Above- and belowground competition between two submersed macrophytes

Several studies have shown that submerged macrophytes provide a refuge for zooplankton against fish predation, whereas the role of emergent and floating-leaved species, which are often dominant in eutrophic turbid lakes, is far less investigated. Zooplankton density in open water and amongst emergent and floating-leaved vegetation was monitored in a small, eutrophic lake (Frederiksborg Slotssø) in Denmark during July–October 2006. Emergent and floating-leaved macrophytes harboured significantly higher densities of pelagic as well as plant-associated zooplankton species, compared to the open water, even during periods where the predation pressure was presumably high (during the recruitment of 0+ fish fry). Zooplankton abundance in open water and among vegetation exhibited low values in July and peaked in August. Bosmina and Ceriodaphnia dominated the zooplankton community in the littoral vegetated areas (up to 4,400 ind l^?1 among Phragmites australis and 11,000 ind l^?1 between Polygonum amphibium stands), whereas the dominant species in the pelagic were Daphnia (up to 67 ind l^?1) and Cyclops (41 ind l^?1). The zooplankton density pattern observed was probably a consequence of concomitant modifications in the predation pressure, refuge availability and concentration of cyanobacteria in the lake. It is suggested that emergent and floating-leaved macrophytes may play an important role in enhancing water clarity due to increased grazing pressure by zooplankton migrating into the plant stands. As a consequence, especially in turbid lakes, the ecological role of these functional types of vegetation, and not merely that of submerged macrophyte species, should be taken into consideration.     

Seasonal fluctuations in macrophyte cover and water transparency of four brown-water lakes: implications for crustacean zooplankton in littoral and pelagic habitats

The dynamics of crustacean zooplankton in the littoral and pelagic zones of four forest lakes having variable water qualities (colour range 130–340 mg Pt l⁻¹, Secchi depth 70–160 cm) were studied. The biomass of zooplankton was higher in the littoral zone than in the pelagic zone only in the lake having the highest transparency. In the three other lakes, biomass was significantly higher in the pelagic zone than in the littoral zone. In the two lakes with highest transparency, the littoral biomass of cladocerans significantly followed the development of macrophyte vegetation, and cladoceran biomass reached the maximum value at the time of highest macrophyte coverage. In lakes with lowest transparency, littoral zooplankton biomass developed independently of macrophyte density and decreased when macrophyte beds were densest. The seasonal development of the littoral copepod biomass did not follow the development of macrophytes in any of the lakes. The mean size of cladocerans in the pelagic zone decreased with increasing Secchi depth of the lake, whereas in the littoral zone no such phenomenon was detected. Seasonally, when water transparency increased temporarily in two of the lakes, the mean size of cladocerans in the pelagic zone decreased steeply. For copepods, no relationship between water transparency and body size was observed. The results suggested that in humic lakes the importance of the littoral zone as a refuge decreases with decreasing transparency of the water and that low water transparency protects cladocerans from fish predation. All the observed between-lake differences could not be explained by fish predation, but were probably attributed to the presence of chaoborid larvae with variable densities. Feeding efficiency of chaoborids is not affected by visibility and thus they can obscure the relationship between water quality, fish density, and the structure of crustacean zooplankton assemblages. Handling editor: S. I. Dodson     

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.     

The influence of macrophyte beds on plankton communities and their export from fluvial lakes in the St Lawrence River

1. To determine the influence of macrophyte beds on plankton abundance within fluvial lakes of the St Lawrence River, planktonic components (macrozooplankton, heterotrophic bacteria, and phytoplankton as chlorophyll- a [Chl- a ]) were sampled in Lake St Francis and Lake St Pierre during summer 1998. We tested the hypothesis that the abundance of planktonic components was higher within macrophyte beds in comparison to the more rapidly flushed open water areas of the fluvial lakes.
2. The large cross channel variation in zooplankton biomass was indeed correlated with the presence of dense beds of submerged macrophytes. Total macrozooplankton biomass was nine-fold greater within the beds (mean=180 μg L−1 dry mass) than in either the open water or areas with only sparse vegetation (mean=20 μg L−1 dry mass).
3. Chl- a and heterotrophic bacterial abundance were also higher in the beds, but only slightly so. There was no difference in total phosphorus or dissolved organic carbon concentrations between areas of dense vegetation, sparse vegetation or open water.
4. Macrophyte beds on the margins of the fluvial lakes allow the development of high planktonic abundance relative to the fast flowing central channel. Macrozooplankton biomass was much higher at the outflows of the lakes (∼50 μg L−1 dry mass) in comparison to the inflows (<20 μg L−1 dry mass). The increase is due to the transfer of organisms from submerged macrophyte beds into the central channel in the downstream quarter of the two lakes where the marginal littoral waters enter central channel waters.
5. Along rivers, shallow fluvial lakes appear to act as sources of plankton which is exported downstream during years of extensive littoral macrophyte development.     

