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.     

Plankton composition and water quality in a pond of spring origin in Turkey

Phyto/zooplankton composition, chlorophyll a, and some water quality parameters were investigated in a spring-originated pond in Central Anatolia between February 2001 and January 2002. Water temperature, pH, dissolved oxygen, Secchi depth, total and calcium hardness, nitrate-nitrogen, nitrite-nitrogen, ammonia-nitrogen, total phosphorus, and soluble reactive phosphorus levels were analyzed. A total of 49 species belonging to Bacillariophyceae, Chlorophyceae, Cyanophyceae, Cryptophyceae, and Dinophyceae were identified. The highest phytoplankton abundance was found in August, whereas the lowest was determined in January. Phytoplankton abundance increased from February to August and declined in the following months. The Bacillariophyceae were dominant in the phytoplankton community. A total of 21 species of Rotifera, 2 species of Cladocera, and 1 genus of Copepoda were found. The zooplankton community was dominated by Rotifera. The highest abundance of zooplankton was recorded in July and the lowest value in November. The annual mean concentration of chlorophyll a was measured as 1.90 μg l⁻¹. In spite of these eutrophic levels (mean values of total phosphorus and nitrate-nitrogen: 0.069 mg P l⁻¹ and 0.68 mg N l⁻¹), phytoplankton cannot grow satisfactorily because of the short water retention time (0.6 day⁻¹). The shallowness of the pond together with the low phytoplankton biomass and the high concentrations of nutrients are discussed.     

Structural and grazing responses of zooplankton community to biomanipulation of some Dutch water bodies

Structure and grazing activities of crustacean zooplankton were compared in five lakes undergoing manipulation with several unmanipulated eutrophic (shallow) and mesotrophic (deep) lakes in The Netherlands. The biomanipulated lakes had lesser number of species and their abundance, both of rotifers and crustaceans, and had much larger mean animal size (3–11 μg C ind.⁻¹) than in the unmanipulated eutrophic lakes (0.65 μG C ind.⁻¹). WhereasD. hyalina (=D. galeata) andD. cucullata generally co-occurred in the unmanipulated lakes, in the manipulated lakes bothD. hyalina and other large-bodied daphnids,D. magna,D. pulex (=D. pulicaria), were the important grazers. In the biomanipulated lakes an increase in the individual crustacean size and of zooplankton mass were reflected in a decrease in seston concentration, higher Secchi-disc depth and a marked decrease in the share in phytoplankton biovolume of cyanobacteria. Biomass relationship between seston (150 μm) and zooplankton indicated a Monod type relationship, with an initial part of the curve in which the zooplankton responds linearly to the seston increase up to aboutca. 2 mg C l⁻¹, followed by a saturation of zooplankton mass (0.39 mg C l⁻¹) at 3–4 mg C l⁻¹ seston, and an inhibitory effect on zooplankton mass at seston levels>4 mg C l⁻¹. This latter is related to predominance in the seston of cyanobacteria. In the biomanipulated lakes, the zooplankton grazing rates often exceeded 100% d⁻¹, during the spring, and food levels generally dropped to <0.5 mg C l⁻¹. The computed specific clearance rate (SCR) of zooplankton of 1.9 l mg⁻¹ Zoop C is well within the range of SCR values (1.7–2.2 l mg⁻¹ Zoop C) from deep and mesotrophic waters, but about an order of magnitude higher than in the eutrophic lakes, with the food levels 10-fold higher. For 25% d⁻¹ clearance of lake seston between 35 and 60 ind. l⁻¹ are needed in the biomanipulated lakes against 1200–1300 ind. l⁻¹ in eutrophic lakes. Similarly, about 10 to 15 times more crustacean grazers are required to eliminate the daily primary production in the eutrophic lakes than in the biomanipulated lakes. These numbers are inversely related to the differences in animal size. The corresponding biomass values of zooplankton needed to clear the daily primary production in the eutrophic waters were 0.1–0.2 mg C l⁻¹ in the biomanipulated lakes, but about 0.45 mg C l⁻¹ in the unmanipulated eutrophic waters. Only if the water was kept persistently clear by zooplankton was there a balanced seston budget between the inputvia primary production and elimination by zooplankton. Mostly, however, the input exceeded the assimilatory removal by zooplankton, such that the estimated seston loss could be attributed to sedimentation and mineralization.     

