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.     

The influence of biomanipulations (fish removal) on the structure of lake foodwebs, case studies using the LakeWeb-model

This paper presents results on how intensive fishing (fish removal) islikely to influence the structure of lake foodwebs. The work is based on acomprehensive dynamic lake ecosystem model, LakeWeb, which accounts forproduction, biomasses, predation and abiotic/biotic interactions of nine keyfunctional groups of organisms: phytoplankton, bacterioplankton, two types ofzooplankton (herbivorous and predatory), two types of fish (prey and predatory),zoobenthos, macrophytes and benthic algae. The model uses ordinary differentialequations, the ecosystem scale and gives seasonal variations (the calculationtime is 1 week). It is designed to account for all fundamental abiotic/bioticinteractions and feedbacks for lakes in general for the nine target groups. TheLakeWeb-model has been calibrated and critically tested using empirical data andregressions based on data from many lakes. It has been shown that the model canclosely capture typical functional and structural patterns in lakes, whichshould give credibility to the results presented in this work. Obtaining suchresults using traditional methods, i.e., extensive field studies in one or a fewlakes, would be very demanding (in terms of money, persons involved and time).In this paper, results are presented for two lakes, one Swedish and oneBelarussian. The intensive fishing operations carried out in LakeBlacksåstjärn, Sweden, to reduce Hg-concentrations in fish did notsucceed. A typical cost of an intensive fishing is about 10,000–30,000 USDper lake of this size ( 0.25 km²). The costs toremove fish would be about 40–120 USD per kg ww fish removed! Intensivefishing simulated for Lake Batorino, Belarus, to reduce the fish biomass willlikely increase the prey fish biomass as long as the predation pressure on preyfish is lower than during the prefishing stage. The biomass of predatory fishwill recover only slowly. However, this operation is not likely to succeed inlowering the algal volume in lakes with a high biomass of predatory zooplankton.This is easy to state qualitatively and the LakeWeb-model offers a practicallyuseful tool to quantify such changes and identify lakes where biomanipulationsare likely to fail or succeed.     

Impacts of environmental scenarios on chlorophyll-a in the management of shallow, eutrophic lakes following biomanipulation: An application of a numerical model

Accurate prediction of species changes in lake ecosystems following biomanipulation measures is of paramount importance in view of water quality management. The temporal variation of phytoplankton biomass as chlorophyll-a and transparency as Secchi depth measurements are studied in the Lake Bleiswijkse Zoom, The Netherlands, with a comprehensive structural dynamic model. In the formulation of the biological model, phytoplankton as several species, zooplankton, detritus, planktivores and benthivores, and piscivores are considered to be major contributing state variables for the model. The primary goal of this paper is to describe the possible impacts of several environmental scenarios on chlorophyll-a biomass qualitatively as it would help lake and environmental managers and relevant authorities elucidate the processes of eutrophication and biomanipulation in a broad way. Some of the scenarios that have been studied by this model are: (1) The effect of fixed stoichiometry in terms of internal nitrogen and phosphorus that are tied up within algal cells; (2) the effects of external phosphorus limitation; (3) light limitation and external nitrogen limitation on algal growth; (4) probable consequences that have taken place within the chlorophyll-a biomass due to change in biomasses of various aquatic organisms; and (5) possible changes of chlorophyll-a biomass due to higher temperatures caused by global warming.     

Zooplankton–phytoplankton relationships in shallow subtropical versus temperate lakes Apopka (Florida,USA) and Trasimeno (Umbria,Italy)

