Lake trophic state and the limnological effects of omnivorous fish

Ecologists have hypothesized that planktivorous fish have greater effects on the plankton and water quality of oligotrophic lakes than eutrophic lakes. We tested this hypothesis in a tank-mesocosm experiment of factorial design in which five biomass levels of filter-feeding omnivorous gizzard shad (Dorosoma cepedianum) were cross-classified with two levels of lake trophic state achieved by filling tank-mesocosms with water and plankton transported by truck from two lakes with different trophic states. The presence of gizzard shad significantly increased total phosphorus, primary productivity, chlorophyll, and particulate phosphorus (PP) 2–20 and 20–200 μm and significantly decreased Secchi depth, cladocerans, copepods and PP > 200 μm. The effects of gizzard shad on chlorophyll, Secchi depth, cladocerans, copepods and PP 2–20 and > 200 μm were dependent on lake trophic state and most intense in the eutrophic lake system. This experiment suggests that filter-feeding omnivorous fish interact synergistically with trophic state so that the limnological effects of omnivorous fish become more intense with increased eutrophication.     

Chemical limnology of two artificial lakes used for both stormwater management and recreation

The aims of this study were to document the mainly chemical behaviour of two linked artificial lakes used for both stormwater management and recreation in the new town of Craigavon. Further, the understanding of their behaviour should help in their management and the design of other similar lakes.The lake mean total phosphorus (73 µg P l^–1), nitrate (0.50 mg N l^–1) and chlorophyll a (25 µg l^–1) concentrations, Secchi depth (1.2 m) and the estimated total phosphorus loading (1.98 g m^–2 a^–1) all classify the main lake as eutrophic. An important source of the phosphorus load on the lakes is the urban area of Craigavon (52% of the total load). The interrelationships between total phosphorus, chlorophyll a and Secchi depth in the main lake are similar to those in natural ones. In addition, the lake follows the total phosphorus load — trophic state relationships (lake total phosphorus and chlorophyll a concentrations and Secchi depth) found to apply elsewhere. These two points indicate that the artificial lakes in Craigavon behave similarly to natural ones.     

Impact of the removal of gizzard shad (Dorosoma cepedianum) on nutrient cycles in Lake Apopka,Florida

1. The St. Johns River Water Management District removed over 5.4 million kg of gizzard shad (Dorosoma cepedianum) from Lake Apopka, FL during 1993–2005, as a means of reducing lake phosphorus and phytoplankton concentrations and improving water clarity. Other steps included reduction of external nutrient inputs and operation of a treatment wetland. We measured nutrient excretion by Lake Apopka gizzard shad to quantify the nutrient effect of this biomanipulation. 2. Both N and P excretion were significantly affected by fish body mass and temperature. Larger fish had lower mass‐specific rates of excretion than smaller fish. 3. High water temperature increased P excretion to a much greater extent than N, resulting in a low N : P of nutrient excretion in midsummer. The N : P of excretion was lower than has been observed in other systems, probably because of higher water temperature. 4. Removal of gizzard shad >200 g prevented the annual release of 45 800 kg N year^?1 (3.46 kg N ha^?1 year^?1) and 7700 kg P year^?1 (0.62 kg P ha^?1 year^?1) on average. The actual impact on the P cycle varied substantially from year to year (range 7900–78 800 kg N year^?1; 1200–14 800 kg P year^?1), primarily because of fluctuations in the catch. 5. On an annual basis, the P directly removed in fish tissues was similar to that removed by the treatment wetland. The P excretion prevented by the removal of fish was approximately 20% of the reduction in external P loading achieved during 1993–2005. 6. In the short term, most of the P demand of planktonic primary producers is met through recycling of P, which greatly exceeds external P loading. Depending on population biomass, phosphorus excretion by the resident gizzard shad population was similar in magnitude to the P release by diffusive flux from the sediments.     

Selective feeding by three native North American freshwater mussels implies food competition with zebra mussels

