Planktonic ciliate community structure in shallow lakes of lowland Western Europe

Temperate shallow meso- to eutrophic lakes can exist in one of two alternative states with contrasting foodwebs, referred to as the clear-water and the turbid state. We describe the planktonic ciliate communities of such lakes based on a survey of 66 northwestern European lakes. Ciliates were enumerated and identified to species level according to the quantitative protargol staining technique. Ciliate biomass was on average twice as high in the turbid than in the clear-water lakes. The ciliate communities were dominated by oligotrichs and protostomatids, and no differences in functional composition or α-diversity could be detected between turbid and clear-water lakes, although β-diversity tended to be higher in the latter. At the species level, however, community structure strongly differed between turbid and clear-water lakes, and several indicator species could be identified for the different lake categories. Variation partitioning showed that nutrient status did not explain ciliate community structure independent of the alternative states, while lake area was identified as an additional structuring factor for the ciliate communities. These results stress the importance of the ecosystem structure in shaping ciliate communities in temperate shallow lakes and suggest that nutrient status has little direct effect on ciliate community structure in such lakes.     

Interannual and between-site variability in the occurrence of clear water phases in two shallow Mediterranean lakes

It is widely accepted that clear water phases constitute a regular stage in the seasonal succession of plankton in dimictic lakes and reservoirs (i.e. PEG Model). The occurrence of such a phenomenon in Mediterranean shallow lakes is characterised by a marked interannual variability, which makes it difficult to establish reliable predictions on the dynamics and functioning of plankton in these ecosystems. In the present paper we analyse the factors influencing the occurrence of the clear water phases in the two shallow lakes of the Albufera of Adra, a coastal wetland region of south-eastern Spain: Lake Honda and Lake Nueva. Despite their geographical proximity, both lakes depicted large hydrological and limnological differences. Lake Honda is an epigenic and recharge lake that is strongly influenced by the hydrological conditions in its watershed, while Lake Nueva can be classified as a hypogenic and discharge lake and, as such, is less affected by the hydrological regime. In contrast, the morphometry, exposure and fetch of Lake Nueva make this ecosystem especially sensitive to wind forcing. Clear water phases in these shallow lakes were linked with periods of low thermal stability and the dominance of small-edible algae in the phytoplankton community, both of which allowed a Daphnia magna population to grow up and induce the algae collapse by grazing. In Lake Honda, those conditions were met during the spring of 2002 under the influence of intense rainfall-events, while in Lake Nueva the clear water phase was induced in the spring of 2003 by the occurrence of strong and frequent wind events. In both lakes, a relatively high water column thermal stability and the abundance of cyanobacteria early in the spring prevented the development of the Daphnia magna population and the occurrence of the clear water phase.     

Relationships among nitrogen and total phosphorus,algal biomass and zooplankton density in the central Amazonia lakes

The relationship among concentrations of total nitrogen (TN), total phosphorus (TP), algal biomass (Chl) and the density and size of individuals of the zooplankton community were studied for the dry season (November 1999–January 2000) at 20 lakes of the Central Amazonia. The study was conducted along a productivity gradient to identify the existence of resource or predator-dependent patterns on the primary producers of the trophic web. A strong positive relationship was observed between the log Chl and TN (r ² = 0.88, P = 0.000) and to log Chl and log TP (r ² = 0.85, P = 0.000) in a simple linear regression. However, when both variables were running together in a multiple regression, TN alone explained every variation of algal biomass (r ² = 0.89, P _TN = 0.022, P _TP = 0.233). The total density of the zooplankton showed a positive correlation with log Chl (r ² = 0.53, P = 0.000) and the large zooplankton (>0.5 mm) was found to be a more positive function of the phytoplankton (r ² = 0.65) than the density of the small ones (<0.5 mm, r ² = 0.44). Results show that complex food web interactions could be responsible for patterns in tropical systems. We contend that Chl variation in tropical lake systems is controlled by TN and TP, but the predictor power of the TN increase the fit of the model in analysis and can be use alone to access the variability in algae biomass to Amazonian tropical lakes. We also agree that the density of large zooplankton individuals is regulated by the biomass of primary producers. Hence we concluded that the resource-dependent hypothesis is supported in these systems. Handling editor: J. Padisak     