This study compares and contrasts the dynamics of phytoplankton, zooplankton, and nutrients in two of the largest shallow lakes in the USA (Lake Apopka, Florida) and Europe (Lago Trasimeno, Umbria, Italy) and considers particularly the biomass ratio of zooplankton to phytoplankton (BZ:BP) in relation to nutrient levels and in the context of data from other subtropical and temperate lakes. Lake Apopka is hypereutrophic with higher concentrations of total phosphorus (TP), nitrogen (TN), and nearly an order of magnitude higher BP than Lago Trasimeno. However, combined data from the two lakes can be fit to a single log–log regression model that explains 72% of the variability in BP based on TP. In contrast, BZ has a significant positive log–log relationship with TP only for Lago Trasimeno, and is much lower than expected based on the TP concentrations observed in Lake Apopka. Lake Apopka has a fish assemblage that includes high densities of gizzard shad (Dorosoma cepedianum) and threadfin shad (D. petenense), similar to other eutrophic Florida lakes that also have extreme low BZ. The ratio BZ:BP is below 0.01 in Lake Apopka, 10-fold lower than in Trasimeno and among the lowest values reported in the literature. Although stress of high water temperature and a greater proportion of inedible cyanobacteria may be contributing factors, the collective results support an emerging view that fish predation limits the biomass of crustacean zooplankton in subtropical lakes. Handling editor: S. I. Dodson     

Predation-driven dynamics of zooplankton and phytoplankton communities in a whole-lake experiment

Summary 1. Species compositions of zooplankton and phytoplankton were followed in Tuesday Lake before and after experimental manipulation of its fish populations (addition of piscivorous largemouth bass, removal of planktivorous minnows). Plankton dynamics were compared to those of adjacent, unmanipulated Paul Lake, where piscivorous fish have been dominant historically. 2. Indices of similarity for the zooplankton communities in the two lakes in 1984 prior to the manipulation were low; however, following the manipulation in spring, 1985, similarity of the zooplankton in the two lakes rose considerably and remained high throughout 1986. This was the result of an increase in Tuesday Lake of previously rare large-bodied cladocerans (Daphnia pulex, Holopedium gibberum) which were the dominants in Paul Lake, and the disappearance in Tuesday Lake of the dominant small-bodied copepod Tropocyclops prasinus, a minor component of the Paul Lake zooplankton. These observations are consistent with prior observations of the effects of size-selective predation on zooplankton communities. 3. Phytoplankton communities also responded strongly to the manipulation, with similarity indices for the two lakes rising from low levels in 1984 to high levels of similarity in 1985 and 1986, reflecting the decrease of formerly dominant Tuesday Lake taxa which were unimportant in Paul Lake and the appearance or increase in Tuesday Lake of several taxa characteristic of the Paul Lake phytoplankton assemblage. these results clearly show that food web structure can have pronounced effects on community composition at all levels of the food web, and that, just as zooplankton communities are structured by sizeselective predation, phytoplankton communities are structured by herbivory. These observations may provide some insight into factors governing the complex distributions of phytoplankton species among various lakes.A contribution from the University of Notre Dame Environmental Research Center, funded by NSF grants BSR-83-08918 and BSR-86-06271     

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     

Plankton community and the relationship with the environment in saline lakes of Onon-Torey plain,Northeastern Mongolia

The plankton community of sixteen saline lakes located on Onon-Torey plain (Northeastern Mongolia) during the filling phase and the raising of the water level was investigated in July 2011. Thirty-five taxa of phytoplankton and thirty-one species of zooplankton were found. For phytoplankton, blue-green algae (Merismopedia elegans, Anabaenopsis elenkinii, Arthrospora fusiformis, Spirulina major, Lyngbya sp., Oscillatoria sp.) and green algae (Monoraphidium minutum, Tetrastrum komarekii, Ankyra ocellata, Oocystis sp.) were dominant. For zooplankton, Filinia longiseta, Brachionus plicatilis, B. variabilis, Hexarthra mira (Rotifera), Daphnia magna, Moina brachiata, M. mongolica (Cladocera), Arctodiaptomus bacillifer, Mixodiaptomus incrassatus, Metadiaptomus asiaticus (Copepoda) dominated. Mineralization, active hydrogen ratio, dissolved oxygen and water temperature were the main factors influencing the diversity, structure and distribution of plankton organisms in the steppe lakes during low water level. The RDA analysis for phytoplankton and zooplankton from different lakes was carried out for selected two groups which included lakes and a subset related species. The first group is of oligohaline and mesohaline lakes in which mostly green algae, rotifers and copepods inhabit. The second group is of mesohaline and polyhaline lakes with mainly blue-green algae, some crustaceans and rotifers inhabiting. High abundance and biomass of Spirulina major, Oscillatoria sp. and Brachionus variabilis were observed in lakes with high mineralization, pH and temperature.     

