State of the art in the functioning of shallow Mediterranean lakes: workshop conclusions

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 semi-arid 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 for data on the ecology of shallow lakes in the region. An appeal is made for international cooperation, development of large-scale research and information exchange to facilitate this and a web-based discussion list has been implemented.     

Climate change effects on nitrogen loading from cultivated catchments in Europe: implications for nitrogen retention, ecological state of lakes and adaptation

Climate change might have profound effects on the nitrogen (N) dynamics in the cultivated landscape as well as on N transport in streams and the eutrophication of lakes. N loading from land to streams is expected to increase in North European temperate lakes due to higher winter rainfall and changes in cropping patterns. Scenario (IPCC, A2) analyses using a number of models of various complexity for Danish streams and lakes suggest an increase in runoff and N transport on an annual basis (higher during winter and typically lower during summer) in streams, a slight increase in N concentrations in streams despite higher losses in riparian wetlands, higher absolute retention of N in lakes (but not as percentage of loading), but only minor changes in lake water concentrations. However, when taking into account also a predicted higher temperature there is a risk of higher frequency and abundance of potentially toxic cyanobacteria in lakes and they may stay longer during the season. Somewhat higher risk of loss of submerged macrophytes at increased N and phosphorus (P) loading and a shift to dominance of small-sized fish preying upon the key grazers on phytoplankton may also enhance the risk of lake shifts from clear to turbid in a warmer North European temperate climate. However, it must be emphasised that the prediction of N transport and thus effects is uncertain as the prediction of regional precipitation and changes in land-use is uncertain. By contrast, N loading is expected to decline in warm temperate and arid climates. However, in warm arid lakes much higher N concentrations are currently observed despite reduced external loading. This is due to increased evapotranspiration leading to higher nutrient concentrations in the remaining water, but may also reflect a low-oxygen induced reduction of nitrification. Therefore, the critical N as well as P loading for good ecological state in lakes likely has to be lower in a future warmer climate in both north temperate and Mediterranean lakes. To obtain this objective, adaptation measures are required. In both climate zones the obvious methods are to change agricultural practices for reducing the loss of nutrients to surface waters, to improve sewage treatment and to reduce the storm-water nutrient runoff. In north temperate zones adaptations may also include re-establishment of artificial and natural wetlands, introduction of riparian buffer zones and re-meandering of channelised streams, which may all have a large impact on, not least, the N loading of lakes. In the arid zone, also restrictions on human use of water are urgently needed, not least on the quantity of water used for irrigation purposes.     

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.     

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.     

The response of periphyton and submerged macrophytes to nitrogen and phosphorus loading in shallow warm lakes: a mesocosm experiment

1. Recent experimental and field studies on temperate shallow lakes indicate that nitrogen may play a greater role in their functioning than previously thought. Several studies document that abundance and richness of submerged macrophytes, both central in shallow lake ecology, may decrease with increasing nitrogen loading, especially at high phosphorus levels. However, the role of nitrogen in warm lakes with fluctuating water regimes remains to be described in detail. 2. The effect of increasing nitrate and phosphate concentrations on submerged macrophyte growth was examined in a 3‐month mesocosm experiment conducted in summer in a shallow freshwater lake on the north western coast of Turkey with a Mediterranean climate. Twenty four field mesocosms, open to the sediment and atmosphere, were stocked with Myriophyllum spicatum shoots and small cyprinid fish. Three nitrate loadings in combination with two phosphate loadings were applied in a fourfold replicated design. 3. Mean ± SD nutrient concentrations maintained throughout the experiment were 0.55 ± 0.17, 2.2 ± 0.97, 9.2 ± 5.45 mg L^?1 total nitrogen and 55 ± 19.2, 73 ± 22.9 μg L^?1 total phosphorus. Mean periphyton biomass increased with increasing nutrient concentrations and peaked at the highest nitrogen and phosphorus loadings, while the mean phytoplankton biomass remained relatively low in all treatments. 4. Percent volume inhabited (% PVI) by macrophytes throughout the experiment and total macrophyte biomass at the end of the experiment did not differ among treatments. In addition to stocked M. spicatum, Ceratophyllum demersum and Potamogeton crispus appeared in the majority of the mesocosms. The plants grew continuously up to 50% PVI throughout the experiment and remained resilient to shading provided by periphyton and phytoplankton. 5. The mean summer air temperature in 2007 was 2.2 °C higher than the average of the last 32 years, which resulted in a water level decrease of 0.3 m in the mesocosms over three months. This might have counteracted the shading of submerged macrophytes provided by phytoplankton and periphyton. The results of the experiment are consistent with observations of higher macrophyte resilience to nutrient loading in Mediterranean lakes compared with northern temperate lakes.     

