Combined Effects of Turbulence and Different Predation Regimes on Zooplankton in Highly Colored Water—Implications for Environmental Change in Lakes

In aquatic ecosystems, predation is affected both by turbulence and visibility, but the combined effects are poorly known. Both factors are changing in lakes in the Northern Hemisphere; the average levels of turbulence are predicted to increase due to increasing wind activities, while water transparency is decreasing, e.g., due to variations in precipitation, and sediment resuspension. We explored experimentally how turbulence influenced the effects of planktivorous fish and invertebrate predators on zooplankton when it was combined with low visibility caused by high levels of water color. The study was conducted as a factorial design in 24 outdoor ponds, using the natural zooplankton community as a prey population. Perch and roach were used as vertebrate predators and Chaoborus flavicans larvae as invertebrate predators. In addition to calm conditions, the turbulent dissipation rate used in the experiments was 10⁻⁶ m² s⁻³, and the water color was 140 mg Pt L⁻¹. The results demonstrated that in a system dominated by invertebrates, predation pressure on cladocerans increased considerably under intermediate turbulence. Under calm conditions, chaoborids caused only a minor reduction in the crustacean biomass. The effect of fish predation on cladocerans was slightly reduced by turbulence, while predation on cyclopoids was strongly enhanced. Surprisingly, under turbulent conditions fish reduced cyclopoid biomass, whereas in calm water it increased in the presence of fish. We thus concluded that turbulence affects fish selectivity. The results suggested that in dystrophic invertebrate-dominated lakes, turbulence may severely affect the abundance of cladocerans. In fish-dominated dystrophic lakes, on the other hand, turbulence-induced changes in planktivory may considerably affect copepods instead of cladocerans. In lakes inhabited by both invertebrates and fish, the response of top-down regulation to turbulence resembles that in fish-dominated systems, due to intraguild predation. The changes in planktivorous predation induced by abiotic factors may possibly cascade to primary producers.     

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.     

轮虫和枝角类的种间竞争及其影响因子研究进展

轮虫和枝角类是浮游动物群落的重要组成部分和优势类群,它们之间的竞争互作是调节水生态系统结构和功能的主要动力之一。普遍的观点认为,轮虫和大型枝角类难以共存,往往被竞争排斥,而和小型枝角类可以共存;实际上,轮虫和枝角类的种间竞争结局存在一定的不确定性。介绍了轮虫和枝角类的种间竞争方式及其相对重要性,对影响轮虫和枝角类种间竞争结局的因素,包括温度、食物、相对起始密度、个体大小、食物临点、耐饥饿能力、捕食及竞争者和捕食者释放的化感物质等,进行了系统的梳理和分析,并提出了今后亟待解决的科学问题和研究切入点。     

Refuge availability and sequence of predators determine the seasonal succession of crustacean zooplankton in a clay-turbid lake

The contribution of predators and abiotic factors to the regulation of the biomass and seasonal succession of crustacean zooplankton was studied in Lake Rehtijärvi (southern Finland). Field data in combination with bioenergetics modeling indicated that invertebrate planktivory by Chaoborus depressed cladoceran populations during early summer. In particular, bosminids that generally form the spring biomass peak of cladocerans in stratified temperate lakes did not appear in the samples until July. In July, predation pressure by chaoborids was relaxed due to their emergence period and cladoceran population growth appeared to be limited by predation by planktivorous fish. The effect of fish predation was amplified by reduced refuge availability for cladocerans. The concentration of dissolved oxygen below the epilimnion was depleted, forcing cladocerans to move upward to less turbid and thus more dangerous water layers. The effect of size selective predation by fish resulted in reduced mean size of cladocerans during the period when refuge thickness (thickness of the water layer with oxygen concentration <1 mg l^?1 and water turbidity >30 NTU) was lowest. The results confirmed that in clay-turbid lakes, invertebrate predators could be the main regulators of herbivorous zooplankton even when cyprinid fish are abundant.     

Longitudinal changes in zooplankton distribution below a reservoir outfall with reference to river planktivory

The fate and interactions with river organisms of zooplankton as they drift downriver from a reservoir on a fourth-order mountain stream (Hiji River, Japan) were investigated. Monthly samples were collected at the reservoir and six river sites, simultaneously, from May 2005 to May 2006. Aquatic macroinvertebrates and fish were colleted, and their stomach contents were analyzed in April and May, 2006, respectively. Drift from the reservoir was the primary source for the river plankton community; the abundance of zooplankton, particularly those of cladocerans and large rotifer, rapidly decreased within several kilometers of the dam. Analysis of the contents of fish stomachs showed that drifting zooplankton was the main food for fish, with strong food selectivity for cladocerans and large rotifers. However, fish and insect planktivores showed longitudinally different stomach contents, with progressively fewer zooplankton found in the stomachs at the downriver sites. The results suggest that the outflow of zooplankton from the reservoir is an important food source for the downstream predators, especially fish, but the drift of zooplankton and consequent food availability for the predators at lower sites are strongly limited by concentrated fish predation just below the reservoir dam.     