Oligotrophication as a result of planktivorous fish removal with rotenone in the small,eutrophic, Lake Mosvatn,Norway

In September 1987 the shallow, eutrophic, Lake Mosvatn was treated with rotenone to eliminate planktivorous fish (mainly whitefish,Coregonus lavaretus, L.), and the effects were studied. The first summer after treatment the zooplankton community changed markedly from rotifer dominance and few grazers, to a community with few rotifers and many grazers. Accordingly there was a fivefold increase in the biomass ofDaphnia galeata. Adult females ofD. galeata approximately doubled in weight. The decrease in rotifer biomass was probably mainly due to a loss of food by competition with the daphnids. The phytoplankton community was also markedly affected. Prior to treatment Secchi depth was 1.7 m and Chl-a 23μg l⁻¹ in the summer. After treatment there was an increase in the proportion of small and gelatinous algae and the mean chlorophyll concentration fell to 7μg Chl-a l⁻¹. Secchi depth increased to>2.3 m (bottom-sight most of the season). After the treatment there were also fewer cyanobacterial blooms. This seems to be related to oligotrophication caused indirectly by increased grazing by the zooplankton. Total nutrient concentrations were affected. Prior to treatment the mean summer concentration of total phosphate was 44μg P l⁻¹. This decreased to 29μg P l⁻¹ in the first summer and 23μg P l⁻¹ the second summer after the treatment. Total nitrogen decreased from 0.68 mg N l⁻¹ before treatment to 0.32 mg N l⁻¹ the first summer after the treatment. The phosphate loading was not reduced, therefor it can be concluded that the fish removal provided a biomanipulation which caused the more oligotrophic conditions.     

The first biomanipulation conference: a synthesis

At the First Biomanipulation Conference held in Amsterdam (8–11 August 1989), studies presented considered mainly trophic interactions in lakes, enclosures and laboratory systems. Studies on the interactions between phytoplankton and zooplankton emphasized the edibility of the phytoplankton in relation to the zooplankton size structure and the trophic state. Most lake experiments involved 50–100% reduction in fish standing stock or alternatively heavy stocking with piscivorous fish. The most dramatic effects of biomanipulation were found in shallow, eutrophic lakes which exhibited radical changes in ecosystem structure because of changes in light climate and consequently, luxuriant development of macrophytes. There was still much controversy about the top-down effects in relation to trophic state, especially those concerning the role of fish and zooplankton in the development and succession of phytoplankton. Many experiments showed indirect effects within the food web, emphasizing the importance of feedbacks and the complexity of the food web rather than the simplicity of the food chain. The stabilizing effects of refugia for zooplankton and fish on the ecosystem were stressed. Shallow lakes responded generally more rapidly to biomanipulation and this was most successfully accomplished when TP concentration was<50μg l⁻¹, even though in a few cases at 10–20 fold higher TP concentrations (mostly PO₄-P lakes) the results achieved could be maintained for two or more summers. For a guaranteed success of the measures an almost complete removal of fish appeared to be indispensible; moreover in many cases removal of benthivorous fish appeared to be even more important than that of planktivorous fish.     