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.     

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.     

Can warm climate‐related structure of littoral predator assemblies weaken the clear water state in shallow lakes?

Shallow lakes, the most abundant lake type in the world, are very sensitive to climatic changes. The structure and functioning of shallow lakes are greatly impacted by submerged plants, and these may be affected by climate warming in various, contrasting, ways. Following a space‐for‐time substitution approach, we aimed to analyse the role of aquatic (submerged and free‐floating) plants in shallow lakes under warm climates. We introduced artificial submerged and free‐floating plant beds in five comparable lakes located in the temperate zone (Denmark, 55–57 °N) and in the subtropical zone (Uruguay, 30–35 °S), with the aim to study the structure and dynamics of the main associated communities. Regardless of differences in environmental variables, such as area, water transparency and nutrient status, we found consistent patterns in littoral community dynamics and structure (i.e. densities and composition of fish, zooplankton, macroinvertebrates, and periphyton) within, but substantial differences between, the two regions. Subtropical fish communities within the macrophyte beds exhibited higher diversity, higher density, smaller size, lower relative abundance of potentially piscivores, and a preference for submerged plants, compared with otherwise similar temperate lakes. By contrast, macroinvertebrates and cladocerans had higher taxon richness and densities, and periphyton higher biomass, in the temperate lakes. Several indicators suggest that the fish predation pressure was much stronger among the plants in the subtropical lakes. The antipredator behaviour of cladocerans also differed significantly between climate zones. Submerged and free‐floating plants exerted different effects on the spatial distribution of the main communities, the effects differing between the climate zones. In the temperate lakes, submerged plants promoted trophic interactions with potentially positive cascading effects on water transparency, in contrast to the free‐floating plants, and in strong contrast to the findings in the subtropical lakes. The higher impact of fish may result in higher sensitivity of warm lakes to external changes (e.g. increase in nutrient loading or water level changes). The current process of warming, particularly in temperate lakes, may entail an increased sensitivity to eutrophication, and a threat to the high diversity, clear water state.     

Effects of temperature, salinity and fish in structuring the macroinvertebrate community in shallow lakes: implications for effects of climate change

Climate warming may lead to changes in the trophic structure and diversity of shallow lakes as a combined effect of increased temperature and salinity and likely increased strength of trophic interactions. We investigated the potential effects of temperature, salinity and fish on the plant-associated macroinvertebrate community by introducing artificial plants in eight comparable shallow brackish lakes located in two climatic regions of contrasting temperature: cold-temperate and Mediterranean. In both regions, lakes covered a salinity gradient from freshwater to oligohaline waters. We undertook day and night-time sampling of macroinvertebrates associated with the artificial plants and fish and free-swimming macroinvertebrate predators within artificial plants and in pelagic areas. Our results showed marked differences in the trophic structure between cold and warm shallow lakes. Plant-associated macroinvertebrates and free-swimming macroinvertebrate predators were more abundant and the communities richer in species in the cold compared to the warm climate, most probably as a result of differences in fish predation pressure. Submerged plants in warm brackish lakes did not seem to counteract the effect of fish predation on macroinvertebrates to the same extent as in temperate freshwater lakes, since small fish were abundant and tended to aggregate within the macrophytes. The richness and abundance of most plant-associated macroinvertebrate taxa decreased with salinity. Despite the lower densities of plant-associated macroinvertebrates in the Mediterranean lakes, periphyton biomass was lower than in cold temperate systems, a fact that was mainly attributed to grazing and disturbance by fish. Our results suggest that, if the current process of warming entails higher chances of shallow lakes becoming warmer and more saline, climatic change may result in a decrease in macroinvertebrate species richness and abundance in shallow lakes.     