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

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

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.     

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)     

Seasonal dynamics of free and attached heterotrophic nanoflagellates in an oligomesotrophic lake

1. The seasonal development of heterotrophic nanoflagellates (HNF), bacteria, rotiferans and crustacean zooplankton was studied in the epilimnion of Lake Pavin, an oligomesotrophic lake in the Massif Central of France.
2. HNF abundance varied from 0.1 to 2.5 × 10³ mL^–1. Free-living HNF reached their highest density in spring when the copepod Acanthodiaptomus denticornis dominated the metazooplankton. They were present in low numbers when rotifers and cladocerans were numerous.
3. Attached HNF, consisting of bicoecids and choanoflagellates, were fixed to large diatoms and to the colonial cyanobacterium Anabaena flos-aquae . The abundance of attached HNF was significantly correlated to bacterial abundance, which fluctuated between 1.1 and 2.7 × 10⁶ mL^–1. Highest abundance of these epiphytic protists was recorded when free-living heterotrophic nanoflagellates declined.
4. The comparison of the dynamics of heterotrophic nanoflagellates, bacteria, and the impact of zooplankton grazing suggested that prey abundance, the presence of suitable attachment sites and limited competition from the free-living forms were the main factors controlling the development of the epiphytic flagellate protists. In contrast, the low abundance of free-living forms during the period of rotiferan and cladoceran development suggests the prevalence of a top-down control by predation of the metazoopankton.     

Water discharge-regulated bacteria/heterotrophic nanoflagellate (HNF) interactions in the water column of the river Rhine

Heterotrophic nanoflagellates (HNF) make up a large fraction of the zooplankton biomass of rivers. Their abundance can be strongly affected by water discharge, but the consequences of this highly dynamic factor for their main prey, the bacteria, is still unknown. The focus of this study was on bacterial/HNF interactions in the Lower River Rhine (Germany) with respect to the discharge-dependent dynamics. The bacterial and HNF abundances and biomasses were determined over the course of 17 months. The potential consumption of bacteria by HNF was calculated based on the biomass data and on data on the HNF production. The mean bacterial abundance in the Rhine at Cologne ranged from 0.3 x 10(6) to 3.5 x 10(6) cells mL(-1), with lowest abundances in winter and highest in late spring. No significant changes in abundance during the downstream passage were found. Neither could a significant correlation be found between bacterial and HNF abundance. The ratio of bacterial to HNF abundance showed high variations which lay between 166 and 19,055 and was negatively dependent on water discharge. Monthly routine calculations on the potential bacterial consumption by HNF revealed a clearance of between 2 and 82% of the bacterial standing stock d(-1). The values increased greatly with water discharge and could exceed 100% d(-1) at times of high water flow. The presented data suggests a change in the top-down control of the planktonic bacteria due to the water discharge: The importance of benthic predation at low water flow (high contact probability to benthic predators) gives way to an increased importance in predation by planktonic HNF at high water flow.     

Factors affecting the bacteria-heterotrophic nanoflagellate relationship in oligo-mesotrophic lakes

The coupling between bacteria and heterotrophic nanoflagellates(HNF) was examined in nine lakes of low productivity for evidenceof the effects of various metazooplankton (i.e. rotifers, cladoceransand copepods) on this relationship. We considered the size ofcladocerans and, in contrast to most previous across-systemstudies, the three strata of the water column (i.e. epilimnion,metalimnion and hypolimnion). Rotifers were numerically dominantin all lakes and accounted for 45–84% of total metazooplanktonabundance, while the abundance of large cladocerans was relativelylow, ranging from 0.066 to 15.2 ind. L^–1. The across-lakerelationship between bacteria and HNF was significant in thedeeper strata (meta- and hypolimnion) but not in the epilimnionand in the two groups of lakes separated on the basis of theiraverage number of large cladocerans (<5 and >5 ind. L^–1,respectively). The results confirmed the negative impacts oflarge cladocerans on HNF, but also showed that rotifers, probablythrough grazing on HNF, may be an important factor causing variationin the bacteria–HNF relationship in unproductive waters.Quadratic models best described the relationships between metazooplanktonand the ratio of bacteria to HNF. This ratio seemed to be aresult of complex interactions between several factors, includingthe zooplankton composition and abundance and the depth of thelake. Indeed, this ratio significantly decreased across lakes,with increase in depth. In addition, shallower lakes (having<5 large cladocerans L^–1 and fewer Polyarthra vulgaris)tended to have more bacteria and HNF and a higher ratio of bacteriato HNF than deeper lakes (which had >5 large cladoceransL^–1 and substantial proportions of P. vulgaris). We suggestthat the epilimnion, metalimnion and hypolimnion of lakes betaken into account when analysing the bacteria–HNF relationshipas well as the cascading effects of zooplankton on microbialcommunities.     