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.     

Spatial patterns in aquatic vegetation composition and environmental covariates along chains of lakes in the Kokemäenjoki watershed (S. Finland)

The diversity and community structure of macrophyte vegetation was studied in 50 boreal lakes forming several upper reaches of lake chains around Lammi, southern Finland. Water chemical parameters and morphometry of the basins were included in a multivariate analysis. Floating-leaved vegetation was the dominant growth form, followed by emergent plants. In downstream lakes, the dominance of floating-leaved macrophytes declined, and emergent species increased in abundance. Species richness was highest in larger lakes, with a wider range of littoral habitats than smaller lakes. Electrical conductivity (range 18–151 mS cm⁻¹, 25 °C) of the water correlated well with patterns in diversity among lakes, but this was not the case for nutrient concentrations. As a whole, morphometrical characteristics of lake basins showed better correlations with vegetation structure than any of the measured chemical parameters. The macrophyte vegetation of neighbouring lake chains differed considerably, depending on the surrounding landscape properties, water quality of the lakes and immigration history of plant species.     

Contrasting Zooplankton Assemblages in Two Oxbow Lakes with Low Transparencies and Narrow Emergent Macrophyte Belts (Krapina River,Croatia)

The aim of this study was to examine the combined effect of water transparency and narrow macrophyte belts on zooplankton assemblages in two oxbow lakes (Krapina River, Croatia). Samples were collected in open water and among helophytes in the littoral zone from April until September 2008. Rotifers were the most abundant group of zooplankton in both lakes, and dominated in the Krapina oxbow lake 1 (KO1). Lake KO1 had significantly lower transparency, lower percentage macrophyte cover and higher chlorophyll a concentration than Krapina oxbow lake 2 (KO2). In lake KO1, variation in the horizontal distribution of cladocerans and rotifers in terms of their abundance seemed to be determined by competition between Bosmina longirostris and Keratella cochlearis, initiated by oscillation in transparency and detritus availability. In lake KO2, with higher transparency and higher percentage macrophyte cover, the abundance of small‐ and large‐bodied cladocerans increased in the littoral zone simultaneously with higher transparency, suggesting fish predation. Results of this study indicated that small differences in transparencies between the two lakes caused significant differences in horizontal distribution of the zooplankton assemblage. Even narrow helophyte belts offered a refuge to zooplankton, although lower transparencies reduced the effectiveness of macrophytes as a refuge from predators. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Body size and substrate association of littoral insects in relation to vegetation structure

Variation in substrate association types and maximum size of aquatic insects were studied in a vegetated littoral zone of three lake basins. The basins differed from each other in trophic status, biomass of benthivorous fish, and abundance of macrophytes. Four types of substrate association – swimmers, crawlers, semisessiles and burrowers, respectively – were assumed to represent decreasing vulnerability to fish predators. Large-sized species were also hypothesised to be more vulnerable to fish predators. The distributions of species traits were examined in relation to vegetation density. Inferring from ``predation hypothesis' opposite selection pressures on the species traits were expected along the vegetation density. Dense macrophyte beds were thought to be dominated by invertebrate predators and open water by fish predators, since the predation efficiency of fish decreases in complex environments. In the case of invertebrate predator domination, large size and higher activity should be favoured traits among the prey species. Distribution patterns of modes of the two studied traits were explored separately for predatory and non-predatory insects. As expected, swimmers and large-sized crawlers were characteristic of the insect assemblages of dense macrophyte beds. The densities of Odonata, Corixidae, Dytiscidae, Ephemeroptera and Sialidae were higher among macrophytes than in open water, where these insect taxa were possibly depleted by fish. On the other hand, the small-sized and fairly immobile Chironomidae were the most abundant group in open water. These results support the existence of a predator transition zone among littoral vegetation, ranging from domination of invertebrate predation among the dense beds to that of fish predation in open water.     

