Abiotic and biotic processes in lakes recovering from acidification: the relative roles of metal toxicity and fish predation as barriers to zooplankton re‐establishment

1. Recovery of acidified aquatic systems may be affected by both abiotic and biotic processes. However, the relative roles of these factors in regulating recovery may be difficult to determine. Lakes around the smelting complexes near Sudbury, Ontario, Canada, formerly affected by acidification and metal exploration, provide an excellent opportunity to examine the factors regulating the recovery of aquatic communities. 2. Substantial recovery of zooplankton communities has occurred in these lakes following declines in acidity and metal concentrations, although toxicity by residual metals still appears to limit survival for many species. Metal bioavailability, not simply total metal concentrations, was very important in determining effects on zooplankton and was associated with a decrease in the relative abundance of cyclopoids and Daphnia spp., resulting in communities dominated by Holopedium gibberum. 3. As chemical habitat quality has improved and fish, initially yellow perch and later piscivores (e.g. smallmouth bass, walleye), have invaded, biotic effects on the zooplankton are also becoming apparent. Simple fish assemblages dominated by perch appear to limit the survival of some zooplankton species, particularly Daphnia mendotae. 4. Both abiotic (residual metal contamination) and biotic (predation from planktivorous fish) processes have very important effects on zooplankton recovery. The re‐establishment of the zooplankton in lakes recovering from stress will require both improvements in habitat quality and the restoration of aquatic food webs.     

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.     

The recovery of acid‐damaged zooplankton communities in Canadian Lakes: the relative importance of abiotic,biotic and spatial variables

1. Acidification has damaged biota in thousands of lakes and streams throughout eastern North America. Fortunately, reduced emissions of sulphur dioxide and nitrogen oxides beginning in the 1960s have allowed pH levels in many affected systems to increase. Determining the extent of biological and pH recovery in these systems is necessary to assess the success of emissions reductions programmes. 2. Although there have been promising signs of biological recovery in many systems, recovery has occurred more slowly than expected for some taxa. Past studies with crustacean zooplankton indicate that a mixture of local abiotic variables, biotic variables and dispersal processes may influence the structure of recovering communities. However, most studies have been unable to determine the relative importance of these three groups of variables. 3. We assessed chemical and biological recovery of acid‐damaged lakes in Killarney Park, Ontario. In addition, we assessed the relative importance of local abiotic variables, biotic variables and dispersal processes for structuring recovering communities. We collected zooplankton community data, abiotic and biotic data from 45 Killarney Park lakes. To assess the recovery of zooplankton communities, we compared zooplankton data collected in 2005 to a survey conducted for the same lakes in 1972–73 using several univariate measures of community structure, as well as multivariate methods based on relative species abundances. To determine the factors influencing the structure of recovering zooplankton communities, we used hierarchical partitioning for univariate measures and spatial modelling and variation partitioning techniques for multivariate analyses. 4. Our survey revealed significant pH increases for the majority of sampled lakes but univariate measures of community structure, such as species richness and diversity, indicated that only minor changes have occurred in many acid‐damaged lakes. Hierarchical partitioning identified several variables that may influence our univariate measures of recovery, including pH, dissolved organic carbon (DOC) levels, fish presence/absence, lake surface area and lake elevation. 5. Multivariate methods revealed a shift in communities through time towards a structure more typical of neutral lakes, providing some evidence for recovery. Variation partitioning suggested that the structure of recovering copepod communities was influenced most by dispersal processes and abiotic variables, while biotic (Chaoborus densities, fish presence/absence) and abiotic variables were more important for cladoceran zooplankton. 6. Our results indicate that the recovery of zooplankton communities in Killarney Park is not yet complete, despite decades of emission reductions. The importance of variables related to acidification, such as pH and DOC, indicates that further chemical recovery may be necessary. The differing importance of abiotic, biotic and dispersal processes for structuring copepod versus cladoceran zooplankton might indicate that different management approaches and expectations for recovery are needed for these groups.     

