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.     

Loss of trophic complexity in saline prairie lakes as indicated by stable-isotope based community-metrics

Variations in climate, watershed characteristics and lake-internal processes often result in a large variability of food-web complexity in lake ecosystems. Some of the largest ranges in these environmental parameters can be found in lakes across the northern Great Plains as they are characterized by extreme gradients in respect to lake morphometry and water chemistry, with individual parameters often varying over several orders of magnitude. To evaluate the effects of environmental conditions on trophic complexity in prairie lake food-webs, we analyzed carbon and nitrogen stable isotopes of fishes, zooplankton and littoral macroinvertebrates in 20 lakes across southern Saskatchewan. Our two-year study identified very diverse patterns of trophic complexity, with was predominantly associated with among-lake differences. Small but significant temporal effects were also detected, which were predominantly associated with changes in productivity. The most influential parameters related to changes in trophic complexity among lakes were salinity, complexity of fish assemblage, and indicators of productivity (e.g. nutrients, Chl a). Generally, trophic diversity, number of trophic levels, and trophic redundancy were highest in productive freshwater lakes with diverse fish communities. Surprisingly, mesosaline lakes that were characterized by very low or no predation pressure from fishes were not colonized by invertebrate predators as it is often the case in boreal systems; instead, trophic complexity was further reduced. Together, prairie lake food-webs appear to be highly sensitive to changes in salinity and the loss of piscivorous fishes, making freshwater and mesosaline lakes most vulnerable to the impacts of climate variability. This is particularly important as global circulation models predict future climate warming to have disproportionate negative impacts on hydrologic conditions in this area.     

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.     

The nature of fish communities: A factor influencing the fishery potential and yields of tropical lakes and reservoirs

The discrepancy between high primary production and poor fish yields in some tropical reservoirs, especially in Souht East Asia, is apparently due to the fact that not all available trophic levels are utilized. Data analysis has shown that this is due to the riverline origin of fish communities which are not well adapted to lacustrine habitats in such lakes and reservoirs. These fishes are able to inhabit only a relatively narrow belt of the shallow inshore area. It has been shown that fish are rare in the deep water and pelagial area of geologically young lakes and reservoirs. Although most riverine fish are also able to inhabit the lacustrine habitat, only a few are preadapted to take advantage of deep or open water environments. In the tropics, Cichlidae and freshwater Clupeidae thrive in lakes and reservoirs. It is apparent that within these families suitable species can be chosen as potential candidates for introduction into young lakes and reservoirs. However, any fish introductions must be scientifically evaluated for desirability. Data from other lakes in the world have indicated that the present theory can be expanded to other than tropical lakes.     

Decoupling of pelagic and littoral food webs in oligotrophic Canadian Shield lakes

The importance of top-down effects of piscivorous fish on phytoplankton in natural oligotrophic lakes is still debated. In this study, we analyzed patterns in phytoplankton and zooplankton abundance in 37 oligotrophic Canadian Shield lakes in relation to variations in both piscivorous fish predation and resources (total phosphorus; TP). Zooplankton community structure (but not total biomass) was partially affected by the variation in fish predation while the phytoplankton community structure and total biomass showed no response. Carbon isotope analyses revealed that the lack of top-down effects is due to the uncoupling of the littoral and the pelagic food webs. We found that the fish community depends mostly on benthic resources, suggesting that only low planktivory occurred in our study lakes. Due to the absence of specialized zooplanktivorous fish, zooplankton is poorly exploited in these lakes and thus able to control phytoplankton by grazing. A comparison of our data with published studies on the TP–chlorophyll a relationships in both natural and manipulated systems shows that the phytoplankton biomass per unit of TP is relatively low in Canadian Shield lakes.     

Invertebrate zooplankton predator composition and diversity in tropical lentic waters

Invertebrate zooplankton predators are generally less diverse in average species numbers in tropical than in temperate lakes and reservoirs. Predatory Copepoda which comprise the majority of limnetic predators are particularly low in species numbers in the tropics. Predatory Cladocera are confined to the North Temperate zone. Chaoborus appears to be cosmopolitan. Among Rotifera, only the cosmopolitan predator Asplanchna occurs in tropical waters while the other common limnetic carnivorous genus Ploesoma is restricted to higher latitudes. Hydracarina, and insects besides Chaoborus, are generally restricted to the littoral and appear to be more diverse in the tropics. Lakes Awasa and Zwai, Ethiopia, are almost devoid of predators in the limnetic, which is invaded by a littoral chydorid Alona diaphana. Low diversity of lake types and low production of tropical zooplankton could restrict predator diversity too. Very low diversity of invertebrate predators in the limnetic and higher diversity in the littoral may characterize tropical lakes in contrast to temperate lakes, which have more invertebrate predators in the limnetic and perhaps relatively less in the littoral. Tropical zooplankton in freshwaters, appears to be a very immature community. Hence opportunistic species can readily invade the limnetic and even dominate in isolated situations as has been shown for Alona davidi, Hydracarina and some other unconventional forms.     