Role of macrophytes in the variability of zooplankton community structure in the littoral zone of shallow lakes

Distribution, diurnal variability, aggregation of zooplankton in the littoral zone of lakes and effect of various macrophyte species on the structure of its community are considered. It is shown that the horizontal migrations of zooplankton, both direct and reverse ones, are caused mainly by the pressure of fish. The effect of predacious zooplankton is less important and is directed mainly at small-sized species. The intensity of horizontal migrations of zooplankton decreases with depth, while the effect of shore avoidance is observed for the large-sized zooplankton species and is caused not only by the pressure of fish but also by other factors, most likely abiotic. The problem of interaction between macrophytes and zooplankton cannot be reduced to the role of macrophytes as a refuge. Allelopathic properties of macrophytes, competitive relations between separate zooplankton species in macrophyte thickets, as well as the effect of predacious invertebrates associated with macrophytes on zooplankton remain unclear. The role of macrophytes as a factor causing changes in hydrodynamic processes in the littoral regions of lakes is also unknown.     

The relationship between zooplankton community structure and lake characteristics in temperate lakes (Northern Wisconsin, USA)

Zooplankton community structure can be affected by within-lakeand by watershed ecological factors, including water chemistry(related to landscape position), lake morphology and human activityin the watershed. We hypothesized that all three groups of driverswould be correlated with zooplankton species richness and speciescomposition for lakes in northern Wisconsin. Data collectedfrom 52 lakes allowed us to explore the relationship of zooplanktoncommunity structure with ecological drivers. We found that crustaceanzooplankton species richness was not significantly correlatedwith independent environmental variables derived from PCA ordination,nor with measures of community structure based on NMS ordination.However, species composition was correlated with environmentalgradients. Larger zooplankton species (Daphnia pulicaria, Epischuralacustris, Skistodiaptomus oregonensis, Mesocyclops americanus)occurred in large and deep lakes low in the landscape gradient,whereas the smaller species Ceriodaphnia dubia, Daphnia retrocurvaand Leptodiaptomus minutus tended to occur high in the landscape.This shift in species composition was correlated with increasedconductivity, primary productivity and the hypolimnetic refugescharacteristic of larger deeper lakes lower in the landscape.Riparian housing development and littoral zone habitat (measuredas building density and by abundance of logs in littoral zones)were not correlated with zooplankton community structure. Inthese relatively low-impact lakes, natural drivers are stillthe most significant determinants of zooplankton community structure.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Reconstructing the past density of planktivorous fish and trophic structure from sedimentary zooplankton fossils: a surface sediment calibration data set from shallow lakes

1. As quantitative information on historical changes in fish community structure is difficult to obtain directly from fish remains in lake sediments, transfer function for planktivorous fish abundance has been developed based on zooplankton remains in surface sediment (upper 1 cm). The transfer function was derived using weighted average regression and calibration against contemporary data on planktivorous fish catch per unit effort (PF-CPUE) in multiple mesh size gill nets. Zooplankton remains were chosen because zooplankton community structure in lakes is highly sensitive to changes in fish predation pressure. The calibration data set consisted of thirty lakes differing in PF-CPUE (range 18–369 fish net^–1), epilimnion total phosphorus (range 0.025–1.28 mg P l^–1) and submerged macrophyte coverage (0–57%). 2. Correlation of log-transformed PF-CPUE, total phosphorus and submerged macrophyte coverage v the percentage abundance in the sediment of the dominant cladocerans and rotifers revealed that the typical pelagic species correlated most highly to PF-CPUE, while the littoral species correlated most highly to submerged macrophyte coverage. Consequently, only pelagic species were taken into consideration when establishing the fish transfer function. 3. Canonical correspondence analysis (CCA) revealed that the pelagic zooplankton assemblage was highly significantly related to PF-CPUE (axis 1), whereas the influence of total phosphorus and submerged macrophyte coverage was insignificant. Predicted PF-CPUE based on weighted average regression without (WA) and with (WA(tol)) downweighting of zooplankton species tolerance correlated significantly with the observed values (r² = 0.64 and 0.60 and RMSE = 0.54 and 0.56, respectively). A marginally better relationship (r² = 0.67) was obtained using WA maximum likelihood estimated optima and tolerance. 4. It is now possible, quantitatively, to reconstruct the historical development in planktivorous fish abundance based on zooplankton fossil records. As good relationships exist between contemporary PF-CPUE data and indicators such as the zooplankton/phytoplankton biomass ratio, Secchi depth and the maximum depth distribution of submerged macrophytes, it is now also possible to derive information on past changes in lake water quality and trophic structure. It will probably prove possible further to improve the transfer function by including other invertebrate remains, e.g. chironomids, Chaoborus, snails, etc., and its scope could be widened by including deeper lakes, more oligotrophic lakes, more acidic lakes and lakes with extensive submerged macrophyte coverage (in the latter case to enable use of the information in the fossil record on plant-associated cladocerans).     