Evidence that fish structure the zooplankton communities of turbid lakes and reservoirs

1. Visually foraging fish typically exclude large zooplankton from clear‐water lakes and reservoirs. Do fish have the same effect in turbid waters, or does turbidity provide a refuge from visual predation? 2. To test the hypothesis that fish exclude large zooplankton species from turbid sites, I searched for populations of medium or large Daphnia species in turbid, fish‐containing reservoirs of south‐central Oklahoma and north‐central Texas, U.S.A., and surveyed the literature for accounts of Daphnia species in turbid habitats worldwide. 3. Only small Daphnia species and the exuberantly spined Daphnia lumholtzi were detected in the turbid reservoirs. The Daphnia species in the reservoirs are smaller than other Daphnia species that occur in the area but were not detected. An extensive survey of the literature suggests that large Daphnia may be found in the lakes of extreme turbidity [Secchi disk depth (SD) < 0.2 m] but that only small and spiny Daphnia are likely to occur in more typical turbid locations (1.0 m > SD > 0.2 m) unless some additional factor reduces the influence of fish predation in such sites. 4. The field samples from Texas and Oklahoma together with the literature review suggest that the effect of visually foraging planktivorous fish on the size structure of turbid‐water zooplankton communities may often be as strong or even stronger than the effect of fish on clear‐water zooplankton communities.     

Cyanobacterial chemical warfare affects zooplankton community composition

1. Toxic algal blooms widely affect our use of water resources both with respect to drinking water and recreation. However, it is not only humans, but also organisms living in freshwater and marine ecosystems that may be affected by algal toxins. 2. In order to assess if cyanobacterial toxins affect the composition of natural zooplankton communities, we quantified the temporal fluctuations in microcystin concentration and zooplankton community composition in six lakes. 3. Microcystin concentrations generally showed a bimodal pattern with peaks in early summer and in autumn, and total zooplankton biomass was negatively correlated with microcystin concentrations. Separating the zooplankton assemblages into finer taxonomic groups revealed that high microcystin concentrations were negatively correlated with Daphnia and calanoid copepods, but positively correlated with small, relatively inefficient phytoplankton feeders, such as cyclopoid copepods, Bosmina and rotifers. 4. In a complementary, mechanistic laboratory experiment using the natural phytoplankton communities from the six lakes, we showed that changes in in situ levels of microcystin were coupled with reduced adult size and diminished juvenile biomass in Daphnia. 5. We argue that in eutrophic lakes, large unselective herbivores, such as Daphnia, are ‘sandwiched’ between high fish predation and toxic food (cyanobacteria). In combination, these two mechanisms may explain why the zooplankton community in eutrophic lakes generally comprise small forms (e.g. rotifers and Bosmina) and selective raptorial feeders, such as cyclopoid copepods, whereas large, unselective herbivores, such as Daphnia, are rare. Hence, this cyanobacterial chemical warfare against herbivores may add to our knowledge on population and community dynamics among zooplankton in eutrophic systems.     

Linking Cascading Effects of Fish Predation and Zooplankton Grazing to Reduced Cyanobacterial Biomass and Toxin Levels Following Biomanipulation

Eutrophication has been one of the largest environmental problems in aquatic ecosystems during the past decades, leading to dense, and often toxic, cyanobacterial blooms. In a way to counteract these problems many lakes have been subject to restoration through biomanipulation. Here we combine 13 years of monitoring data with experimental assessment of grazing efficiency of a naturally occurring zooplankton community and a, from a human perspective, desired community of large Daphnia to assess the effects of an altered trophic cascade associated with biomanipulation. Lake monitoring data show that the relative proportion of Daphnia spp. grazers in June has increased following years of biomanipulation and that this increase coincides with a drop in cyanobacterial biomass and lowered microcystin concentrations compared to before the biomanipulation. In June, the proportion of Daphnia spp. (on a biomass basis) went from around 3% in 2005 (the first year of biomanipulation) up to around 58% in 2012. During months when the proportion of Daphnia spp. remained unchanged (July and August) no effect on lower trophic levels was observed. Our field grazing experiment revealed that Daphnia were more efficient in controlling the standing biomass of cyanobacteria, as grazing by the natural zooplankton community never even compensated for the algal growth during the experiment and sometimes even promoted cyanobacterial growth. Furthermore, although the total cyanobacterial toxin levels remained unaffected by both grazer communities in the experimental study, the Daphnia dominated community promoted the transfer of toxins to the extracellular, dissolved phase, likely through feeding on cyanobacteria. Our results show that biomanipulation by fish removal is a useful tool for lake management, leading to a top-down mediated trophic cascade, through alterations in the grazer community, to reduced cyanobacterial biomass and lowered cyanobacterial toxin levels. This improved water quality enhances both the ecological and societal value of lakes as units for ecosystem services.     

