Lake responses to reduced nutrient loading – an analysis of contemporary long-term data from 35 case studies

1. 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 μg L^?1 before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate 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 μg 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 changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.     

Response of a shallow Mediterranean lake to nutrient diversion: does it follow similar patterns as in northern shallow lakes?

1. In view of the paucity of data on the response of warm shallow lakes to reductions in nutrient loading, this paper presents a long‐term limnological data set to document changes in the food‐web of a shallow Mediterranean lake (Lake Albufera, Valencia, Spain) that has experienced reductions in phosphorus (P) (77%) and nitrogen (N) (24%) loading following sewage diversion. 2. Nine years after sewage diversion, P concentration in the lake was reduced by 30% but remained high (TP = 0.34 mg L^?1), although the mean water retention time in the lake was only 0.1 years. Nitrate concentrations did not significantly change, probably because the lake continued to receive untreated effluents from ricefields. 3. Chlorophyll a concentration was reduced by half (annual mean of 180 μg L^?1). Cyanobacteria abundance remained high but its composition changed towards smaller species, both filamentous and chroococcal forms. 4. Cladocera abundance increased and reached peaks twice a year (December to March and July to September). After nutrient reduction, short‐term clear‐water phases (up to 5 weeks) occurred during February to March in several years, concomitant with annual flushing of the lake and lower fish densities. The abundance of Cladocera in winter contrasted with the spring peaks observed in northern restored shallow lakes. The zooplankton to phytoplankton biomass ratio remained lower than in northern temperate shallow lakes, probably because of fish predation on zooplankton. 5. Improvement of the water quality of Lake Albufera remained insufficient to counteract littoral reed regression or improve underwater light allowing submerged plants re‐colonise the lake. 6. Sewage diversion from Lake Albufera impacted the food web through the plankton, but higher trophic levels, such as fish and waterfowl, were affected to a lesser degree. Although the fish species present in the lake are mainly omnivorous, long‐term data on commercial fish captures indicated that fish communities changed in response to nutrient level and trophic structure as has been observed in restored shallow lakes at northern latitudes. 7. Phosphorus concentrations produced similar phytoplankton biomass in Lake Albufera as in more northern shallow lakes with abundant planktivorous fish and small zooplankton. However, in Lake Albufera, high average concentrations were maintained throughout the year. Overall, results suggest that nutrient control may be a greater priority in eutrophicated warm shallow lakes than in similar lakes at higher latitudes.     

Phosphorus decrease and climate variability: mediators of synchrony in phytoplankton changes among European peri-alpine lakes

1. In an attempt to discern long‐term regional patterns in phytoplankton community composition we analysed data from five deep peri‐alpine lake basins that have been included in long‐term monitoring programmes since the beginning of the 1970s. Local management measures have led to synchronous declines in phosphorus concentrations by more than 50% in all four lakes. Their trophic state now ranges from mesotrophic to oligotrophic. 2. No coherence in phytoplankton biomass was observed among lakes, or any significant decrease in response to phosphorus (P)‐reduction (oligotrophication), except in Lakes Constance and Walen. 3. Multivariate analyses identified long‐term changes in phytoplankton composition, which occurred coherently in all lakes despite the differing absolute phosphorus concentrations. 4. In all lakes, the phytoplankton species benefiting from oligotrophication included mixotrophic species and/or species indicative of oligo‐mesotrophic conditions. 5. A major change in community composition occurred in all lakes at the end of the 1980s. During this period there was also a major shift in climatic conditions during winter and early spring, suggesting an impact of climatic factors. 6. Our results provide evidence that synchronous long‐term changes in geographically separated phytoplankton communities may occur even when overall biomass changes are not synchronous.     

Eutrophication,recovery and temperature in Lake Mjøsa: detecting trends with monitoring data and sediment records