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     

Intraspecific variation in diet, growth, and morphology of landlocked Galaxias maculatus during its larval period: the role of food availability and predation risk

Food availability and predation risk have been shown to affect phenotypes during early life history of fishes. Galaxias maculatus, a small fish widely distributed around the southern hemisphere, clearly exhibits a complex trade-off between feeding and predation avoidance during growth over the larval period. We studied the effect of different environmental variables on diet, growth, mortality, and morphology through field surveys and data revision in the literature for limnetic G. maculatus larvae in five oligotrophic lakes of Patagonia. Both number of food categories and prey ingested by larvae were directly related to zooplankton density. Larval growth rate was related with zooplankton density and temperature. Lakes with high zooplankton densities and low predation risk had larvae with deeper bodies and shorter caudal peduncles, while in lakes with less food and high predation risk larvae were slender with shallower bodies and longer peduncles. Food availability and predation risk seem to operate on the swimming performance of G. maculatus larvae through the slenderness of the body and the length of the caudal peduncle. The observed phenotypic variation in growth and morphology could be a key feature that has allowed this species to successfully colonize a wide variety of environments in the southern hemisphere.     

Subtle top‐down control of a freshwater meiofaunal assemblage by juvenile fish

1. Top‐down control of prey assemblages by fish predators has been clearly demonstrated in lakes (for zooplankton prey) and rivers (for macroinvertebrate prey). Fish predation can have a significant impact on the body size of prey assemblages; often large‐bodied prey are reduced in abundance, and indirect facilitation of small‐bodied prey occurs potentially initiating a trophic cascade. 2. Benthic communities in aquatic ecosystems also include a numerous and functionally important meiofaunal‐sized component, but in freshwaters the impact of fish predation on meiofaunal assemblages is unknown. We used a laboratory microcosm study to explore the impact of juvenile fish predation on the abundance and size structure of a riverine meiofaunal assemblage. 3. The presence of fish in our microcosms had no significant effect on overall meiofaunal (temporary and permanent) abundance. However, for the Copepoda, we found the first evidence of top‐down control of freshwater meiofaunal assemblages; in microcosms with juvenile fish, the abundance of large‐bodied Copepoda was significantly reduced, whereas small‐bodied Copepoda were significantly more abundant suggesting indirect facilitation. 4. We conclude that predation by juvenile fish can alter the structure of freshwater meiofaunal assemblages, although we do not yet know whether these relatively subtle changes are overwhelmed by large‐scale events such as flow disturbances.     

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.     

Distribution of the freshwater fishes of Japan: an historical overview

Japanese freshwater fishes, including lampreys, comprise 15 orders, 35 families, and 96 genera, with 211 species and subspecies. Most belong to the families Cyprinidae (29% of species and subspecies), Gobiidae (21%), Salmonidae (10%), and Cobitidae (8%). Cyprinids and cobitids presumably originated from east Asia, gobiids from southeast Asia, and cottids and salmonids from the north Pacific. Japanese freshwater fishes include 88 endemic species and subspecies, of which three have been extirpated. Fishes introduced into natural rivers and lakes for inland commercial fisheries and sport fishing, and by accident, include many exotic species, of which 23 now inhabit natural freshwaters. These often have destroyed the local fish fauna by predation, and caused genetic pollution by hybridization with local strains. Destruction of freshwater environments by land development also poses a threat to Japanese freshwater fish communities. In addition Japanese freshwater systems have been markedly altered by development of rice paddy fields which have caused some species to decline but others to flourish, and changed the distribution patterns of fishes between upstream and downstream areas. To conserve endangered species and declining communities of Japanese freshwater fishes, we need to clarify the characteristics of their original habitats and the effects of developing paddy fields, from both the ecological and historical points of view.     

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...     

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.     

Trophic complexity of small fish in nearshore food webs

Small nearshore fishes are an important part of lacustrine and functional diversity and link pelagic and benthic habitats by serving as prey for larger nearshore and offshore fishes. However, the trophic complexity of these small nearshore fishes is often unrecognized and detailed studies of their role in food webs are lacking. Here, we examined niche space patterns of small nearshore fish species using Bayesian analyses of carbon and nitrogen stable isotope data in nine freshwater lakes that are among the largest lakes in Minnesota. We found considerable variability in niche areas within species and high variability in niche overlap across species. At the assemblage level, niche overlap (average diet overlap of all species pairs at a lake) decreased as whole-lake species richness increased, possibly indicating a greater degree of resource specialization in more speciose lakes. Overall fish niche space was weakly but significantly related to niche space of their invertebrate prey. Although nearshore benthic resources contributed to fish diets in all lakes, all fish species also had non-negligible and variable contributions from pelagic zooplankton. This inter- and intraspecific variability in trophic niche space likely contributes to the multi-level trophic complexity, functional diversity, and potentially food web resilience to ecosystem changes.

