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.     

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.     

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.     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L?l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: <0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,> 0.4 mg P L?1).
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating‐leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L?1. The Shannon–Wiener and the Hurlbert probability of inter‐specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish.
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti‐benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large‐bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high.
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10‐fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15‐fold from class 1 to 5 and submerged macrophytes disappeared from most lakes.
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.     

Total Mercury in Sediments,Macrophytes, and Fish from a Shallow Steppe Lake in Eastern Austria

During summer 2011, samples of sediment, macrophytes, and fish tissues from the shallow, slightly alkaline Lake Neusiedl, Austria, were evaluated for their total Hg content. This is the first report of Hg levels from this lake. Sediments displayed Hg contents between 0.025 and 0.113 μg g^?1 dw (dry weight), significantly correlating with the proportion of organic components pointing to a small anthropogenic impact on the lake's Hg content. Hg Levels in plants and fish were unexpectedly high: both investigated submerged plant species, Potamogeton pectinatus and Myriophyllum spicatum, showed mean values of 0.245±0.152 and 0.298±0.115 μg g^?1 dw, respectively. Biomagnification was evident when comparing muscle samples of the planktivorous fish species rudd Scardinus erythrophthalmus (n=10, mean=0.084 μg g^?1 ww (wet weight)) with the piscivorous perch Perca fluviatilis (n=21, mean=0.184 μg g^?1 ww) or pike‐perch Sander lucioperca (n=9, mean=0.205 μg g^?1 ww). Significantly lower values were found in the muscle of the piscivorous pike Esox lucius (n=25, mean=0.135 μg g^?1 ww), pointing to a specific Hg metabolism of this fish, presumably under the particular physicochemical properties of the lake. Hg Concentrations in fish could pose a risk to piscivorous birds in this protected wetland system.     

Reconstructing the past density of planktivorous fish and trophic structure from sedimentary zooplankton fossils: a surface sediment calibration data set from shallow lakes

1. As quantitative information on historical changes in fish community structure is difficult to obtain directly from fish remains in lake sediments, transfer function for planktivorous fish abundance has been developed based on zooplankton remains in surface sediment (upper 1 cm). The transfer function was derived using weighted average regression and calibration against contemporary data on planktivorous fish catch per unit effort (PF-CPUE) in multiple mesh size gill nets. Zooplankton remains were chosen because zooplankton community structure in lakes is highly sensitive to changes in fish predation pressure. The calibration data set consisted of thirty lakes differing in PF-CPUE (range 18–369 fish net^–1), epilimnion total phosphorus (range 0.025–1.28 mg P l^–1) and submerged macrophyte coverage (0–57%). 2. Correlation of log-transformed PF-CPUE, total phosphorus and submerged macrophyte coverage v the percentage abundance in the sediment of the dominant cladocerans and rotifers revealed that the typical pelagic species correlated most highly to PF-CPUE, while the littoral species correlated most highly to submerged macrophyte coverage. Consequently, only pelagic species were taken into consideration when establishing the fish transfer function. 3. Canonical correspondence analysis (CCA) revealed that the pelagic zooplankton assemblage was highly significantly related to PF-CPUE (axis 1), whereas the influence of total phosphorus and submerged macrophyte coverage was insignificant. Predicted PF-CPUE based on weighted average regression without (WA) and with (WA(tol)) downweighting of zooplankton species tolerance correlated significantly with the observed values (r² = 0.64 and 0.60 and RMSE = 0.54 and 0.56, respectively). A marginally better relationship (r² = 0.67) was obtained using WA maximum likelihood estimated optima and tolerance. 4. It is now possible, quantitatively, to reconstruct the historical development in planktivorous fish abundance based on zooplankton fossil records. As good relationships exist between contemporary PF-CPUE data and indicators such as the zooplankton/phytoplankton biomass ratio, Secchi depth and the maximum depth distribution of submerged macrophytes, it is now also possible to derive information on past changes in lake water quality and trophic structure. It will probably prove possible further to improve the transfer function by including other invertebrate remains, e.g. chironomids, Chaoborus, snails, etc., and its scope could be widened by including deeper lakes, more oligotrophic lakes, more acidic lakes and lakes with extensive submerged macrophyte coverage (in the latter case to enable use of the information in the fossil record on plant-associated cladocerans).     