Studies on zooplankton in two acidified high mountain lakes in the Alps

Being located in remote areas, alpine lakes are good indicators of regional and global pollution, and are particularly sensitive to atmospheric depositions. When situated in areas where acidic rocks dominate, they are sensitive to acidification. In the framework of an international project partially funded by the EU, a two-year study on zooplankton was carried out on two lakes, Lake Paione Inferiore (LPI) and Lake Paione Superiore (LPS), selected because of their susceptibility to acidification. In particular, LPS is permanently acidified, with pH ranging between 5.3 and 6.2, and LPI is acidified during the ice melt, when pH drops to 6.1 units. In addition, LPI is subjected to further anthropogenic interference, since fish (Onchorynchus mykiss) have been repeatedly introduced into the lake during the last thirty years. Literature information is available on the species composition and the seasonal dynamics of the zooplankton communities of the two lakes before these anthropogenic disturbances took place. Previously, the two lakes had a similar species composition and abundant zooplankton. An analysis of the present-day situation reveals significant changes compared to the past. Both acidification and the introduction of fish are responsible for the marked alterations observed. By reconstructing of the past cladocera assemblages and by analysing the literature on lakes with different impacts of the two factors (pH and fish), we can estimate the relative importance of the two different anthropogenic disturbances on species composition and abundance of the zooplankton communities of the two lakes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Changes in the biota of Chany Lake along a salinity gradient

Relationships among salinity and diversity, abundance, biomass of major biological components of Chany Lake (western Siberia, Russia) are examined across a salinity gradient. As salinity increased from 0.8 to 6.4 g l⁻¹, the species richness of aquatic vascular plants decreased from 16 to 2 species, of phytoplankton from 98 to 52 species, and of zooplankton from 61 to 16 species, but changes in species diversity of zoobenthos were negligible. Guest Editor: John M. Melack Saline Waters and their Biota     

The role of solid waste materials as habitats for macroinvertebrates in a lowland dam reservoir

Eight hypereutrophic phytoplankton dominated ponds from the Brussels Capital Region (Belgium) were biomanipulated (emptied with fish removal) to restore their ecological quality and reduce the risk of cyanobacterial bloom formation. Continuous monitoring of the ponds before and after the biomanipulation allowed the effects of the management intervention on different compartments of pond ecosystems (phytoplankton, zooplankton, submerged vegetation and nutrients) to be assessed. Fish removal resulted in a drastic reduction in phytoplankton biomass and a shift to the clear-water state in seven out of eight biomanipulated ponds. The reduction in phytoplankton biomass was associated with a marked increase in density and size of large cladocerans in six ponds and a restoration of submerged macrophytes in five ponds. The phytoplankton biomass in the ponds with extensive stands of submerged macrophytes was less affected by planktivorous fish recolonisation of some of the ponds later in the summer. The two non-vegetated ponds as well as one pond with sparse submerged vegetation showed a marked increase in phytoplankton biomass associated with the appearance of fish. Phytoplankton biomass increase coincided with the decrease in large Cladocera density and size. One pond lacking submerged macrophytes could maintain very low phytoplankton biomass owing to large Cladocera grazing alone. The results of this study confirmed the importance of large zooplankton grazing and revegetation with submerged macrophytes for the maintenance of the clear-water state and restoration success in hypereutrophic ponds. They also showed that large Cladocera size is more important than their number for efficient phytoplankton control and when cladocerans are large enough, they can considerably restrain phytoplankton growth, including bloom-forming cyanobacteria, even when submerged vegetation is not restored. The positive result of fish removal in seven out of eight biomanipulated ponds clearly indicated that such management intervention can be used, at least, for the short-term restoration of ecological water quality and prevention of noxious cyanobacterial bloom formation. The negative result of biomanipulation in one pond seems to be related to the pollution by sewage water. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

Biological indicators track differential responses of pelagic and littoral areas to nutrient load reductions in German lakes

