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...
Dominance by cyanobacteria hampers human use of lakes and reservoirs worldwide. Previous studies indicate that excessive nutrient loading and warmer conditions promote dominance by cyanobacteria, but evidence from global scale field data has so far been scarce. Our analysis, based on a study of 143 lakes along a latitudinal transect ranging from subarctic Europe to southern South America, shows that although warmer climates do not result in higher overall phytoplankton biomass, the percentage of the total phytoplankton biovolume attributable to cyanobacteria increases steeply with temperature. Our results also reveal that the percent cyanobacteria is greater in lakes with high rates of light absorption. This points to a positive feedback because restriction of light availability is often a consequence of high phytoplankton biovolume, which in turn may be driven by nutrient loading. Our results indicate a synergistic effect of nutrients and climate. The implications are that in a future warmer climate, nutrient concentrations may have to be reduced substantially from present values in many lakes if cyanobacterial dominance is to be controlled.
10.1007/s10750-012-1182-1Contact CEH NORA team at noraceh@ceh.ac.ukThe NERC and CEH trademarks and logos ('the Trademarks') are registered trademarks of NERC in the UK and other countries, and may not be used without the prior written consent of the Trademark owner. form, nor will it be during the first three months after its submission to Hydrobiologia."Corresponding author: Erik Jeppesen (ej@dmu.dk)We dedicate this paper to the late Prof. Jürgen Benndorf, a true pioneer and mentor in lake and reservoir management oriented research, who inspired a number of us to initiate longterm comprehensive experimental ecological studies on lakes and reservoirs. AbstractFish play a key role in the trophic dynamics of lakes. With climate warming, complex changes in fish assemblage structure may be expected owing to direct effects of temperature and indirect effects operating through eutrophication, water level changes, stratification and salinisation. We reviewed published and new long-term (10-100 years) fish data series from 24 European lakes (area: 0.04-5648 km 2 ; mean depth: 1-177m; a north-south gradient from 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 4 Sweden to Spain). Along with an annual temperature increase of about 0.15-0.3 °C per decade profound changes have occurred in either fish assemblage composition, body size and/or age structure during recent decades and a shift towards higher dominance of eurythermal species.These shifts have occurred despite a reduction in nutrient loading in many of the lakes that should have benefited the larger-sized individuals and the fish species typically inhabiting cold-water, low-nutrient lakes. The cold-stenothermic Arctic charr has been particularly affected and its abundance has decreased in the majority of the lakes where its presence was recorded. The harvest of cool-stenothermal trout has decreased substantially in two southern lakes. Vendace, whitefish and smelt show a different response depending on lake depth and latitude. Perch has apparently been stimulated in the north, with stronger year classes in warm years, but its abundance has declined in the southern Lake Maggiore, Italy. Where introduced, roach seems to take advantage of the higher temperature after years of low population densities. Eurythermal species such as common bream, pike-perch and/or shad are apparently on the increase in several of the lakes. The response of fish to the warming has been surprisingly strong and fast in recent decades, making them ideal sentinels for detecting and documenting climate-induced modifications of freshwater ecosystems.
1. The presence of contiguous beds of submerged (Myriophyllum spicatum, Ceratophyllum demersum and Najas marina) and floating‐leaved (Trapa natans) vegetation in a north Italian lake allowed us to test the effect of the different host architecture on epiphytic algae and invertebrates and to predict the consequences for the lake of changes in the predominant vegetation. 2. Epiphyton development, measured as carbon, nitrogen, phosphorus, chlorophyll a (Chl a), phaeophytin and as algal and macroinvertebrate density, was significantly higher on submerged plants than on T. natans. The C : Chl a ratio, a proxy of the ratio of heterotrophs to autotrophs, was higher on the floating‐leaved plants. The elemental (C : N : P) and pigment (Chl a : phaeophytin) ratios were not significantly different between the two vegetation types. 3. The taxonomic composition of epiphytic algae and invertebrates was similar on the different plants. The more varied morphology of the floating‐leaved T. natans resulted in a higher diversity of epiphytic algae, however, but not of macroinvertebrates. 4. There was a significant inverse relationship between epiphyton biomass and the standing crop of the host plant, suggesting a key role for light and water exchange in epiphyton development. 5. Replacement of floating‐leaved by submerged plants would increase the total biomass of epiphytic algae and invertebrates.
SUMMARY1. 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 PO 4 -P and 0.6 mg L )1 NO 3 -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.
