Impact of Holoshestes heterodon Eigenmann (Pisces: Characidae) on the plankton community of a subtropical reservoir: the importance of predation by Chaoborus larvae

We analyzed the effects of planktivorous Holeshestes heterodon Eigenmann (Characidae) predation on the plankton community of a small subtropical reservoir, using four enclosures (volume about 17.5 m³), open to the sediment, established in the littoral zone. Two enclosures were stocked with fish (mean TL 5.7 cm), at a density of about 4–5 fish m^–3 (approx. 8 g m^–3), whereas two remained fishless. The experiment lasted a little longer than one month. In the fish enclosures, most Crustacea and Chaoborus larvae remained scarce, probably as a result of visually selective fish predation. In both fishless enclosures, Chaoborus larvae became abundant. However, in only one of these did large individuals become relatively numerous; this discrepancy in the demographic structure of the Chaoborus populations between the two fishless enclosures is unexplained. Only in the fishless enclosure without appreciable numbers of large Chaoborus did densities of Crustacea increase greatly. It is suggested that in the enclosure containing large Chaoborus individuals, crustacean populations were prevented from developing due to predation pressure, while the small Chaoborus larvae of the other enclosure could not readily consume these prey. Rotifers were low in abundance in the absence of fish, probably as a consequence of Chaoborus predation. Phytoplankton density increased in all four enclosures, due probably to the lack of water flow. Only in the fishless enclosure with high densities of crustaceans did phytoplankton abundance decrease markedly at the end of the experiment, perhaps because of grazing losses.     

Experimental study of trophic cascade effect of silver carp (Hypophthalmichthys molitrixon) in a subtropical lake, Lake Donghu: on plankton community and underlying mechanisms of changes of crustacean community

An enclosure experiment was carried out to test trophic cascade effect of filter-feeding fish on the ecosystem: growth of crustacean zooplankton, and possible mechanism of changes of crustacean community structure. Four fish biomass levels were set as follows: 0, 116, 176 and 316 g m^-2, and lake water (containing ca. 190 g m^-2 of filter-feeding fishes) was comparatively monitored. Nutrient levels were high in all treatments during the experiment. Lowest algal biomass were measured in fishless treatment. Algal biomass decreased during days 21–56 as a function of fish biomass in treatments of low (LF), medium (MF) and high (HF) fish biomass. Crustaceans biomass decreased with increasing fish biomass. Small-bodied cladocerans, Moina micrura, Diaphanosoma brachyurum and Scapholeberis kingii survived when fish biomass was high whilst, large-bodied cladocerans Daphnia spp. and the cyclopoids Theromcyclops taihokuensis, T. brevifuratus, Mescyclops notius and Cyclops vicinus were abundant only in NF enclosures. Evasive calanoid Sinodiaptomus sarsi was significantly enhanced in LF, but decreased significantly with further increase of fish biomass. Demographic data indicated that M. micrura was well developed in all treatments. Our study indicates that algal biomass might be controlled by silver carp biomass in eutrophic environment. Changes of crustacean community are probably affected by the age of the first generation of species. Species with short generation time were dominant and species with long generation time survived less with high fish biomass. Evasive calanoids hardly developed in treatments with high fish biomass because of the (bottle neck) effect of nauplii. Species abundance were positively related to fish predation avoidance. Other than direct predation, zooplankton might also be suppressed by filter-feeding fish via competition.     

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: changes in nutrient concentrations,phytoplankton and zoobenthos

Large bag-type (75 m³) and tube-type (105 m³) enclosures were set up in the shallow eutrophic Lake Suwa and were each stocked with exotic planktivorous whitefish (Coregonus lavaretus maraena). The release of whitefish caused the increase in nutrient concentration in the tube-type enclosure whereas no such increase was observed in the bag-type enclosure. Bottom sediment seemed to be an important source of chironomid food for whitefish. The proportion of phytoplankton measuring<10μm and 20–40μm, which respectively corresponded toOchromonas spp. andCryptomonas sp., were lower in the fish enclosures than in the control, which might have been caused by high grazing pressure by rotifers. The predation by whitefish might have affected the species composition of phytoplankton through reducing copepod predation on rotifers, not through reducing the densities of cladocerans which directly feed on phytoplankton as many investigators have reported. The phytoplankton biomass was not affected much by the release of fish. Possible reasons are that the increase in density of rotifers reduced the biomass of available phytoplankton and also that inedible Cyanophyceae were in the decreasing phase of their seasonal succession and could not increase successfully in spite of elevated nutrient levels.     

