Does the impact of nutrients on the biological structure and function of brackish and freshwater lakes differ?

The effects of nutrients on the biological structure of brackish and freshwater lakes were compared. Quantitative analysis of late summer fish, zooplankton, mysid and macrophyte populations was undertaken in 20–36 shallow brackish lakes of various trophic states and the findings compared with a similar analysis of shallow freshwater lakes based on either sampling (fish) or existing data (zooplankton, mysids and macrophytes). Special emphasis was placed on differences in pelagic top-down control. Whereas the fish biomass (CPUE, multiple mesh-size gill nets) rose with increasing P-concentration in freshwater lakes, that of brackish lakes was markedly reduced at P-concentrations above ca. 0.4 mg P l^-1 and there was a concomitant shift to exclusive dominance by the small sticklebacks (Gasterosteus aculeatus and Pungitius pungitius); as a result, fish density remained relatively high. Mysids (Neomysis integer) were found at a salinity greater than 0.5 and increased substantially with increasing P-concentration, reaching levels as high as 13 ind. l^-1. This is in contrast to the carnivorous zooplankton of freshwater lakes, which are most abundant at intermediate P levels. The efficient algal controller, Daphnia was only found at a salinity below 2 and N. integer in lakes with a salinity above 0.5. Above 2 the filter-feeding zooplankton were usually dominated by the less efficient algal controllers Eurytemora and Acartia. In contrast to freshwater lakes, no shift to a clearwater state was found in eutrophic brackish lakes when submerged macrophytes became abundant. We conclude that predation pressure on zooplankton is higher and algal grazing capacity lower in brackish eutrophic-hypertrophic lakes than in comparable freshwater lakes, and that the differences in trophic structure of brackish and freshwater lakes have major implications for the measures available to reduce the recovery period following a reduction in nutrient loading. From the point of view of top-down control, the salinity threshold dividing freshwater and brackish lakes is much lower than the conventionally defined 5.     

How does the vertical distribution of chlorophyll vary in littoral sediments of small lakes?

1. In this study, sediment chlorophyll profiles at twenty littoral stations in three oligo‐mesotrophic lakes were compared to test whether the vertical distribution of chlorophyll is related to site characteristics (light availability, temperature, physical disturbances) and whether these profiles differ between shallow and deep portions of the littoral zone.
2. The magnitude of chlorophyll peaks at the sediment surface did not vary with light intensity. Chlorophyll peaks in the shallow littoral zone had a weak tendency to decrease with increasing effective fetch. The magnitude of chlorophyll peaks at deeper sites was more closely related to water temperature than to substrate slope.
3. High chlorophyll concentrations were measured down to 1–3 cm in the sediments, both at shallow (< 2.5 m) and deep (4–10 m) stations. The depth to which high chlorophyll was found in sediments did not vary with effective fetch or sediment water content, two indices of wave disturbance in the shallow littoral zone, or with substrate slope, an index of sediment stability in the deep littoral zone. Sediment mixing is apparently not related to common indices of physical disturbances.
4. Between 8 and 100% of sediment surface chlorophyll was 'retained' 4–5 cm into the sediments. The proportion of chlorophyll 'retained' in littoral sediments increased with increasing depth, increasing lake productivity (total phosphorus concentration) and increasing lake pH.
5. Among‐core variability (standard error/mean) in chlorophyll concentration at the sediment surface ranged from less than 1% to 33% at different stations and was highest at shallow, exposed sites. These levels of variability are similar to those found in other periphytic communities.     

Eutrophication of moderately deep Dutch lakes during the past century: flaws in the expectations of water management?