Elevated metal concentrations inhibit biological recovery of Cladocera in previously acidified boreal lakes

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Sub-fossils of cladocerans in the surface sediment of 135 lakes as proxies for community structure of zooplankton,fish abundance and lake temperature

To elucidate the possibilities of using zooplankton remains in the surface sediment to describe present-days community structure and population dynamics of zooplankton, fish abundance and temperature, we compared contemporary data sampled in the pelagial during summer with the sediment record from the upper 1 cm of the sediment in 135 lakes covering a latitude gradient from Greenland in the north to New Zealand in the south. The abundance of three genera Bosmina, Daphnia and Ceriodaphnia of the total pool of ephippia was significantly related to the total abundance of the same taxa in the pelagic zone. However, in most lakes the abundance of Ceriodaphnia was higher in the sediment than in the water, which may be attributed to the overall preference by this genus for the littoral habitat. Using contemporary data from 27 Danish lakes sampled fortnightly during summer for 10 years, we found substantial inter-annual variations in the abundance of Daphnia spp., Ceriodaphnia spp., B. longirostris and B. coregoni. Yet, the sediment record mimicked the medium level well for most of the lakes, which suggests that the sediment record provides an integrated picture of the pelagic cladoceran community, which otherwise can be obtained only by long-term frequent contemporary sampling for several years. The contribution of Daphnia to the sum of Daphnia and Bosmina ephippia was negatively correlated with the abundance of fish expressed as catch per night in multi-mesh sized gill nets (CPUE). Yet, region-specific differences occurred, which partly could be eliminated by including nutrient state expressed as total phosphorus (TP) in a multiple regression. The average ratio of ephippia to the sum of ephippia and carapaces of Bosmina varied 40-fold between the sampling regions and was significantly negatively related to summer mean air temperature, and for Danish lakes also, albeit weakly, to fish CPUE but not to chlorophyll a. Apparently, temperature is the most important factor determining the ratio of parthenogenetic to ephippia producing specimens of Bosmina. We conclude that the sediment record of cladocerans is a useful indicator of community structure of pelagic cladocerans and the abundance of fish and temperature.     

The potential of artificial refugia for maintaining a community of large-bodied cladocera against fish predation in a shallow eutrophic lake

The Norfolk Broads are a series of shallow lakes which are highly eutrophic and typified by dense populations of phytoplankton and an absence of submerged aquatic plants. The zooplankton community is subject to intense predation pressure by young fish and is dominated by small-bodied organisms which have a low potential for reducing phytoplankton populations through grazing. Various designs and densities of artificial refugia for zooplankton against fish predation were established in Hoveton Great Broad in order to enhance populations of large-bodied Cladocera. Initially some of the refuges contained higher densities and larger individuals ofDaphnia andCeriodaphnia than the surrounding open water. However, towards the end of the first season and throughout the subsequent two years, population densities and size-structure were similar both within and outside the refuges, although there was still evidence of enhanced body-size ofDaphnia within the refuges compared with the open water. The provision of habitat structures designed as refugia from fish predation did not enhance large-bodied cladoceran populations enough to promote this restoration strategy as feasible for eutrophic and shallow lakes.     

Trophic structure in the pelagial of 25 shallow New Zealand lakes: changes along nutrient and fish gradients