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.     

Nutrient ratios, differential retention, and the effect on nutrient limitation in a deep oligotrophic lake

Nutrient ratios have been related to nutrient limitation of algal growth in lakes. Retention of nutrients in lakes, by sedimentation and by denitrification, reduces the nutrient concentrations in the water column, thereby enhancing nutrient limitation. Differential retention of nitrogen and phosphorus alters their ratios in lakes and thereby contributes to determine whether nitrogen or phosphorus limits algal growth. We examined the relationships between differential nutrient retention, nutrient ratios, and nutrient limitation in Lake Brunner, a deep oligotrophic lake. The observed retention of nitrogen (20%) and phosphorus (47%) agreed with predictions by empirical equations from literature. As a result of differential retention with a much larger proportion of phosphorus retained than that of nitrogen, the nitrogen:phosphorus ratio was higher in the lake (69) than in the inflows (46). While the mean ratio in the inflows suggested no or only moderate phosphorus limitation, the lake appeared to be severely phosphorus limited. Combining empirical equations from literature that predict nitrogen and phosphorus retention suggests that the nitrogen:phosphorus ratio is enhanced by greater retention of phosphorus compared to nitrogen only in deep lakes with relatively short residence times, such as Lake Brunner. In contrast, in most lakes differential retention is expected to result in lower nitrogen:phosphorus ratios.     

Substantial differences in littoral fish community structure and dynamics in subtropical and temperate shallow lakes

1. Fish play a key role in the functioning of temperate shallow lakes by affecting nutrient exchange among habitats as well as lake trophic structure and dynamics. These processes are, in turn, strongly influenced by the abundance of submerged macrophytes, because piscivorous fish are often abundant at high macrophyte density. Whether this applies to warmer climates as well is virtually unknown. 2. To compare fish community structure and dynamics in plant beds between subtropical and temperate shallow lakes we conducted experiments with artificial submerged and free‐floating plant beds in a set of 10 shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N), paired along a gradient of limnological characteristics. 3. The differences between regions were more pronounced than differences attributable to trophic state. The subtropical littoral fish communities were characterised by higher species richness, higher densities, higher biomass, higher trophic diversity (with predominance of omnivores and lack of true piscivores) and smaller body size than in the comparable temperate lakes. On average, fish densities were 93 ind. m⁻² (±10 SE) in the subtropical and 10 ind. m⁻² (±2 SE) in the temperate lakes. We found a twofold higher total fish biomass per unit of total phosphorus in the subtropical than in the temperate lakes, and as fish size is smaller in the former, the implication is that more energy reaches the littoral zone fish community of the warmer lakes. 4. Plant architecture affected the spatial distribution of fish within each climate zone. Thus, in the temperate zone fish exhibited higher densities among the artificial free‐floating plants while subtropical fish were denser in the artificial submerged plant beds. These patterns appeared in most lakes, regardless of water turbidity or trophic state. 5. The subtropical littoral fish communities resembled the fish communities typically occurring in temperate eutrophic and hypertrophic lakes. Our results add to the growing evidence that climate warming may lead to more complex and omnivory‐dominated food webs and higher density and dominance of smaller‐sized fish. This type of community structure may lead to a weakening of the trophic cascading effects commonly observed in temperate shallow lakes and a higher risk of eutrophication.     

Retention of nitrogen, phosphorus and silicon in a large semi-arid riverine lake system