The short-term effects of perch predation on a zooplankton prey community

An enclosure experiment was performed in 1981 to determine if predation by perch (Perca flavescens) affected the density, depth distribution or size structure of members of the crustacean zooplankton community in Lac Choiniere, Quebec. Perch predation reduces the density of all but the smallest zooplankton in enclosures, and the relative rate of predation is influenced by both size and depth distribution of prey species. More individuals and proportionally more large individuals are found in deep water when perch are present. These results are discussed in relation to theories that size-selective predation alters numerous characteristics of zooplankton prey community structure.     

Influence of benthic and interstitial processes on nutrient changes along a regulated reach of a large river (Rhône River,France)

An enclosure study was conducted in Ranger Lake in south-central Ontario, Canada from 4 July to 5 August 1997 to determine predation effects of the larvae of the phantom midge fly Chaoboruson the zooplankton community. Zooplankton assemblages were established in 12 enclosures (2 m in diameter, 7.5 m deep). Three densities of fourth-instar Chaoborus trivittatus (0 l^–1, 0.1 l^–1 and 0.5 l^–1) were introduced as predator treatments to the enclosures. Temperature, dissolved oxygen and zooplankton community composition were monitored for six weeks. To determine if the zooplankton community composition changed, a repeated measures multivariate analysis was performed on percent biomass of Bosmina and calanoid copepods. There were no significant differences in mean taxon percent biomass among predator treatments. There were significant differences in mean taxon percent biomass between water layers (epilimnion and metalimnion). There were also significant differences in lengths of Bosmina and calanoid copepods among predator treatments at the end of the experiment. Crop content analysis of C. trivittatusshowed that Bosmina constituted 88–98% of the prey items found in the crops. These results demonstrate that the use of deep enclosures, a Chaoborus species which vertically migrates, and lower natural densities of Chaoborus may provide prey with an important natural refuge from predation and so allow a more accurate determination of the predation impact of Chaoborus trivittatusin temperate lakes where fish control Chaoborus densities.     

Plankton dynamics in a tropical floodplain lake: fish, nutrients, and the relative importance of bottom-up and top-down control

1. Two enclosure experiments were carried out in Laguna Bufeos, a neotropical várzea lake located in the floodplain of River Ichilo (Bolivia). The experiments aimed (i) to assess the relative importance of bottom‐up and top‐down control on the plankton community, (ii) to assess the relative impact of direct and indirect effects of planktivorous fish on the zooplankton, and (iii) to attempt to identify the mechanisms responsible for these effects. 2. During the first experiment, bottom‐up control seemed to dominate the planktonic food web. Compared with fishless enclosures, oxygen concentrations, chlorophyll a levels and the population densities of all cladoceran zooplankton taxa increased in enclosures with fish. Birth rates of Moina minuta, the dominant taxon, were substantially higher in the presence than in the absence of fish, whereas death rates did not differ between treatments. These results are the first to suggest that the positive effects of fish on crustacean zooplankton via effects on nutrient cycling and the enhancement of primary production can compensate for losses because of fish‐related mortality. 3. During the second experiment, the direction of control appeared to vary between trophic levels: the phytoplankton appeared to be bottom‐up controlled whereas the zooplankton was mainly top‐down controlled. Chlorophyll a concentrations were enhanced by both fish and nutrient additions. The majority of the zooplankton taxa were reduced by the presence of fish. Birth rates of most cladoceran taxa did not differ between treatments, whereas death rates were higher in the enclosures with fish than in the fishless enclosures. Bosminopsis deitersi reached higher densities in the presence of fish, probably because of a release from predation by Chaoborus. 4. We convincingly showed strong deviations from trophic cascade‐based expectations, supporting the idea that trophic cascades may be weak in tropical lakes.     