Fish manipulation as a lake restoration tool in shallow,eutrophic temperate lakes 1: cross-analysis of three Danish case-studies

The use of fish manipulation as a tool for lake restoration in eutrophic lakes has been investigated since 1986 in three shallow, eutrophic Danish lakes. The lakes differ with respect to nutrient loading and nutrient levels (130–1000 μg P l⁻¹, 1–6 mg N l⁻¹). A 50% removal of planktivorous fish in the less eutrophic cyanobacteria-diatom dominated Lake V?ng caused marked changes in lower trophic levels, phosphorus concentration and transparency. Only minor changes occurred after a 78% removal of planktivorous fish in eutrophic cyanobacteria dominated Frederiksborg Castle Lake. In the hypertrophic, green algae dominated Lake S?byg?rd a low recruitment of all fish species and a 16% removal of fish biomass created substantial changes in trophic structure, but no decrease in phosphorus concentration. The different response pattern is interpreted as (1) a difference in density and persistence of bloomforming cyanobacteria caused by between-lake variations in nutrient levels and probably also mixing- and flushing rates, (2) a difference in specific loss rates through sedimentation of the algal community prevaling after the fish manipulation, (3) a decreased impact of planktivorous fish with increasing mean depth and (4) a lake specific difference in ability to create a self-increasing reduction in the phosphorus level in the lake water. This in turn seems related to the phosphorus loading.     

Responses to food web manipulation in a shallow waterfowl lake

The effects of fish stock reduction have been studies in 3 Dutch lakes (Lake Zwemlust, Lake Bleiswijkse Zoom and Lake Noorddiep) and 1 Danish lake (Lake Væng) during 4–5 years. A general response id described. The fish stock reduction led in general to a low fish stock, low chlorophyii-a, high transparency and high abunuance of macrophytes. Large Daphnia became abundant, but their density decreased, due to food limitation and predation by fish. The total nitrogen concentration became low due to N-uptake by macrophytes and enhanced denitrification. In Lake Bleiswijkse Zoom the water transparency deteriorated and the clear water state was not stable. The fish stock increased and the production of young fish in summer was high. lear water occurred only in spring. Large daphnids were absent in summer and the macrophytes decreased.In Lake Zwemlust, Lake Væng and Lake Noorddiep the water remained clear during the first five years. In summer of the sixth year (1992) transparency decreased in Lake Zwemlust (with high P-concentration of 1.0 mg P l^-1). Also in Lake Væng (with a low nutrient concentration of 0.15 mg P.^-1) a short term turbid stage (1.5 month) occurred in summer 1992 after a sudden collapse of the macrophytes. Deterioration of the water quality seems to start in summer and seems related to a collapse in macrophytes. At a low planktivorous fishstock (e.g. Lake Væng)thhe duration of the turbid state is shorter. than in presence of a high planktivorous fish biomass (e.g. Lake Zwemlust, and later years of Lake Bleiswijkse Zoom).     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

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.     

Phytoplankton and primary production in clear-vegetated,inorganic-turbid,and algal-turbid shallow lakes from the pampa plain (Argentina)

Shallow lakes often alternate between two possible states: one clear with submerged macrophytes, and another one turbid, dominated by phytoplankton. A third type of shallow lakes, the inorganic turbid, result from high contents of suspended inorganic material, and is characterized by low phytoplankton biomass and macrophytes absence. In our survey, the structure and photosynthetic properties (based on ¹⁴C method) of phytoplankton were related to environmental conditions in these three types of lakes in the Pampa Plain. The underwater light climate was characterized. Clear-vegetated lakes were more transparent (K _d 4.5–7.7 m⁻¹), had high DOC concentrations (>45 mg l⁻¹), low phytoplankton Chl a (1.6–2.7 μg l⁻¹) dominated by nanoflagellates. Phytoplankton productivity and photosynthetic efficiency (α ~ 0.03 mgC mgChla ⁻¹ h⁻¹ W⁻¹ m²) were relatively low. Inorganic-turbid lakes showed highest K _d values (59.8–61.4 m⁻¹), lowest phytoplankton densities (dominated by Bacillariophyta), and Chl a ranged from 14.6 to 18.3 μg l⁻¹. Phytoplankton-turbid lakes showed, in general, high K _d (4.9–58.5 m⁻¹) due to their high phytoplankton abundances. These lakes exhibited the highest Chl a values (14.2–125.7 μg l⁻¹), and the highest productivities and efficiencies (maximum 0.56 mgC mgChla ⁻¹ h⁻¹ W⁻¹ m²). Autotrophic picoplankton abundance, dominated by ficocianine-rich picocyanobacteria, differed among the shallow lakes independently of their type (0.086 × 10⁵–41.7 × 10⁵ cells ml⁻¹). This article provides a complete characterization of phytoplankton structure (all size fractions), and primary production of the three types of lakes from the Pampa Plain, one of the richest areas in shallow lakes from South America. Handling editor: J. Padisak     

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.     

