Dominant species, functional assemblages and frequency of equilibrium phases in late summer phytoplankton assemblages in Hungarian small shallow lakes

Late summer phytoplankton associations were studied qualitatively and quantitatively in 80 Hungarian lakes altogether (mostly shallow salt lakes, reservoirs, oxbows, gravel pit lakes). Equilibrium phases sensu Sommer et al. (1993) were found only in 17 lakes. Most of them were under some kind (high salt content or very low level of nutrients) of stress factor. It is concluded that environmental stress forces phytoplankton communities towards equilibrium. No relationship between occurrence of equilibria and trophic state was found. Species number of non-equilibrated lakes was almost three times as high as those in equilibrium. Of the 31 recently described (Reynolds et al., 2002) phytoplankton assemblages most of those were recognized that are likely to occur in shallow lakes. Separation of a functional group W _S from W2 for Synura dominated lakes is suggested. It seemed also necessary to raise a group (Y _Ph) for lakes dominated by Phacotus. Sorting of Dinophyta species into different already described functional groups is desirable.     

Connectivity and cladoceran species richness in a metacommunity of shallow lakes

SUMMARY 1. The shallow ponds of the nature reserve 'De Maten' form a metacommunity, in which individual ponds are highly interconnected via a system of overflows and rivulets. This study reports on the relations between cladoceran species richness and (a) connectivity patterns and (b) local environmental variables.
2. No relation was found between local species richness and three connectivity variables or dispersal pathways.
3. Spatial configuration was related to richness, but was confounded by environmental variables for 2 of 3 years. In those 2 years, there was a significant linear relation between Secchi disc depth and species richness, suggesting an important impact of the clearwater/turbid state alternative equilibria in shallow lakes in determining cladoceran richness. Only in the year in which environmental variables were unimportant did connectivity between the ponds influence species richness.
4. These results suggest that local environmental variables related to the clearwater/turbid state alternative equilibria in shallow lakes are important in determining cladoceran species richness. Connectivity and dispersal of individuals between the different ponds only act secondarily by increasing the general species richness within a pond through dispersal from ponds with different environmental conditions.     

The effect of turbidity state and microhabitat on macroinvertebrate assemblages: a pilot study of six shallow lakes

Shallow lakes can occur in two alternative stable states, a clear-water state and a turbid state. This is associated with separate assemblages of fish, zooplankton and plants. Little is known about whether macroinvertebrate assemblages differ across both stable states. This study investigated this in a connected set of three turbid and three clear-water shallow lakes. To overcome confounding effects of differences in spatial structure of macrophytes in turbid and clear-water lakes, we sampled three microhabitats that occurred in both alternative stable states: open water, sago pondweed (Potamogeton pectinatus) and reed (Phragmites australis). Univariate analyses indicated no differences in the number of organisms, taxon richness or diversity between turbid and clear-water lakes. Multivariate analysis, however, showed significant differences in the macroinvertebrate community structure of both stable states. Nine taxa explained a significant amount of the variation between both lake types, of which seven preferred the clear-water lakes. The number of organisms and the taxon richness were higher in reed than in the other microhabitats, but diversity and evenness did not differ among the microhabitats. Multivariate analyses could separate all three microhabitats. Eight taxa, mainly detritus feeders and collector–gatherers, explained most of the variation in the data and preferred the reed microhabitat. The effects of stable state (6.8% explained variance) and microhabitat (13.1% explained variance) on the macroinvertebrate assemblages were largely independent from each other (1.5% shared variance). Although macroinvertebrates are not implemented in the initial theory of stable states, our results show clearly different assemblages across both stable states.     

Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance

A number of mechanisms result in a feedback between water clarity and macrophytes and, consequently, the occurrence of alternative stable states in shallow lakes. We hypothesize that bottom-up mechanisms and interactions within the benthic food web are more important in a charophyte-dominated clear-water state, while top-down mechanism and interactions in the planktonic food web prevail at angiosperm dominance. Charophytes, which dominate at lower nutrient concentrations and develop higher densities than most angiosperms, can have a higher influence on sedimentation, resuspension, and water column nutrients. During dominance of dense submerged vegetation like charophytes, zooplankton can be hampered by low food quality and quantity and by high predation pressure from juvenile fish, which in turn are favoured by the high refuge potential of this vegetation. Grazing pressure from zooplankton on phytoplankton can therefore be low in charophytes, but the main feedback in angiosperm-dominated ecosystems. Charophytes offer a higher surface than most angiosperms to periphyton, which favors benthic invertebrates. These support macrophytes by grazing periphyton and constitute a central link in a trophic cascade from fish to periphyton and macrophytes. To test these hypotheses, more experiments and field measurements comparing the effect of charophytes and angiosperms on water clarity are needed.     

Sediment phosphorus release in phytoplankton dominated versus macrophyte dominated shallow lakes: importance of oxygen conditions

Increased discharges of organic matter from different sources in Morales Stream, one of the main tributaries of the Matanza-Riachuelo River, caused not only an increase in its primary production but also drastic changes in the composition of its sediments, thus favoring eutrophication processes. An in situ study was carried out in order to assess the effects of an organic point source contamination (from intensive cattle rearing) on the sediments of Morales Stream. Surface water and sediment samples were analysed to determine the chemical characteristics of the water–sediment system. The amounts and forms of sediment phosphorus were determined using the `EDTA method' (Golterman, 1996) at two sites of the stream having different nutrient loads. The increase in the organic load of Morales Stream waters influences the dynamics of sediment P, producing two main effects: (1) an increase in the organic matter amount of the sediment that leads to an increase in the amount of P associated to organic fractions, which may be released by bacterial activity under anoxic conditions; and (2) a decrease in the concentration of P in the fraction bound to iron. Morales Stream sediments may act as a potential source of P, which can release this nutrient to water under the reducing conditions originated by uncontrolled discharges of organic residues to this water body.     

The impact of bird herbivory on macrophytes and the resilience of the clear-water state in shallow lakes: a model study

Land-use change associated with human development can alter aquatic habitat and imperil aquatic species. Fish are challenged when urban streams are altered, for example for stormwater conveyance, but little is known about how such activities influence the space use of individual fish. Electronic tagging and experimental displacement of fish can be used to explore site fidelity and homing behaviour of fish and can therefore be useful for testing hypotheses about space use and habitat selection. In this study, we used experimental displacement to determine how longnose dace (LND, Rhinichthys cataractae) utilize reaches within a watershed that have varying degrees of degradation. LND were tagged using passive integrated transponders (PIT tags), transported upstream, and released either into the natural stream reach, impaired stormwater drain reach, or at their confluence. Fixed PIT antennas were used to monitor movement of the PIT-tagged fish among the three reaches for a period of 3 weeks. LND exhibited dramatic and rapid selection against the stormwater drain. No LND moved into the drain and 97% of fish transported to the drain left within 24 h. LND were actively avoiding the stormwater drain, emphasizing the need for enhancement work to improve the biological connectivity of the system.     

Empirical relationships between cladoceran fauna and trophic state in thirteen northern German lakes: analysis of surficial sediments

The cladoceran faunas from 13 northern German lakes (the Plön district) of different trophic levels were analysed using the bosminid and chydorid remains in the surficial sediment as an integrated sample of the total lake fauna. In the dimictic lakes, a shift occurred in the composition of planktonic Cladocera from Bosmina coregoni, including morphs with a mucro, to predominance of Chydorus sphaericus and Bosmina c. thersites as trophic state increased. In the benthic Cladocera (Chydoridae), there was a decrease in species diversity, in the percentages of phytophilous species and in the portion of species that are typical of clear water lakes, but there was an increase both in the number of mud dwellers and species that are typical of polluted lakes. No positive correlation was found between mean depth of the lakes and the planktonic/littoral ratio.     

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.     

