Empirical and dynamical models of production and biomass of benthic algae in lakes

This work presents new empirical and dynamical models for benthic algae in lakes. The models were developed within the framework of a more comprehensive lake ecosystem model, LakeWeb, which also accounts for phytoplankton, bacterioplankton, two types of zooplankton (herbivorous and predatory), macrophytes, prey fish and predatory fish. The new dynamic model provides seasonal variations (the calculation time is 1 week). It is meant to account for all factors regulating the production and biomass of benthic algae for lakes in general. This work also presents and uses a new data-base established by us from published sources. Many of the lakes included in this study are situated in the former Soviet Union. They were investigated during the Soviet period and the data and results have up until now been largely unknown in the West. We present empirical models for benthic algae, and show that the biomass of benthic algae in whole lakes can be estimated from the ratio between the lake area above the Secchi depth to the total lake area and the primary production of phytoplankton. We also present several critical tests of the dynamical model. The dynamical and empirical models give corresponding results over a wide limnological domain. We provide algorithms for (1) the production rate of benthic algae (2) the elimination rate (related to the turnover time of benthic algae), (3) the rate of benthic algae consumption by zoobenthos, and (4) the rate of physical erosion of benthic algae. Our results indicate that the production of benthic algae is highly dependent on lake morphometry and sediment character, as well as water clarity, and less dependent on nutrient (phosphorus) concentrations in water and sediments. This work provides new quantitative support to such conclusions and also a useful model for predictions of production and biomass of benthic algae.     

Modelling Production and Biomasses of Prey and Predatory Fish in Lakes

This work presents a new dynamic model to predict two fundamental functional categories of fish in lakes, prey and predatory fish. The model has been developed within the framework of a more comprehensive lake ecosystem model, LakeWeb, which also accounts for phytoplankton, bacterioplankton, two types of zooplankton (herbivorous and predatory), zoobenthos, macrophytes and benthic algae. The new fish model gives seasonal variations (the calculation time is 1 week). It is meant to account for all factors regulating the production of fish for lakes in general. The model has not been calibrated and tested in the traditional way using data from a few well investigated lakes. Instead, it has been tested using empirical data and regressions based on data from many lakes. The basic aim of the model is that it should capture typical functional and structural patterns in many lakes. It accounts in a relatively simple manner for many complicated processes, like fishing (by birds, animals and man), fish migration to and from lakes and how macrophyte cover gives shelter to small fish and reduces predation pressure. Food choices are handled by distribution coefficients regulating how much of the different available food sources a given organism would consume. Beside these distribution coefficients, and the way the food choices are structured (the food choice panel), fundamental concepts in the fish model are: (1) metabolic efficiency ratios, which express how much of the food consumed by the predator that will increase the biomass of the predator and how much that will be lost by respiration and faeces, (2) actual consumption rates, which are defined from the ratio between the actual biomass of the predator and the normal biomass of the predator, and the normal consumption rates, which are related to the turnover time of the predator. We have demonstrated that the new model gives predictions which agree well with the values given by the empirical regressions, and also expected and requested divergences from these regression lines when they do not provide sufficient resolution.     

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.     

A mass-balance model for phosphorus in lakes accounting for biouptake and retention in biota

SUMMARY 1. A new mass-balance model was developed to predict phosphorus fluxes to, within and from lakes and lake concentrations of phosphorus.
2. The model is driven by data easily accessed from standard monitoring programmes or maps, so it should be useful in many contexts of lake management.
3. The phosphorus model gives seasonal (weekly) variations and accounts for all fundamental abiotic/biotic interactions and feedbacks for lakes in general for phosphorus. The model has been developed within the framework of a more comprehensive ecosystem model, LakeWeb, which accounts for production values and biomasses of nine functional groups of organisms (phytoplankton, bacterioplankton, herbivorous and predatory zooplankton, prey and predatory fish, zoobenthos, macrophytes and benthic algae).
4. The LakeWeb-model has been critically tested in a wide limnological domain and shown to predict very well. By using this model, it is possible for the first time to predict realistic values of biouptake and retention of phosphorous in biota in lakes on a weekly basis.     