Phosphorus dynamics at multiple time scales in the pelagic zone of a large shallow lake in Florida,USA

Phosphorus (P) dynamics in large shallow lakes are greatly influenced by physical processes such as wind-driven sediment resuspension, at times scales from hours to years. Results from long-term (30 year) research on Lake Okeechobee, Florida (area 1,730 km², mean depth 2.7 m) illustrate key features of these P dynamics. Variations in wind velocity result in changes in water column transparency, suspended solids, and total P (TP). In summer there are diurnal changes in TP associated with afternoon winds, and in winter, when strong winds occur for multiple days, monthly average TP remains high compared to summer. The magnitude of daily and seasonal TP changes can exceed 100 μg l⁻¹. Hurricanes and tropical storms also cause extreme changes in TP that are superimposed on seasonal dynamics. When a hurricane passed 80 km south of the lake in October 1999, mean pelagic TP increased from 88 to 222 μg l⁻¹. During large resuspension events, light attenuation is substantially increased, and this influences the biomass and spatial extent of submerged plants, as well as water column TP. In Lake Okeechobee, TP concentrations typically are ∼20 μg l⁻¹ when submerged plants are dense, and soluble reactive P concentrations are reduced below detection, perhaps by the periphyton and plant uptake and by precipitation with calcium at high pH. In contrast, TP exceeds 50 μg l⁻¹ when submerged plants and periphyton are absent due to prolonged deep water, and phytoplankton biomass and algal bloom frequency both are increased. In Lake Okeechobee and other large shallow lakes, complex models that explicitly consider wind-wave energy, hydrodynamics, and sediment resuspension, transport, and key biological processes are needed to accurately predict how lake water TP will respond to different management options.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

THE INFLUENCE OF SUBMERGED MACROPHYTES ON SEDIMENTARY DIATOM ASSEMBLAGES1

Submerged macrophytes are a central component of lake ecosystems; however, little is known regarding their long‐term response to environmental change. We have examined the potential of diatoms as indicators of past macrophyte biomass. We first sampled periphyton to determine whether habitat was a predictor of diatom assemblage. We then sampled 41 lakes in Quebec, Canada, to evaluate whether whole‐lake submerged macrophyte biomass (BiomEpiV) influenced surface sediment diatom assemblages. A multivariate regression tree (MRT) was used to construct a semiquantitative model to reconstruct past macrophyte biomass. We determined that periphytic diatom assemblages on macrophytes were significantly different from those on wood and rocks (ANOSIM R = 0.63, P < 0.01). A redundancy analysis (RDA) of the 41‐lake data set identified BiomEpiV as a significant (P < 0.05) variable in structuring sedimentary diatom assemblages. The MRT analysis classified the lakes into three groups. These groups were (A) high‐macrophyte, nutrient‐limited lakes (BiomEpiV ≥525 μg · L^?1; total phosphorus [TP] <35 μg · L^?1; 23 lakes); (B) low‐macrophyte, nutrient‐limited lakes (BiomEpiV <525 μg · L^?1; TP <35 μg · L^?1; 12 lakes); and (C) eutrophic lakes (TP ≥35 μg · L^?1; six lakes). A semiquantitative model correctly predicted the MRT group of the lake 71% of the time (P < 0.001). These results suggest that submerged macrophytes have a significant influence on diatom community structure and that sedimentary diatom assemblages can be used to infer past macrophyte abundance.     

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.     