The response of a benthic macroinvertebrate assemblage to whole-lake biomanipulation was studied in a small Finnish mesotrophic lake. From 1993 to 1997, over 200 kg ha^–1 of fish, mainly roach (Rutilus rutilus (L.)) and bream (Abramis brama (L.)) were caught and the fish biomass was reduced by nearly 80%. The biomass and density of benthic invertebrates were investigated during the years of fish removal and for the following three years. The decrease in benthivorous fish stock led to a higher biomass and density of all major groups of benthic invertebrates during the early years of fish removal. Non-biting midges (Chironomidae), water mites (Hydrachnellae), mayfly nymphs (Ephemeroptera), sphaeriid clams (Bivalvia: Sphaeridae) and biting midges (Ceratopogonidae) seemed to respond most profoundly to changes in fish biomass. The biomass of most invertebrate groups correlated negatively with the catch-per-unit-effort (CPUE). The total biomass and density of invertebrates had strong negative correlations with the CPUE (r= -0.85, p = 0.016, r = -0.84, p = 0.019, respectively), but they did not correlate significantly with total phosphorus, chlorophyll a, or temperature. However, the variation in total biomass that was not explained by the CPUE, was significantly associated with total phosphorus.The fish stock recovered to almost its initial level within three years after fish removal had been discontinued. As an apparent response to increased predation pressure, the biomasses of many invertebrate groups decreased again in the years 1999–2000. The strong relationship between macrozoobenthos and fish populations in the studied lake is likely to be a consequence of the open and sparsely vegetated bottom, which offers minimal shelter to invertebrate prey. An additional factor behind the recent low biomass levels may be changes in primary production. Phosphorus and chlorophyll a concentrations started to decrease markedly after three years of fishing and they have remained at a low level.     

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 .     

Impacts of environmental scenarios on chlorophyll-a in the management of shallow, eutrophic lakes following biomanipulation: An application of a numerical model

Accurate prediction of species changes in lake ecosystems following biomanipulation measures is of paramount importance in view of water quality management. The temporal variation of phytoplankton biomass as chlorophyll-a and transparency as Secchi depth measurements are studied in the Lake Bleiswijkse Zoom, The Netherlands, with a comprehensive structural dynamic model. In the formulation of the biological model, phytoplankton as several species, zooplankton, detritus, planktivores and benthivores, and piscivores are considered to be major contributing state variables for the model. The primary goal of this paper is to describe the possible impacts of several environmental scenarios on chlorophyll-a biomass qualitatively as it would help lake and environmental managers and relevant authorities elucidate the processes of eutrophication and biomanipulation in a broad way. Some of the scenarios that have been studied by this model are: (1) The effect of fixed stoichiometry in terms of internal nitrogen and phosphorus that are tied up within algal cells; (2) the effects of external phosphorus limitation; (3) light limitation and external nitrogen limitation on algal growth; (4) probable consequences that have taken place within the chlorophyll-a biomass due to change in biomasses of various aquatic organisms; and (5) possible changes of chlorophyll-a biomass due to higher temperatures caused by global warming.     

Factors related to gizzard shad and the threadfin shad occurrence and abundance in Florida lakes

Gizzard shad Dorosoma cepedianum were collected in 23 and threadfin shad D. petenense were collected in 22 of the 60 Florida lakes sampled. Logistic regression equations were 94% effective for predicting gizzard shad occurrence from chlorophyll and lake surface area, and 84% effective for predicting threadfin shad occurrence from lake surface area and lake volume inhabited (PVI). Occurrence of both shad species was related positively to lake size. In lakes where gizzard shad or threadfin shad were collected, shad density and biomass of both shad species were related positively to chlorophyll. Gizzard shad populations were generally vulnerable to predation in lakes, with the per cent of gizzard shad ≤200mm L _T values exceeding 60% with few exceptions. Effects of gizzard shad and threadfin shad on fish community dynamics may be confined to relatively large (>100 ha) and fertile (chlorophyll >20–30μg l⁻¹) Florida lakes.     

Stochastic modelling of nutrient loading and lake ecosystem response in relation to submerged macrophytes and benthivorous fish

SUMMARY 1. The strong stabilising effect of increased submerged macrophytes (charophytes) and benthivorous fish reduction on the clear water state was shown for shallow Lake Veluwe and Lake Wolderwijd. 2. The first two links in the chain of relationships from external phosphorus (P) loading to in‐lake total‐P concentrations to chlorophyll a concentrations to water transparency, showed a significant correlation with the areal fraction of coverage with charophytes. Higher coverages lead to (i) lower ratios of the in‐lake total‐P concentration compared with the volume weighted average concentration in the inlet water, indicating a higher retention of P in the presence of charophytes (ii) lower chlorophyll a to total‐P ratios, indicating a positive effect of charophytes on top‐down control of algae, and (iii) higher water transparency because of lower algal turbidity. Transparency further improved as a result of benthivorous fish reduction and a significant positive correlation between non‐algal turbidity and benthivorous fish biomass. 3. A model was developed taking into account the inherent variability in precipitation and uncertainties in the empirical relationships determining phosphorus export from stream catchments and other sources and eutrophication variables in the receiving lakes. The model was used to compute (i) probability distributions for in‐lake total‐P, chlorophyll a and Secchi Disc transparency in relation to the coverage with charophytes and benthivorous fish biomass, and (ii) exceedence probabilities with respect to critical values for in‐lake total‐P and water transparency for several management scenarios. 4. The effects of an expected rise in external nutrient loading on the in‐lake total‐P and chlorophyll a concentrations and on water transparency can be compensated for by two proposed control measures: (i) extended treatment at a waste water treatment plant directly discharging into Lake Veluwe, and (ii) diverting the outlet of a stream draining a catchment with high fertilisation. The minimal internal charophyte coverage needed to sufficiently stabilise the clear water state and to meet with the objective of a summer mean water transparency of at least 1 m was estimated at well over 30% of the lake area, while the benthivorous fish stock should be maintained at the present level of c. 20 kg ha^?1.     