The trophic state of shallow lakes in The Netherlands

A study has been carried out on the development of an ecological assessment method for shallow lakes in The Netherlands. Analyses of eutrophication characteristics of 93 lakes with, in total, 127 sampling localities gave insight into some of the steering variables, such as total-P, total-N, chlorophyll-a and transparency. On the basis of phytoplankton species composition, three main groups of lakes could be distinguished. The first group, characterized by persistent filamentous Cyanobacteria, had the highest summer means of chlorophyll-a and total-P. The second group was characterized by non-persistent filamentous Cyanobacteria, occurring in low abundancies, and at lower chlorophyll-a and total-P concentrations. The third group of lakes was characterized by the absence or very low abundance of the filamentous Cyanobacteria. As a result of these analyses, criteria for the abatement of eutrophication in shallow lakes could be refined.     

Differential endozoochory of aquatic invertebrates by two duck species in shallow lakes

Animal vectors are essential for the movement of invertebrate resting eggs between water bodies. However, differences in habitat preferences and feeding behaviour between bird species may result in variations in the dispersal of invertebrates via these birds, even if the different bird species live in the same lake. To test such effects, faecal samples from Anas platyrhynchos (collected in autumn and spring) and Tadorna tadorna (collected in spring) were cultured in water at 20° C and 12 L: 12 D conditions in order to quantify the resting eggs which could be internally transported by these birds. One half of each faecal sample was initially cultured at a conductivity of 0.6 mS cm⁻¹ and the other half at 6 mS cm⁻¹. 1054 invertebrates hatched from a total of 60 faecal samples, including cladocerans, copepods, ostracods, rotifers and ciliates, with a wide variability among faeces. Autumn yielded a low proportion of samples with hatchlings (12.5%) compared to spring (90%). Significant differences were observed between birds, but not between conductivity treatments. Thus, our results imply different hatching dynamics affected by disperser and season, but most species transported as resting eggs by birds seem to have a wide tolerance to hatch under variable salinity conditions. These differences may largery influence the metacommunity dynamics of lake networks, and could be a key factor to consider in wetland conservation planning.     

Zooplankton, phytoplankton and the microbial food web in two turbid and two clearwater shallow lakes in Belgium

Components of the pelagic food web in four eutrophic shallow lakes in two wetland reserves in Belgium (Blankaart and De Maten) were monitored during the course of 1998–1999. In each wetland reserve, a clearwater and a turbid lake were sampled. The two lakes in each wetland reserve had similar nutrient loadings and occurred in close proximity of each other. In accordance with the alternative stable states theory, food web structure differed strongly between the clearwater and turbid lakes. Phytoplankton biomass was higher in the turbid than the clearwater lakes. Whereas chlorophytes dominated the phytoplankton in the turbid lakes, cryptophytes were the most important phytoplankton group in the clearwater lakes. The biomass of microheterotrophs (bacteria, heterotrophic nanoflagellates and ciliates) was higher in the turbid than the clearwater lakes. Biomass and community composition of micro- and macrozooplankton was not clearly related to water clarity. The ratio of macrozooplankton to phytoplankton biomass – an indicator of zooplankton grazing pressure on phytoplankton – was higher in the clearwater when compared to the turbid lakes. The factors potentially regulating water clarity, phytoplankton, microheterotrophs and macrozooplankton are discussed. Implications for the management of these lakes are discussed.     

Response of waterbirds to alternating clear and turbid water phases in two shallow Mediterranean lakes

Albufera de Adra (Southern Spain) constitutes an internationally-recognised marsh for waterbirds; important populations of some endangered species such as White-headed Duck and Red-crested Pochard overwinter and breed in its two shallow permanent lakes (Lake Honda and Lake Nueva). In a recently published article, we revealed the factors responsible for the irregular alternation between phytoplankton-dominated turbid phases and macrophyte-dominated clear water phases in Lake Honda and Lake Nueva. In this note, we try to clarify the impact of such an alternation of equilibrium states on the waterbird dynamics. Marked increments in abundance, and brood recruitment of dabbling and diving waterbirds were recorded during the clear water phases in contrast with the turbid water phases, as the increase of macrophytes associated with increased water transparency attract waterbirds for available food. Implications for ecosystem management, restoration and conservation are identified.     