We analyzed data from 81 shallow European lakes, which were sampled with standardized methods, for combined effects of climatic, physical, and chemical features of food‐web interactions, with a specific focus on zooplankton biomass and community structure. multiple‐regression analysis showed that total phosphorus (TP) generally was the most important predictor of zooplankton biomass and community structure. Climate was the next most important predictor and acted mainly through its effect on pelagic zooplankton taxa. Benthic and plant‐associated taxa (typically almost half the total zooplankton biomass) were, however, affected mainly by macrophyte coverage. Neither climate nor TP affected the relation between small and large taxa, and we found only a weak trend with increasing TP of increasing mean crustacean body mass. Dividing the data set into three climate zones revealed a pronounced difference in response to lake productivity between cold lakes, with long periods of ice cover, and the two warmer lake types. These ÂÂice lakes differed from the others with respect to the effect of TP on chlorophyll a, the zooplankton : chlorophyll a ratio, the chlorophyll a :TP ratio, and the proportion of cyclopoids in the copepod community. Our data suggest that bottom‐up forces, such as nutrient concentration, are the most important predictors of zooplankton biomass. In addition, climate contributes significantly—possibly by affecting top‐down regulation by fish—and may interact with productivity in determining the zooplankton standing biomass and community composition. Hence, the present study suggests that food‐web dynamics are closely linked to climatic features.
Studies on shallow lakes from the north temperate zone show that they alternate between clear and turbid water states in response to control factors. However, the ecology of semiarid to arid shallow Mediterranean lakes is less explored. Hydrological effects (e.g. water level fluctuations, water residence time) on major ions and nutrient dynamics and processes, and ecology of submerged macrophytes appear to have a crucial role for food webs in shallow Mediterranean lakes. Nutrient control may be of greater priority in eutrophicated warm shallow lakes than in similar lakes at higher latitudes. This will be relevant for the implementation of the European Water Framework Directive, and conservation and management of these ecosystems. Strong trophic cascading effects of fish resulting from dominance of omnivorous and benthivorous fish species, whose diversity is usually high, together with frequent spawning and absence of efficient piscivores, seem to be the reason for the lack of large-bodied grazers that could control phytoplankton. However, such effects may vary within the region depending on fish distribution and community. These factors need elaboration in order to allow shallow lake ecologists and managers to develop better restoration strategies for eutrophicated shallow Mediterranean lakes. Consequently, modifications for the implementation of the European Water Framework Directive for determining ecological status in shallow Mediterranean lakes appear to be necessary. Furthermore, the implications of climate warming may be even more challenging than in high latitude lakes since shallow lakes in the Mediterranean region are among the most sensitive to extreme climate changes. There is an urgent need cooperation, development of large-scale research and information exchange to facilitate this and a web-based discussion list has been implemented.
1. Results are analysed from 11 experiments in which effects of fish addition and nutrient loading on shallow lakes were studied in mesocosms. The experiments, five in 1998, six in 1999, were carried out in six lakes, distributed from Finland to southern Spain, according to a standard protocol. 2. Effects of the treatments on 29 standard chemical, phytoplankton and zooplankton variables are examined to assess the relative importance of bottom-up (nutrient enrichment) and top-down (fish predation) effects. For each year, the experiments in different locations are treated as replicates in a meta-analysis. Results of individual experiments are then compared in terms of the patterns of significant influences of nutrient addition and fish predation with these overall results (the baseline), and between years in the same location. 3. The overall meta-analysis gave consistent results across the 2 years, with nutrient loading influencing all of the chemical variables, and on average 31% of primary producer and 39% of zooplankton variables. In contrast, fish influenced none of the chemical variables, 11% of the primary producer and 44% of the zooplankton variables. Nutrient effects on the system were thus about three times greater than fish effects, although fish effects were not inconsiderable. 4. The relative importance of nutrients and fish in individual experiments often differed between years at the same location and effects deviated to varying degrees from the baseline. These deviations were treated as measures of consistency (predictability) of conclusions in repeat experiments. Consistency increased southwards and this is interpreted as a consequence of more variable annual weather northwards. 5. The influence of nutrient loading was greater southwards and this was probably manifested through naturally greater annual macrophyte abundance in warmer locations in consequence of the longer plant growing-season. There was no trend in the relative importance of fish effects with latitude but this may partly be an artefact of the simple fish Correspondence: Brian Moss,
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