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.     

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.     

Goals,methods and possibilities for directing development of limnic ecosystems

Case studies illustrating lake and wetland ecosystem problems as well as restoration methods are given. Among these methods, aeration of deep lakes, sediment removal from shallow, polluted lakes, sediment manipulation in polluted and acidified lakes, biomanipulation and wetland management methods are considered. A treatment program for directing ecosystem development is designed in each individual case according to lake type, degradation problems and goals of restoration. The most common goal in treatment of lakes is to meiotrophicate (oligotrophicate) hypertrophicated ecosystems. In the case of wetlands, conservation and restoration aspects are combined with a growing interest focused on biomass production by emergent macrophytes. Within practical frames of applied limnology, basic limnological research and training of doctorands have been organized as team-work for ecosystem-oriented investigations.     

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.     

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.     

Effects of a filter-feeding fish [silver carp, Hypophthalmichthys molitrix (Val.)] on phyto- and zooplankton in a mesotrophic reservoir: results from an enclosure experiment

SUMMARY 1. Silver carp, Hypophthalmichthys molitrix (Val.), feeds on both phyto- and zooplankton and has been used in lake biomanipulation studies to suppress algal biomass. Because reports on the effects of silver carp on lake food webs have been contradictory, we conducted an enclosure experiment to test how a moderate biomass of the fish (10 g wet weight m⁻³) affects phytoplankton and crustacean zooplankton in a mesotrophic temperate reservoir.
2. Phytoplankton biomass <30 μm and particulate organic carbon (POC) <30 μm were significantly higher in enclosures with silver carp than in enclosures without fish, whereas Secchi depth was lower. Total copepod biomass declined strongly in both treatments during the experiment, but it was significantly higher in fish-free enclosures. Daphnid biomass was also consistently higher in enclosures without fish, although this effect was not significant. However, the presence of fish led to a fast and significant decrease in the size at maturity of Daphnia galeata Sars. Thus, the moderate biomass of silver carp had a stronger negative effect on cladoceran zooplankton than on phytoplankton.
3. Based on these results and those of previous studies, we conclude that silver carp should be used for biomanipulation only if the primary aim is to reduce nuisance blooms of large phytoplankton species (e.g. cyanobacteria) that cannot be effectively controlled by large herbivorous zooplankton. Therefore, stocking of silver carp appears to be most appropriate in tropical lakes that are highly productive and naturally lack large cladoceran zooplankton.     

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.     

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.

     

Effects of Biomanipulation on Fish and Plankton Communities in Ten Eutrophic Lakes of Southern Finland

The effects of biomanipulation were studied in ten Finnish lakes to determine responses in fish and plankton communities and water quality after mass removal of cyprinids. From 1997 to 2001, the fish communities shifted from the dominance of large cyprinids to an explosion of small cyprinids and a higher proportion of piscivores in effectively biomanipulated lakes (>200 kg ha⁻¹ 3 yr⁻¹). The biomass of cyanobacteria decreased, and the duration of the blooms shortened and shifted towards the autumn. Decreased concentrations and slower cycling of nutrients and increased grazing by cladocerans probably affected the declined biomass of cyanobacteria. Less intensive sediment disturbance and increased phosphorus-retention in fast growing fish biomass may have turned the role of the fish assemblage from ‘nutrient recycler’ to ‘nutrient storage’. Increased potential grazing pressure, higher proportion of edible algae, and lower chlorophyll a:total phosphorus ratio indicated strengthened herbivore control. A high mass removal catch in relation to trophic state, low background turbidity, and bearable external loading favoured the successful biomanipulation, whereas intensive cyprinid reproduction, high nutrient loading and non-algal turbidity hindered the recovery. Three important issues should be noticed before biomanipulation in Finland: (1) careful selection of target lake, (2) well-planned, effective and long-lasting biomanipulation and (3) sustainable management of piscivores. An erratum to this article is available at .     