External nutrient loading was reduced over the past decades as a measure for improving the water quality of eutrophic lakes in western Europe, and has since been accelerated by the adoption of the European Water Framework Directive (WFD) in 2000 (EC, 2000). A variety of eutrophication-related metrics have indicated that the response of biological communities to this decreased nutrient loading has been diverse. Phytoplankton, a major component of the pelagic community, often responded rapidly, whereas a significant delay was observed for submerged macrophytes colonizing littoral areas. In this study we tested whether assessment methods developed for phytoplankton and macrophytes in lakes during Germany's implementation of the WFD reflect this differential response. An assessment of 263 German lakes confirmed that a lower ecological state was recorded when based on the biological quality element (BQE) for macrophytes than the BQE for phytoplankton during the investigated period (2003–2012). On average, lakes had a moderate ecological status for both phytoplankton and macrophyte BQEs, but differences of up to three classes were observed in single cases. Long-term data were available for five lowland lakes subject to strong reductions in phosphorus loading. Their phytoplankton-based assessments indicated a constant improvement of the ecological status in parallel to decreasing water phosphorus concentrations. In contrast, macrophyte-based assessments indicated a 10–20 year delay in their ecological recovery following nutrient load reduction. This delay was confirmed by detailed data on the temporal development of macrophyte species diversity and maximum colonization depths of two lakes after nutrient load reduction. We conclude that the available WFD assessment methods for phytoplankton and macrophyte BQEs are suitable to track the differential response of pelagic and littoral areas to nutrient load reductions in German lakes.     

Asymmetrical food web responses in trophic-level richness, biomass, and function following lake acidification

We tested for disproportional changes in annual and seasonal species richness and biomass among five trophic levels (phytoplankton, herbivorous, omnivorous, and carnivorous zooplankton, and fish) as well as altered trophic structure and ecosystem function following the 5-year experimental acidification of Little Rock Lake (Wisconsin, USA) from pH 6.1 to 4.7. Abiotic and biotic controls of trophic level response during acidification were also identified. Asymmetric reductions of species richness among trophic levels, separated by life stage and feeding type, were evident and changes in trophic structure were most pronounced by the end of the acidification period. Relative declines in richness of fish and zooplankton were greater than phytoplankton, which were generally unaffected, leading to a reduction of upper trophic level diversity. Each of the lower four trophic levels responded to a distinct combination of abiotic and biotic variables during acidification. pH was identified as a direct driver of change for only carnivorous zooplankton, while all other trophic levels were affected more by indirect interactions caused by acidification. Fluctuations in ecosystem function (zooplankton biomass and primary production) were also evident, with losses at all trophic levels only detected during the last year of acidification. The acidified basin displayed a tendency for greater variation in biomass for upper trophic levels relative to reference conditions implying greater unpredictability in ecosystem function. Together, these results suggest that trophic asymmetry may be an important and recurring feature of ecosystem response to anthropogenic stress.     

Some aspects of the seasonal distribution of flagellates in mountain lakes

In a larger regional survey in Tyrol, phytoplankton species composition and biovolume of mid-altitude and high-mountain lakes was studied. Results from eight lakes showed that flagellates (mainly Chrysophyceae, Dinophyceae, and Cryptophyceae) are important components of the phytoplankton.In the mid-altitude lakes a spring and an autumn maximum of Chrysophyceae as well as a summer maximum of large dinoflagellates are observed, whereas Cryptophyceae and Dinophyceae show irregular distributions. In the high-mountain lakes the seasonal variations of phytoplankton, including flagellates, are limited by the long duration of the winter situation. However similar sequences of phytoplankton assemblages as in the midaltitude lakes can be observed. Flagellates in high-mountain lakes are important to sustain phytoplankton standing crop under the winter snow and ice cover.In order to show similarities and differences of high-mountain and mid-altitude lakes, vertical profiles of phytoplankton from three lakes and seasonal patterns of Gymnodinium uberrimum from two lakes are compared. In addition the patterns of cryptomonads differing in their ecological requirements (Cryptomonas spp. and Rhodomonas minuta) are shown for a meromictic mid-altitude lake.     