Piscivore enhancement effects on food webs depend on planktivore body size and species composition in replicated whole lake experiments

Recent syntheses of trophic cascade and biomanipulation research have suggested that the effects of piscivores on planktivorous fish populations are reduced, when planktivores are capable of outgrowing predator gape limitation and in systems with complex food web interactions. These hypotheses, however, have not been tested in long-term, whole-lake, experiments where processes such as fish recruitment and compensatory food web responses may be important. We conducted a replicated whole-lake experiment to test for the effects of supplemental piscivore introductions on food webs of eutrophic lakes dominated by deep-bodied planktivores. Responses to piscivore enhancement were compared between lakes differing in food web structure due to the presence of omnivorous gizzard shad (Dorosoma cepedianum). A significant decrease in the relative abundance of juvenile planktivorous fish, and an increase in total benthic macroinvertebrate density was observed in lakes containing mainly bluegills (Lepomus machrochirus). In contrast, lakes containing gizzard shad exhibited no significant responses to piscivore manipulation. Our results support the hypothesis that food webs in lakes dominated by deep-bodied planktivorous fish species respond weakly to piscivore enhancement. In addition, our findings support the hypothesis that cascading trophic interactions are weaker in lake ecosystems with more complex food web interactions such as those containing gizzard shad.     

Plant community structure determines primary productivity in shallow,eutrophic lakes

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Effects of elevated turbidity on shallow lake fish communities

Synopsis We compared the fish communities of two shallow lakes in the lower Waikato River basin, North Island, New Zealand, to determine the effects of elevated suspended solids (SS) and collapse of submerged macrophytes. Lake Waahi was turbid (20–40 g m^-3 SS) and devoid of submerged macrophytes whereas Lake Whangape was clearer (5 g m^-3 SS) and dominated by submerged macrophytes. The lakes had similar fish species richness and had nine major species in common; representing eight families including Anguillidae, Retropinnidae, Galaxiidae, Eleotridae, Mugilidae, Ictaluridae, Poeciliidae, and Cyprinidae (two species). The only major fish that was absent from Lake Waahi was a lacustrine form of the common smelt, Retropinna retropinna, which disappeared after the lake became turbid in the late 1970s. CPUE, condition, and size of most species in Lake Waahi were similar to, or greater than, those in Lake Whangape. Lake Whangape clearly exceeded Lake Waahi only for CPUE of two species. Within Lake Whangape two species displayed significantly greater condition, and one species greater size, in a turbid arm of the lake than in the main basin. Apart from lacustrine Retropinna retropinna, the fish in these lakes appear well adapted to cope with, or to avoid, the direct toxic effects of suspended and settleable solids on sensitive early developmental stages. In Lake Waahi loss of cover and food provided by submerged macrophytes appears to have been compensated for by increased turbidity and an associated increase in the biomass of the mysid, Tenagomysis chiltoni (a major prey item).     

Responses of four submerged macrophytes to freshwater snail density (Radix swinhoei) under clear‐water conditions: A mesocosm study

Macrophytes play a key role in stabilizing clear‐water conditions in shallow freshwater ecosystems. Their populations are maintained by a balance between plant grazing and plant growth. As a freshwater snail commonly found in shallow lakes, Radix swinhoei can affect the growth of submerged macrophytes by removing epiphyton from the surface of aquatic plants and by grazing directly on macrophyte organs. Thus, we conducted a long‐term (11‐month) experiment to explore the effects of snail density on macrophytes with distinctive structures in an outdoor clear‐water mesocosm system (with relatively low total nitrogen (TN, 0.66 ± 0.27 mg/L) and total phosphorus (TP, 36 ± 20 μg/L) and a phytoplankton chlorophyll a (Chla) range of 14.8 ± 4.9 μg/L) based on two different snail densities (low and high) and four macrophyte species treatments (Myriophyllum spicatum, Potamogeton wrightii, P. crispus, and P. oxyphyllus). In the high‐density treatment, snail biomass and abundance (36.5 ± 16.5 g/m² and 169 ± 92 ind/m², respectively) were approximately twice that observed in the low‐density treatment, resulting in lower total and aboveground biomass and ramet number in the macrophytes. In addition, plant height and plant volume inhabited (PVI) showed species‐specific responses to snail densities, that is, the height of P. oxyphyllus and PVI of M. spicatum were both higher under low‐density treatment. Thus, compared with low‐density treatment, the inhibitory effects of long‐term high snail density on macrophytes by direct feeding may be greater than the positive effects resulting from epiphyton clearance when under clear‐water conditions with low epiphyton biomass. Thus, under clear‐water conditions, the growth and community composition of submerged macrophytes could be potentially modified by the manual addition of invertebrates (i.e., snails) to lakes if the inhibitory effects from predatory fish are minor.