1. How climate warming may interact with other pressures on aquatic ecosystems is an important issue for research and management. We combined lake monitoring data with a palaeolimnological study to explore the combined effects of eutrophication and subsequent oligotrophication with a long‐term temperature increase in epilimnetic waters. Our goals were (i) to evaluate how well sediment‐based reconstructions reflect the instrumental observations, (ii) to use the palaeo‐record to characterise a reference state for the lake and (iii) to explore whether data from the sediment record can aid in separating the effects of nutrient load and temperature in a large and deep lake. 2. Lake Mjøsa is a large and deep lake in south‐eastern Norway. Eutrophication symptoms peaked in the 1970s, which led to extensive measures to reduce the phosphorus load. A monitoring programme has run continuously from 1972. Monitoring has documented a marked decrease in phosphorus load and algal biomass and also revealed an increase in epilimnetic temperature and extended summer stratification. 3. Records of algal pigments and diatoms were extracted from sediment cores taken from 236 m depth. The pigment record documented dramatic changes in lake production consistent with the monitoring record. The diatom record reflected well the eutrophication history of the lake and also demonstrated that the assemblage of the recent recovery stage differs from that of the pre‐eutrophication period. 4. Ordination of diatom assemblages over time constrained by proxies for nutrient load and temperature indicated that the diatom assemblage correlated with both factors, which together accounted for 60% of the variation in diatom composition. No interaction was detected between these factors. The results suggest that the diatom assemblage has responded to varying nutrient loads as well as to changes in temperature and/or factors that correlate with temperature. 5. Reconstructions of algal biomass and total phosphorus content mirrored known changes through the monitoring period, although the absolute phosphorus estimates were too high relative to the instrumental record. The sediment record from Lake Mjøsa provides a baseline for lake production in terms of algal pigments and organic contents, and for the diatom assemblage composition in a pristine stage.     

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.     

Consequences of Fish Kills for Long-Term Trophic Structure in Shallow Lakes: Implications for Theory and Restoration

Fish kills are a common occurrence in shallow, eutrophic lakes, but their ecological consequences, especially in the long term, are poorly understood. We studied the decadal-scale response of two UK shallow lakes to fish kills using a palaeolimnological approach. Eutrophic and turbid Barningham Lake experienced two fish kills in the early 1950s and late 1970s with fish recovering after both events, whereas less eutrophic, macrophyte-dominated Wolterton Lake experienced one kill event in the early 1970s from which fish failed to recover. Our palaeo-data show fish-driven trophic cascade effects across all trophic levels (covering benthic and pelagic species) in both lakes regardless of pre-kill macrophyte coverage and trophic status. In turbid Barningham Lake, similar to long-term studies of biomanipulations in other eutrophic lakes, effects at the macrophyte level are shown to be temporary after the first kill (c. 20 years) and non-existent after the second kill. In plant-dominated Wolterton Lake, permanent fish disappearance failed to halt a long-term pattern of macrophyte community change (for example, loss of charophytes and over-wintering macrophyte species) symptomatic of eutrophication. Important implications for theory and restoration ecology arise from our study. Firstly, our data support ideas of slow eutrophication-driven change in shallow lakes where perturbations are not necessary prerequisites for macrophyte loss. Secondly, the study emphasises a key need for lake managers to reduce external nutrient loading if sustainable and long-term lake restoration is to be achieved. Our research highlights the enormous potential of multi-indicator palaeolimnology and alludes to an important need to consider potential fish kill signatures when interpreting results.     

Combining contemporary ecology and palaeolimnology to understand shallow lake ecosystem change

1. Palaeolimnology and contemporary ecology are complementary disciplines but are rarely combined. By reviewing the literature and using a case study, we show how linking the timescales of these approaches affords a powerful means of understanding ecological change in shallow lakes. 2. Recently, palaeolimnology has largely been pre‐occupied with developing transfer functions which use surface sediment‐lake environment datasets to reconstruct a single environmental variable. Such models ignore complex controls over biological structure and can be prone to considerable error in prediction. Furthermore, by reducing species assemblage data to a series of numbers, transfer functions neglect valuable ecological information on species’ seasonality, habitat structure and food web interactions. These elements can be readily extracted from palaeolimnological data with the interpretive assistance of contemporary experiments and surveys. For example, for one shallow lake, we show how it is possible to infer long‐term seasonality change from plant macrofossil and fossil diatom data with the assistance of seasonal datasets on macrophyte and algal dynamics. 3. On the other hand, theories on shallow lake functioning have generally been developed from short‐term (<1–15 years) studies as opposed to palaeo‐data that cover the actual timescales (decades–centuries) of shallow lake response to stressors such as eutrophication and climate change. Palaeolimnological techniques can track long‐term dynamics in lakes whilst smoothing out short‐term variability and thus provide a unique and important means of not only developing ecological theories, but of testing them. 4. By combining contemporary ecology and palaeolimnology, it should be possible to gain a fuller understanding of changing ecological patterns and processes in shallow lakes on multiple timescales.     

