Restoration by biomanipulation in a small hypertrophic lake: first-year results

Biomanipulation was carried out in order to improve the water quality of the small hypertrophic Lake Zwemlust (1.5 ha; mean depth 1.5 m). In March 1987 the lake was drained to facilitate the elimination of fish. Fish populations were dominated by planktivorous and benthivorous species (total stock c. 1500 kg) and were collected by seine- and electro-fishing. The lake was subsequently re-stocked with 1500 northern pike fingerlings (Esox lucius L.) and a low density of adult rudd (Scardinius erythrophthalmus). The offspring of the rudd served as food for the predator pike. Stacks of Salix twigs, roots of Nuphar lutea and plantlets of Chara globularis were brought in as refuge and spawning grounds for the pike, as well as shelter for the zooplankton.The impact of this biomanipulation on the light penetration, phytoplankton density, macrophytes, zooplankton and fish communities and on nutrient concentrations was monitored from March 1987 onwards. This paper presents the results in the first year after biomanipulation.The abundance of phytoplankton in the first summer (1987) after this biomanipulation was very low, and consequently accompanied by increase of Secchi-disc transparency and drastic decline of chlorophyll a concentration.The submerged vegetation remained scarce, with only 5 % of the bottom covered by macrophytes at the end of the season.Zooplankters became more abundant and there was a shift from rotifers to cladocerans, comprised mainly of Daphnia and Bosmina species, the former including at least 3 species.The offspring of the stocked rudd was present in the lake from the end of August 1987. Only 19% of the stocked pike survived the first year.Bioassays and experiments with zooplankton community grazing showed that the grazing pressure imposed by the zooplankton community was able to keep chlorophyll a concentrations and algal abundance to low levels, even in the presence of very high concentrations of inorganic N and P. The total nutrient level increased after biomanipulation, probably due to increased release from the sediment by bioturbation, the biomass of chironomids being high.At the end of 1987 Lake Zwemlust was still in an unstable stage. A new fish population dominated by piscivores, intended to control the planktivorous and benthivorous fish, and the submerged macrophytes did not yet stabilize.     

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: a review of biological control measures in eutrophic waters and the potential for Murray cod Maccullochella peelii peelii to promote water quality in temperate Australia

Biomanipulation is a method of controlling algal blooms in eutrophic freshwater ecosystems. The most common approach has been to enhance herbivores through a reduction of planktivorous fish and introduction of piscivorous fish. The method was originally intended to reduce grazing pressure on zooplankton, thereby increasing grazing pressure on phytoplankton to increase water clarity and promote the growth of aquatic macrophytes. Biomanipulation has received considerable attention since it was proposed in 1975 where innovative approaches and explanations of the processes have been developed. Although many successful biomanipulation exercises have been conducted internationally, it has received comparatively little attention in the Southern Hemisphere and has not been trialled in the southern temperate climate of South Australia. This is a review to speculate upon the criteria for and against the application of biomanipulation in southern temperate Australia using the native species Murray cod (Maccullochella peelii peelii) and to suggest future research.     

Whole-lake food-web manipulation as a means to study community interactions in a small ecosystem

Whole-lake food-web manipulation was carried out in the hypertrophic Lake Zwemlust (The Netherlands), with the aim of studying the effects on the lake's trophic status and to gain an insight into complex interactions among lake communities. Before manipulation this small (1.5 ha) and shallow (1.5 m) lake was characterized byMicrocystis blooms in summer and high chlorophyll-a concentrations were common (ca. 250 μg 1⁻¹). In March 1987 the planktivorous and benthivorous fish species in the lake were completely removed (ca. 1000 kg ha⁻¹), a new simple fish community (pike and rudd) was introduced and artificial refuges were created. The effects of this manipulation on the light climate, nutrient concentrations, phytoplankton, zooplankton, fish, macrophytes, and macrofauna were monitored during 1987, 1988 and 1989. Community interactions were investigated in phytoplankton bioassays and zooplankton grazing experiments. After the manipulation, despite the still high P and N loads to the lake (ca. 2.2 g P m⁻² y⁻¹ andca. 5.3 g N m⁻² y⁻¹), the phytoplankton density was low (Chl-a<5μg l⁻¹), due to control by large-sized zooplankton in spring and N-limitation in summer and autumn. A marked increase in the abundance of macrophytes and filamentous green algae in 1988 and 1989, as well as N loss due to denitrification, contributed to the N limitation of the phytoplankton. Before manipulation no submerged macro-vegetation was present but in 1988, the second year after manipulation, about 50% of the lake bottom was covered by macrophytes increasing to 80% in 1989. This led to substantial accumulation of both N and P, namely 76% and 73% respectively of the total nutrients in the lake in particulate matter. Undesirable features of the increase in macrophytes were: 1) direct nuisance to swimmers; and, 2) the large scale development of snails, especiallyL. peregra, which may harbour the parasite causing ‘swimmers' itch’. But harvesting of only about 3% of the total macrophyte biomass from the swimmers' area, twice a year, reduced the nuisance for swimmers without adversely affecting the water clarity.     