Effects of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community of a deep,tropical reservoir: an enclosure experiment

1. An in situ enclosure experiment was conducted in a deep reservoir of southern China to examine (i) the effects of a low biomass (4 g wet weight m^?3) of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community and (ii) the response of Daphnia to eutrophication. 2. In the absence of fish, Daphnia galeata dominated the zooplankton community, whereas calanoids were dominant in the fish treatments, followed by D. galeata. Silver carp stocking significantly reduced total zooplankton biomass, and that of D. galeata and Leptodorarichardi, but markedly increased the biomass of smaller cladocerans, copepod nauplii and rotifers. In contrast, nutrient enrichment had no significant effect on the plankton community except for cyclopoids. 3. Chlorophyta, Cryptophyta and Bacillariophyta were dominant phytoplankton groups during the experiment. Chlorophyta with high growth rates (mainly Chlorella vulgaris in the fish enclosures and Ankyra sp. in the fishless enclosures) eventually dominated the phytoplankton community. Total phytoplankton biomass and the biomass of edible phytoplankton [greatest axial linear dimension (GALD) < 30 μm], Chlorophyta, Cryptophyta, Bacillariophyta and Cyanobacteria showed positive responses to fish stocking, while inedible phytoplankton (GALD ≥ 30 μm) was significantly reduced in the fish enclosures. However, there was no significant effect on the plankton community from the interaction of fish and nutrients. 4. Overall, the impact of fish on the plankton community was much greater than that of nutrients. High total phosphorus concentrations in the control treatment and relatively low temperatures may reduce the importance of nutrient enrichment. These results suggest it is not appropriate to use a low biomass of silver carp to control phytoplankton biomass in warmer, eutrophic fresh waters containing large herbivorous cladocerans.     

Predation by a water mite (Piona exigua) on enclosed populations of zooplankton

Short term, replicated experimental alteration of densities of a predatory water mite Piona exigua Viets, inside 0.45–0.475 m³ litre enclosures, revealed little evidence of the effects of predation on the number and relative abundance of the enclosed zooplankton species. Predation rates more closely approximated those estimated from single prey functional response experiments in the second experimental period (December) than in the first (March). In December Daphnia was the only susceptible taxon present in large numbers, whereas in March, Ceriodaphnia and Chydorus were also present. This result is consistent with laboratory findings that predation rates are lowered in the presence of more than one prey type.The difficulty of obtaining evidence for significant effects of these planktonic predators is in part due to changes in the preferred prey species in the diet of Piona depending on stage and sex of the mite and to aspects of experimental design. The wide variability between replicate enclosures at each predator density reduced the power of the statistical analyses used to test the null hypothesis. Enclosures with no predators are necessary to investigate the effects of enclosure on the zooplankton prey, since these effects may outweigh those due to predator consumption.     

Seasonal dynamic of phototrophic epibionts on crustacean zooplankton in a eutrophic reservoir with cyanobacterial bloom

Seasonality of burden and prevalence of phototrophic (microalgal) epibionts Characidiopsis ellipsoidea, Colacium vesiculosum and Colacium sp. on dominating crustacean zooplankton (Daphnia longispina, Cyclops vicinus and Mesocyclops leuckarti) were studied in a small reservoir Bugach with cyanobacterial bloom. The correlations between the seasonal dynamics of prevalence and the dynamics of others biotic and abiotic factors were calculated. The conclusions were as follows. The substrate species, that determined the development of the epibionts on the three studied crustacean zooplankton, was Daphnia longispina (Cladocera). Despite intensive epibiotic infestation of crustacean zooplankton, epibionts did not appear to have caused non-consumptive mortality of the crustacean zooplankton. But they could have contributed to the Daphnia summer decline by increasing mortality due to its consumption by planktivorous fishes. The phototropic epibionts may successfully coexist with cyanobacterial bloom. The possible role of the epibionts in changing nutrient fluxes in pelagic food web is discussed.     