Strength of phytoplankton–nutrient relationship: evidence from 13 biomanipulated ponds

Phytoplankton biomass–nutrient relationship is widely used by lake managers to assess the eutrophication impact and to set the nutrient targets. Submerged vegetation and large zooplankton grazing have long been identified as factors weakening the relationship by decoupling phytoplankton from nutrients. Proving this decoupling unambiguously is difficult because, in natural systems, many factors act together, blurring each other’s effect. In this article, we present the results of continuous monitoring of 13 ponds where the effects of submerged vegetation and zooplankton grazing were enhanced by biomanipulation (fish removal). The monitoring allowed these effects to be assessed and compared with the pre-biomanipulation situations when phytoplankton biomass was mainly nutrient driven. The comparison showed a strong weakening effect of submerged vegetation and large zooplankton grazing on the chlorophyll a–total phosphorus relationship suggesting that a considerable degree of ecological quality of ponds affected by eutrophication can be restored even when nutrient-loading reduction is not feasible.     

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.     

Do the effects of piscivorous largemouth bass cascade to the plankton?

Ecologists have hypothesized that an increase in the biomass of piscivorous fish in lakes will cause a decrease in populations of planktivorous fish, an increase in the size of herbivorous zooplankton and a decrease in the biomass of phytoplankton. Here we present an experimental test of whether the effects of largemouth bass (Micropterus salmoides) cascade to the planktivorous fish, zooplankton and phytoplankton of a 15-ha water storage reservoir. A pilot study indicated that the reservoir was eutrophic with dense populations of planktivorous fish dominated by threadfin shad (Dorosoma petenense). No piscovorous fish were present in the reservoir. We conducted a one-month mesocosm experiment using water and plankton from the reservoir showing that the presence of threadfin shad reduced large-sized zooplankton and increased the productivity and biomass of phytoplankton. To test whether the effects of piscivorous fish could cascade to the plankton, we assessed the effects of the addition of piscivorous largemouth bass on the planktivorous fish, zooplankton and biomass of phytoplankton of the reservoir by monitoring the reservoir during the year before and the two years after largemouth bass were stocked. In the second year after the addition of largemouth bass, the number of planktivorous fish decreased and the relative abundance of threadfin shad declined. Although the abundance of cladocerans increased after the addition of largemouth bass, the average size of zooplankton did not change. We did not detect changes in chlorophyll a, Secchi depth, or concentrations of total phosphorus and total nitrogen as a result of the addition of largemouth bass.     

Effects of planktivore abundance on chlorophyll-a and Secchi depth

We used two analyses to test the hypothesis that planktivore abundances contribute to the residual variations of Secchi depth or chlorophyll-a plotted with respect to mean summer epilimnetic total phosphorus. The first analysis involved 15 lake years of data from six lakes. The data set comprised mark-recapture assessments of piscivore and planktivore numbers and estimates of mean summer chlorophyll-a, total phosphorus and Secchi depth. We found that residual chlorophyll-a variation was not significantly (p>0.05) correlated with planktivore densities, but that planktivore densities did contribute (p<0.02) to the residual variation of Secchi depth on mean total phosphorus. The second analysis included all of the data used in the first plus an additional 13 lake years of data from the literature. These data showed that the percentage of the total fish community comprising planktivores did not significantly (p>0.05) contribute to the residual variation in chlorophyll-a with respect to mean summer total phosphorus. Together, our results suggest that planktivore abundance has a significant cascading impact on water clarity, but no long term statistically significant impact on mean summer chlorophyll-a concentration.     