We studied the trophic development of the past 30–100 years in eight moderately deep Dutch lakes based on their sedimentary fossil diatom assemblages. The dominant diatoms indicating meso- to eutrophic conditions were Aulacoseira subarctica, Cyclotella ocellata, C. cyclopuncta, C. meneghiniana, Puncticulata bodanica, Aulacoseira granulata, Cyclostephanos dubius, C. invisitatus, Stephanodiscus hantzschii, S. medius, and S. parvus. Ordination of diatom data separated the lakes into four groups according to their total phosphorus concentrations (TP), water supply, water management, and origin. The first group consists of dike-breach lakes, which were in stable eutrophic to hypertrophic conditions throughout the past century with diatom-inferred TP (DI-TP) concentrations of between 70 and 300 μg l⁻¹. The main factors influencing these dike-breach lakes are river management, ground water supply of riverine origin, and local land use. The second group are artificial lakes of fluctuating oligo- to mesotrophic conditions and DI-TP concentrations of 10–30 μg l⁻¹. Only one of the artificial lakes showed a DI-TP increase due to changes in catchment agricultural practice. A third group includes an artificial moat and an inland dike-breach lake with DI-TP concentrations of 50–100 μg l⁻¹. The fourth group contains an individual dike-breach lake with stable mesotrophic conditions of 50 μg l⁻¹ throughout the past century. Rather than showing a regional pattern, the studied lakes behave very individualistically with regard to their trophic history, reflecting changes in the local hydrology and in their nutrient sources.     

Habitat variability of the Littorelletea uniflorae plant communities in Polish Lobelia lakes

Research was carried out to determine aquatic oligochaete habitat preferences in the Rokytná River (Thaya River basin), a sixth order highland stream in the Czech Republic during the period of April 1999–April 2001. Quantitative samples were collected and current velocity and basic physico-chemical variables were measured monthly in four typical habitats in the Rokytná River. During this study, 28, 842 individuals representing 44 oligochaete species or higher taxa were collected. Temporal variability of proportional occurrence of trophic groups found on selected habitats (gathering collectors and grazers) was analysed. Habitat preferences of oligochaete species were evaluated by Canonical Correspondence Analysis (CANOCO). Biochemical oxygen demand (BOD) and nitrate (NO₃⁻) ion concentration were the most important variables explaining the distribution of Oligochaeta along the first axis. Current velocity (at 40% of the depth) and presence of oligochaetes associated with the habitat where gravel bars never formed were the most important variables along the second axis. Both axes were correlated with the temperature, reflecting the sampling in summer along the first axis and sampling in late spring along the second axis. The amount of organic matter (BOD) and concentrations of NO₃⁻ ions represented both oligochaete food source and decomposition products contributing to the growth of algae. Current velocity and preferred habitat explained the spatial pattern of oligochaete distributions.     

Phytoplankton of the extremely acidic mining lakes of Lusatia (Germany) with pH ≤3

Most of the flooded, open-cast lignite mining lakes of Lusatia (Germany) impacted by the oxidation of iron sulphides (pyrite and marcasite) are extremely acidic. Of 32 lakes regularly studied from 1995 to 1998, 14 have a pH <3 (median pH 2.3–2.9). These lakes are typically buffered by high concentrations of Fe (III) and have high conductivity (1000–5000 S cm^–1). Concentrations of dissolved inorganic carbon (DIC) and phosphorus are typically extremely low. These factors result in a very different environment for algae than found in neutral and acid-rain impacted lakes. The planktonic algal flora is generally dominated by flagellates belonging to genera of Chlorophyta (Chlamydomonas), Heterokontophyta of the class Chrysophyceae (Ochromonas, Chromulina), Cryptophyta (Cyathomonas) and Euglenophyta (Lepocinclis, Euglena mutabilis). Near-spherical non-motile Chlorophyta (Nanochlorum sp.), Heterokontophyta of the class Bacillariophyceae (Eunotia exigua, Nitzschia), Dinophyta (Gymnodinium, Peridinium umbonatum), other Chlorophyta (Scourfieldia cordiformis) and Cryptophyta (Rhodomonas minuta) are also found.     