Lakes and reservoirs (impoundments) are often viewed as a sink for nutrients within the river continuum. To date, most studies on nutrient retention within impoundments are derived from the temperate climate zones of Europe and North America, only consider one nutrient, and are often short-term (1–2 years). Here, we present a long-term (17 year) data set and nutrient (nitrogen, phosphorus and silica) budget for two connected semi-arid lakes (the Lower Lakes) at the terminus of the River Murray, Australia. Most of the filterable reactive phosphorus and nitrate entering the lakes were retained (77 and 92%, respectively). Total phosphorus (TP) was also strongly retained (55% of the annual TP load on average) and the annual TP retention rates could be predicted as a function of the areal hydraulic loading rate (annual lake outflow/lake surface area). On average, there was a slight net retention (7%) of the annual total nitrogen (TN) load but a slight net export (6% of the load) of organic N. TN retention as function of the areal hydraulic loading rate was lower than expected from existing models, possibly because of high nitrogen fixation rates in the Lower Lakes. Silica was retained (39%) at similar rates to those observed in previous studies. There was also a marked increase in the TN:TP and TN:Si ratios within the lake (TN:TP~30 and TN:Si~0.67) compared to those entering (TN:TP~15, TN:Si~0.45), as a consequence of the relatively low net retention of nitrogen.     

Climate-related differences in the dominance of submerged macrophytes in shallow lakes

It has been suggested that shallow lakes in warm climates have a higher probability of being turbid, rather than macrophyte dominated, compared with lakes in cooler climates, but little field evidence exists to evaluate this hypothesis. We analyzed data from 782 lake years in different climate zones in North America, South America, and Europe. We tested if systematic differences exist in the relationship between the abundance of submerged macrophytes and environmental factors such as lake depth and nutrient levels. In the pooled dataset the proportion of lakes with substantial submerged macrophyte coverage (> 30% of the lake area) decreased in a sigmoidal way with increasing total phosphorus (TP) concentration, falling most steeply between 0.05 and 0.2 mg L⁻¹. Substantial submerged macrophyte coverage was also rare in lakes with total nitrogen (TN) concentrations above 1–2 mg L⁻¹, except for lakes with very low TP concentrations where macrophytes remain abundant until higher TN concentrations. The deviance reduction of logistic regression models predicting macrophyte coverage from nutrients and water depth was generally low, and notably lowest in tropical and subtropical regions (Brazil, Uruguay, and Florida), suggesting that macrophyte coverage was strongly influenced by other factors. The maximum TP concentration allowing substantial submerged macrophyte coverage was clearly higher in cold regions with more frost days. This is in agreement with other studies which found a large influence of ice cover duration on shallow lakes' ecology through partial fish kills that may improve light conditions for submerged macrophytes by cascading effects on periphyton and phytoplankton. Our findings suggest that, in regions where climatic warming is projected to lead to fewer frost days, macrophyte cover will decrease unless the nutrient levels are lowered.     

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.     

Impacts of climate warming on lake fish community structure and potential effects on ecosystem function

Fish play a key role in the trophic dynamics of lakes, not least in shallow systems. With climate warming, complex changes in fish community structure may be expected owing to the direct and indirect effects of temperature, and indirect effects of eutrophication, water-level changes and salinisation on fish metabolism, biotic interactions and geographical distribution. We review published and new data supporting the hypotheses that, with a warming climate, there will be changes in: fish community structure (e.g. higher or lower richness depending on local conditions); life history traits (e.g. smaller body size, shorter life span, earlier and less synchronised reproduction); feeding mode (i.e. increased omnivory and herbivory); behaviour (i.e. stronger association with littoral areas and a greater proportion of benthivores); and winter survival. All these changes imply higher predation on zooplankton and macroinvertebrates with increasing temperatures, suggesting that the changes in the fish communities partly resemble, and may intensify, the effects triggered by eutrophication. Modulating factors identified in cold and temperate systems, such as the presence of submerged plants and winter ice cover, seem to be weaker or non-existent in warm(ing) lakes. Consequently, in the future lower nutrient thresholds may be needed to obtain clear-water conditions and good ecological status in the future in currently cold or temperate lakes. Although examples are still scarce and more research is needed, we foresee biomanipulation to be a less successful restoration tool in warm(ing) lakes without a strong reduction of the nutrient load.     