Factors influencing taxonomic composition and abundance of macrozoobenthos in extralittoral zone of shallow eutrophic lakes

Zoobenthos is an essential part of shallow lake ecosystems, exerting a considerable impact upon their functioning. We studied 13 eutrophic, shallow, polymictic lakes from Northern Poland to find out which environmental factors influence taxonomic composition, abundance and biodiversity of their zoobenthos. The Canonical Correspondence Analysis allowed to distinguish three lake types: (1) macrophyte-dominated lakes, with high plant cover and well illuminated bottom, inhabited by abundant, diverse benthic taxa; (2) deeper phytoplankton-dominated lakes, with shaded bottom, high sediment oxygen demand (SOD) and rather sparse zobenthos community, dominated by Chironomus and Chaoborus larvae; (3) shallower phytoplankton-dominated lakes, with intermediate amount of light at the bottom and lower SOD values and comparatively diverse zoobenthos, but with lower number of taxa than in the first group. Apart from plant presence, distinguishing between macrophyte-dominated lakes and the other types, the most important variable in the CCA was amount of light reaching the bottom. Probably the impact of light on the bottom fauna was indirect: light stimulated development of macrophytes or phytobenthos (depending on its intensity) and thus improved food and oxygen conditions. Zoobenthos was also affected by oxygen conditions (mainly SOD), presumably by short-time oxygen depletions occurring in the deep phytoplankton-dominated lakes and preventing survival of some benthic taxa.     

Driving forces shaping phytoplankton assemblages in two subtropical plateau lakes with contrasting trophic status

1. We studied driving forces shaping phytoplankton assemblages in two subtropical plateau lakes with contrasting trophic status, the oligotrophic deep Lake Fuxian and the eutrophic shallow Lake Xingyun. 2. Phytoplankton samples were taken monthly for a year and phytoplankton species were sorted into the main taxonomic groups and associations proposed by Reynolds. A canonical correspondence analysis (CCA) was used to test the occurrence of these classification schemes and to determine their discriminatory power. 3. The results suggest that the major driving forces in Lake Fuxian were physical variables, and particularly the underwater light climate, whereas, nutrients were the important driving force in Lake Xingyun. 4. Top–down control through zooplankton grazing in Lake Fuxian was hardly ever a significant determinant itself, because of the scarcity of zooplankton and their low grazing efficiency of predation while a dominance of inedible cyanobacteria throughout the year rendered top–down controls ineffective failing in Lake Xingyun. Hence phytoplankton communities in both lakes appear to be regulated primarily by bottom–up controls.     

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.     

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.     

Size‐based interactions across trophic levels in food webs of shallow Mediterranean lakes

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Prediction of phosphorus and nitrogen concentrations in lakes from both internal and external loading rates

A model predicting phosphorus and nitrogen concentrations from loading rates was tested using data collected from a shallow, nutrient-rich lake where both internal and external loading occurred. Predicted nutrient concentrations agreed closely with obaerved values and it is suggested that the model could be used to predict the reduction in loading rate required to effect lake restoration.     

Cascading trophic effects in pampean shallow lakes: results of a mesocosm experiment using two coexisting fish species with different feeding strategies

Planktivorous fish, both visual predators and filter feeders, enhance eutrophication processes in lakes. In pampean shallow lakes several planktivorous species may coexist but often two species dominate: silverside (Odontesthes bonariensis), a visual planktivorous fish when young adult, and sabalito (Cyphocharax voga), an omnivorous filter feeder. To assess the relative strength of the cascading trophic effects of the two species, a mesocosm experiment was conducted using different proportions of both species as treatments. Differences were found in water transparency, phytoplankton biomass, crustacean abundance, individual size and biomass. Our results suggest that visual predators intensify eutrophication effects more strongly than filter feeders do.     

The light : nutrient ratio in lakes: a test of hypothesized trends in bacterial nutrient limitation

The light-to-nutrient hypothesis explores how the balance between energy (as light energy) and nutrients (as total phosphorus) shapes aquatic ecosystem structure and process. The balance of energy and nutrients is thought to regulate ecosystem structure and process such that, in a "high" light-to-nutrient environment, bacteria would probably be driven towards phosphorus (P) limitation, whereas, in a "low" light-to-nutrient environment, bacteria would be driven towards carbon (C) limitation. We assessed the growth limitation of bacteria in two reservoirs of the southern U.S.A. using a mortality-corrected dilution-growth approach. We compared the frequency of P and C growth limitation with the intralake variation in the light-to-nutrient environment. As a metric of the light-to-nutrient environment, we used the ratio of the mean light in the surface mixed layer ( I _m) to the total phosphorus concentration ( TP ). In each lake, bacterial growth was more often P-limited when the I _m :  TP ratio was above the median ratio than below. We believe our data provide the first evidence supporting this aspect of the light-to-nutrient hypothesis.     