Predator-prey relationships within the pelagic community of Neusiedler See

Neusiedler See, a shallow alkaline lake, has become increasingly eutrophic; this enrichment improved the nutritive situation of the herbivorous zooplankton leading to a higher standing stock. A multiple regression analysis of the long-term development of the crustacean plankton indicates that abiotic factors (i.e. wind, temperature) have the most important impact on the community in spring and autumn, biotic factors (i.e. food, predation) during the summer months. Currently an invertebrate (Leptodora kindti) and two planktivorous fish (Pelecus cultratus and Alburnus alburnus) control the population development of Diaphanosoma mongolianum during summer. L. kindti predation acts on immature stages, whereas the fishes consume adult stages. L. kindti densities of 100 to 200 ind. × m^-3 affect the prey population to a negligible extent; densities between 300 and 500 ind. × m^-3 result in elimination rates of 5% to >40% of the juvenile Diaphanosoma stock. The impacts by the invertebrate predator are pronounced but short-term events. Juvenile and underyearling fish eliminate 4–13% of the zooplankton in the open lake and 5–33% inshore; their predation pressure acts on all younger stages of the crustaceans. Planktivorous fish older than 0 + concentrate on the large food items (adult stages of the crustaceans). A. alburnus is able to consume 6–16% of the zooplankton standing stock during summer. P. cultratus eliminates about 1–49% of D. mongolianum, 1–4% of A. spinosus and 1–31% of L. kindti. From May until August the position of the dominant predator changes, beginning with juvenile fish which are then followed by P. cultratus and then by A. alburnus. Predation by L. kindti is of importance in July and August; at that time it causes suppression of the juveniles of D. mongolianum, but, on the other hand, the older stages of L. kindti are endangered themselves by the predatory impact of planktivorous fish.     

Use of enclosures to detect the contribution of particular zooplankton to growth of young-of-the-year yellow perch (Perca flavescens Mitchell)

Summary Densities of the cladoceran, Holopedium gibberum, were manipulated in 18 enclosures containing juvenile (age 0+) yellow perch (Perca flavescens) and mean-lake densities of other zooplankton. In enclosures, where nearlake densities of all zooplankton species including Holopedium were maintained, young-of-the-year perch grew significantly heavier and longer than in experimental enclosures where Holopedium was excluded. Holopedium comprised between 15–45% of the diet (wet weight) of perch in the first 2 weeks of July in the control treatment (Holopedium at or near ambient lake densities) and only 3–7% of total biomass ingested in the experimental treatment (Holopedium density selectively reduced). Predation on Holopedium decreased dramatically after the 2nd week of July in the control treatment after which Chaoborus, chironomids, and Sida became dominant prey items (by weight) of juvenile perch. These findings suggest that growth and survivorship of age 0+ perch in Precambrian Shield lakes may be coupled to Holopedium abundance. Thus, utilization of Holopedium by young-of-the-year yellow perch may affect recruitment of this species since overwintering survivorship, range of accessible prey sizes or species, and vulnerability of juvenile perch to predation by larger fish depend on body size, which is reduced when Holopedium is excluded from the diet.     

Is coarse woody debris in lakes a refuge or a trap for benthic invertebrates exposed to fish predation?

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Fish-zooplankton interactions and their effects on water quality of a tropical Brazilian reservoir

The influence of zooplanktivorous fishes on the plankton community and water quality of Americana Reservoir, Brazil was studied experimentally in 4 floating enclosures during the dry seasons (July–September) of 1982 and 1983. Two enclosures were stocked with adult fish (Astyanax bimaculatus in 1982;A. fasciatus in 1983) at near maximal densities measured in the reservoir upper surface waters (35 m^–2) and two were fish-free during each experiment lasting about one month. Marked differences were evident between the fish and fish-free enclosures after a 2–3 week period in each experiment, particularly with respect to water transparency, phytoplankton biomass, and zooplankton abundance as well as species and size composition. By the end of each experiment water transparencies were lower and phytoplankton biomass higher in the fish enclosures compared to those without fish. Also at that time Rotifera were the prominent zooplankters in the fish enclosures and Cladocera in the fish-free ones. Larger or more conspicuous species of Cladocera asDaphnia gessneri, D. ambigua, andMoina micrura were present in the fish-free enclosures but not in the fish enclosures. The interactions between fish predation, zooplankton grazing, phytoplankton biomass and water quality conditions are discussed in relation to eutrophication of a tropical aquatic ecosystem.     