Lake Muzahi,Rwanda: limnological features and phytoplankton production

Lake Muhazi, a small lake of Rwanda (East Africa) was studied from 1986 to 1990. A dramatic decrease of the catch of Oreochromis niloticus (350 T y⁻¹ in the fifties vs 30 T y⁻¹ in 1982) suggested a loss of productivity or overfishing. In the same period, other ecological changes occurred: the submerged macrophytes regressed and there was a decrease in Secchi depth (0.65 m in 1987 vs 1.5 m in the fifties). Compared to other lakes of the same area, the plankton production seemed low. The results of the present study characterize lake Muhazi as a shallow lake with a rather unstable diurnal stratification and with slight differences in mixing regime between its eastern, deepest part and its western, shallowest part. Secchi disk depth does not vary seasonally to a large extent. The water has a rather high mineral content (conductivity of about 500 μS cm⁻¹ at 25 °C) and low concentrations of dissolved N and P, except in the hypolimnion, where NH _inf4 ^sup+ -N can be high. Two species, Microcystis aeruginosa and Ceratium hirundinella, account for most of the phytoplankton biomass, which is about 50–80 mg chlorophyll a m⁻² in the euphotic zone, usually with little seasonal variation. Daily gross production estimates amount to about 6 to 9.5 g O₂ m⁻² d⁻¹ with a significant difference between the two parts of the lake. Data on C:N and C:P ratio in the phytoplankton suggest that some N deficiency might occur in the eastern part. Moreover, the Zm:Zc ratio could also lead to rather low net production rates (0.21–0.25 d⁻¹ for a mixed layer of 4 m) In conclusion, the primary production of lake Muhazi is medium for African lakes and the hypothesis that decreased planktonic production could account for a reduced fish production should be discarded. Whereas the present yield of the fishery is only 20 kg ha⁻¹ y⁻¹, the yield estimated from primary production ranges between 46 and 64 kg ha⁻¹ y⁻¹. This could be reached through proper management. Finally, some hypotheses are given to explain the ecological changes which occurred in the lake.     

The effects of planktivorous fishes on the plankton community in a eutrophic lake

Stocking silver carp, a phytoplankton feeder, and bighead carp, an omnivorous plankton feeder, into an eutrophic lake at high densities caused a dramatic change in the lake ecosystem.Microcystis, which had been dominant in summer and a main food of the fishes decreased markedly, and green algae smaller than 10 μm then became dominant. Consequently, chlorophyll-a per unit area decreased slightly, while the rate of production was higher than that in the previous years. As the total density of the fishes increased (from 0.09 to 0.11 fish m⁻²), the growth of silver carp was retarded, while that of bighead carp increased.Microcystis, was unable to become dominant due to increased grazing pressure by the fishes, and small green algae became dominant. The lake conditions thus became more favourable for zooplankters which selectively consumed small green algae, and accordingly, the production of zooplankton rose. Bighead carp consumed more animal food, which they assimilate at a higher rate than plant food, and grew better in spite of the fact that the fish density increased. The feeding rate of silver carp was greatly reduced because the green algae were too small to ingest, and the fish therefore grew poorly. Results of a computer simulation of a model consisting of five compartments, representing the blue-green algae, green algae, zooplankton, silver and bighead carp, support the food-web change observed in the lake.     

Effects of planktivorous fish mass mortality on the plankton community of Lake Mendota,Wisconsin: implications for biomanipulation

Massive mortality of planktivorous fish had a dramatic impact on plankton community dynamics of Lake Mendota, Wisconsin, USA. After fish mortality, the largerDaphnia pulicaria replaced the smallerDaphnia galeata mendotae, resulting in greater grazing pressure on phytoplankton. This was accompanied by a much longer spring clear-water period and lower summer phytoplankton biomass compared to years before the fish mortality. Analysis of historical data (from the mid-1970's) showed that previous fluctuations in planktivorous abundance had similar effects onDaphnia abundance and species composition, and on spring phytoplankton biomass. However, the mid-1970's fish fluctuations had no detectable effect on summer phytoplankton. Concentrations of phosphorus were much higher in the 1970's (spring P 80–135 μg 1⁻¹) than in the 1980's (spring P 19–36 μg 1⁻¹) and it is possible that high P concentrations may reduce trophic cascade effects on summer phytoplankton communities. This suggests that the success of biomanipulation programs may be dependent on lake nutrient status.     