Macroinvertebrates on Eichhornia crassipes roots in two lakes of the Paraná River floodplain

Changes in the abundance, biomass and in the relativeproportions of functional feeding groups ofmacroinvertebrates were studied for 17 months duringlow and high water periods in two floodplain lakeswith indirect connections to the Paraná River.At low water, Eichhornia crassipes root clusterswere colonized by comparatively low densities ofmacroinvertebrates, especially in the lake moredistant to the river. Collector-gatherers andpredators were the most abundant functional feedinggroups, but different taxa of these groups dominatedin the two lakes during low water.At high water, macroinvertebrates densities increasedas a result of increased flow through the roots. Inthis period, collector-filterers dominated at bothlakes and constituted over 34% of themacroinvertebrates. Four independent variables(conductivity, hydrological periods, dissolved oxygenand site) explain 87% of the variability inmacroinvertebrates density expressed as the number ofindividuals per m2. No significant relationshipwere found between macroinvertebrate biomass and anyof the independent variables.     

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.     

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

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Is reduction of the benthivorous fish an important cause of high transparency following biomanipulation in shallow lakes?

Experimental reduction of the fish stock in two shallow lakes in The Netherlands shows that such a biomanipulation can lead to a substantial increase in transparency, which is caused not only by a decrease in algal biomass, but also by a decrease in resuspended sediment and detritus. A model was developed to describe transparency in relation to chlorophyll-a and inorganic, suspended solids (resuspended sediment). With the use of this model it is shown that more than 50% of the turbidity in these shallow lakes before biomanipulation was determined by the sediment resuspension, mainly caused by benthivorous fish. Another analysis reveals that the concentration of inorganic suspended solids and the biomass of benthivorous fish are positively correlated, and that even in the absence of algae a benthivorous fish biomass of 600 kg ha⁻¹ can reduce the Secchi depth to 0.4 m in shallow lakes. In addition, it is argued that algal biomass is also indirectly reduced by removal of benthivorous fish. Reduction of benthivorous fish is necessary to get macrophytes and macrophytes seem to be necessary to keep the algal biomass low in nutrient-rich shallow lakes. It is concluded that the impact of benthivorous fish on the turbidity can be large, especially in shallow lakes.     

Metabolic versatility of haloalkaliphilic bacteria from soda lakes belonging to the Alkalispirillum–Alkalilimnicola group

Four new isolates were obtained from denitrifying enrichments with various electron donors using sediment samples from hypersaline soda lakes. Based on 16S rRNA gene analysis and DNA-DNA hybridization results, they were all identified as members of the Gammaproteobacteria closely associated with the Alkalispirillum–Alkalilimnicola group. Two isolates were obtained from samples enriched with nitrate as electron acceptor and H₂ or polysulfide as electron donors, and another two strains were obtained with N₂O as the electron acceptor and sulfide or acetate as electron donors. All four new isolates, together with the type strains of the genera Alkalispirillum and Alkalilimnicola originally described as obligate aerobes, were capable of anaerobic growth with acetate using either nitrate or N₂O as electron acceptors. Their denitrification pathway, however, was disrupted at the level of nitrite. RuBisCO form I gene was detected and sequenced in the new isolates and in Alkalilimnicola halodurans but not in Alkalispirillum mobile. These data, together with the evidence of Oremland et al. (Appl Environ Microbiol 68:4795–4802, 2002) on the potential of Alkalilimnicola sp. MLHE-1 for autotrophic growth with arsenite as electron donor and nitrate as electron acceptor, demonstrate much higher metabolic diversity of this specific group of haloalkaliphilic Gammaproteobacteria than was originally anticipated.     

Reaction of large and shallow lakes Peipsi and Võrtsjärv to the changes of nutrient loading