A stable carbon and nitrogen isotope study of lake food webs in Canada's Boreal Plain

1.  Stable carbon and nitrogen isotope and fish stomach content analyses were used to investigate food webs in five relatively undisturbed lakes on the Boreal Plain of Canada. Stable isotope analysis was also used to determine the importance of external and internal carbon sources.
2.  Overlap in the carbon and nitrogen signatures of primary producers made it difficult to determine unambiguously the feeding habits of many invertebrates. However, isotope analysis suggested that external carbon inputs were detectable in the aquatic food chains of the one lake with a short water residence time («1 year). In the other four lakes, with water residence times ≥1 year, autochthonous carbon was the only detectable carbon source in the food webs.
3.  Food webs in these lakes spanned a range of four to five trophic levels. Both invertebrates and fish appeared to eat a variety of food, often feeding at more than one trophic level.
4.  With the exception of one lake (SPH20), top predators in these lakes, northern pike ( Esox lucius ) and fathead minnows ( Pimephales promelas ), occupied similar trophic positions despite large differences in body size and trophic morphology. In SPH20, where there were two additional fish species, pike occupied a higher trophic position. However, all the top predators in each lake appeared to be omnivores and generalists.
5.  The prevalence of omnivory and the apparent generalist feeding habits of fish in these lakes suggest that organisms are flexible in their feeding habits and that these food webs will be resilient to disturbance.     

The use of structural dynamic models to explain successes and failures of biomanipulation

A development of a structural dynamic model, i.e. a model with current change of the most important parameters according to a goal function, is presented with the aim to explain the structural changes observed in lakes, when the nutrient concentration is increased or decreased. This type of models may be important in lake management as it may be possible qualitatively to predict the success or failure of biomanipulation. The answer to the crucial question: àt which phosphorus level will the success of biomanipulation be most probable?' will probably require the development of model which takes into account site specific processes and properties, i.e., a more complicated model. As goal function is proposed the thermodynamic function, exergy, which is defined as the work content of the system (model) compared with the system at thermodynamic equilibrium. It is shown that the structural dynamic modelling approach has been able to explain the shift from large to small zooplankton species at a certain level of phosphorus concentration, accompanied by a shifts from a dominance of zooplankton, and predatory fish to a system dominated by planktivorous fish and phytoplankton. The shift in zooplankton species cannot be explained by application of catastrophe theoretical models, which have been used to explain the hysteresis reaction. The results show that the shift should be expected at approximately 0.12 mg P l-1 and that a typical hysteresis reaction occurs at this concentration in accordance with the expectations. These results are consistent with many observations but should be interpreted with great caution, as the model is simple and general and don't account for a number of processes which may influence the results significantly in specific lake studies. The structural dynamic approach has previously been used in ten case studies with good agreement with the observations, but more case studies are needed before a general recommendation of the use of this type of models can be given. The results from this study point toward to apply this type of models for lake management where biomanipulation is involved, although it should be recommended to improve the presented general model with introduction of site specific properties for a considered lake study.     

Comparison of losses of planktivorous fish by predation and seine-fishing in a lake undergoing long-term biomanipulation

SUMMARY 1. Piscivore stocking at artificially high densities and fishing are the two common approaches to reduce the amount of planktivorous and benthivorous fish in lake biomanipulation programmes. Both measures have advantages and disadvantages, but their relative efficacy has not previously been directly compared.
2. We calculated the average annual catch of roach and bream in a lake undergoing long-term biomanipulation (Feldberger Haussee, Germany) by seining each year between 1992 and 1998. We compared this value with a bioenergetics estimate of annual consumption rates of the dominant cohorts of piscivores, pikeperch and pike, in 1997 and 1998. We also determined species composition and length distribution of prey fish in stomachs of the piscivores.
3. Roach was the dominant prey species of both pikeperch and pike, whereas bream was rarely taken by either piscivorous species. Seining removed on average larger specimens of roach than were found in the stomachs of the piscivores.
4. Based on stocking densities of the piscivores, published mortality rates, and individual consumption rates, feeding of pikeperch and pike on roach exceeded the manual removal of roach by seining by a factor of 4–15 (biomass) in 1997 and 1998.
5. Based on these results, a combination of fishing and piscivore enhancement is recommended. Whereas the stocks of adult roach and bream have to be reduced mainly by fishing, the predation of piscivores should be directed predominantly towards the juvenile zooplanktivorous fish. Therefore, small size-classes of piscivorous fish should be promoted by fisheries management, including stocking and harvest regulations.     