Combining limnological and palaeolimnological data to disentangle the effects of nutrient pollution and climate change on lake ecosystems: problems and potential

1. As long‐term observational lake records continue to lengthen, the historical overlap with lake sediment records grows, providing increasing opportunities for placing the contemporary ecological status of lakes in a temporal perspective. 2. Comparisons between long‐term data sets and sediment records, however, require lake sediments to be accurately dated and for sediment accumulation rates to be sufficiently rapid to allow precise matching with observational data. 3. The critical role of the sediment record in this context is its value in tracking the changing impact of human activity on a lake from a pre‐disturbance reference through to the present day. 4. Here, we use data from a range of lakes across Europe presented as case studies in this Special Section. The seven sites considered all possess both long‐term observational records and high‐quality sediment records. Our objective is to assess whether recent climate change is having an impact on their trophic status and in particular whether that impact can be disentangled from the changes associated with nutrient pollution. 5. The palaeo‐data show clear evidence for the beginning of nutrient pollution varying from the mid‐nineteenth century at Loch Leven to the early and middle twentieth century at other sites. The monitoring data show different degrees of recovery when judged against the palaeo‐reference. 6. The reason for limited recovery is attributed to continuing high nutrient concentrations related to an increase in diffuse nutrient loading or to internal P recycling, but there is some evidence that climate change may be playing a role in offsetting recovery at some sites. If this is the case, then lake ecosystems suffering from eutrophication may not necessarily return to their pre‐eutrophication reference status despite the measures that have been taken to reduce external nutrient loading. 7. The extent to which future warming might further limit such recovery can be evaluated only by continued monitoring combined with the use of palaeo‐records that set the pre‐eutrophication reference.     

Shallow lake restoration by nutrient loading reduction—some recent findings and challenges ahead

Shallow lakes respond to nutrient loading reductions. Major findings in a recent multi-lake comparison of data from lakes with long time series revealed: that a new state of equilibrium was typically reached for phosphorus (P) after 10–15 years and for nitrogen (N) after <5–10 years; that the in-lake Total N:Total P and inorganic N:P ratios increased; that the phytoplankton and fish biomass often decreased; that the percentage of piscivores often increased as did the zooplankton:phytoplankton biomass ratio, the contribution of Daphnia to zooplankton biomass, and cladoceran size. This indicates that enhanced resource and predator control often interact during recovery from eutrophication. So far, focus has been directed at reducing external loading of P. However, one experimental study and cross-system analyses of data from many lakes in north temperate lakes indicate that nitrogen may play a more significant role for abundance and species richness of submerged plants than usually anticipated when total phosphorus is moderate high. According to the alternative states hypothesis we should expect ecological resistance to nutrient loading reduction and P hysteresis. We present results suggesting that the two alternative states are less stable than originally anticipated. How global warming affects the water clarity of shallow lakes is debatable. We suggest that water clarity often will decrease due to either enhanced growth of phytoplankton or, if submerged macrophytes are stimulated, by reduced capacity of these plants to maintain clear-water conditions. The latter is supported by a cross-system comparison of lakes in Florida and Denmark. The proportion of small fish might increase and we might see higher aggregation of fish within the vegetation (leading to loss of zooplankton refuges), more annual fish cohorts, more omnivorous feeding by fish and less specialist piscivory. Moreover, lakes may have prolonged growth seasons with a higher risk of long-lasting algal blooms and at places dense floating plant communities. The effects of global warming need to be taken into consideration by lake managers when setting future targets for critical loading, as these may well have to be adjusted in the future. Finally, we highlight some of the future challenges we see in lake restoration research.     

Restoration of shallow lakes by nutrient control and biomanipulation—the successful strategy varies with lake size and climate

Major efforts have been made world-wide 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 km²), 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. Although fewer data are available on tropical lakes these seem to respond to external loading reductions, an example being Lake Paranoá, Brazil (38 km²). However, differences in biological interactions between cold temperate versus warm temperate-subtropical-tropical lakes make transfer of existing biological restoration methods to warm lakes difficult. Warm lakes often have prolonged growth seasons with a higher risk of long-lasting algal blooms and dense floating plant communities, smaller fish, higher aggregation of fish in vegetation (leading to loss of zooplankton refuge), more annual fish cohorts, more omnivorous feeding by fish and less specialist piscivory. The trophic structures of warm lakes vary markedly, depending on precipitation, continental or coastal regions locations, lake age and temperature. Unfortunately, little is known about trophic dynamics and the role of fish in warm lakes. Since many warm lakes suffer from eutrophication, new insights are needed into trophic interactions and potential lake restoration methods, especially since eutrophication is expected to increase in the future owing to economic development and global warming.     

