SUMMARY1. 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: B 0.05, 0.05-0.1,). 2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floatingleaved 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 mg in class 1 to 1.5 mg 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 clado- ceran 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.

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SUMMARY1. This synthesis examines 35 long-term (5-35 years, mean: 16 years) lake re-oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 lg L )1 before loading reduction), subtropical to temperate (latitude: 28-65°), and lowland to upland (altitude: 0-481 m). Shallow northtemperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in-lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP was reached after 10-15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially. 3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in-lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables. 4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria. 5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100-150 lg L )1 . This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters. 6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity. 7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental cha...

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Phytoplankton dominance (as biomass) by heterocystous cyanobacteria, nonheterocystous cyanobacteria, and chlorophytes was studied along a trophic gradient (0.011–2.2 mg P∙L−1) by analyzing regularly collected semiquantitative data from 178 shallow Danish lakes (mean depth < 3 m) and quantitative data from 32 lakes. Heterocystous cyanobacteria were dominant at low total P (TP) (< 0.25 mg P∙L−1) and nonheterocystous cyanobacteria at intermediate TP (0.25–0.8 mg P∙L−1), while chlorophytes often were dominant at high TP (> 1 mg P∙L−1). In contrast with many earlier findings, heterocystous cyanobacteria were not dominant at low total N (TN):TP or low inorganic N concentrations; chlorophytes were dominant at extremely high pH, and the shift from cyanobacterial to chlorophyte dominance could not be explained by a change in the photic zone to mixing zone ratio. We suggest that chlorophyte dominance in hypertrophic shallow lakes is attributable to continuous input of nutrients and carbon from the sediment and external sources. This renders the fast-growing chlorophytes a superior competitor compared with the relatively slow-growing cyanobacteria, even when inorganic nutrient concentration is low and pH high. New predictive models relating phytoplankton dominance to TP in shallow lakes were developed, as former models failed to predict our observations satisfactorily.

Major efforts have been made worldwide to improve the ecological quality of shallow lakes by reducing external nutrient loading. These have often resulted in lower in-lake total phosphorus (TP) and decreased chlorophyll a levels in surface water, reduced phytoplankton biomass and higher Secchi depth. Internal loading delays recovery, but in north temperate lakes a new equilibrium with respect to TP often is reached after <10-15 years. In comparison, the response time to reduced nitrogen (N) loading is typically <5 years. Also increased top-down control may be important. Fish biomass often declines, and the percentage of piscivores, the zooplankton:phytoplankton biomass ratio, the contribution of Daphnia to zooplankton biomass and the cladoceran size all tend to increase. This holds for both small and relatively large lakes, for example, the largest lake in Denmark (40 km 2 ), shallow Lake Arresø, has responded relatively rapidly to a ca. 76% loading reduction arising from nutrient reduction and top-down control. Some lakes, however, have proven resistant to loading reductions. To accelerate recovery several physico-chemical and biological restoration methods have been developed for north temperate lakes and used with varying degrees of success. Biological measures, such as selective removal of planktivorous fish, stocking of piscivorous fish and implantation or protection of submerged plants, often are cheap versus traditional physico-chemical methods and are therefore attractive. However, their long-term effectiveness is uncertain. It is argued that additional measures beyond loading reduction are less cost-efficient and often not needed in very large lakes. J. P. Jensen was deceased.

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