Diel vertical and horizontal distribution of crustacean zooplankton and young of the year fish in a sub-alpine lake: an approach based on high frequency sampling

Understanding the spatial dynamics of predators and their preyis one of the most important goals in aquatic ecology. We studiedspatial and temporal onshore–offshore distribution patternsin young of the year (YOY) Eurasian perch (Perca fluviatilis)and crustacean zooplankton (Daphnia hyalina, Cyclops prealpinus)along a transect in Lake Annecy (France). Our study representsa first attempt at coupling hydroacoustic fish survey and highfrequency zooplankton recording to assess simultaneously thelarge-scale distribution patterns of YOY fish and their zooplanktonprey over a diel cycle (day, dusk and night sampling). We hypothesizedthat the spatial distribution of zooplankton could be shapedby both anti-predator behaviour (horizontal and vertical migrations)and predation losses. Fish biomass, size structure and dietwere assessed from split-beam echosounding and net trawlingsamples, whereas crustacean abundances were estimated with asmall modified Longhurst–Hardy continuous plankton recorder.We evaluated the diel changes in the spatial distribution patternsof fish and zooplankton and determined the overlap between theirdistributions. Fish biomass was dominated by YOY perch in upperwarmer layers and salmonids (Coregonus lavaretus and Salvelinusalpinus) in the colder and oxygenated deep layers. YOY perchwere aggregated in dense schools in the epilimnion during theday and dispersed at night. Fish biomass was distributed alonga strong increasing onshore–offshore gradient at night,whereas crustacean prey showed a decreasing gradient. This onshore–offshorenegative gradient in crustacean distribution, expressed on ashorter scale during the day, shifted toward the surface watersat night. A distinct kinetic of diel vertical migration (DVM)patterns was exhibited by daphnid and cyclopoid populationsand resulted in distinct vulnerability to perch predation. Spatio-temporaldistribution of crustaceans in Lake Annecy during the diel cyclestudy was probably shaped both by predation loss to YOY perchand by anti-predator behaviour (DVM, DHM) by zooplankton. Theimplications for fine-scale studies of fish-zooplankton interactionsare discussed.     

富营养化对小型湖泊浮游甲壳动物群落结构及多样性的影响

2017年3月到2018年2月研究了临涣湖浮游甲壳动物群落结构的季节变化。临涣湖共记录浮游甲壳动物13种,其中枝角类8属8种,桡足类5属5种。短尾秀体溞(Diaphanosoma brachyurum)、广布中剑水蚤(Mesocyclops leuckarti)和象鼻溞(Bosmina sp.)等小型富营养种类是温暖季节的优势种,而近邻剑水蚤(Cyclops vicinus)是冬季的优势种。盔形溞(Daphnia galeata)等大型种类仅在少数月份中被观察到。临涣湖浮游甲壳动物的年平均密度和生物量分别为28.3个/L和0.33 mg/L。营养状态指数(TSI_M)的年平均值为62.6。浮游甲壳动物的Shannon指数、Pielou指数和Simpson指数的年平均值分别为0.86、0.74和0.49,且3种多样性指数均具有显著的季节差异。营养盐水平、营养状态指数和物种多样性指数均表明,临涣湖水体处于富营养化状态。冗余分析结果表明,水温、总磷浓度和叶绿素a浓度是影响临涣湖浮游甲壳动物群落结构变化的上行效应因子。鲢、鳙鱼的捕食压力是临涣湖浮游甲壳动物群落结构小型化的下行效应因子。     

Predation impact of Cyclops vicinus on the rotifer community in Lake Constance in spring

The predation impact of Cyclops vicinus on rotifers was studiedunder near-natural conditions in small enclosures to evaluatewhether copepod predation is responsible for the decline ofrotifers in Lake Constance in spring. Cyclops vicinus fed selectivelyon Synchaeta spp.; Keratella and Polyarthra spp. were not selectedfor. Predation rates increased with prey density up to a maximumof 37 Synchueta day^–1 at a density of 1.6 x 10⁶ Synchaetam^–2, i.e. at -1200 Synchaeta l^–1. Calculation ofcropping rates suggests that Cyclops alone can control the abundanceof Synchaeta in spring, i.e. that mainly Cyclops is responsiblefor the decline of Synchaeta species in Lake Constance in May.     