More than 20-year monitoring of Estonian rivers reveals that the loading of nitrogen to large shallow lakes Peipsi (3,555 km², mean depth 7.1 m) and Võrtsjärv (270 km², mean depth 2.8 m) decreased substantially in the 1990s. Phosphorus loading decreased to a much smaller extent than nitrogen loading. In L. Võrtsjärv both N and P concentrations followed the decreasing trends of loading, which show the high sensitivity of large shallow lakes to catchment processes. Our study showed a positive relationship between P content in sediments and the relative depth of the lake. Assumingly the resilience of a lake in responding to the reduction of nutrient loading decreases together with the decrease of its relative depth. In L. Peipsi the concentration of P has not decreased since the 1990s. Our data show indirectly that P loading from Russia to L. Peipsi may have increased. The N/P ratio has decreased in both lakes. Cyanobacterial blooms have been common in both lakes already at the beginning of the 20th century. The blooms disappeared during heavy nitrogen loading in the 1980s but started again in L. Peipsi in recent years together with the drop of the N/P ratio. In L. Võrtsjärv the N/P ratio is higher and the ecosystem is more stable although the share of N₂-fixing cyanobacteria increased from the 1990s. Reappearing cyanobacterial blooms in L. Peipsi have caused fish-kills in recent years. In L. Peipsi summer/autumn fish-kills during water-blooms are a straightforward consequence of reduced nitrogen level at remaining high phosphorus level while in L. Võrtsjärv the climatic factors affecting water level are more critical––at low water level winter fish-kills may occur. In L. Võrtsjärv nutrient loading has decreased and water quality has improved, present ecological status seems to be mostly controlled by climatic factors through changes of water level. The most important measure to improve water quality in L. Peipsi would be the reduction of phosphorus loading from both Estonian and Russian subcatchments.     

How important is the crustacean plankton for the maintenance of water clarity in shallow lakes with abundant submerged vegetation?

• 1 We measured the abundance and biomass of filter‐feeding microcrustacean zooplankton and calculated their grazing impact on phytoplankton biomass during summer in five shallow, mesotrophic to eutrophic lakes. For three of the lakes data exist both from years with dense submerged vegetation and low turbidity (the clearwater state), as well as from years characterised by sparse vegetation and high turbidity (the turbid state). In the other two lakes data are available only for clearwater conditions.
• 2 In all lakes conditions of dense vegetation and clear water coincided with a low abundance of crustacean plankton during summer. In the three lakes that shifted, the calculated biovolume ingested by crustacean plankton (filtering rate) was 3–11 times lower during clearwater conditions compared with turbid conditions. Because phytoplankton biomass was lower during clearwater conditions, however, daily grazing pressure from microcrustacea (expressed as percentage of phytoplankton biomass) did not differ between states. In three of the five lakes, grazers were estimated to take less than 10% of the phytoplankton biomass per day, indicating filtration by zooplankton was not the most important mechanism to maintain clearwater conditions.
• 3 High densities of Cladocera were found in three of the lakes within dense stands of Charophyta. However, these samples were dominated by plant‐associated taxa that even during the night were rarely found outside the vegetation. This indicates that plant‐associated zooplankton has no major influence on the maintenance of water clarity outside the vegetation.
• 4 Spring peak abundance of Cladocera was observed in three of the lakes. In two of these, where seasonal development was studied in both the clearwater and the turbid state, spring peaks were lower during the clearwater state.
• 5 Predation, low food availability or a combination of both may explain the low zooplankton densities. Phytoplankton may be limited by low phosphorus availability in the lakes dominated by Charophyta. Our results indicate that the importance of zooplankton grazing may have minor importance for the maintenance of the clearwater state in lakes with dense, well‐established submerged vegetation.

     

Mass development of the flagellate Gonyostomum semen (Raphidophyta) in Swedish forest lakes - an effect of acidification?

During the last 20–30 years the flagellate Gonyostomum semen has become more abundant in lakes, especially small humic lakes, in Scandinavia. Mass development of the alga has been reported from areas affected by anthropogenic acid deposition, and reports from bathers of health problems are becoming frequent. Although there is an apparent connection between the appearance of Gonyostomum and acidification, it seems not to be the low pH per se that is the cause, but rather interlinked factors. The present tendency towards a wider distribution of the alga in non-humic lakes, emphasizes the importance of a better understanding of small humic lakes.     