Changes in bacterial and ciliate densities with trophic status in Mediterranean shallow lakes

Ciliate and bacterial densities and their link with eutrophication were studied in fourteen shallow lakes in northwest Spain. Total phosphorus (TP) in these lakes varied between 30 μg l⁻¹ and 925 μg l⁻¹ and chlorophyll a concentration (chla) between 0.5 μg l⁻¹ and 107 μg l⁻¹. Bacterial abundance ranged from 1 × 10⁶ to 14 × 10⁶ cells ml⁻¹, while ciliate abundance ranged from 0.6 cells ml⁻¹ to 229 cells ml⁻¹. Lakes were classified into three trophic types from their TP and chla concentrations. Bacterial abundance was significantly correlated with trophic type, as well as with TP and with chla separately, whereas ciliate abundance was only correlated with chla. No significant relationship could be established between bacterial and ciliate abundance across the trophic gradient. A general pattern was observed in the ratios of bacterial abundance to TP and chla concentrations, of decreasing ratios with increases in the nutrient loading. This pattern was not found for ciliates. The dominant zooplankton group in 13 of the 14 lakes studied was Rotifera, which accounted for a mean of 71% of total zooplankton abundance (41% of zooplankton biomass). The positive correlation between bacteria and ciliates with this group, and the absence of any relationship with Cladocera suggest that top down control by cladocerans was weaker in our lakes than previously shown in northern European shallow lakes. Rotifers could be important predators of bacteria in the high-nutrient lakes of our study. Higher slopes of regressions on bacterial abundance towards the hypertrophic range indicate that top-down control was weaker in our lakes than in northern European shallow lakes.     

REBECCA databases: experiences from compilation and analyses of monitoring data from 5,000 lakes in 20 European countries

Chemical and biological data from more than 5,000 lakes in 20 European countries have been compiled into databases within the EU project REBECCA. The project’s purpose was to provide scientific support for implementation of the EU Water Framework Directive (WFD). The databases contain the biological elements phytoplankton, macrophytes, macroinvertebrates and fish, together with relevant chemistry data and station information. The common database strategy has enabled project partners to perform analyses of chemical–biological relationships and to describe reference conditions for large geographic regions in Europe. This strategy has obvious benefits compared with single-country analyses: results will be more representative for larger European regions, and the statistical power and precision will be larger. The high number of samples within some regions has also enabled analysis of type-specific relationships for several lake types. These results are essential for the intercalibration of ecological assessment systems for lakes, as required by the WFD. However, the common database approach has also involved costs and limitations. The data process has been resource-demanding, and the requirements for a flexible database structure have made it less user-friendly for project partners. Moreover, there are considerable heterogeneities among datasets from different countries regarding sampling methods and taxonomic precision; this may reduce comparability of the data and increase the uncertainty of the results. This article gives an overview of the contents and functions of the REBECCA Lakes databases, and of our experiences from constructing and using the databases. We conclude with recommendations for compilation of environmental data for future international projects.     

Morpho-Functional Groups and phytoplankton development in two deep lakes (Lake Garda,Italy and Lake Stechlin,Germany)

Phylogenetic classifications of plants often do not reflect their ecological functions. In fact, the functional mechanisms of biological communities may be better understood if species are pooled into groups having similar characteristics. The objective of this work is to evaluate, with the use of multivariate methods, classifications based on the morphological and functional characteristics (size and form, mobility, potential mixotrophy, nutrient requirements, presence of gelatinous envelopes) of cyanobacteria and eukaryotic algae to explain the seasonal dynamic of the phytoplankton community. The analyses involve data from two deep lakes: Lake Garda, southern Alps, z _max = 350 m; biennium 2002–2003) and Lake Stechlin (north-east Germany, z _max = 67 m; 1995, 1998 and 2001). In both lakes, the temporal evolution of the phytoplankton communities within individual years followed a regular annual cycle, with the exception of Lake Stechlin in 1998, when an irregular phytoplankton pattern was caused by a sudden mass appearance of Planktothrix rubescens in the spring and summer months, resulting in a collapse of the whole community in autumn. Overall, the temporal developments of the phytoplankton communities obtained on the basis of patterns of the morpho-functional groups appeared highly comparable with those obtained, in the single years, on the basis of the original phytoplankton species matrices. The comparison of the morpho-functional groups of the lakes Garda and Stechlin showed important differences in the abundance and seasonality of the dominant phytoplankton types. The results obtained in this study underline that the use of classifications based on the adaptive strategies of the single species may represent a useful tool to investigate the community evolution and to compare phytoplankton assemblages of different lakes, overcoming problems related to possible differences of taxonomic accuracy and identification.     