Proposals for macrophyte restoration in eutrophic coastal lagoons

Based on the comparison of environmental requirements forRuppia cirrhosa andPotamogeton pectinatus growth, macrophyteversus phytoplankton biomass and production features, and differences in hydrological and nutrient balances between Tancada lagoon (where macrophytes form dense beds) and Enca?izada lagoon (with no macrophytes at all), several proposals for macrophyte restoration are presented. The highest photosynthetic efficiency ofR. cirrhosa takes place at high irradiance and it grows over a wide range of salinity.P. pectinatus is better adapted to lower light intensity and salinity thanRuppia. R. cirrhosa transplanted from Tancada to Enca?izada was successful in enclosures, where light availability increased (μ=0.013 cm⁻¹), but not in open waters where light extinction coefficient was 0.032 cm⁻¹. Phytoplankton biomass (0.11–2.15 g C m⁻²) is much lower than macrophyte biomass (16–200 g C m⁻²) in Tancada lagoon. However, phytoplankton production (165 g C m⁻² yr⁻¹ in Tancada, 480 g C m⁻² yr⁻¹ in Enca?izada) is the same order of magnitude as macrophyte production (244–467 g C m⁻² yr⁻¹). Turnover rates are 0.3–0.9 day⁻¹ for phytoplankton and 1.2–2.5 yr⁻¹ for macrophytes. Phytoplankton and inorganic particles are responsible for high turbidity of the water in Enca?izada lagoon. Phytoplankton blooms in Enca?izada lagoon are supported by high freshwater inflows from rice field drains from May to November. The Q_s (seawater discharge)/Q_F (Freshwater discharge) ratios are, respectively, 0.24 and 0.48, which denotes a higher seawater influence in Tancada than in Enca?izada lagoon. Decreasing freshwater inputs to Enca?izada lagoon both in May and November thus allowing greater inputs of sea water, is proposed as the most effective way to restore this eutrophic coastal lagoon. The objective being to reduce nutrient loadings to the lagoon and phytoplankton in order to favour macrophyte re-colonization.     

Trophic status of Tilitso,a high altitude Himalayan lake

The trophic status and water quality of Lake Tilitso (4920 m above sea level) in a high altitude region in central Nepal were surveyed in September, 1984. The lake is rather large with a maximum depth of 95 m and a surface area of 10.2 km². The lake water was turbid due to glacier silt and the euphotic layer was only 5 m deep. The nutrient concentration was very low with total phosphorus concentration 1–6 μg l⁻¹, and DTN concentration 0.10–0.22 mg l⁻¹. The phytoplankton biomass and chlorophyll-a concentration were also low. Primary production was estimated to be about 12 mg C m⁻² d⁻¹. The concentrations of particulate matter and most cations and bacterial number were higher in the epilimnion than in the hypolimnion. The trophic status of this lake was estimated as ultraoligotrophic.     

First attempt to apply whole-lake food-web manipulation on a large scale in The Netherlands

Lake Breukeleveen (180 ha, mean depth 1.45 m), a compartment of the eutrophic Loosdrecht lakes system, was selected to study the effects of whole-lake foodweb manipulation on a large scale. In Lake Loosdrecht (dominated by filamentous cyanobacteria), due to water management measures taken from 1970–1984 (sewerage systems, dephosphorization) the external P load has been reduced from 1.2 g m⁻² y⁻¹ to 0.35 g m⁻² y⁻¹. The water transparency (Secchi-depthca. 30 cm), however, has not improved. The aim of the food-web manipulation in Lake Breukeleveen was not only to improve the light climate of the lake, but also to study if the successfull effects observed in small lakes (a few ha) can be upscaled. In March 1989 the standing crop of planktivorous and bentivorous fish populations was reduced by intensive fishery, fromca. 150 kg ha⁻¹ toca. 57 kg ha⁻¹. The lake was made unaccessible to fish migrating from the other lakes and it was stocked with large-sized daphnids and 0⁺ pike. However, water transparency did not increase in the following summer and autumn 1989, which is in contrast with great improvement in the light conditions previously observed in smaller lakes. The main explanations for the negative outcome in Lake Breukeleveen are: 1) the rapid increase of the planktivorous fish biomass and carnivorous cladocerans, predating on the zooplankton community; 2) suppression of the large daphnids by the high concentrations of filamentous cyanobacteria; 3) high turbidity of the lake due to resuspension of bottom material induced by wind, unlike in smaller lakes, and thus inability of submerged macrophytes to develop and to stabilize the ecosystem.     