The impact of vertebrate and invertebrate predators on a stream benthic community

I assessed the impact of both vertebrate and invertebrate predators on a lotic benthic community in a 1-month-long experiment, using enclosures containing cobble/gravel bottoms, with large-mesh netting that allowed invertebrates to drift freely. Brown trout (Salmo trutta) and leeches (Erpobdella octoculata) were used as predators and four treatments were tested: a predator-free control, leeches only, trout only, and leeches and trout together. A density of 26.7 leeches/m² (20 leeches/enclosure) and 1.3 trout/m² (one trout per enclosure) was stocked into the enclosures. The total biomass of invertebrate prey was significantly lower in the trout and trout plus leech treatments than in the leech and control treatments, which were due to strong negative effects of trout on Gammarus. On the individual prey taxon level, both trout and leeches affected the abundance of Asellus , Baetis and Ephemerella, whereas the abundance of Gammarus was only affected by trout, and the abundance of Orthocladiinae and Limnephilidae was only affected by leeches. In the treatment with trout and leeches together, the abundance of Ephemerella and Baetis was higher than when trout or leeches were alone, which was probably due to predator interactions. Leeches and trout had no effects on prey immigration but did affect per capita emigration rates. Both trout and leeches indirectly increased periphyton biomass in enclosures, probably due to their strong effects on grazers. Both trout and leeches were size-selective predators, with trout selecting large prey, and leeches selecting small prey. Size-selective predation by trout and leeches affected the size structure of five commonly consumed prey taxa. Trout produced prey populations of small sizes owing to consumption of large prey as well as increased emigration out of enclosures by these large prey. Leech predation produced prey assemblages of larger size owing to consumption and increased emigration of small prey. These results suggest that in lotic habits, predatory invertebrates can be as strong interactors as vertebrate predators. Received: 23 June 1997 / Accepted: 4 May 1998     

Plankton community of a polyhumic lake with and without Daphnia longispina (Cladocera)

The impact of Daphnia longispina (Cladocera) on the plankton food web was studied in a polyhumic lake where this species comprised almost all zooplankton biomass. Plastic enclosures (volume 7 m³) were inserted into the lake retaining the initial water stratification except that in one enclosure zooplankton was removed. After the removal of Daphniaa rotifer, Keratella cochlearis, ciliates and heterotrophic nanoflagellates increased markedly and the density and biomass of bacteria decreased. Edible algal species, Cryptomonas rostratiformisand three small chrysophytes,Ochromonas, Pedinella and Spinifermonas, took advantage of the removal of Daphnia, while more grazing-resistant species declined. In spite of the changes in the species composition of phytoplankton, the removal of Daphnia did not affect the biomass, primary production or respiration of plankton. The results implied that the density of heterotrophic flagellates and ciliates was controlled by Daphnia, but in its absence the former took its role as the bacterial grazers.     

Density and size distribution of Daphnia populations at different fish predation levels

Density and size structure changes of natural daphniids populations were studied in enclosures with a different level of fish predation. Daphnia pulicaria was totaly removed in high predation variants, and the differences of the mean body length both in adults and juveniles are apparent between low predation enclosure and enclosure without fish. Daphnia galeata was replaced by D. magna and D. pulicaria in the enclosure without fish. The decline of densities and the substantial (30–50%) and fast (during 12 days) shift of mean body length both in adults and juveniles of D. galeata was induced by the fish (carp fry) introduction to the high predation enclosures.     

Different Predation Impacts of Two Cyclopoid Species on a Small-sized Zooplankton Community: An Experimental Analysis with Mesocosms

We used mesocosms to analyze predation impacts on the prey populations and prey community structures by two cyclopoid copepod species, the larger Mesocyclops pehpeiensis and the smaller Thermocyclops taihokuensis, who coexist with small-sized herbivorous zooplankton species in a fish-abundant lake. The overall predation impact on the prey populations was stronger for Mesocyclops than for Thermocyclops. Mesocyclops had a strong and less selective impact on the rotifer community but a selective impact on the crustaceans. In contrast, Thermocyclops had a selective predation impact on rotifers but a weak and less selective impact on the crustacean community. As a result, the former predator reduced the diversity of the crustacean community but not the rotifer community, while the latter had an opposite impact on the diversities of the two communities. It has been suggested that fish induce development of a zooplankton community dominated by the small-sized zooplankton species in fish-abundant lakes. Our results demonstrated that cyclopoid copepods altered species composition and diversity of the small-sized zooplankton community in such lakes. Thus, the results have given an important suggestion on the role of the invertebrate predator cyclopoid copepods, which often coexist with fish, that they determine population dynamics and community structures of small-sized zooplankton in fish-abundant lakes.