Characteristics of caddis larvae assemblages from shallow lakes in the Bükk Mountains,North Hungary

Seventeen caddisfly species of four families (Limnephilidae, Sericostomatidae, Phryganeidae, and Leptoceridae) with a total of 1047 individuals were collected from four sampling sites in the littoral zone of Disznóskút Lakes in the Bükk Mountains. Three of these four families of Trichoptera were also recorded in the Central European lakes. From among the four substrates (sand, silt, large stones, and aquatic plants) of the littoral zone, silt and aquatic plants were dominant. Besides species of Limnephilidae found in higher than the average abundance in the silt, species of Phryganeidae and Leptoceridae also occurred. The trophic relationships of the species of these families can be considered as follows: shredders- predators- scrapers, shredders- detritivores- predators, and predators- shredders- scrapers. From the aquatic plants the species of Limnephilidae, Phryganeidae and Leptoceridae, belonging to the trophic relationships of detritivores- shredders, scrapers- predators, shredders- detritivores- predators, and predators- shredders- scrapers- detritivores were collected. The analysis (PCA) of the spatial and seasonal distribution of Trichopteran larvae at the four littoral zone sites shows that the assemblages collected near the inflow of the stream into the lake in late summer and near the inflow of the small channel into the lake in summer are very different from the others, for which Limnephilus elegans, Micropterna lateralis, and Stenophylax permistus are responsible at the previous site, while Halesus digitatus, Halesus tessellatus, and Mystacides nigra are responsible at the latter one. The other assemblages (mostly those in winter) are very similar to each other, which can be explained by the decreasing number of detectable species and their presence in low abundance. For the variance, Limnephilus rhombicus (57%) and Potamophylax nigricornis (16%) are essentially responsible. Considering the species composition of the larval assemblages, the site at the inflow of the stream into the lake was the most different (38%) from the others, and was 62% similar to them. This dissimilarity can possibly be explained by the presence of a mixed assemblage with typical stream and lake inhabitants of species of Limnephilidae present in high abundance at this silt dominated site and other caddis species that are typical stream inhabitants or common in streams and small rivers, and may have drifted into the lake.

     

Influence of submerged macrophytes,temperature, and nutrient loading on the development of redox potential around the sediment–water interface in lakes

Redox potential is a significant factor in aquatic systems to regulate the availability of nutrients and some metals. To assess the driving variables regulating redox potential, background parameters (dissolved oxygen, pH, temperature, chlorophyll-a, soluble reactive and total phosphorus content of water, coverage and height of submerged macrophytes) and redox potential profiles around the sediment–water interface (SWI) were measured in simulated shallow lake ecosystems. There were two nutrient regimes (enriched and non-enriched) and three temperature scenarios (unheated; +3.5°C; +5°C) installed in the experimental setups, which were constructed to study the effects of global climate change. Temperature did not have any detectable effect on redox potentials, and we presume that nutrient addition had only indirect positive effects through triggering phytoplankton dominance which causes macrophyte absence. When submerged macrophytes were present in high density (80–100% coverage), redox potentials at the SWI varied between 60–215 mV and the mean redox potential was 133 ± 34 mV (mean ± 1 SD). In contrast to this, when phytoplankton dominance was coupled to low macrophyte density (0–20% coverage), the range of redox potentials at the SWI was 160–290 mV and the mean redox potential was 218 ± 34 mV. The results revealed the primary importance of submersed macrophytes; macrophyte coverage determined alone the redox potential of the sediment–water interface by 81%. This study suggests that possible positive effects of macrophytes on redox potential can be suppressed by their negative effects in case of 80–100% coverage and total inhabitation of the water column.     

Variability of the molecular mass of α-subunit in phycocyanins of colonial Microcystis aeruginosa f. aeruginosa in different eutrophic pond and lakes

Electrospray ionization mass spectra of phycocyaninsof eight samples from natural blooms of Microcystis aeruginosa f. aeruginosa collectedat different eutrophic pond and lakes indicated thatthe subunits of phycocyanins in the samespecies have different molecular masses, whereas the subunits have almost constant molecularmasses. A negative linear relationship between themolecular mass of subunit of phycocyanin andthe concentration of chlorophyll of natural andcolonial cyanobacterial samples was found. Aninterannual similarity of the subunitmolecular masses of phycocyanins was observed fromsamples collected at the same sampling site at LakeKasumigaura during 1994 and 1995. A locationalvariability of the subunit molecular massesof phycocyanins was also observed among samplescollected at three eutrophic pond and lakes.     