Macrophyte-related shifts in the nitrogen and phosphorus contents of the different trophic levels in a biomanipulated shallow lake

Lake Zwemlust, a small highly eutrophic lake, was biomanipulated without reducing the external nutrient loading, and the effects were studied for four years. In this paper we pay special attention to the shifts in relative distribution of nitrogen and phosphorus in the different trophic levels and to the changes in growth limitation of the autotrophs.Despite of the high external nutrient loads to the lake (ca 2.4 g P m^–2 y^–1 and 9.6 g N m^–2 y^–1), the effects of biomanipulation on the lake ecosystem were pronounced. Before biomanipulation no submerged vegetation was present in the lake and P and N were stored in the phytoplankton (44% N, 47% P), fish (33% N, 9% P) and in dissolved forms (23% N, 44% P). P and N contents in sediments were not determined. In the spring and summer following the biomanipulation (1987), zooplankton grazing controlled the phytoplankton biomass and about 90% of N and P were present in dissolved form in the water. From 1988 onwards submerged macrophyte stands continue to thrive, reducing the ammonium and nitrate concentrations in the water below detection levels. In July 1989 storage of N and P in the macrophytes reached 86% and 80%, respectively. Elodea nuttallii (Planchon) St.John, the dominant species in 1988 and 1989, acted as sink both for N and P during spring and early summer, withdrawing up to ca 60% of its N and P content from the sediment. At the end of the year only part of the N and P from the decayed macrophytes (ca 30% of N and 60% of P) was recovered in the water phase of the ecosystem (chiefly in dissolved forms). The rest remained in the sediment, although some N may have been released from the lake by denitrification.In summer 1990 only 30% of the N and P was found in the macrophytes (dominant species Ceratophyllum demersum L.), while ca 30% of N and P was again stored in phytoplankton and fish.     

The influence of plant-associated filter feeders on phytoplankton biomass: a mesocosm study

Low phytoplankton biomass usually occurs in the presence of submerged macrophytes, possibly because submerged macrophytes enhance top-down control of phytoplankton by offering a refuge for efficient grazers like Daphnia against fish predation. However, other field studies also suggest that submerged macrophytes suppress phytoplankton in the absence of Daphnia. In order to investigate these mechanisms further, we conducted an outdoor mesocosm experiment to study the effect of submerged macrophytes (Elodea nuttallii) on phytoplankton and zooplankton biomass. The experiment combined four nutrient addition levels (0, 10, 100, and 1000 μg P l⁻¹; N/P ratio: 16) with three macrophyte levels (no macrophytes, artificial macrophytes, and real macrophytes). We inoculated the tanks with species-rich inocula of phytoplankton and zooplankton but excluded fish or macro-invertebrates. Probably due to the lack of predators in the mesocosms, potential grazing rates of pelagic zooplankton (estimated from zooplankton biomass) did not differ between the macrophyte treatment combinations. Compared to the treatment combinations without macrophytes, lower phytoplankton biomass occurred in the treatment combinations with real macrophytes at all the nutrient addition levels and in those with artificial macrophytes at all the nutrient levels except the highest. Significantly, higher abundances of plant-associated filter feeders (Simocephalus vetulus and Ceriodaphnia spp.) occurred in the treatment combinations with real and artificial macrophytes. The estimated potential grazing rate of these plant-associated filter feeders indicated that these filter feeders could be responsible for the lower phytoplankton biomass in the presence of real and artificial macrophytes. Our results suggest that the plant-associated filter feeders may be significant grazers in vegetated shallow lakes.     