Study of biocomplexity in an aquatic ecosystem through ascendency

The ascendancy concept aims at quantitatively describing the growth and development of an ecosystem as whole. Growth is an increase in the total system throughflow, while development is taken to be a rise in the average mutual information inherent in the network flow structure. As an ecosystem matures and goes through a series of successional stages, its ascendancy exhibits a propensity to increase. In any ecosystem the equilibrium condition may gradually turn into a chaotic situation for different reasons. In this paper a model is proposed of an aquatic ecosystem comprising of three groups, viz., phytoplankton, zooplankton and fish. Rate parameters are changed according to the change of the size of the organisms. The model is run in different conditions with gradual decrement of the body sizes of zooplankton. Allometric principle of the relationship of body size of zooplankton and two rate parameters (growth rate and half saturation constant) are incorporated in this model. According to allometric principle gradual decrement of body sizes of zooplankton consequently increases the grazing rate and decreases the half-saturation constant of this organisms. The system exhibits different states (equilibrium point--stable limit cycle--doubling and ultimately chaos) by gradual increase of zooplankton grazing rate and decrease of half-saturation constant. This paper tests the high level of ascendancy of the systems at the edge of oscillation before starting of the chaos. This high level of throughflow and mutual information, i.e. Ascendency supports the hypothesis that the system can coordinate the most complex behavior and shows maximum biocomplexity in this situation.     

Indirect effects of fish community structure on submerged vegetation in shallow,eutrophic lakes: an alternative mechanism

The loss of submerged macrophytes during eutrophication of shallow lakes is a commonly observed phenomenon. The proximate reason for this decline is a reduction of available light due to increasing phytoplankton and/or epiphyton biomass. Here we argue that the ultimate cause for the transition from a macrophyte-dominated state to a phytoplankton-dominated state is a change in fish community structure. A catastrophic disturbance event (e.g. winterkill) acting selectively on piscivores, cascades down food chains, eventually reducing macrophyte growth through shading by epiphyton, an effect that is reinforced by increasing phytoplankton biomass. The transition back from the phytoplankton to the macrophyte state depends on an increase in piscivore standing stock and a reduction of planktivores. A conceptual model of these mechanisms is presented and supported by literature data and preliminary observations from a field experiment.     

Modelling production and biomasses of zoobenthos in lakes

This work presents a dynamic model to predict zoobenthos in lakes. The model has been developed within the framework of a more comprehensive lake ecosystem model, LakeWeb, which also accounts for the following functional groups of organisms, phytoplankton, bacterioplankton, two types of zooplankton (herbivorous and predatory), macrophytes, prey fish and predatory fish. This work also presents a new data-base for zoobenthos in lakes. Many of the lakes included in this study are situated in the former Soviet Union. They were investigated during the Soviet period and those results have been largely unknown in the West. Using this data-base, this work also presents new empirical models for zoobenthos. The new dynamic model gives seasonal variations (the calculation time, dt, is 1 week using Euler's method and enough iterations to get stable solutions). The basic aim of the dynamic model is that it should capture general functional and structural patterns in lakes. We have demonstrated by several model tests along limnological gradients (total phosphorus concentrations, pH, lake colour, latitude and lake size) that the dynamic model gives predictions that agree well with the values given by the empirical regressions, and also expected and requested divergences from these regressions when they do not provide sufficient resolution. It would have been very difficult indeed to carry out such tests regarding ecosystem responses using traditional methods with extensive field studies in a few lakes. We have given algorithms for (1) production of zoobenthos from eating macrophytes, benthic algae and sediments, (2) elimination (related to the turnover time of zooplankton), and (3) zoobenthos consumption by prey fish, and the factors influencing these processes/rates. The model is driven by data easily accessed from standard monitoring programs or maps a prerequisite for practical utility in contexts of lake management.