To gain better insight into the importance of predator and resourcecontrol in New Zealand lakes we surveyed the late summer trophicstructure of 25 shallow South Island lakes with contrastingnutrient levels (6–603 µg TP l^–1) and fishdensities. Total catch of fish per net (CPUE) in multi-meshgillnets placed in the open water and the littoral zones waspositively related with the nutrient level. Trout CPUE was negativelycorrelated with total phosphorus (TP) and total nitrogen (TN).Zooplankton seemed largely influenced by fish, as high fishCPUE coincided with low zooplankton and Daphnia biomass, lowaverage weight of cladocerans, low contribution of Daphnia tototal cladoceran biomass, low ratio of calanoids to total copepodbiomass and low ratio of zooplankton biomass to phytoplanktonbiomass. However, chlorophyll a was only slightly negativelyrelated to Daphnia biomass and not to zooplankton biomass ina multiple regression that included TN and TP. Ciliate abundancewas positively related to chlorophyll a and negatively to Daphniabiomass, but not to total zooplankton biomass, while no relationshipswere found between heterotrophic nanoflagellates and zooplankton.The relationships between fish abundance and nutrients and fishabundance and zooplankton:phytoplankton ratio and between chlorophylla and TP largely followed the pattern obtained for 42 northtemperate Danish lakes. We conclude that fish, including trout,have a major effect on the zooplankton community structure andbiomass in the pelagial of the shallow oligotrophic to slightlyeutrophic New Zealand lakes, but that the cascading effectson phytoplankton and protist are apparently modest.     

Effects of emission reductions from the Sudbury smelters on the recovery of acid- and metal-damaged lakes

Aquatic ecosystems in a 17,000 km² area around Sudbury, Ontario, Canada, have been affected by the atmospheric deposition of pollutants from nearly a century of operations at the Sudbury area metal smelters. Effects were most severe in the lakes closest to the smelters which historically received very high deposition of both acid and metal particulates. After smelter emissions were greatly reduced in the 1970‘s, evidence began to emerge of improvements in lake water quality, and some recovery of biological communities, and the emphasis of Sudbury area monitoring programs changed from the assessment of damage to the tracking of recovery patterns. Further reductions in smelter emissions during the1990‘s have been accompanied by continuing improvements in aquatic habitat quality, but the evaluation of lake responses to emission controls is complicated by the interaction of lake acidity and metal concentrations with other factors. Weather-related variations in storage and release of sulphur from lake catchments appear to greatly influence chemical recovery. Despite the general water quality improvements observed to date, some lakes are still highly acidic and elevated levels of copper and nickel persist in the water and sediments of many lakes. Severely damaged biological communities have been slow to recover, probably reflecting a combination of continuing habitat quality limitations and restricted opportunities for dispersal. This revised version was published online in July 2006 with corrections to the Cover Date.     

Recovery of copepod, but not cladoceran, zooplankton from severe and chronic effects of multiple stressors

In the mid‐twentieth century, many lakes near Sudbury, Canada, were severely contaminated by acid and metal emissions from local smelters. For example, in the early 1970s, Middle Lake had pH of 4.2, and Cu and Ni levels both >0.5 mg L^?1 . To determine if crustacean zooplankton could recover from such severe and chronic damage, Middle Lake was neutralized in 1973. A comparison of its zooplankton with that of 22 reference (pH > 6) lakes indicates that the planktonic Copepoda completely recovered by 2001. In contrast, the cladoceran assemblage improved but did not recover. Colonist sources existed – Cladocera and Copepoda occurred with equal frequency in area lakes – but six separate colonizations by cladoceran species failed. We argue that local factors, metal toxicity and predation by yellow perch, have, to date, prevented cladoceran recovery. Nonetheless, the complete copepod recovery is encouraging, given the severity and duration of pre‐neutralization stress.     

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.     

Zooplankton seasonal dynamics in a recently filled mine pit lake: the effect of non-indigenous <Emphasis Type="Italic">Daphnia</Emphasis> establishment

We examined the temporal and vertical dynamics of zooplankton in Weavers Lake, New Zealand, between October 2004 and October 2005, at a time when it was colonised by a non-indigenous Daphnia species. Zooplankton community composition changed during the study from one of rotifer dominance (e.g. Asplanchna, Polyarthra, Brachionus and Keratella species) to cladoceran (Daphnia dentifera) dominance. Temporal changes in zooplankton community composition were strongly associated with a gradual increase in lake water clarity, and were attributable to the highly efficient filter feeding of D. dentifera. The corresponding reduction in rotifer densities may have resulted from the superior competitive abilities of the newly established Daphnia. As Daphnia were rare inhabitants of New Zealand lakes before 1990, the arrival and rapid spread of the non-indigenous D. dentifera has lead to widespread changes in both water clarity and zooplankton community composition. An apparent lack of mixing in the lake was facilitated by the lake’s extremely small surface area:depth ratio. However, we conclude that physical features of the lake had minimal influence on water clarity relative to the invasion of D. dentifera.     