Major changes in trophic dynamics in large, deep sub-alpine Lake Maggiore from 1940s to 2002: a high resolution comparative palaeo-neolimnological study

1. We describe the changes in trophic dynamics in Lake Maggiore from c. 1943 to 2002 using subfossil cladoceran data from a high resolution sediment record, long‐term contemporary data series and historical information. During this period the lake went through a eutrophication phase until 1980 followed by oligotrophication. 2. During the eutrophication period a major increase occurred in the abundance of Chydorus sphaericus, the proportion of planktonic cladocerans and total abundance of cladocerans in the sediment. Since 1980 the abundance declined again and subfossil Eubosmina mucro length and contemporary Daphnia body length increased, most probably as a result of higher abundance of invertebrate predators. 3. Changes in the fish stock composition caused by the introduction of exotic fish during the pre‐eutrophication period and a complete ban on fishing because of Dichloro‐diphenil‐ethanes (DDTs) pollution of the lake (during oligotrophication) could also be detected in the community assemblage and size structure of the sediment zooplankton. 4. We found good correspondence between trophic changes inferred from cladoceran subfossils (community composition, size and predation pressure) and contemporary data, suggesting that sediment samples can be used to infer past development in trophic dynamics, including predation by fish and pelagic invertebrates in lakes with scarce neolimnological data. 5. Furthermore, by combining palaeolimnological cladoceran data rarely obtained from contemporary samples (e.g. benthic and plant‐associated cladocerans, mucro length of bosminids) with contemporary data of organisms poorly represented in the sediment record (e.g. remains of Bythotrephes and fishes) a more complete understanding of changes in trophic dynamics was obtained. 6. The detection in the sediments of meteorological events whose effects on zooplankton had been recorded in the long‐term studies also provided evidence that eutrophication tends to override climate signals. 7. We conclude that a combined palaeo‐neolimnological approach can be a powerful tool for elucidating past changes in the trophic dynamics of lakes and the interaction with climate induced changes, not least when high resolution sediment records are available.     

Hydrological regulation drives regime shifts: evidence from paleolimnology and ecosystem modeling of a large shallow Chinese lake

Quantitative evidence of sudden shifts in ecological structure and function in large shallow lakes is rare, even though they provide essential benefits to society. Such ‘regime shifts’ can be driven by human activities which degrade ecological stability including water level control (WLC) and nutrient loading. Interactions between WLC and nutrient loading on the long‐term dynamics of shallow lake ecosystems are, however, often overlooked and largely underestimated, which has hampered the effectiveness of lake management. Here, we focus on a large shallow lake (Lake Chaohu) located in one of the most densely populated areas in China, the lower Yangtze River floodplain, which has undergone both WLC and increasing nutrient loading over the last several decades. We applied a novel methodology that combines consistent evidence from both paleolimnological records and ecosystem modeling to overcome the hurdle of data insufficiency and to unravel the drivers and underlying mechanisms in ecosystem dynamics. We identified the occurrence of two regime shifts: one in 1963, characterized by the abrupt disappearance of submerged vegetation, and another around 1980, with strong algal blooms being observed thereafter. Using model scenarios, we further disentangled the roles of WLC and nutrient loading, showing that the 1963 shift was predominantly triggered by WLC, whereas the shift ca. 1980 was attributed to aggravated nutrient loading. Our analysis also shows interactions between these two stressors. Compared to the dynamics driven by nutrient loading alone, WLC reduced the critical P loading and resulted in earlier disappearance of submerged vegetation and emergence of algal blooms by approximately 26 and 10 years, respectively. Overall, our study reveals the significant role of hydrological regulation in driving shallow lake ecosystem dynamics, and it highlights the urgency of using multi‐objective management criteria that includes ecological sustainability perspectives when implementing hydrological regulation for aquatic ecosystems around the globe.     

Using palaeolimnological and limnological data to reconstruct the recent history of European lake ecosystems: introduction

1. As future climate change is expected to have a major impact on freshwater lake ecosystems, it is important to assess the extent to which changes taking place in freshwater lakes can be attributed to the degree of climate change that has already taken place. 2. To address this issue, it is necessary to examine evidence spanning many decades by combining long‐term observational data sets and palaeolimnological records. 3. Here, we introduce a series of case studies of seven European lakes for which both long‐term data sets and sediment records are available. Most of the sites have been affected by eutrophication and are now in recovery. 4. The studies attempt to disentangle the effects of climate change from those of nutrient pollution and conclude that nutrient pollution is still the dominant factor controlling the trophic state of lakes. 5. At most sites, however, there is also evidence of climate influence related in some cases to natural variability in the climate system, and in others to the trend to higher temperatures over recent decades attributed to anthropogenic warming. 6. More generally and despite some problems, the studies indicate the value of combining limnological and palaeolimnological records in reconstructing lake history and in disentangling the changing role of different pressures on lake ecosystems.     

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.     