A comparison of zooplankton densities and biomass in Lakes Peipsi and Võrtsjärv (Estonia): rotifers and crustaceans versus ciliates

The abundance and biomass of ciliates, rotifers, cladocerans and copepods were studied in Lake Peipsi and Lake Võrtsjärv, both of which are shallow, turbid and large. Our hypothesis was that in a large shallow eutrophic lake, the ciliates could be the most important zooplankton group. The mean metazooplankton biomass was higher in Peipsi than in Võrtsjärv (mean values and SD, 1.8 ± 0.7 and 1.3 ± 0.6 mg WM l^?1). In Peipsi, the metazooplankton biomass was dominated by filtrators that feed on large-sized phytoplankton and are characteristic of oligo-mesotrophic waters. In Võrtsjärv, the metazooplankton was dominated by species characteristic of eutrophic waters. The planktonic ciliates in both lakes were dominated by oligotrichs. The biomass of ciliates was much greater in Võrtsjärv (mean 2.3 ± 1.4 mg WM l^?1) than in Peipsi (0.1 ± 0.08 mg WM l^?1). Ciliates formed about 60% of the total zooplankton biomass in Võrtsjärv but only 6% in Peipsi. Thus, the food chains in the two lakes differ: a grazing food chain in Peipsi and a detrital food-chain in Võrtsjärv. Consequently, top-down control of phytoplankton can be assumed to be much more important in Peipsi than in Võrtsjärv. When the detrital food chain prevails, the planktonic ciliates become the most important zooplankton group in shallow, eutrophic and large lake. Neglecting protozooplankton can result in serious underestimates of total zooplankton biomass since two-thirds of the zooplankton biomass in Võrtsjärv comprises ciliates.     

Comparative analysis of bacterioplankton assemblages from maritime Antarctic freshwater lakes with contrasting trophic status

The bacterioplankton assemblages of eight maritime Antarctic lakes with a wide range of trophic status and geographic span (six lakes from Hope Bay, Antarctic Peninsula and two from Potter Peninsula, King George Island) were described using denaturing gradient gel electrophoresis and band sequencing during two consecutive austral summers (2003–2004). Analyses of the gels identified a total of 230 bands spread across 57 different positions. Among those bands, 14 were shared between lakes from Hope Bay and Potter Peninsula, 17 were observed only in particular lakes, and 17 were registered both years in the same lake. We successfully reamplified and sequenced 43 bands located in 36 different positions belonging to Bacteroidetes, Actinobacteria, Betaproteobacteria and Cyanobacteria. The closest matches for 63% of the sequenced bands were from Antarctic or from other cold environment clones and sequences already in the databases, suggesting the widespread dominance of microbial communities adapted to cold habitats. The results of the multivariate analyses (Cluster Analysis and CCA) indicated that the nutrient status of the lake influences the bacterioplankton assemblages. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Modeling the role of wind and warming on Microcystis aeruginosa blooms in shallow lakes with different trophic status

This study focuses on the role of wind exposure and storm events, in interaction with trophic status and temperature, on the competition between two species: Microcystis aeruginosa and a typical green alga. It is based on a water column model containing ecological and fluid mechanic features including mixing and shear stress at the bottom. This model addresses for the first time the impact of storm events (inducing sediment and nutrient resuspension) on algal dynamics. Simulations with realistic environmental forcings were performed with different sets of wind, temperature, and trophic conditions. With normal temperatures, conditions for dominance and bloom formation of M. aeruginosa in summer are restricted to hypertrophic waters with low wind exposure. Higher wind exposure (above 2 m s^?1) impairs the formation blooms even in favorable trophic conditions and enhances the dominance of green algae. Hotter temperatures allow the dominance of M. aeruginosa for lower phosphorus conditions and higher wind exposure and lead to the exclusion of green algae for high phosphorus content and low wind exposure. Nevertheless, high wind exposure (above 3 m s^?1) still prevents dense bloom formation and allows the coexistence of both species. Storm events bring two counterbalancing features: sediment and nutrient resuspension. The first leads to a decrease of phytoplankton density in response to high turbidity, and the second leads to an increase and better maintenance of M. aeruginosa blooms due to high phosphorus concentration released in the water. This result depends on the timing of the event and on general wind exposure as phosphorus release only benefits M. aeruginosa if exposure to wind is low.     

