenced by the River Rhine and loadings in The Netherlands. Two important Action Plans (the Rhine Action Plan, the North Sea Action Plan) are now in operation to reduce nutrient emissions. The targets set are not likely to be fully reached, so that supplementary reduction of phosphorus supplied to inland fresh waters will be required. In several shallow lakes such a reduction has been achieved recently, but without leading to discernible recovery. The main causes of delay are phosphorus storage and its subsequent release from sediments and foodweb; however, the remaining extraneous phosphorus supply is often still too high. Supplementary actions are, therefore, called for. A further reduction of phosphorus inputs is suggested, besides supplementary measures proposed, viz. dredging, flushing, biomanipulation, chemomanipulation. Restoration to the past situation via upwelling groundwater appeares to be feasible in some cases. There is a common consensus that each lake behaves differently depending to its morphology, hydrology and history of eutrophication. Therefore each lake has to be studied before restoration measures can be applied. Besides, the ecosystem should not only be studied as a separate entity, but as a part of systems of a higher integration level.