Effect of river straightening on the hydrochemical properties of floodplain lakes: Observations from the Łyna and Drwęca Rivers,N Poland

The aim of the paper is to present two independent approaches to the issue of preservation of former meander loops based on examples of hydrotechnical works undertaken along meandering sections of the Drw?ca and ?yna Rivers (N Poland) in the 20th century. Except for significant changes in morphology of both rivers (shortened lengths, decrease in sinuosities and increase in channel slopes), the direct effect of their straightening is numerous cut-off channels. Hydrotechnical works performed along the middle section of the ?yna River maintained the connections between the active river channel and downstream arms of former meander loops. Thanks to that connection, the new water bodies successfully function as biogeochemical filters in the river valley and create favorable conditions for biodiversity. The consequence of the channelization of the Drw?ca River is numerous cut-offs separated from the river channel and characterized by advanced processes of eutrophication.The water quality data collected within the period of 2004–2006 showed significant consequences of the performed works for the man-made ecosystems. The comparison of water quality in semi-lotic and lentic cut-offs in relation to water in their parent rivers showed statistical differences. Significantly high concentrations of orthophosphates, total phosphorus and sulphates as well as the lowest concentrations of nitrates characterized lentic environments along the Drw?ca River. Besides hydrological connectivity, an important factor responsible for transformation processes of the man-made ecosystems is hydrological regime. It influences ecological integrity in floodplain ecosystems during high water levels and largely controls their chemical characteristics and aeration. Flood disturbances for lentic water bodies may renew connectivity and reset succession, can hold up the trend towards terrestrialization and disappearance of man-made water bodies in the floodplain. However, in light of the conducted studies, the maintenance of the connection between the downstream arm of the cut-off channel and the river is highly recommended.     

Does N2 fixation meet the nitrogen requirements of heterocystous blue-green algae in shallow eutrophic lakes?

Summary Phytoplankton net carbon uptake and nitrogen fixation were studied in two shallow, eutrophic lakes in South Sweden. Ranges of diurnal net carbon uptake were estimated by subtracting 24-h respiration rates corresponding to 5–20% of P _max, respectively, from daytime carbon uptake values. total nitrogen requirement of the phytoplankton assemblage was determined from the diurnal net carbon uptake, assuming a phytoplankton C:N ratio of 9.5:1. Nitrogen supplied by nitrogen fixation only occasionally corresponded to the demands of the total phytoplankton assemblage. When heterocystous algae made up a substantial proportion (10%) of the total phytoplankton biomass, nitrogen fixation could meet the requirements of heterocystous blue-green algae on c. 50% of the sampling occasions. Nitrogen deficiencies in heterocystous algae were most probably balanced by the simultaneous or sequential assimilation of dissolved inorganic nitrogen. It was concluded that uptake of ammonium or nitrate, regenerated from lake seston and sediment, is the main process by which growth of phytoplankton is maintained during summer in the lake ecosystems studied.     

Can small water level fluctuations affect the biomass of Nymphaea alba in large lakes?

In this study we investigated above-ground biomass and morphological responses of a floating-leaved plant species, Nymphaea alba, to small spring water level manipulations (0.1–0.5 m) in a large, shallow lake over a 9-year period (1995–2003). A year effect was found in mean annual above-ground plant biomass with higher values found in years of low water levels, 275–339 g DW m⁻² in 1995 and 2003 against 143–198 g DW m⁻² in 1996–2002 (no data transformation). No significant changes in biomass patterns were observed within each season (one summer peak), except in 1995 when a summer decline in biomass occurred. The amplitude and duration of exposure to high water levels affected the spring and annual above ground biomass of N. alba. The plant responded to high spring water levels by producing longer and thinner petioles to preserve leaves from flooding while no significant changes in leaf surface area (except in May) and leaf/petiole biomass ratio were obtained. The results are interpreted with regard to plant adaptations to changing environments (biomass allocation patterns in the different plant organs and stem density) and the effects of other abiotic factors relevant to the size of the system. We concluded that small deviations in spring water level can be driving forces in a large system in controlling the above-ground biomass of this floating-leaved plant.