Food web manipulation in Lake Zwemlust: Positive and negative effects during the first two years

The hypertrophic Lake Zwemlust, a small water body used as a swimming pool, was characterized by algal blooms in summer, reducing the Secchi disk transparency to less than 0.3 m. Since in The Netherlands a Secchi disk transparency of 1 m is obligatory for swimming waters, corrective measures were called for to improve the light climate of the lake. In March, 1987, as an experiment, the lake was drained by pumping out the water to facilitate fish elimination. Planktivorous and benthivorous fish species, which were predominant, were removed by seine- and electro-fishing. After the lake had refilled by seepage it was restocked by a new simple fish community comprising pike (Esox lucius) and rudd (Scardinius erythrophthalmus) only. Stacks of willow twigs (Salix) and macrophytes (roots ofNuphar lutea and seedlings ofChara globularis) were introduced into the lake as spawning grounds and refuges for the pike against cannibalism and as shelter for the zooplankton. The effects of this food web manipulation on the light climate, phytoplankton, zooplankton, fish, macrophytes, macrofauna and on the nutrient concentrations were monitored during 1987 and 1988. In summer 1987, despite of high nutrient concentrations, the phytoplankton density was low, due to control by zooplankton, causing a Secchi disk transparency of 2.5 m, the maximum depth. Chlorophyll-a concentrations were low (<5 g Chl.l^–1), blooms of cyanobacteria did not occur and a shift from rotifers to cladocerans took place. In 1988, however, also some negative effects were noticed. Macrophytes and filamentous green algae reached a much higher biomass (50–60% cover of the lake bottom) than in 1987; some species, growing through the entire water column, interfered with the lake's recreational use. Associated with the macro-vegetation and possibly with the absence of larger cyprinids, the diet of which also comprises snails, a large scale development of the snail population, among themLymnaea peregra var.ovata took place. This species is known to act as an intermediate host of the bird-parasitizing trematodeTrichobilharzia ocellata, the cercariae of which cause an itching sensation at the spot of penetration of the human skin, accompanied by rash (schistosome dermatitis or swimmers' itch); in July, 1988, about 40% of the bathers complained about this itching. A positive effect of the macrophytes and filamentous green algae was the high uptake of nitrogen, resulting in a low nitrogen concentration in the lake and growth limitation of the phytoplankton population by nitrogen in the summer of 1988. In 1988 the cladocerans were abundant in April only; and unlike in 1987, in the summer of 1988 there was a shift from cladocerans to rotifers. Therefore, only in early spring (April) zooplankton grazing controlled phytoplankton growth and in summer nitrogen limitation was the major controlling factor, keeping chlorophyll-a concentrations low.     

Can macrophytes be useful in biomanipulation of lakes? The Lake Zwemlust example

Lake Zwemlust (area 1.5 ha, Z_m 1.5 m) has been the object of an extensive limnological study since its biomanipulation involving removal of planktivorous fish (bream) in March 1987 and emptying of the lake. In the subsequent summer period of 1987 the Secchi depth increased to the lake bottom (2.5 m), compared withca 30 cm in the earlier summers. The reaction of submerged macrophytes to improving under-water light climate was rapid. In summer 1987, besides the introducedChara globularis, 5 species of submerged macrophytes occurred and colonized 10% of the lake area. In 1988 and 1989 only quantitative changes were observed; new species did not appear, but the area colonized by macrophytes increased by 7 and 10 times, respectively.Elodea nuttallii was dominant among the macrophytes andMougeotia sp. among the filamentous green algae. Their abundance, contributed to transient N-limination of phytoplankton causing a persistent clear water phase in 1988 and 1989, unlike in 1987 when zooplankton grazing contributed chiefly to the water clarity. Laboratory bioassays on macrophytes confirmed nitrogen limitation.     