Bottom-up versus top-down effects on ciliate community composition in four eutrophic lakes (China)

Previous studies have shown that ciliate plankton is generally controlled by food resources (e.g., algae) and predators (e.g., metazooplankton). Among lakes with similar trophic levels but different distributions of phyto- and metazooplankton, the main forces acting on ciliate assemblages may be different. We investigated the relationship between ciliate communities and bottom-up versus top-down variables based on a survey of four subtropical eutrophic lakes (China). Two of the lakes (Chaohu, Taihu) are located on the Mid-lower Yangtze Plain near sea level, and the other two (Dianchi, Xingyunhu) on the Yunnan-Kweichow Plateau at 1700 m above sea level. Blooms of cyanobacteria developed during summer in Lakes Chaohu and Taihu and throughout the year in Lakes Dianchi and Xingyunhu. Ciliate functional feeding groups differed significantly between lakes. The results of canonical correspondence analysis (CCA) and variation partitioning showed that cyanobacteria significantly influence ciliate species, whereas ‘edible’ algae (cryptophytes, diatoms) and cladocerans were the important variables in explaining the ciliate community structure of Lakes Dianchi and Xingyunhu compared with Lakes Taihu and Chaohu. Our results highlight the importance of consistent cyanobacterial blooms in shaping the ciliate community in subtropical eutrophic shallow lakes by interacting with top-down and bottom-up factors.     

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.     

Copepod and cladoceran success in an oligotrophic lake

Cladocerans have proved to be much less successful members ofthe Lake Tahoe limnetic community than the calanoid copepods.An analysis of temporal and spatial heterogeneity of the cladoceranBosmina longirostris and the herbivorous calanoid copepod Diaptomustyrrelli revealed important differences between the two typesof zooplankton. Although the productivity of both species islimited to varying degree by the abundance of certain algalspecies and the availability of particulate nitrogen, they differstrongly in the mechanisms responsible for mortality. Correlationanalysis suggests that B. longirosris is severely limited temporallyand spatially by predation from the omnivorous calanoid copepodEpischura nevadensis. In contrast, D. tyrrelli is not noticeablylimited by predators. Instead, abiotic seasonal events and starvationcontrol its population dynamics. Mortality and natality differencesbetween Bosmina and Diaptomus in Lake Tahoe help to explainthe general lack of success of cladocerans in oligotrophic lakes.     

Spatiotemporal food web dynamics along a desert riparian–upland transition

Increased awareness of spatiotemporal variation in species interactions has motivated the study of temporally-resolved food web dynamics at the landscape level. Empiricists have focused attention on cross-habitat flows of materials, nutrients, and prey, largely ignoring the movement of predators between habitats that differ in productivity (and how predators integrate pulses in resource availability over time). We set out to study seasonal variation in food web interactions between mammalian carnivores and their rodent prey along a riparian–upland gradient in semi-arid southeastern Arizona which features both spatial and temporal heterogeneity in resource availability. Specifically, we tested whether mammalian carnivores spill over from productive, near-river habitats into adjacent, desert-scrub habitats; and if they do, to document the effects of this spillover on rodent communities. Furthermore, we examined seasonal variation in top-down effects by measuring changes in carnivore diet and distribution patterns and rodent populations over time. The results indicate that carnivores track seasonally-abundant resources across the landscape, varying both their diet and movement patterns. In turn, desert-scrub rodent population dynamics track seasonal shifts in carnivore habitat use but not resource availability, suggesting that predation plays a role in structuring rodent communities along the San Pedro River. Further evidence comes from data on rodent community composition, which differs between desert-scrub habitats near and far from the river, despite similarities in resource availability. Our data also suggest that seasonal omnivory helps predators survive lean times, increasing their effects on prey populations. Taken together, these results underscore the importance of spatiotemporal variation in species interactions, highlighting the complexity of natural systems and the need for further detailed studies of food web dynamics.     