The Importance of Lake Morphometry for the Structureand Function of Lakes

This work demonstrates quantitatively and in a comprehensive way that the size and form of lakes regulate many general transport processes, such as sedimentation, resuspension, diffusion, mixing, burial and outflow, which in turn regulate many abiotic state variables, such as concentrations of phosphorus, suspended particulate matter, many water chemical variables and water clarity, which in turn regulate primary production, which regulate secondary production, for example of zooplankton and fish. Such relationships are discussed not qualitatively but quantitatively using a new generation of validated dynamic ecosystem models (LakeWeb and LakeMab) based on mechanistic principles. It has been shown by critical model tests (including blind tests using data covering wide limnological ranges) that these models give predictions that agree well with empirical data. This should lend credibility to the results presented in this work, which would have been very difficult to obtain using traditional methods with extensive field studies in a few lakes. Simulations have been carried out where the inflow of phosphorus is held constant and the consequences simulated for small, large, shallow and deep lakes. There are striking differences in total phosphorus (TP) concentrations and trophic state (from 10 to 100 µg TP/l) and hence also in changes in many variables characterizing lake structure and function, such as Secchi depth, suspended particulate matter, pH, water temperature, chlorophyll, algal volume, macrophyte cover; as well as production and biomasses of benthic algae, bacterioplankton, macrophytes, herbivorous zooplankton, predatory zooplankton, zoobenthos, prey fish and predatory fish. These changes have been quantified in a comprehensive manner in this work and the approach to calculate such changes are basic for an understanding of how different lakes react to changes in nutrient loading (eutrophication). (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The role of macroinvertebrates and fish in regulating the provision by macrophytes of refugia for zooplankton in a warm temperate shallow lake

1. The zooplankton often undergoes diel horizontal migration (DHM) from the open water to the littoral of shallow lakes, thus avoiding predators in the former. This behaviour has functional impacts within the lake, as it enhances zooplankton survival, increases their control of phytoplankton and tends to stabilise the clear water state. However, most of the evidence supporting this migration pattern comes from cold north temperate lakes, and more evidence from tropical and subtropical areas, as well as from southern temperate areas, is needed. 2. We conducted a field study of the diel horizontal and vertical migration of zooplankton, and the horizontal distribution of potential predatory macroinvertebrates and fish, over two consecutive days in the summer in a temperate lake in the southern hemisphere. We took zooplankton samples at two depths, at three sampling stations (inside beds of aquatic macrophytes, at their edge and in open water) along three transects running from the centre of a bed of Ceratophyllum demersum to open water. At each sampling station, we also took samples of macroinvertebrates and fish and measured physical and chemical environmental variables. 3. Zooplankton (pelagic cladocerans, calanoid copepods and rotifers) avoided the shore, probably because of the greater risk from predators there. Larger and more vulnerable cladocerans, such as Diaphanosoma brachyurum and Moina micrura, were two to four times more abundant in open water than at the edge of or inside beds of macrophytes, respectively, by both day and night. Less vulnerable zooplankton [i.e. of medium body size (Ceriodaphnia dubia) or with the ability to swim fast (calanoid copepods)] were distributed evenly between open water and the edge of the plant beds. Small zooplankton, Bosmina huaronensis and pelagic rotifers, showed an even distribution among the three sampling stations. Accordingly, no DHM of zooplankton occurred, although larger organisms migrated vertically inside C. demersum stands. 4. Macrophytes contained high densities of predatory macroinvertebrates and fish. The predator assemblage, composed of large‐bodied macroinvertebrates (including odonates and shrimps) and small littoral fish, was permanently associated with submerged macrophytes. None of these groups moved outside the plant beds or changed their population structure (fish) over the diel cycle. 5. Submerged macrophyte beds do not represent a refuge for zooplankton in lakes where predators are numerous among the plants, implying a weaker top‐down control of phytoplankton biomass by zooplankton and, consequently, a more turbid lake. The effectiveness of macrophytes as a refuge for zooplankton depends on the associated assemblage of predatory macroinvertebrates and fish among the plants.     