Small habitat size and isolation can promote species richness: second-order effects on biodiversity in shallow lakes and ponds

Contemporary ecological landscape planning is often based on the assumption that small isolated habitat patches sustain relatively few species. Here, we suggest that for shallow lakes and ponds, the opposite can be true for some groups of organisms. Fish communities tend to be poor or even absent in small isolated lakes. However, submerged vegetation is often more abundant in such waterbodies. As a consequence of low fish biomass and high vegetation abundance, the richness of aquatic birds, plants, amphibians and invertebrates is often relatively high in small, shallow, isolated lakes. Although the rarity of fish is in line with expectations from the ruling paradigms about effects of habitat fragmentation in landscape ecology, the relative richness of various other groups of organisms in small ponds is opposite to these expectations. The case of shallow lakes illustrates that incorporating ecological interactions is essential to understanding the potential effects of habitat fragmentation. Single-species meta-population approaches may be misleading if ecological interactions are strong. A meta-community approach that explicitly incorporates biotic interactions, also those involving different trophic levels, is needed. Our diagnosis suggests that connection of isolated habitat fragments may in some cases reduce, rather than enhance, landscape-level biodiversity, and implies that biodiversity at the regional level will be maximized if the local habitat patches vary widely in size and degree of connectivity.     

Comparative analysis of rotifer community structure in five subtropical shallow lakes in East China: role of physical and chemical conditions

Worldwide, there have been few comparative studies on rotifer communities in subtropical lakes. We studied changes in rotifer community structure over 1 year and its relationship to several physicochemical variables in five subtropical shallow lakes in East China, covering a nutrient gradient from mesotrophy to moderate eutrophy. In these lakes, the genera Brachionus, Lecane, and Trichocerca dominated the rotifer species composition, and Polyarthra dolichoptera, Keratella cochlearis, Filinia longiseta, T. pusilla, and Anuraeopsis fissa were the dominant species. With increased nutrient loading, total rotifer abundance and species dominance increased, indicating that rotifer abundance might be a more sensitive indicator of trophic state than species composition. Comparative analyses of the six rotifer community indices calculated in this study and redundancy analysis (RDA) revealed that the two slightly eutrophic lakes and the other two moderately eutrophic lakes exhibited a high degree similarity in community structure. This suggests that the trophic state of a lake determines the rotifer community structure. In contrast, in the two moderately eutrophic lakes, the mass ratios of TN:TP and the contents of TP suggested N-limitation and cyanobacteria dominance in phytoplankton communities might be possible. In these lakes TN played a more important role in shaping the rotifer community according to stepwise multiple regression and RDA. RDA analysis also suggested that rotifer species distribution was strongly associated with trophic state and water temperature, with water temperature being the most important factor in determining seasonality.     

Zooplankton (Cladocera) in assessments of biologic integrity and reference conditions: application of sedimentary assemblages from shallow boreal lakes

Zooplankton are potentially powerful proxies for the assessments of biologic integrity. The paleolimnological perspective and use of fossil Cladocera also provide the means to reconstruct reference conditions and natural long-term community dynamics. Unfortunately, the use of zooplankton in lake quality assessments is currently underexploited. We studied a surface sediment dataset of 41 lakes in Finland to examine the relationship between Cladocera remains and environmental variables. Of the examined environmental variables, total phosphorus availability was found to be the most important variable in explaining the Cladocera community composition. Following the tests on species environment relations, we selected a lake trophic typology as the most suitable environmental variable for developing a new tool for limnoecological quality assessments. A test of the model on a modern and historic sample from a eutrophied lake showed that the test lake has proceeded from “mesotrophic/poor” to “eutrophic/bad” limnoecological state in agreement with previous independent evidence. The model developed here showed favorable performance that can be used to provide reliable estimates of ecological and environmental state of lakes.     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Effect of summer weather on internal loading and chlorophyll a in a shallow lake: a modeling approach

The effect of weather on the eutrophication of a shallow lake was estimated by a hydrodynamic lake model coupled with a simple water quality module. The model was applied to Lake Villikkalanjärvi in southern Finland. This shallow, agriculturally loaded lake may stratify during warm and calm periods in summer and as a result oxygen is often consumed from the hypolimnion, causing high internal loading of phosphorus. Vertical mixing and temperature distribution in the lake were simulated by a one-dimensional, horizontally integrated hydrodynamic model. State variables included in the water quality model were dissolved reactive phosphorus, chlorophyll a and dissolved oxygen. The model was first calibrated against observations from 1989 and 1990. Thereafter, simulations were carried out using weather data from the years 1961 to 1988. The results indicated that warm summer periods may cause high chlorophyll a concentrations due to high internal loading. In four years with exceptionally warm summers the model predicted maximum chlorophyll a concentrations almost twice as high as in years without remarkable internal loading. The model simulates accurately temperature and mixing but the reliability of water quality predictions could be improved by adding more factors regulating algal biomass and sediment phosphorus release.     