Changes in benthic macroinvertebrate abundance and lake isotope (C,N) signals following biomanipulation: an 18-year study in shallow Lake Vaeng,Denmark

Change in the abundance of benthic macroinvertebrates and the stable isotope composition (C, N) of benthic invertebrates and zooplankton in Lake Vaeng, Denmark, was investigated over an 18-year period following biomanipulation (removal of cyprinids). During the first nine years after biomanipulation, the lake was clear and submerged macrophytes were abundant; after this period, a shift occurred to low plant abundance and high turbidity. Two years after the biomanipulation, total density of benthic macroinvertebrates reached a maximum of 17042 (±2335 SE) individuals m⁻² and the density was overall higher when the lake was in a clear state. Redundancy analysis (RDA) suggested macrophyte abundance and total nitrogen (TN) concentration were the dominant structuring forces on the benthic macroinvertebrate assemblage. Stable isotope analysis revealed that δ¹³C of macroinvertebrates and zooplankton was markedly higher in years with high submerged macrophyte abundance than in years without macrophytes, most likely reflecting elevated δ¹³C of phytoplankton and periphyton mediated by a macrophyte-induced lowering of lake water CO₂ concentrations. We conclude that the strong relationship between macrophyte coverage and δ¹³C of macroinvertebrates and cladocerans may be useful in paleoecological studies of past changes in the dynamics of shallow lakes, as change in macrophyte abundance may be tracked by the δ¹³C of invertebrate remains in the sediment.     

Bacterioplankton in the littoral and pelagic zones of subtropical shallow lakes

We measured bacterioplankton (phylotypes detected by fluorescent in situ hybridisation, morphometric forms, abundance and production) in samples collected in summer in the littoral and pelagic zones of 10 subtropical shallow lakes of contrasting area (from 13 to 80,800 ha). Compared to the pelagic zones, the littoral zones were overall characterised by higher macrophyte dominance and lower concentrations of total phosphorus and alkalinity and higher concentrations of dissolved organic carbon (DOC) and humic substances. Similarities of bacterial production and biomass turnover and density of active phylotypes and morphotype proportions were related to similarities in a set of environmental variables (including nutrients, humic substances content, predator density and phytoplankton biomass), and some additionally to lake area. Horizontal heterogeneity in bacterioplankton variables (littoral versus pelagic) increased with lake area. Bacterioplankton biomass and production tended to be lower in the littoral zone than in the pelagic zone despite higher concentrations of DOC and humic substances. A likely explanation is higher predation on bacterioplankton in the littoral zone, although allelophatic effects exerted by macrophytes cannot be excluded. Our results indicate that organic cycling via bacterioplankton may be less efficient in the littoral zone than in the pelagic zone of shallow lakes.     

Macrophytes as refuge or risky area for zooplankton: a balance set by littoral predacious macroinvertebrates

1. Zooplankton use macrophytes as day-time refuge areas when trying to escape from pelagic predators. But macrophytes can also host a diverse and abundant macroinvertebrate assemblage and zooplankton are also likely to face predacious macroinvertebrates once they enter the littoral zone. This study aimed to elucidate the role of macroinvertebrates in determining the refuge capacity of macrophytes.
2. We conducted a field enclosure experiment using plastic bags and complementary laboratory feeding trials to test how macroinvertebrates counteract the benefits to zooplankton of the macrophyte refuge. The field experiment consisted of three treatments with different macroinvertebrate assemblages: without predators (WP), low abundance and diversity (LAD) and high abundance and diversity of predators (HAD – which represents lake conditions).
3. Populations of Diaphanosoma brachyurum , Bosmina huaronensis and Moina micrura (Cladocera) and of both male and female Notodiaptomus incompositus (Copepoda, Calanoida) declined (by nearly 80%) in the presence of HAD in comparison to WP and LAD treatments.
4. Feeding trials revealed that Buenoa sp. (backswimmer), adults of Palaemonetes argentinus (grass shrimp) and Cyanallagma interruptum (damselfly) had a significant negative impact on cladocerans ( D. brachyurum, B. huaronensis ) and the calanoid copepod population (males, females and copepodites). These predators showed a strong predation effect ranging from 75% to 100% reductions of zooplankton populations.
5. The refuge effect offered by macrophytes to zooplankton depends on and is balanced by the predacious macroinvertebrate assemblage that plants host. The risk of confronting littoral predators is high and macroinvertebrate presence can turn the macrophytes into risky areas for zooplankton.     