Biomanipulation of Lago di Candia (Northern Italy): a three-year experience of aquatic macrophyte management

A long-term experimental aquatic plant management programme has been in progress since 1985 in the eutrophic Lago di Candia (Northern Italy). 7.7 ha of water chestnut (Trapa natans L.) were harvested in 1986, 8.2 ha in 1987 and 11.0 ha in 1988, from August to October, removing 334, 290 and 418 tons of fresh plant material, respectively, from the lake. This annual harvesting of about 50% of the total water chestnut cover resulted in the removal from the lake of about 70 kg yr⁻¹ of phosphorus, a significant portion of the external annual loading originating mainly from run-off and precipitation. In addition, up to 38t yr⁻¹ of organic matter and 1t yr⁻¹ of nitrogen were removed. First results of the effect of the harvesting on Secchi-disc transparency, oxygen, total phosphorus, phytoplankton and zooplankton are discussed also in connection with a study of fish manipulation (Giussaniet al., 1990).     

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.     

Impact of submerged macrophytes on fish-zooplanl phytoplankton interactions: large-scale enclosure experiments in a shallow eutrophic lake

1. The impact of changes in submerged macrophyte abundance on fish-zooplankton-phytoplankton interactions was studied in eighteen large-scale (100 m²) enclosures in a shallow eutrophic take. The submerged macrophytes comprised Potamategon pectinatus L., P. pusillus L. and Callitriche hermaphroditica L. while the fish fry stock comprised three-spined sticklebacks, Gasterosteus acuteatus L., and roach, Rutilus rutilus L. 2. In the absence of macrophytes zooplankton biomass was low and dominated by cyclopoid copepods regardless of fish density, while the phytoplankton biovolume was high (up to 38 mm³1) and dominated by small pennate diatoms and chlorococcales. When the lake volume infested by submerged macrophytes (PVI) exceeded 15–20% and the fish density was below a catch per unit effort (CPUE) of 10 (approx. 2 fry m^?2), planktonic cladoceran biomass was high and dominated by relatively large-sized specimens, while the phytoplankton biovolume was low and dominated by small fast-growing flagellates. At higher fish densities, zooplankton biomass and average biomass of cladocerans decreased and a shift to cyclopoids occurred, while phytoplankton biovolume increased markedly and became dominated by cyanophytes and dinoflagellates. 3. Stepwise multiple linear regressions on log-transformed data revealed that the biomass of Daphnia, Bosmina, Ceriodaphmia and Chydorus were all significantly positively related to PVI and negatively to the abundance of fish or PVI x fish. The average individual biomass of cladocerans was negatively related to fish, but unrelated to PVI. Calculated zooplankton grazing pressure on phytoplankton was positively related to PVI and negatively to PVI x fish. Accordingly the phytoplankton biovolume was negatively related to PVI and to PVI x zooplankton biomass. Cyanophytes and chryptophytes (% of biomass) were positively and Chlorococcales and diatoms negatively related to PVI, while cyanophytes and Chlorococcales were negatively related to PVI x zooplankton biomass. In contrast diatoms and cryptophytes were positively related to the zooplankton biomass or PVI x zooplankton. 4. The results suggest that fish predation has less impact on the zooplankton community in the more structured environment of macrophyte beds, particularly when the PVI exceeds 15–20%. They further suggest that the refuge capacity of macrophytes decreases markedly with increasing fish density (in our study above approximately 10 CPUE). Provided that the density of planktivorous fish is not high, even small improvements in submerged macrophyte abundance may have a substantial positive impact on the zooplankton, leading to a lower phytoplankton biovolume and higher water transparency. However, at high fish densities the refuge effect seems low and no major zooplankton mediated effects of enhanced growth of macrophytes are to be expected.