Elevated temperature prolongs long‐term effects of a pesticide on Daphnia spp. due to altered competition in zooplankton communities

Considerable research efforts have been made to predict the influences of climate change on species composition in biological communities. However, little is known about how changing environmental conditions and anthropogenic pollution can affect aquatic communities in combination. We investigated the influence of short warming periods on the response of a zooplankton community to the insecticide esfenvalerate at a range of environmentally realistic concentrations (0.03, 0.3 and 3 μg L^?1) in 55 outdoor pond microcosms. Warming periods increased the cumulative water temperature, but did not exceed the maximum temperature measured under ambient conditions. Under warming conditions alone the abundance of some zooplankton taxa increased selectively compared to ambient conditions. This resulted in a shift in the community composition that had not recovered by the end of the experiment, 8 weeks after the last warming period. Regarding the pesticide exposure, short‐term effects of esfenvalerate on the community structure and the sensitive taxa Daphnia spp. did not differ between the two temperature regimes. In contrast, long‐term effects of esfenvalerate on Daphnia spp., a taxon that did not benefit from elevated temperatures, were observed twice as long under warming than under ambient conditions. This resulted in long‐term effects on Daphnia spp. until 4 months after contamination at 3 μg L^?1 esfenvalerate. Under both temperature regimes, we identified strength of interspecific competition as the mechanism determining the time until recovery. However, enhanced interspecific competition under warming conditions was prolonged and explained the delayed recovery of Daphnia spp. from esfenvalerate. These results show that, for realistic prediction of the combined effects of changing environmental factors and toxicants on sensitive taxa, the impacts of stressors on the biotic interactions within the community need to be considered.     

Daphnia community analysis in shallow Kenyan lakes and ponds using dormant eggs in surface sediments

1. Water fleas of the genus Daphnia are considered rare in tropical regions, and information on species distribution and community ecology is scarce and anecdotal. This study presents the results of a survey of Daphnia species distribution and community composition in 40 standing waterbodies in southern Kenya. The study sites cover a wide range of tropical standing aquatic habitats, from small ephemeral pools to large permanent lakes between approximately 700 and 2800 m a.s.l. Our analysis combines data on Daphnia distribution and abundance from zooplankton samples and dormant eggs in surface sediments. 2. Nearly 70% (27 of 40) of the sampled waterbodies were inhabited by Daphnia. Although their abundance in the active community was often very low, this high incidence shows that Daphnia can be equally widespread in tropical regions as in temperate regions. 3. Analysis of local species assemblages from dormant eggs in surface sediments was more productive than snapshot sampling of zooplankton communities. Surface‐sediment samples yielded eight Daphnia species in total, and allowed the detection of Daphnia in 25 waterbodies; zooplankton samples revealed the presence of only four Daphnia species in 16 waterbodies. 4. Daphnia barbata, D. laevis, and D. pulex were the most frequently recorded and most abundant Daphnia species. Canonical correspondence analysis of species–environment relationships indicates that variation in the Daphnia community composition of Kenyan waters was best explained by fish presence, temperature, macrophyte cover and altitude. Daphnia barbata and D. pulex tended to co‐occur with each other and with fish. Both species tended to occur in relatively large (>10 ha) and deeper (>2 m) alkaline waters (pH 8.5). Daphnia laevis mainly occurred in cool and clear, macrophyte‐dominated lakes at high altitudes.     