Influence of nutrients, submerged macrophytes and zooplankton grazing on phytoplankton biomass and diversity along a latitudinal gradient in Europe

In order to evaluate latitudinal differences in the relationship of phytoplankton biomass and diversity with environmental conditions in shallow lakes, we sampled 98 shallow lakes from three European regions: Denmark (DK), Belgium/The Netherlands (BNL) and southern Spain (SP). Phytoplankton biomass increased with total phosphorus (TP) concentrations and decreased with submerged macrophyte cover across the three regions. Generic richness was significantly negatively related to submerged macrophyte cover and related environmental variables. Zooplankton:phytoplankton biomass ratios were positively related to submerged macrophyte cover and negatively to phytoplankton generic richness in DK and BNL, suggesting that the low generic richness in lakes with submerged macrophytes was due to a higher zooplankton grazing pressure in these regions. In SP, phytoplankton generic richness was not influenced by zooplankton grazing pressure but related to conductivity. We observed no relationship between phytoplankton generic richness and TP concentration in any of the three regions. The three regions differed significantly with respect to mean local and regional generic richness, with BNL being more diverse than the other two regions. Our observations suggest that phytoplankton diversity in European shallow lakes is influenced by submerged macrophyte cover indirectly by modulating zooplankton grazing. This influence of submerged macrophytes and zooplankton grazing on phytoplankton diversity decreases from north to south.     

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.     

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.     

Reciprocal Relationships Between Exotic Rusty Crayfish,Macrophytes, and <Emphasis Type="Italic">Lepomis</Emphasis> Species in Northern Wisconsin Lakes

Abstract The non-native rusty crayfish (Orconectes rusticus) has invaded many lakes of northern Wisconsin, profoundly changing littoral zones in the process. There are other lakes that have been invaded, but do not exhibit these changes. We hypothesized that endogenous feedbacks could form involving rusty crayfish, the macrophytes they destroy, and Lepomis species whose abundance is positively related to macrophyte abundance and also consume juvenile crayfish. We assessed this proposal with long-term data from one lake, a regional comparative study, and a case study of Lepomis predation on crayfish. Through time and across lakes, abundances of rusty crayfish, littoral macrophytes and species of the genus Lepomis were related in a fashion that indicated a set of feedbacks that regulate the abundance of all three. Intense predation on juvenile crayfish by abundant Lepomis is capable of maintaining some crayfish populations at low abundance. Thus, some lakes display profound ecological changes where crayfish achieve high abundance, and others sustain crayfish at low abundance. Consequently, lakes invaded by rusty crayfish may take on the appearance of alternative ecological regimes. Direct experimentation is necessary to determine if, and under what conditions, a lake can exist in either regime. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Phytoplankton communities in the littoral zone of lakes: Observations on structure and dynamics in oligotrophic and eutrophic systems

Summary The structure and seasonal dynamics of phytoplankton communities in the littoral zone were compared between oligotrophic and eutrophic lakes in the southeastern United States. Differences in diversity and species composition between lakes could be ascribed to long-term variation in nutrients corresponding to trophic status. However, significant within-lake variation could not be accounted for by microstratification of nutrients or other abiotic variables. Local biotic factors, perhaps dominated by the spawning activities of centrarchid fishes, resuspend periphyton and generate tychoplankton which becomes a persistent and integral part of the phytoplankton community in eutrophic systems. The patchy distribution of these biotic factors and resultant tychoplankton may lead to the observed variation. Grazing by herbivorous zooplankton was considered to be the major factor affecting the relative abundance of phytoplankton in the littoral zone, completely overriding the effects of nutrient concentration and biotic interactions between phytoplankton species during spring and summer.The relative importance of tychoplankton and grazing as regulatory factors operates independently of the trophic status or geographical location of a lake, making comparisons of different studies difficult and perhaps meaningless if traditional analyses based only on nutrients and interactions between species of phytoplankton are used. Limnetic as well as littoral components must be considered in future studies of phytoplankton communities in the littoral zone.