Comparative and experimental approaches to top-down and bottom-up regulation of bacteria

The regulation of bacterial community biomass and productivity by resources and predators is a central concern in the study of microbial food webs. Resource or bottom-up regulation refers to the limitation of bacteria by carbon and nutrients derived from allocthonous inputs, primary production, and heterotrophic production. Predatory or top-down regulation refers to the limitation of bacteria below levels supportable by resources alone. Large scale comparative studies demonstrate strong correlations between bacterial productivity and biomass, suggesting significant resource regulation. Comparisons of the abundances of heterotrophic flagellates and bacteria, however, imply that in some cases there may be top-down regulation of bacteria in eutrophic environments. Experimental studies in lakes support the importance of resource regulation and reveal little top-down control from protozoans. Increases in bacterial abundance and production with nutrient enrichment were limited in enclosure experiments with high abundances of the cladoceran, Daphnia. Regulation of bacteria by Daphnia may occur in many lakes seasonally and prevail in some lakes throughout the year where these animals sustain dense populations. In most situations, however, bacteria appear to be limited primarily by resources.     

Habitat selection by predators and prey in communities with asymmetrical intraguild predation

Competition and predation have broad ecological consequences as they may influence individual behavior and community structure. In some cases, they are linked and predator and prey are also competitors (intraguild predation). I present a game theoretic model of habitat use by predators and prey under conditions of asymmetrical intraguild predation. This model predicts that when the diet of intraguild predators is restricted to intraguild prey and the resource for which predators and prey compete (the basal resource), co-occurrence is only stable when dietary overlap is low and productivity of the basal resource is not high. The addition of alternative resources for predators results in co-occurrence under all conditions. Variation in alternative resource productivity produces a continuum of intraguild prey distributions from matching relative habitat safety, to one that reflects both food and predation risk. When there is a substantial alternative resource for predators, the distribution of predators matches that of alternative resource availability while the distribution of prey is influenced by both habitat riskiness and food availability. The density and distribution of the predator's alternative resource thus influence habitat selection by the intraguild prey. This stresses the importance of indirect interactions in structuring habitat use in communities and the need to view habitat selection in a community context.     

Drivers assessment of zooplankton grazing on phytoplankton under different scenarios of fish predation and turbidity in an in situ mesocosm experiment

Zooplankton play a key role in energy transfer within lake food webs, but we have a poor knowledge concerning their role as phytoplankton grazers in shallow subtropical lakes. In this study, we aimed to determine how zooplankton grazing upon phytoplankton is altered in different scenarios of fish predation and turbidity, and we explored the relevance of grazing compared to other environmental variables, to explain phytoplankton biomass changes. A mesocosm experiment was conducted by including the following treatments: fish, turbidity, fish + turbidity, and a control (without fish or varying turbidity). The experiment lasted 21 days, and samples were taken four times. Zooplankton grazing was only effective for the microphagous group upon Cryptophyceae, while large Chlorophyceae and small pennate Bacillariophyceae biomass were benefited in the presence of copepods and cladocerans, being negatively affected by depletions in nitrogen availability. In the turbidity treatment, a reduction in phytoplankton biomass was obtained, artificially increasing zooplankton grazing on phytoplankton, while fish presence inhibited grazing of adult copepods and cladocerans. The other groups of phytoplankton were only influenced by the environment. This experiment suggests that phytoplankton biomass variations would be more affected by the environment than by zooplankton grazing in shallow lakes from the Paraná River.

     

High predation is of key importance for dominance of small-bodied zooplankton in warm shallow lakes: evidence from lakes, fish exclosures and surface sediments

The mean body size of limnetic cladocerans decreases from cold temperate to tropical regions, in both the northern and the southern hemisphere. This size shift has been attributed to both direct (e.g. physiological) or indirect (especially increased predation) impacts. To provide further information on the role of predation, we compiled results from several studies of subtropical Uruguayan lakes using three different approaches: (i) field observations from two lakes with contrasting fish abundance, Lakes Rivera and Rodó, (ii) fish exclusion experiments conducted in in-lake mesocosms in three lakes, and (iii) analyses of the Daphnia egg bank in the surface sediment of eighteen lakes. When fish predation pressure was low due to fish kills in Lake Rivera, large-bodied Daphnia appeared. In contrast, small-sized cladocerans were abundant in Lake Rodó, which exhibited a typical high abundance of fish. Likewise, relatively large cladocerans (e.g. Daphnia and Simocephalus) appeared in fishless mesocosms after only 2 weeks, most likely hatched from resting egg banks stored in the surface sediment, but their abundance declined again after fish stocking. Moreover, field studies showed that 9 out of 18 Uruguayan shallow lakes had resting eggs of Daphnia in their surface sediment despite that this genus was only recorded in three of the lakes in summer water samples, indicating that Daphnia might be able to build up populations at low risk of predation. Our results show that medium and large-sized zooplankton can occur in subtropical lakes when fish predation is removed. The evidence provided here collectively confirms the hypothesis that predation, rather than high-temperature induced physiological constraints, is the key factor determining the dominance of small-sized zooplankton in warm lakes.     

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

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