Predators, resources, and trophic chains in the regulation of plankton population and biomass in oligothrophic lakes

The relative strength of "top-down" versus "bottom-up" control of plankton community structure and biomass in two small oligotrophic lakes (with and without fish), located near the Polar circle (Russia), has been investigated for two years, 1996 and 1997. The comparative analyses of zooplankton biomass and species abundance showed strong negative effect of fish, stickeback (Pungitius pungitius L.), on the zooplankton community species, size structure and biomass of particular prey species but no effect on the biomass of the whole trophic level. An intensive predation in Verkhneye lake has lead to: 1) sixfold decline in biomass of large cladoceran Holopedium gibberum comparing to the lake lacking predator, 2) shift in the size mode in zooplankton community and the replacement of the typical large grazers by small species--Bosmina longirostris and rotifers. Their abundance and biomass even increased, demonstrating the stimulating effect of fish on the "inefficient" and unprofitable prey organisms. The analysis of contributions of different factors into the cladoceran's birth rate changes was applied to demonstrate the relative impact of predators and resources on zooplankton abundance. An occasional introduction of the stickleback to Vodoprovodnoye lake (the reference lake in 1996) in summer 1997 lead to drastic canges in this ecosystem: devastating decrease of zooplankton biomass and complete elimination of five previously dominant grazer species. The abundance of edible phytoplankton was slightly higher in the lake with fish in 1996 and considerably higher in the lake where fish has appeared in 1997 showing the prevailing "top-down" control of phytoplankton in oligotrophic ecosystem. The reasons of trophic cascade appearance in oligotrophic lakes are also discussed.     

高营养级捕食者在浅水湖泊沿岸带与敞水区能量耦合的维持作用

研究利用保安湖沿岸带与敞水区两种生境中高营养级捕食者(达氏鲌与红鳍原鲌)的碳(δ13C)、氮(δ15N)稳定同位素数据,通过稳定同位素质量平衡混合模型、非度量多维尺度分析(NMDS)等方法,分析了两种生境中鲌类食物来源的差异.结果表明,两种生境中鲌类食物来源基本一致,且食物来源较为广泛,包括沿岸带饵料鱼类、敞水区饵料鱼...     

Enhanced Input of Terrestrial Particulate Organic Matter Reduces the Resilience of the Clear-Water State of Shallow Lakes: A Model Study

The amount of terrestrial particulate organic matter (t-POM) entering lakes is predicted to increase as a result of climate change. This may especially alter the structure and functioning of ecosystems in small, shallow lakes which can rapidly shift from a clear-water, macrophyte-dominated into a turbid, phytoplankton-dominated state. We used the integrative ecosystem model PCLake to predict how rising t-POM inputs affect the resilience of the clear-water state. PCLake links a pelagic and benthic food chain with abiotic components by a number of direct and indirect effects. We focused on three pathways (zoobenthos, zooplankton, light availability) by which elevated t-POM inputs (with and without additional nutrients) may modify the critical nutrient loading thresholds at which a clear-water lake becomes turbid and vice versa. Our model results show that (1) increased zoobenthos biomass due to the enhanced food availability results in more benthivorous fish which reduce light availability due to bioturbation, (2) zooplankton biomass does not change, but suspended t-POM reduces the consumption of autochthonous particulate organic matter which increases the turbidity, and (3) the suspended t-POM reduces the light availability for submerged macrophytes. Therefore, light availability is the key process that is indirectly or directly changed by t-POM input. This strikingly resembles the deteriorating effect of terrestrial dissolved organic matter on the light climate of lakes. In all scenarios, the resilience of the clear-water state is reduced thus making the turbid state more likely at a given nutrient loading. Therefore, our study suggests that rising t-POM input can add to the effects of climate warming making reductions in nutrient loadings even more urgent.     

Macrophyte refuges, prey behaviour and trophic interactions: consequences for lake water clarity

Macrophytes may enhance grazing on phytoplankton by providing a refuge for zooplankton against fish predation. Loss of macrophytes can trigger sudden degradation of water clarity (regime shift) in lakes. However, the presence of piscivores may drive planktivorous fish to take refuge amongst littoral macrophytes. To address the possibility of regime shifts, I here constructed an empirically based model that combined population dynamics of organisms with game theory for optimal habitat selection, taking into consideration the trophic structure, lake size and eutrophication. The model showed that macrophytes generally acted as a refuge for zooplankton, rather than for fish. The model predicted that regime shifts were more likely in small, shallow lakes and that the presence of macrophytes raised the possibility of regime shifts. The present study demonstrated that the fast dynamics of animal behaviour could lead to regime shifts, in connection with slower variables such as nutrient loading.     