Fish and nutrient enrichment effects on rotifers in a Mediterranean shallow lake: a mesocosm experiment

A mesocosm experiment was conducted “in situ” in a Chara dominated shallow lake near Valencia (Spain) to study top–down and bottom–up effects on rotifers by means of nutrient and fish additions. Both processes were important in determining rotifer abundance, biomass and diversity. A total of 36 mesocoms were established with triplicate treatment combinations of three fish levels (from no fish to 45 individuals of Gambusia holbrooki males) and four nutrient enrichment levels (from no additions to 10 mg l⁻¹ nitrate-N and 1 mg l⁻¹ phosphate-P). The main effect was a notable increase of planktonic and plant associated rotifers densities with fish. Rotifers benefited from mosquitofish predation on microcrustaceans and chironomids. The results showed a marked negative relationship between rotifer and cyclopoid abundances, indicating the importance of the predatory pressure of cyclopoids on rotifers. Effects on rotifer diversity were also evident, in general rotifer diversity decreased with nutrients and increased with fish. The effects of nutrients analysed at species level showed two contrasting density responses: an increase or a decrease with nutrients, which levelled off at high nutrient concentrations. High-level nutrient additions (from 5 mg l⁻¹ nitrate-N and 0.5 mg l⁻¹ phosphate-P) induced a switch to a turbid state with macrophyte disappearance. Most planktonic rotifer species, as well as plant associated ones, diminished when the turbid state was well established, especially in the mesocosms without fish. In the turbid mesocosms, relative abundance of plant-associated rotifers (as a whole) was higher than that of planktonic rotifers. The changes in rotifer species composition after the switch from a clear to a turbid water state are also described. Species of the genus Anuraeopsis, Trichocerca and Hexarthra, dominant in the clear water state, practically disappeared in the turbid water state, in which Proalides tentaculatus and Lecane nana were the main species. Guest editors: S. S. S. Sarma, R. D. Gulati, R. L. Wallace, S. Nandini, H. J. Dumont & R. Rico-Martínez Advances in Rotifer Research     

The role of cladocerans reflecting the trophic status of two large and shallow Estonian lakes

The role of pelagic cladoceran communities is discussed on the basis of a comparative study conducted in two Estonian lakes, the moderately eutrophic Lake Peipsi (N_tot 700, P_tot 40 μg l^?1 as average of ice-free period of 1997–2003) and in a strongly eutrophic Lake Võrtsjärv (N_tot 1600, P_tot 54 μg l^?1). The cladoceran community was found to reflect the differences in the trophic state of these lakes. In L. Peipsi, characteristic species of oligo-mesotrophic and eutrophic waters co-dominated (making up 20% or more of total zooplankton abundance or biomass), whereas in L. Võrtsjärv only species of eutrophic waters occurred. In L. Peipsi, the dominant cladocerans were Bosmina berolinensis and Daphnia galeata, while Chydorus sphaericus was the most abundant cladoceran in L. Võrtsjärv. The cladocerans of L. Peipsi (mean individual wet weight 25 μg) were significantly (threefold) larger than those of L. Võrtsjärv (8 μg). The mean wet biomass of cladocerans was higher and total cladoceran abundance was lower in L. Peipsi compared to L. Võrtsjärv (biomass varied from 0.133 to 1.570 g m^?3; mean value 0.800 g m^?3 in L. Peipsi and from 0.201 to 0.706 g m^?3, mean 0.400 g m^?3 in L. Võrtsjärv; the corresponding data for abundances were: 8,000–43,000 ind. m^?3, mean 30,000 ind. m^?3 for L. Peipsi, 50,000–100,000, mean 52,000 ind. m^?3 for L. Võrtsjärv). Based upon differences in body size, cladocerans were more effective transporters of energy in L. Peipsi than in L. Võrtsjärv. Cladocerans proved to be informative indicators of the trophic status and of the efficiency of the food web in studied lakes.