     

The effects of predation by juvenile fish on the meiobenthic community structure in a natural pond

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Trading safety for food: evidence from gut contents in roach and bleak captured at different distances offshore from their daytime littoral refuge

1. Regular diel habitat shifts in roach were detected by hydro‐acoustics in five moderately eutrophic, stratifying (maximum depth 24–27 m) and approximately circular lakes (of surface area 15, 75, 125, 300 and 900 ha and diameters 250, 600, 1000, 1700 and 2600 m) in north‐eastern Poland in the years 1998–2000, when the lakes were free of smelt and other typical offshore planktivores, and their offshore areas were completely free of fish during the day. 2. The diel change in roach distribution was shown to assume a similar pattern in each lake: fish migrated from a daytime littoral refuge towards the centre of the lake at dusk, and returned to the littoral refuge at dawn. After sunset, fish gradually dispersed offshore until they covered the entire lake area in each of the three smaller lakes. In each of the two larger lakes, only small numbers of fish were seen in the central area at night, implying that the centre of the lake retained high food availability throughout the summer. 3. Inshore–offshore gradients in zooplankton prey density, body size, and numbers of eggs per clutch were weak or undetectable in the two smallest lakes, but strong and persistent in the three larger lakes, with Daphnia densities 5–30 times as high and body length 1.2–1.5 times as great in the central area as inshore. 4. The likely increase in the potential predation risk with distance from the littoral daytime refuge was found to be compensated by increased food gains in those fish which moved offshore at dusk to feed within a short time window, when light intensity was lower to make the risk reduced, but still high enough to see zooplankton prey. The benefit because of increased prey acquisition was greatest in the centre of the largest lake (at 1300 m from the shore), as revealed from gut inspections of roach and bleak trawl‐sampled at different distances from the edge of the reed belt, and seen as a gradual, order‐of‐magnitude increase in the volume of food in the foregut, The food volume against distance‐from‐shore regression was highly significant on each of the four sampling dates in the largest lake, in spite of the wide variability of food volume in individual fish.     

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.     

Clay-Turbid Interactions May Not Cascade—A Reminder for Lake Managers

Food web management is a frequently used lake restoration method, which aims to reduce phytoplankton biomass by strengthening herbivorous zooplankton through reduction of planktivorous fish. However, in clay‐turbid lakes several factors may reduce the effectivity of food web management. Increasing turbidity reduces the effectivity of fish predation and weakens the link between zooplankton and phytoplankton. Therefore, the effects of fish stock manipulations may not cascade to lower trophic levels as expected. Additionally, in clay‐turbid conditions invertebrate predators may coexist in high densities with planktivorous fish and negate the effects of fish reductions. For instance, in the stratifying regions of the clay‐turbid Lake Hiidenvesi, Chaoborus flavicans is the main regulator of cladocerans and occupies the water column throughout the day, although planktivorous Osmerus eperlanus is very abundant. The coexistence of chaoborids and fish is facilitated by a metalimnetic turbidity peak, which prevents efficient predation by fish. In the shallow parts of the lake, chaoborids are absent despite high water turbidity. We suggest that, generally, the importance of invertebrate predators in relation to vertebrate predators may change along turbidity and depth gradients. The importance of fish predation is highest in shallow waters with low turbidity. When water depth increases, the importance of fish in the top‐down regulation of zooplankton declines, whereas that of chaoborids increases, the change along the depth gradient being moderate in clear‐water lakes and steep in highly turbid lakes. Thus, especially deep clay‐turbid lakes may be problematic for implementing food web management as a restoration tool.     

Trade-offs in the vertical distribution of zooplankton: ideal free distribution with costs?

Zooplankton vertical migratory patterns are a classic example of optimal habitat choice. We hypothesize that zooplankton distribute themselves vertically in the water column according to an ideal free distribution (IFD) with costs such as to optimize their fitness. In lakes with a deep-water chlorophyll maximum, zooplankton are faced with a trade-off, either experiencing high food (high reproductive potential) but low temperature (slow development) in the hypolimnion or high temperature and low food in the epilimnion. Thus, in the absence of fish predation (e.g. at night) they should allocate the time spent in the different habitats according to fitness gain dependent on the temperature gradient and distribution of food. We tested this hypothesis with a Daphnia hyalina x galeata clone in large indoor columns (Plön Plankton Towers) and with a dynamic energy budget model. In the tower experiments, we simulated a deep-water algal maximum below the thermocline with epilimnetic/hypolimnetic temperature differences of 2, 5 and 10 degrees C. Experimental data supported the model. We found a significantly larger proportion of daphniids in the hypolimnion when the temperature difference was smaller. Our results are consistent with the concept of IFD with costs originally developed for stream fishes. This concept can be applied to predict the vertical distribution of zooplankton in habitats where fish predation is of minor importance.     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.