Subfossil Cladocera in relation to contemporary environmental variables in 54 Pan-European lakes

1. Changes in cladoceran subfossils in the surface sediments of 54 shallow lakes were studied along a European latitude gradient (36–68°N). Multivariate methods, such as regression trees and ordination, were applied to explore the relationships between cladoceran taxa distribution and contemporary environmental variables, with special focus on the impact of climate. 2. Multivariate regression tree analysis showed distinct differences in cladoceran community structure and lake characteristics along the latitude gradient, identifying three groups: (i) northern lakes characterised by low annual mean temperature, conductivity, nutrient concentrations and fish abundance, (ii) southern, macrophyte rich, warm water lakes with high conductivity and high fish abundance and (iii) Mid‐European lakes at intermediate latitudes with intermediate conductivities, trophic state and temperatures. 3. Large‐sized, pelagic species dominated a group of seven northern lakes with low conductivity, where acid‐tolerant species were also occasionally abundant. Small‐sized, benthic‐associated species dominated a group of five warm water lakes with high conductivity. Cladoceran communities generally showed low species‐specific preferences for habitat and environmental conditions in the Mid‐European group of lakes. Taxon richness was low in the southern‐most, high‐conductivity lakes as well as in the two northern‐most sub‐arctic lakes. 4. The proportion of cladoceran resting eggs relative to body shields was high in the northern lakes, and linearly (negatively) related to both temperature and Chl a, indicating that both cold climate (short growing season) and low food availability induce high ephippia production. 5. Latitude and, implicitly, temperature were strongly correlated with conductivity and nutrient concentrations, highlighting the difficulties of disentangling a direct climate signal from indirect effects of climate, such as changes in fish community structure and human‐related impacts, when a latitude gradient is used as a climate proxy. Future studies should focus on the interrelationships between latitude and gradients in nutrient concentration and conductivity.     

Sediment resuspension: rescue or downfall of a thermally stratified eutrophic lake?

The aim of this study was to estimate the effect of sediment resuspension, a common phenomenon in many lakes, on the phosphorus budget of a eutrophicated lake. We used two different approaches, mass balance calculations and spatially comprehensive resuspension measurements, to determine the level of phosphorus loading from which rehabilitation action is started in a dimictic north temperate lake. The effect of resuspension was assumed to be significant, since it often is a governing process for cycling of material in lakes. Internal loading was multifold to that of external loading as determined by the budget calculation. Spatially comprehensive sedimentation and resuspension measurements were necessary, since deep site versus spatially comprehensive measurements had a marked difference in their results. Resuspension of P slightly exceeded the internal loading assessed by budget calculations and thereby proved its significance as a governing in-lake process that influences P cycling strongly. The shallow areas were of importance, since most of the total P load originated from there. The fate of P after resuspension depends on the retention capacity of resuspended particles in addition to prevailing biological and physico-chemical conditions. Therefore, sediment resuspension can either strengthen or diminish internal nutrient load and the processes of the shallow zones are of importance.     

A model study on the role of wetland zones in lake eutrophication and restoration

Shallow lakes respond in different ways to changes in nutrient loading (nitrogen, phosphorus). These lakes may be in two different states: turbid, dominated by phytoplankton, and clear, dominated by submerged macrophytes. Both states are self-stabilizing; a shift from turbid to clear occurs at much lower nutrient loading than a shift in the opposite direction. These critical loading levels vary among lakes and are dependent on morphological, biological, and lake management factors. This paper focuses on the role of wetland zones. Several processes are important: transport and settling of suspended solids, denitrification, nutrient uptake by marsh vegetation (increasing nutrient retention), and improvement of habitat conditions for predatory fish. A conceptual model of a lake with surrounding reed marsh was made, including these relations. The lake-part of this model consists of an existing lake model named PCLake. The relative area of lake and marsh can be varied. Model calculations revealed that nutrient concentrations are lowered by the presence of a marsh area, and that the critical loading level for a shift to clear water is increased. This happens only if the mixing rate of the lake and marsh water is adequate. In general, the relative marsh area should be quite large in order to have a substantial effect. Export of nutrients can be enhanced by harvesting of reed vegetation. Optimal predatory fish stock contributes to water quality improvement, but only if combined with favourable loading and physical conditions. Within limits, the presence of a wetland zone around lakes may thus increase the ability of lakes to cope with nutrients and enhance restoration. Validation of the conclusions in real lakes is recommended, a task hampered by the fact that, in the Netherlands, many wetland zones have disappeared in the past.