The dynamics and role of limnetic zooplankton in Loosdrecht lakes (The Netherlands)

The paper summarizes the results of a ten-year (1981–1991) zooplankton research on the Lake Loosdrecht, a highly eutrophic lake. The main cause of the lake's eutrophication and deteriorating water quality was supply up to mid 1984 of water from the River Vecht. This supply was replaced by dephosphorized water from the Amsterdam-Rhine Canal in 1984. The effects of this and other restoration measures on the lake's ecosystem were studied. Despite a reduction in the external P-load from ca. 1.0 g P m^–2 y^–1 to ca. 0.35 g m^–2 y^–1 now, the filamentous prokaryotes, including cyanobacteria and Prochlorothrix, continue to dominate the phytoplankton.Among the crustacean plankton Bosmina spp, Chydorus sp. and three species of cyclopoid copepods and their nauplii are quite common. Though there was no major change in the composition of abundant species, Daphnia cucullata, which is the only daphnid in these lakes, became virtually extinct since 1989. Among about 20 genera and 40 species of rotifers the important ones are: Anuraeopsis fissa, Keratella cochlearis, Filinia longiseta and Polyarthra. The rotifers usually peak in mid-summer following the crustacean peak in spring. The mean annual densities of crustaceans decreased during 1988–1991. Whereas seston (< 150 µm) mean mass in the lake increased since 1983 by 20–60%, zooplankton (> 150 µm) mass decreased by 15–35%.The grazing by crustacean community, which was attributable mainly to Bosmina, had mean rates between 10 and 25% d^–1. Between 42 and 47% of the food ingested was assimilated. In spring and early summer when both rotifers and crustaceans have their maximal densities the clearance rates of the rotifers were much higher. Based on C/P ratios, the zooplankton (> 150 µm) mass contained 2.5 times more phosphorus than seston (< 150 µm) mass so that the zooplankton comprised 12.5 % of the total-P in total particulate matter in the open water, compared with only 4.5% of the total particulate C. The mean excretion rates of P by zooplankton varied narrowly between 1.5 and 1.8 µg P 1^– d^–1, which equalled between 14 and 28% d^–1 of the P needed for phytoplankton production.The lack of response to restoration measures cannot be ascribed to one single factor. Apparently, the external P-loading is still not low enough and internal P-loading, though low, may be still high enough to sustain high seston levels. Intensive predation by bream is perhaps more important than food quality (high concentrations of filamentous cyanobacteria) in depressing the development of large-bodied zooplankton grazers, e.g. Daphnia. This may also contribute to resistance of the lake's ecosystem to respond to rehabilitation measures.     

Trophic structure of nine Czech reservoirs regularly stocked with piscivorous fish

For nearly 20 years, most Czech reservoirs supplying drinking water have been under statutory protection which permitted reservoir managers to manipulate fish stocks in order to maintain a sustainable water quality. The most common biomanipulative measure adopted was stocking with piscivorous fish (mostly 5 cm fry) using an annual stocking level of approx. 25 000 fish per reservoir. Nine reservoirs were studied for signs of top-down food web effects, as predicted by the trophic cascade hypothesis based on levels of total phosphorus (TP), chlorophyll a (Chl a), zooplankton biomass (ZB) and zooplankton community structure. In all nine reservoirs, only small Daphnia species were recorded, such as D. galeata and D. cucullata. The proportion of large-bodied daphnids retained on a 0.71 mm sieve was less than 10% of the total crustacean biomass in all reservoirs. The relationship of Chl a level – TP, and of ZB – Chl a, was positive under enhanced piscivory and did not differ statistically from the relationships in other reservoirs with natural fish stocks. This implies that bottom-up forces remained stronger than top-down ones in the studied reservoirs, despite the stocking of piscivorous fish. The failure of this attempt at biomanipulation may be due to an insufficient stocking rate of predatory fish and/or inadequate data on the resident planktivorous fish levels.     