Long‐term changes in hypoxia and soluble reactive phosphorus in the hypolimnion of a large temperate lake: consequences of a climate regime shift

The (Lower) Lake of Zurich provides an ideal system for studying the long‐term impact of environmental change on deep‐water hypoxia because of its sensitivity to climatic forcing, its history of eutrophication and subsequent oligotrophication, and the quality and length of its data set. Based on 39 years (1972–2010) of measured profiles of temperature, oxygen concentration and phosphorus (P) concentration, the potentially confounding effects of oligotrophication and climatic forcing on the occurrence and extent of deep‐water hypoxia in the lake were investigated. The time‐series of Nürnberg's hypoxic factor (HF) for the lake can be divided into three distinct segments: (i) a segment of consistently low HF from 1972 to the late‐1980s climate regime shift (CRS); (ii) a transitional segment between the late‐1980s CRS and approximately 2000 within which the HF was highly variable; and (iii) a segment of consistently high HF thereafter. The increase in hypoxia during the study period was not a consequence of a change in trophic status, as the lake underwent oligotrophication as a result of reduced external P loading during this time. Instead, wavelet analysis suggests that changes in the lake's mixing regime, initiated by the late‐1980s CRS, ultimately led to a delayed but abrupt decrease in the deep‐water oxygen concentration, resulting in a general expansion of the hypoxic zone in autumn. Even after detrending to remove long‐term effects, the concentration of soluble reactive P in the bottom water of the lake was highly correlated with various measures of hypoxia, providing quantitative evidence supporting the probable effect of hypoxia on internal P loading. Such climate‐induced, ecosystem‐scale changes, which may result in undesirable effects such as a decline in water quality and a reduction in coldwater fish habitats, provide further evidence for the vulnerability of large temperate lakes to predicted increases in global air temperature.     

Weedbeds and big bugs: the importance of scale in detecting the influence of nutrients and predation on macroinvertebrates in plant‐dominated shallow lakes

1. The scale of investigations influences the interpretation of results. Here, we investigate the influence of fish and nutrients on biotic communities in shallow lakes, using studies at two different scales: (i) within‐lake experimental manipulation and (ii) comparative, among‐lake relationships. 2. At both scales, fish predation had an overriding influence on macroinvertebrates; fish reduced macroinvertebrate biomass and altered community composition. Prey selection appeared to be size based. Fish influenced zooplankton abundance and light penetration through the water column also, but there was no indication that fish caused increased resuspension of sediment. 3. There were effects of nutrients at both scales, but these effects differed with the scale of the investigation. Nutrients increased phytoplankton and periphyton at the within‐lake scale, and were associated with increased periphyton at the among‐lake scale. No significant effect of nutrients on macroinvertebrates was observed at the within‐lake scale. However, at the among‐lake scale, nutrients positively influenced the biomass and density of macroinvertebrates, and ameliorated the effect of fish on macroinvertebrates. 4. Increased prey availability at higher nutrient concentrations would be expected to cause changes in the fish community. However, at the among‐lake scale, differences were not apparent in fish biomass among lakes with different nutrient conditions, suggesting that stochastic events influence the fish community in these small and relatively isolated shallow lakes. 5. The intensity of predation by fish significantly influences macroinvertebrate community structure of shallow lakes, but nutrients also play a role. The scale of investigation influences the ability to detect the influence of nutrients on the different components of shallow lake communities, particularly for longer lived organisms such as macroinvertebrates, where the response takes longer to manifest.     

Lake‐type‐specific seasonal patterns of nutrient limitation in German lakes,with target nitrogen and phosphorus concentrations for good ecological status

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Ecological interactions in a shallow sand-pit lake (Lake Créteil,Parisian Basin,France): a modelling approach

A large data set (n = 154) of phytoplankton production and biomass in relation to physico-chemical environmental factors was collected from 1979 to 1986 in a recently created sand-pit lake (Paris suburbs). These data are well suited to interpret the oligotrophication observed along the 8 years period, characterized by a regular decrease in chlorophyll (from 16 to 4 µg l^-1 as annual averages).A model describing the ecological functioning of the lake has been established. Biological processes related to phyto-, bacterio- and zooplankton as well as sediment-water interactions, are described within several submodels. Most of the parameters involved were determined by in situ measurements in this or similar environments The model provides a good simulation of observed data and confirms that the reduction of nutrient loading, resulting from the diversion — in 1981 — of a sewer previously discharging into the lake, was responsible for the oligotrophication of the system. The model allows to explore the response of planktonic compartments accross a gradient of nutrient loading. The role of hydrology is also tested. The systematic run of the model with and without zooplankton leads to a better understanding of top-down control.     

Is the island biogeography model a poor predictor of biodiversity patterns in shallow lakes?

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