Dynamics of submerged macrophyte populations in response to biomanipulation

1. A 6‐year study (1992–97) of changes in submerged vegetation after biomanipulation was carried out in the eutrophicated Lake Finjasjön, Southern Sweden. Ten sites around the lake were revisited each year. At each site five samples of above‐ground biomass were taken at 10 cm water depth intervals. An investigation of the seed bank at the 10 sites, and a grazing experiment where birds and large fish were excluded was also conducted. 2. Between 1992 and 1996, in shallow areas (water depth < 3 m), vegetation cover increased from < 3 to 75% and above‐ground biomass from < 1 to 100 g DW m^–2. Mean outer water depth increased from 0.3 to 2.5 m. Elodea canadensis and Myriophyllum spicatum accounted for > 95% of the increase in biomass and plant cover. The following year (1997), however, cover and above‐ground biomass decreased, mainly attributable to the total disappearance of E. canadensis. Secchi depth increased after biomanipulation until 1996, but decreased again in 1997. 3. Total and mean number of submerged species increased after biomanipulation, probably as a result of the improved light climate. However, after the initial increase in species number there was a decrease during the following years, possibly attributed to competition from the rapidly expanding E. canadensis and M. spicatum. The lack of increase in species number after the disappearance of E. canadensis in 1997 implies that other factors also affected species richness. 4. A viable seed bank was not necessary for a rapid recolonization of submerged macrophytes, nor did grazing by waterfowl or fish delay the re‐colonization of submerged macrophytes. 5. Submerged macrophytes are capable of rapid recolonization if conditions improve, even in large lakes such as Finjasjön (11 km²). Species that spread by fragments will increase rapidly and probably outcompete other species. 6. The results indicate that after the initial Secchi depth increase, probably caused by high zooplankton densities, submerged vegetation further improved the light climate. The decrease in macrophyte biomass in 1997 may have caused the observed increase in phosphorus and chlorophyll a, and the decrease in Secchi depth. We suggest that nutrient competition from periphyton, attached to the macrophytes, may be an important factor in limiting phytoplankton production, although other factors (e.g. zooplankton grazing) are also of importance, especially as triggers for the shift to a clear‐water state.     

Identification of a biomanipulation‐driven regime shift in Lake Vesijärvi: implications for lake management

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Stocking piscivores to improve fishing and water clarity: a synthesis of the Lake Mendota biomanipulation project

SUMMARYY 1. A total of 2.7 × 10⁶ walleye fingerlings and 1.7 × 10⁵ northern pike fingerlings were stocked during 1987–99 in eutrophic Lake Mendota. The objectives of the biomanipulation were to improve sport fishing and to increase piscivory to levels that would reduce planktivore biomass, increase Daphnia grazing and ultimately reduce algal densities in the lake. The combined biomass of the two piscivore species in the lake increased rapidly from < 1 kg ha^?1 and stabilised at 4–6 kg ha^?1 throughout the evaluation period. 2. Restrictive harvest regulations (i.e. increase in minimum size limit and reduction in bag limit) were implemented in 1988 to protect the stocked piscivores. Further restrictions were added in 1991 and 1996 for walleye and northern pike, respectively. These restrictions were essential because fishing pressure on both species (especially walleye) increased dramatically during biomanipulation. 3. Commencing in 1987 with a massive natural die‐off of cisco and declining yellow perch populations, total planktivore biomass dropped from about 300–600 kg ha^?1 prior to the die‐off and the fish stocking, to about 20–40 kg ha^?1 in subsequent years. These low planktivore biomasses lasted until a resurgence in the perch population in 1999. 4. During the period prior to biomanipulation when cisco were very abundant, the dominant Daphnia species was the smaller‐bodied D. galeata mendotae, which usually reached a biomass maximum in June and then crashed shortly thereafter. Beginning in 1988, the larger‐bodied D. pulicaria dominated, with relatively high biomasses occurring earlier in the spring and lasting well past mid‐summer of many years. 5. In many years dominated by D. pulicaria, Secchi disc readings were greater during the spring and summer months when compared with years dominated by D. galeata mendotae. During the biomanipulation evaluation period, phosphorus (P) levels also changed dramatically thus complicating our analysis. Earlier research on Lake Mendota had shown that Daphnia grazing increased summer Secchi disc readings, but P concentrations linked to agricultural and urban runoff and to climate‐controlled internal mixing processes were also important factors affecting summer readings. 6. The Lake Mendota biomanipulation project has been a success given that high densities of the large‐bodied D. pulicaria have continued to dominate for over a decade, and the diversity of fishing opportunities have improved for walleye, northern pike and, more recently, yellow perch. 7. Massive stocking coupled with very restrictive fishing regulations produced moderate increases in piscivore densities. Larger increases could be realised by more drastic restrictions on sport fishing, but these regulations would be very controversial to anglers. 8. If the lake's food web remains in a favourable biomanipulation state (i.e. high herbivory), further improvements in water clarity are possible with future reductions in P loadings from a recently initiated non‐point pollution abatement programme in the lake's drainage basin.     