Top-down or Bottom-up Effects by Fish: Issues of Concern in Biomanipulation of Lakes

A large-scale biomanipulation trial was carried out on Lake Vesijärvi in Finland during 1989–1993. Following the mass removal of coarse fish the biomass of cyanobacteria collapsed from 1.4 g/m^?3 to below 0.4 g/m^?3, while total phosphorus concentration declined from 45 μ g/L to 30 μ g/L. No relevant changes in zooplankton communities were observed. The results suggest that the success of food web manipulation as a tool for lake restoration is not necessarily dependent on the grazing rate of zooplankton. The effects of reduced fish-mediated internal loading and recycling of nutrients are in many cases stronger than those of reduced planktivory. Alternative stable states of water quality may also exist in lakes not covered by macrophytes, owing to the changes in the behavior of fish stocks. Year-to-year variation in the littoral zone may cause large oscillations in lake ecosystems—for example, through the recruitment of fish. In addition, the nutrients translocated by fish from the littoral zone may affect the nutrient dynamics of the pelagial plankton community. In terms of phytoplankton species composition and the ratio of phosphorus to chlorophyll a, the water quality in Lake Vesijärvi has improved in a stepwise fashion within the last 10 years. This is probably due to the fact that the five-year mass removal of fish in Enonselkä fulfilled the requirement of sustained management of fish stocks in order to maintain nonequilibrial conditions between alternate stable states. The prediction of the water quality development is obscured, however, by spatial and temporal within-lake variation, which sets high requirements for sampling programs.     

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

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

The role of light for fish–zooplankton–phytoplankton interactions during winter in shallow lakes – a climate change perspective

1. Variations in the light regime can affect the availability and quality of food for zooplankton grazers as well as their exposure to fish predation. In northern lakes light is particularly low in winter and, with increasing warming, the northern limit of some present-day plankton communities may move further north and the plankton will thus receive less winter light.
2. We followed the changes in the biomass and community structure of zooplankton and phytoplankton in a clear and a turbid shallow lake during winter (November–March) in enclosures both with and without fish and with four different light treatments (100%, 55%, 7% and <1% of incoming light).
3. In both lakes total zooplankton biomass and chlorophyll- a were influenced by light availability and the presence of fish. Presence of fish irrespective of the light level led to low crustacean biomass, high rotifer biomass and changes in the life history of copepods. The strength of the fish effect on zooplankton biomass diminished with declining light and the effect of light was strongest in the presence of fish.
4. When fish were present, reduced light led to a shift from rotifers to calanoid copepods in the clear lake and from rotifers to cyclopoid copepods in the turbid lake. Light affected the phytoplankton biomass and, to a lesser extent, the phytoplankton community composition and size. However, the fish effect on phytoplankton was overall weak.
5. Our results from typical Danish shallow eutrophic lakes suggest that major changes in winter light conditions are needed in order to have a significant effect on the plankton community. The change in light occurring when such plankton communities move northwards in response to global warming will mostly be of modest importance for this lake type, at least for the rest of this century in an IPCC A2 scenario, while stronger effects may be observed in deep lakes.     