Sediments,not plants,offer the preferred refuge for Daphnia against fish predation in Mediterranean shallow lakes: an experimental demonstration

1. Different behavioural responses of planktonic animals to their main predators, fish, have been reported from shallow lakes. In north temperate lakes, large‐bodied zooplankton may seek refuge from predation among macrophytes, whereas in subtropical lakes, avoidance of macrophytes has been observed. The prevalent behaviour probably depends on the characteristics of the fish community, which in Mediterranean lakes is typically dispersed in both the open water zone and in the littoral, as in temperate lakes, and is dominated by small size classes, as in subtropical lakes. 2. We performed ‘habitat choice’ experiments to test the response of Daphnia magna to predation cues at both the horizontal and vertical level by mimicking a ‘shallow littoral’ zone with plants and a ‘deeper pelagic’ zone with sediments. 3. Initial separate response experiments showed that natural plants, artificial plants and predation cues all repelled D. magna in the absence of other stimuli, while sediments alone did not trigger any significant response by D. magna. 4. The habitat choice experiments showed that, in the presence of predation cues and absence of plants, Daphnia moved towards areas with sediment. In the presence of both plants and sediments, Daphnia moved away from the plants towards the sediments under both shallow and deep water treatment conditions. 5. Based on these results, we suggest that Daphnia in Mediterranean shallow lakes avoid submerged macrophytes and instead prefer to hide near the sediment when exposed to predation risk, as also observed in subtropical shallow lakes. This pattern is not likely to change with water level alterations, a common feature of lakes in the region, even if the effectiveness of the refuge may be reduced.     

Zooplankton community structure and environmental conditions in a set of interconnected ponds

We studied the zooplankton community structure in a set of 33 interconnected shallow ponds that are restricted to a relatively small area (`De Maten', Genk, Belgium, 200 ha). As the ponds share the same water source, geology and history, and as the ponds are interconnected (reducing chance effects of dispersal with colonisation), differences in zooplankton community structure can be attributed to local biotic and abiotic interactions. We studied zooplankton community, biotic (phytoplankton, macrophyte cover, fish densities, macroinvertebrate densities), abiotic (turbidity, nutrient concentrations, pH, conductivity, iron concentration) and morphometric (depth, area, perimeter) characteristics of the different ponds. Our results indicate that the ponds differ substantially in their zooplankton community structure, and that these differences are strongly related to differences in trophic structure and biotic interactions, in concordance with the theory of alternative equilibria. Ponds in the clear-water state are characterised by large Daphnia species and species associated with the littoral zone, low chlorophyll-a concentrations, low fish densities and high macroinvertebrate densities. Ponds in the turbid-water state are characterised by high abundances of rotifers, cyclopoid copepods and the opposite environmental conditions. Some ponds show an intermediate pattern, with a dominance of small Daphnia species. Our results show that interconnected ponds may differ strongly in zooplankton community composition, and that these differences are related to differences in predation intensity (top-down) and habitat diversity (macrophyte cover).     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Tropical lakes — functional ecology and future development:. The need for a process-orientated approach

The major classes of tropical lakes include shallow, lowland lakes; deep, tertiary lakes; high altitudinal lakes; rainforests lakes; and man-made lakes at all latitudes and altitudes. Basic ecological processes are similar in temperate and tropical lakes, including grazing, competition, predation and abiotic adaptation. Small tropical lakes of intermediate age are probably not biotically more complicated than similar-sized temperate lakes. The structure of the areas of adaptative radiation and the dispersal ability of the species are important for the present distribution of taxa. Fish play a key role in the tropics since many species both consume zooplankton and compete with them for algal and pelagic sestonic food. This important co-evolution between fish and algae, leaving a fraction of the algal community with a predation refuge, may have decreased the ability of zooplankton to exploit algae. In addition, heavy predation from juvenile and adult fish may greatly simplify the zooplankton community, and have resulted in the scarcity of Cladocera, notably the efficient filter-feeder Daphnia. Little is known of possible physiological constraints to cladoceran distribution, however. Thus similar co-evolution as hypothesized between fish and algae seems not to have occurred to such a great extent between fish and zooplankton. Diurnal patterns in habitat selection of fish may also influence nutrient re-distribution in the tropics as in many temperate lakes. Serious environmental problems threaten tropical lakes, including eutrophication, clear-cutting of the rain forest, unwise introduction of new species not adapted to prevailing conditions, overfishing, extensive use of biocids, and probably acidic rain in areas with poorly buffered waters. Important processes in tropical lakes could be elucidated by concentrating research upon the fate of phytoplankton successional production, involving competition, grazing, sinking, fungi and bacterial attack. Co-evolution of fish and algae should be further investigated as it could in part explain the general scarcity and simplicity of the zooplankton community. Limnocorral experiments should also be used for further assessing processes in tropical lakes.