Effects of macrophytes on lake‐water quality across latitudes: a meta‐analysis

Macrophytes are widely recognized for improving water quality and stabilizing the desirable clear‐water state in lakes. The positive effects of macrophytes on water quality have been noted to be weaker in the (sub)tropics compared to those of temperate regions. We conducted a global meta‐analysis using 47 studies that met our set criteria to assess the overall effects of macrophytes on water quality (measured by phytoplankton chlorophyll a concentration, total nitrogen concentration, total phosphorus concentration, Secchi depth and the trophic state index) and to investigate how these effects correlate with latitude using meta‐regressions. We also examined if the effects of macrophytes on lake‐water quality differ with growth form and study design in (sub)tropical and temperate areas by grouping the data and then comparing the effect sizes. We found that macrophytes significantly reduced phytoplankton chlorophyll a concentration, total nitrogen concentration, total phosphorus concentration, as well as the trophic state index, but they did not have a significant overall effect on Secchi depth. The effects of macrophytes on reducing phytoplankton chlorophyll a concentration, total nitrogen concentration and the trophic state index did not differ with latitude. However, the reduction of total phosphorus concentration was greater at lower latitudes. We showed that at lower latitudes, the positive effects of macrophytes on water quality are similar to or greater than those at higher latitudes, thus challenging the prevailing paradigm of macrophytes being less effective at enhancing lake‐water quality in the (sub)tropics. Furthermore, our data showed that the macrophyte effects vary by growth forms, and the growth forms that positively affect water quality differ between the (sub)tropical and temperate areas. We showed a lack of significant macrophyte effects in surveys within and outside macrophyte stands, suggesting difference in the sensitivities of study designs or possibly weaker effects of macrophytes in lakes compared to experimental settings.     

Thresholds and Stability of Alternative Regimes in Shallow Prairie–Parkland Lakes of Central North America

Numerous studies have demonstrated alternative regimes in shallow lake ecosystems around the world, with one state dominated by submerged macrophytes and the other by phytoplankton. However, the stability of each regime, and thresholds at which lakes shift to the alternative regime, are poorly known. We used a cross-sectional analysis of 72 shallow lakes located in prairie and parkland areas of Minnesota, USA, during 2005 and 2006 to assess the occurrence of alternative regimes and shifts between them. Cluster analysis revealed two distinct groups of lakes characterized not only by different macrophyte abundance and chlorophyll a levels but also by different total phosphorus–chlorophyll a relationships. Thirty-nine lakes were macrophyte- and 23 lakes phytoplankton-dominated in both years, whereas 10 sites shifted sharply between those regimes. We failed to detect a universal shifting threshold in terms of chlorophyll a or total phosphorus. However, 95% of the lakes with chlorophyll a concentrations less than 22 μg l⁻¹ were in a clear-water regime, whereas 95% of the lakes with chlorophyll a higher than 31 μg l⁻¹ were in a turbid regime. Total phosphorus less than 62 μg l⁻¹ was an accurate predictor of lakes in a stable clear-water regime, whereas a large change in biomass of planktivores and benthivores between years was the only variable weakly related to regime shifts. Our results support the theoretical prediction that regime thresholds vary among lakes. We recommend that lake managers focus on improving resilience of clear regimes in shallow lakes by reducing nutrient loading, rather than attempting to identify and manage complex triggers of regime shifts. Author contributions KDZ, MAH, BRH, and MLK all contributed to the design of the study, performed the research, analyzed data, and helped write the article.     

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

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

Stable isotope evidence of ontogenetic changes in the diet of gizzard shad Dorosoma cepedianum

Stable sulphur isotopic composition (δ³⁴S) of gizzard shad Dorosoma cepedianum was used to investigate the seasonal and ontogenetic variation in the diet of young and adult fish. This study evaluated fish from a hypereutrophic lake that had recently undergone a 40% reduction of large (>300 mm total length, L _T) D. cepedianum biomass as part of a biomanipulation experiment, which aimed at reducing internal nutrient loading. Dorosoma cepedianum δ³⁴S values showed evidence of ontogenetic changes with young fish (<200 mm L _T) depending more on benthic food sources than adults (>200 mm L _T). The δ³⁴S composition of the adult fish suggested an increasing importance of zooplankton in the diet, although benthic food sources remained part of the diet of all D. cepedianum collected in this study. The results indicated that benthic feeding is used by D. cepedianum of all sizes, suggesting that biomanipulation efforts may need to target all sizes of fish to realize benefits.