Long-Term Changes in the Diet of Gymnogobius isaza from Lake Biwa,Japan: Effects of Body Size and Environmental Prey Availability

Body size and environmental prey availability are both key factors determining feeding habits of gape-limited fish predators. However, our understanding of their interactive or relative effects is still limited. In this study, we performed quantitative dietary analysis of different body sizes of goby (Gymnogobius isaza) specimens collected from Lake Biwa between 1962 and 2004. First, we report that the diet was composed mainly of zooplankton (cladocerans and copepods) before the 1980s, and thereafter, shifted to zoobenthos (gammarids). This foraging shift coincided with, and thus can be linked to, known historical events in the lake at that time: decrease in zooplankton abundance with the alleviation of eutrophication, increase in fish body size resulting from fish population collapse, and increase in gammarid abundance due to reduced fish predation pressure. Supporting this view, our data analyses revealed how the long-term changes in the diet composition would be co-mediated by changes in fish body size and environmental prey availability. Specifically, while zoobenthos abundance strongly affected the fish diet composition, larger (smaller) fish preferred zoobenthos (zooplankton). Furthermore, the body size effects were stronger than those of prey availability. These results provide the best long-term evidence that fish feeding habits vary over decades with its body size and prey community due to anthropogenic disturbances.     

Effects of habitat complexity on community structure and predator avoidance behaviour of littoral zooplankton in temperate versus subtropical shallow lakes

1. Structural complexity may stabilise predator–prey interactions and affect the outcome of trophic cascades by providing prey refuges. In deep lakes, vulnerable zooplankton move vertically to avoid fish predation. In contrast, submerged plants often provide a diel refuge against fish predation for large‐bodied zooplankton in shallow temperate lakes, with consequences for the whole ecosystem. 2. To test the extent to which macrophytes serve as refuges for zooplankton in temperate and subtropical lakes, we introduced artificial plant beds into the littoral area of five pairs of shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N). We used plants of different architecture (submerged and free‐floating) along a gradient of turbidity over which the lakes were paired. 3. We found remarkable differences in the structure (taxon‐richness at the genus level, composition and density) of the zooplankton communities in the littoral area between climate zones. Richer communities of larger‐bodied taxa (frequently including Daphnia spp.) occurred in the temperate lakes, whereas small‐bodied taxa characterised the subtropical lakes. More genera and a higher density of benthic/plant‐associated cladocerans also occurred in the temperate lakes. The density of all crustaceans, except calanoid copepods, was significantly higher in the temperate lakes (c. 5.5‐fold higher). 4. Fish and shrimps (genus Palaemonetes) seemed to exert a stronger predation pressure on zooplankton in the plant beds in the subtropical lakes, while the pelagic invertebrate Chaoborus sp. was slightly more abundant than in the temperate lakes. In contrast, plant‐associated predatory macroinvertebrates were eight times more abundant in the temperate than in the subtropical lakes. 5. The artificial submerged plants hosted significantly more cladocerans than the free‐floating plants, which were particularly avoided in the subtropical lakes. Patterns indicating diel horizontal migration were frequently observed for both overall zooplankton density and individual taxa in the temperate, but not the subtropical, lakes. In contrast, patterns of diel vertical migration prevailed for both the overall zooplankton and for most individual taxa in the subtropics, irrespective of water turbidity. 6. Higher fish predation probably shapes the general structure and dynamics of cladoceran communities in the subtropical lakes. Our results support the hypothesis that horizontal migration is less prevalent in the subtropics than in temperate lakes, and that no predator‐avoidance behaviour effectively counteracts predation pressure in the subtropics. Positive effects of aquatic plants on water transparency, via their acting as a refuge for zooplankton, may be generally weak or rare in warm lakes.     

OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

Water level and fish-mediated cascading effects on the microbial community in eutrophic warm shallow lakes: a mesocosm experiment

Information on the effects of water level changes on microbial planktonic communities in lakes is limited but vital for understanding ecosystem dynamics in Mediterranean lakes subjected to major intra- and inter-annual variations in water level. We performed an in situ mesocosm experiment in an eutrophic Turkish lake at two different depths crossed with presence/absence of fish in order to explore the effects of water level variations and the role of top-down regulation at contrasting depths. Strong effects of fish were found on zooplankton, weakening through the food chain to ciliates, HNF and bacterioplankton, whereas the effect of water level variations was overall modest. Presence of fish resulted in lower biomass of zooplankton and higher biomasses of phytoplankton, ciliates and total plankton. The cascading effects of fish were strongest in the shallow mesocosms as evidenced by a lower zooplankton contribution to total plankton biomass and lower zooplankton:ciliate and HNF:bacteria biomass ratios. Our results suggest that a lowering of the water level in warm shallow lakes will enhance the contribution of bacteria, HNF and ciliates to the plankton biomass, likely due to increased density of submerged macrophytes (less phytoplankton); this effect will, however, be less pronounced in the presence of fish.     