Composition,density and feeding of crustacean zooplankton community in a shallow,temperate lake (Lake Balaton,Hungary)

Composition density and filtering rates of crustacean zooplankton were studied in the open water and among the macrophytes of the oligo-mesotrophic part of Lake Balaton from 1981 to 1983. From the individual filtering rates of the different populations and the densities community grazing rates were derived.Copepoda made up 79–90% of crustacean plankton community in the open lake and 95–97% of it in littoral zone. Among them the nauplii dominated. At the end of summer 1982, when Anabaenopsis was in bloom, the filter-feeding species (Eudiaptomus, Daphnia) practically disappeared, being replaced by cyclopoids. Daphnia had the highest filtering rates followed by those of Eudiaptomus and Diaphanosoma. Among copepods, the filtering rates in decreasing order were: ovigerous > all adults > copepodites > nauplii. > . The filtering rates of the different species varied both seasonally as well as from year to year. In 1983, when the concentration of organic seston decreased, filtering rate increased compared with those in the earlier years. During the water bloom in 1982, the rates decreased by 70% on the average.The community grazing rate was very low (3% per day) in the open lake and among macrophytes, both in 1981 and 1982; also the share of crustacean zooplankton in grazing was very low. In 1983, together with the improving of water quality, the community grazing rate increased 4-fold. In 1981 and 1983 the rates were influenced by water temperature but in 1982 by seston concentration.     

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.     

Zooplankton and their relationships with water quality and fisheries

Zooplankton population structure in Rutland Water changed over the first five years of the reservoir's existence. Daphnia pulex, initially dominant was replaced by D. hyalina in late 1975 and since 1979 this latter cladoceran has coexisted for part of each year with Bosmina longirostris. Population fluctuations of all the main crustacean zooplankton — D. hyalina, Cyclops spp. and Diaptomus gracilis — were erratic in the first three years (1975–1977) but have since become more regular. The relationships of the zooplankton with their potential algal food supply and with their potential predators, the fish population, are discussed.     

Low Genotypic Diversity in a Daphnia Pulex Population in a Biomanipulated Lake: The Lack of Vertical and Seasonal Variability

Allozyme diversity of a population of D. pulex was studied over two seasons in a small lake (Gräfenhain, Saxony, Germany). In this eutrophic lake, accumulation of hydrogen sulfide during winter ice covers leads to an annual killing-off of cladocerans. Hence, the population must be re-established every year by hatchlings from sexual eggs. The absence of planktivorous fish due to strong biomanipulation ensures high abundances of Daphnia from June until September. Besides the investigation of seasonal changes of genotype diversity, vertical differences between individuals from the epilimnion and from a deep chlorophyll maximum (DCM) were studied. Specimens were classified by allozyme electrophoresis using six allozyme loci. Except for aldehyde oxidase (AO) all tested allozyme loci were monomorphic. Two alleles were found at the AO-locus in the overall study, allowing the population to be subdivided into three clonegroups (ss, fs, ff). The proportion of each clonegroup both in the epilimnion and in the DCM did not vary over the two seasons and no differences were found in the year-to-year comparison. Although two completely different habitats (epilimnion, DCM) were found to be used by D. pulex, clonal structure in both depth strata was similar at all times indicating no intraspecific segregation of the population. In spite of a non-permanent population with periodical sexual reproduction, a constant deviation from Hardy-Weinberg-equilibrium was detected in the overall study. This remarkable genetic structure is interpreted to be the result of both founder effects and a strong and long-lasting selection pressure caused by 15 years of stationary conditions due to intense biomanipulation.