Linking Cascading Effects of Fish Predation and Zooplankton Grazing to Reduced Cyanobacterial Biomass and Toxin Levels Following Biomanipulation

Eutrophication has been one of the largest environmental problems in aquatic ecosystems during the past decades, leading to dense, and often toxic, cyanobacterial blooms. In a way to counteract these problems many lakes have been subject to restoration through biomanipulation. Here we combine 13 years of monitoring data with experimental assessment of grazing efficiency of a naturally occurring zooplankton community and a, from a human perspective, desired community of large Daphnia to assess the effects of an altered trophic cascade associated with biomanipulation. Lake monitoring data show that the relative proportion of Daphnia spp. grazers in June has increased following years of biomanipulation and that this increase coincides with a drop in cyanobacterial biomass and lowered microcystin concentrations compared to before the biomanipulation. In June, the proportion of Daphnia spp. (on a biomass basis) went from around 3% in 2005 (the first year of biomanipulation) up to around 58% in 2012. During months when the proportion of Daphnia spp. remained unchanged (July and August) no effect on lower trophic levels was observed. Our field grazing experiment revealed that Daphnia were more efficient in controlling the standing biomass of cyanobacteria, as grazing by the natural zooplankton community never even compensated for the algal growth during the experiment and sometimes even promoted cyanobacterial growth. Furthermore, although the total cyanobacterial toxin levels remained unaffected by both grazer communities in the experimental study, the Daphnia dominated community promoted the transfer of toxins to the extracellular, dissolved phase, likely through feeding on cyanobacteria. Our results show that biomanipulation by fish removal is a useful tool for lake management, leading to a top-down mediated trophic cascade, through alterations in the grazer community, to reduced cyanobacterial biomass and lowered cyanobacterial toxin levels. This improved water quality enhances both the ecological and societal value of lakes as units for ecosystem services.     

Community concordance between the fish and aquatic birds of lakes in northern Alberta, Canada: the relative importance of environmental and biotic factors

1. Community concordance measures the degree to which patterns in community structure in a set of sites are similar between two different taxonomic groups. Although seldom incorporated into studies of lake ecosystems, aquatic birds can be influenced by the same environmental features of lakes which affect fish and invertebrates, and can interact with these organisms directly as predators, competitors or prey. We surveyed lakes in north-central Alberta, Canada, to determine if co-occurring fish and aquatic bird assemblages displayed concordance, and assessed the relative importance of environmental and biotic factors in contributing to observed concordance.
2. In 41 lakes (3–305 ha), we encountered seven species of fish and thirty-one avian taxa which subsequently were used in multivariate analyses. Fish assemblages dominated by large piscivores were in large deep lakes, whereas fishless lakes and lakes with only small-bodied fish were small and shallow, and thus, prone to winter hypoxia. Bird assemblages displayed three general patterns: (a) small shallow lakes supported a 'core' of widespread species (between three and eight species per lake); (b) large, deep lakes supported more species (between 11 and 16), including large, aerially foraging piscivores; and (c) large, shallow lakes supported the most species (between 15 and 23), including many ducks.
3. Randomization tests of matrix concordance and Mantel tests both showed that fish and bird assemblages were significantly concordant. Concordance reflected the fact that both groups were strongly affected by the same key environmental factors, principally lake size and maximum depth, and to a lesser extent, productivity and geographic isolation. Direct interactions between birds and fish, such as predation and competition, appeared to play much smaller roles in shaping the two assemblages.     

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

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

Applicability of planktonic biomanipulation for managing eutrophication in the subtropics

Although large-bodied cladocerans such asDaphnia can reduce algal biomass significantly in temperate lakes if freed from fish predation, the applicability of such biomanipulation techniques for eutrophication management in the subtropics and tropics has been examined only recently. Subtropical cladoceran assemblages differ from those of temperate lakes by their low species richness, early summer gameogenesis, and greatly reduced body size. Eutrophic Florida lakes are dominated by pump-filter feeding fish rather than by size selective planktivores as a temperate lakes. Cladocerans in Florida lakes can increase in abundance significantly if freed from fish but fail to have an impact on algal biomass or composition. The greatest potential for using biomanipulation to manage phytoplankton-dominated lakes in the subtropics and tropics lies with phytophagous fish. Future research should concentrate on defining the role of individual fish taxa on phytoplankton composition and community structure, nutrient cycling, and planktonic productivity before embarking on whole lake manipulation projects.