Impact of fish predation on cladoceran body weight distribution and zooplankton grazing in lakes during winter

1. It is well accepted that fish, if abundant, can have a major impact on the zooplankton community structure during summer, which, particularly in eutrophic lakes, may cascade to phytoplankton and ultimately influence water clarity. Fish predation affects mean size of cladocerans and the zooplankton grazing pressure on phytoplankton. Little is, however, known about the role of fish during winter. 2. We analysed data from 34 lakes studied for 8–9 years divided into three seasons: summer, autumn/spring and winter, and four lake classes: all lakes, shallow lakes without submerged plants, shallow lakes with submerged plants and deep lakes. We recorded how body weight of Daphnia and then cladocerans varied among the three seasons. For all lake types there was a significant positive correlation in the mean body weight of Daphnia and all cladocerans between the different seasons, and only in lakes with macrophytes did the slope differ significantly from one (winter versus summer for Daphnia). 3. These results suggest that the fish predation pressure during autumn/spring and winter is as high as during summer, and maybe even higher during winter in macrophyte‐rich lakes. It could be argued that the winter zooplankton community structure resembles that of the summer community because of low specimen turnover during winter mediated by low fecundity, which, in turn, reflects food shortage, low temperatures and low winter hatching from resting eggs. However, we found frequent major changes in mean body weight of Daphnia and cladocerans in three fish‐biomanipulated lakes during the winter season. 4. The seasonal pattern of zooplankton : phytoplankton biomass ratio showed no correlation between summer and winter for shallow lakes with abundant vegetation or for deep lakes. For the shallow lakes, the ratio was substantially higher during summer than in winter and autumn/spring, suggesting a higher zooplankton grazing potential during summer, while the ratio was often higher in winter in deep lakes. Direct and indirect effects of macrophytes, and internal P loading and mixing, all varying over the season, might weaken the fish signal on this ratio. 5. Overall, our data indicate that release of fish predation may have strong cascading effects on zooplankton grazing on phytoplankton and water clarity in temperate, coastal situated eutrophic lakes, not only during summer but also during winter.     

Community structure and diel migration of zooplankton in shallow brackish lakes: role of salinity and predators

Diel horizontal migration (DHM), where zooplankton moves towards macrophytes during daytime to avoid planktivorous fish, has been reported as a common migration pattern of zooplankton in shallow temperate freshwater lakes. However, in shallow eutrophic brackish lakes, macrophytes seem not to have the same refuge effect, as these lakes may remain turbid even at relatively high macrophyte abundances. To investigate the extent to which macrophytes serve as a refuge for zooplankton at different salinities, we introduced artificial plants mimicking submerged macrophytes in the littoral zone of four shallow lakes, with salinities ranging from almost freshwater (0.3) to oligohaline waters (3.8). Furthermore, we examined the effects of different salinities on the community structure. Diel samples of zooplankton were taken from artificial plants, from areas where macrophytes had been removed (intermediate areas) and, in two of the lakes, also in open water. Fish and macroinvertebrates were sampled amongst the artificial plants and in intermediate areas to investigate their influence on zooplankton migration. Our results indicated that diel vertical migration (DVM) was the most frequent migration pattern of zooplankton groups, suggesting that submerged macrophytes were a poor refuge against predation at all salinities under study. Presumably, this pattern was the result of the relatively high densities of small planktivorous fish and macroinvertebrate predators within the submerged plants. In addition, we found major differences in the composition of zooplankton, fish and macroinvertebrate communities at the different salinities and species richness and diversity of zooplankton decreased with increasing salinity. At low salinities both planktonic/free-swimming and benthic/plant-associated cladocerans occurred, whilst only benthic ones occurred at the highest salinity. The low zooplankton biomass and overall smaller-bodied zooplankton specimens may result in a lower grazing capacity on phytoplankton, and enhance the turbid state in nutrient rich shallow brackish lakes.     