Nitrogen fixation in a large shallow lake: rates and initiation conditions

The fixation of molecular nitrogen (N₂fix) by cyanobacteria in situ and in PO₄-P enrichment experiments was investigated in large shallow Lake Võrtsjärv in 1998–2000. In this lake, N₂fix started when TN/TP mass ratio was about 20, which is much higher than Redfield mass ratio 7. The rate of N₂fix varied between 0.81 and 2.61 gN l^–1 d^–1 and maximum rate (2.61 gN l^–1 d^–1) was measured in 15.08.2000. In L. Võrtsjärv a lag period of a couple of weaks occurred between the set-up of favourable conditions for N₂fix as the appearance of N₂-fixing species and depletion of mineral nitrogen, and the real N₂fix itself. However, if the favorable conditions for N₂fix occurred in the lake, N₂fix started after enrichment with PO₄-P in mesocosms even then when no N₂fix was detected in the lake. N₂fix in mesocosms was also more intensive than in lake water. In our experiments PO₄-P concentrations higher than 100 gP l^–1started to inhibit N₂fix.     

Major changes in CO<Subscript>2</Subscript> efflux when shallow lakes shift from a turbid to a clear water state

Lakes can be sources or sinks of carbon, depending on local conditions. Recent studies have shown that the CO₂ efflux increases when lakes recover from eutrophication, mainly as a result of a reduction in phytoplankton biomass, leading to less uptake of CO₂ by producers. We hypothesised that lake restoration by removal of coarse fish (biomanipulation) or invasion of mussels would have a similar effect. We studied 14–22 year time series of five temperate Danish lakes and found profound effects on the calculated CO₂ efflux of major shifts in ecosystem structure. In two lakes, where limited colonisation of submerged macrophytes occurred after biomanipulation or invasion of zebra mussels (Dreissena polymorpha), the efflux increased significantly with decreasing phytoplankton chlorophyll a. In three lakes with major interannual variation in macrophyte abundance, the efflux declined with increasing macrophyte abundance in two of the lakes, while no relation to macrophytes or chlorophyll a was found in the third lake, likely due to high groundwater input to this lake. We conclude that clearing water through invasive mussels or lake restoration by biomanipulation may increase the CO₂ efflux from lakes. However, if submerged macrophytes establish and form dense beds, the CO₂ efflux may decline again.     

A comparative study on planktonic ciliates in two shallow mesotrophic lakes (China): species composition, distribution and quantitative importance

The species richness and seasonal development of planktonic ciliates were studied and compared in two shallow mesotrophic lakes, one covered with dense submerged macrophytes, the other macrophyte poor. Considerable differences in ciliate species composition, dominant taxa, abundance and biomass were observed. Ciliates were much more species rich in the macrophyte-rich lake, while they were more abundant numerically in the macrophyte-poor lake. Altogether, 96 species, included in 53 genera, 14 orders were identified. Among them, 80 species (included in 45 genera, 14 orders) observed from the macrophyte-rich lake, against 49 species (36 genera, 12 orders) were from the macrophyte-poor lake. In the macrophyte-rich lake, the mean abundance and biomass were 13.5 cells ml^-1 and 547.10 g l^-1 f.w.; abundance and biomass were higher in spring and winter; naked oligotrichs dominated total ciliate abundance and Peritrichida dominated the biomass. In the macrophyte-poor lake, ciliate mean abundance and biomass were 35.5 cells ml^-1 and 953.39 g l^-1 f.w.; abundances peaked in autumn; Scuticociliates dominated the abundance and Tintinnids dominated the biomass. Possible causes for the observed differences are discussed.     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

The role of charophytes in increasing water transparency: a case study of two shallow lakes in Estonia

Hydrobiologia - The hydrochemical regime and the biota in two lakes of Vooremaa landscape protection area, Central Estonia, were studied in 2000–2001 within the frames of the EC project...