Modeling the foodweb in coastal areas: a case study of Ringkøbing Fjord,Denmark

This work utilizes the CoastWeb model, a foodweb model for coastal areas that also includes a mass-balance model (CoastMab) for phosphorus and many abiotic/biotic interactions, to study the development in Ringkøbing Fjord, Denmark, from 1985 to 2004. This shallow coastal lagoon has an area of 300 km² and a mean depth of 1.9 m. The water exchange between the lagoon and the North Sea is regulated by a sluice. In 1996 there was a major regime shift in this lagoon with drastic reductions in chlorophyll-a concentrations, significant increases in water clarity (Secchi depth) and major changes in the number and biomass of clams as well as in macrophyte cover. Regime shifts is a “hot” topic in aquatic ecology and in this work the CoastWeb model is used as a tool to understand and quantify the causes behind this regime shift. The CoastWeb model is general and can also be used for other coastal areas. The basic model calculates monthly production values and changes in biomasses of ten functional groups of organisms (phytoplankton, bacterioplankton, herbivorous, and predatory zooplankton, benthic algae, macrophytes, jellyfish, zoobenthos and prey and predatory fish) and in Ringkøbing Fjord, also for clams (Mya arenaria). In spite of its complexity, the model is relatively simple to use, since all driving variables may be readily accessed from maps or monitoring programs. The model includes much abiotic/biotic feedback and it can also be used to address other causes for regime shifts other than the changes in salinity and nutrient inflow, which have caused the changes in Ringkøbing Fjord. The model has previously been tested for more than 20 smaller coastal areas and was shown to predict variations in foodweb characteristics very well. The focus of this paper is on temporal variations within one well-studied coastal area. The paper compares modeled values to empirical data for Ringkøbing Fjord and discusses fundamental ecosystem features such as regime shifts and compensatory effects in a way that is not practically feasible without the use of quantitative models.     

Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

Changes in phosphorus and nitrogen cycling following food web manipulations in a shallow Dutch lake

1. The flow of phosphorus and nitrogen through the food web of the shallow, eutrophic lake Wolderwijd was analysed for 2 different years before and for 1 year after food web manipulation.
2. After fish removal the water became clear and the growth of macrophytes began. Fish removal resulted in a significant reduction of the total nutrient pool in the water, but differences between the nutrient cycles before and after the experiment were mainly caused by a gradual change driven by a reduced phosphorus input.
3. The zooplankton biomass before and after food web manipulation did not change significantly. Unfavourable food conditions and predation by young fish limited zooplankton biomass after the food web manipulation.
4. After fish removal benthic algae, fish, zoobenthos and macrophytes form the largest pools of nutrients apart from the sediment top layer. However, they contribute only little to nutrient cycles in the water column.     

A LIMNOLOGICAL SYNOPSIS OF BHANGAZI SOUTH,A DYSTROPHIC COASTAL LAKE IN THE GREATER ST LUCIA WETLAND PARK (KWAZULU/NATAL), WITH COMMENTS ON ITS CONSERVATION VALUE.

Summary

Base-line limnological and biological data are given for Lake Bhangazi South, a small lake on the coastal plain of Kwazulu-Natal, South Africa. This shallow (Z_max < 6 m), subtropical lake is a warm and seemingly continuously polymictic system, but experiences severe (continuous?) deoxygenation of its deeper muddy sediments. Nutrient status (of N in particular), light attenuation, phytoplankton productivity and zooplankton biomass are high relative to other regionally comparable coastal lakes investigated. While the ichthyofauna is quite rich, zooplankton and especially zoobenthos communities are species-poor, and lack the relict estuarine components which often dominate these latter assemblages in comparable coastal lakes. The zooplankton is typically freshwater in composition, and contains a new species of copepod (Tropodiaptomus bhangazii Rayner); the lake may be older, or seen longer evolutionary divergence than its proximate counterparts. Benthic species diversity in the lake is especially low amongst the Crustacea and Mollusca, possibly reflecting the effect of relatively acid waters (pH ca 6.5) on the calcium budgets of these groups. Scanty observational data indicative of ecosystem changes over the past 35 years are reported and briefly evaluated from a conservation/management perspective. Along with the endemic copepod, several rare fish species give the lake added conservation status.