The relative importance of algae and bacteria as food for Daphnia longispina (Cladocera) in a polyhumic lake

SUMMARY. 1. Grazing on algae and bacteria by the planktonic cladoceran, Daphnia longispina , was studied in a small polyhumic lake with low phytoplankton primary production in southern Finland.
2. D. longispina filtered algae at average rates of 0.09-0.82 ml ind⁻¹ h⁻¹. The filtering rates on bacteria were 26-92% of those on algae in parallel experiments.
3. From June to August algae, including mixotrophic and heterotrophic forms, comprised 56-93% of the food ingested by D. longispina. In mid-September and early October, when the Daphnia population was declining and the algal biomass was low, bacteria comprised 73% and 55% respectively of the food of Daphnia.
4. For D. longispina , an energy pathway via bacteria and bactivorous flagellates is probably a more important link to allochthonous organic matter than direct utilization of epilimnetic bacteria.     

The effect of small zooplankton on the microbial loop and edible algae during a cyanobacterial bloom

SUMMARY 1. We studied the effect of the small crustacean zooplankton on heterotrophic micro-organisms and edible phytoplankton in a eutrophic lake during a cyanobacterial bloom.
2. Small (15 L) enclosures were filled with natural or screened (100 μm) lake water and incubated for 5 days in the lake. Screening removed crustacean zooplankton but the initial density of rotifers and phytoplankton remained the same in control and removal treatments. Changes in the abundance and biomass of bacteria, autotrophic picoplankton (APP), heterotrophic nanoflagellates (HNF) and ciliates were measured daily.
3. The crustacean zooplankton, dominated by the small cladoceran Chydorus sphaericus , did not affect cyanobacteria, the main phytoplankton group during the experiment.
4. The removal of the crustacean zooplankton induced a higher abundance of ciliates and reduced that of the HNF, indicating the importance of ciliates in controlling HNF in this system.     

Image analysis of Daphnia populations: non-destructive determination of demography and biomass in cultures

SUMMARY 1. An image analysis technique was developed for the semiautomatic determination of abundance, size distribution and biomass in Daphnia cultures. This allowed detailed observations of growth, demography and biomass accumulation in live populations, avoiding artifacts caused by subsampling and sampling losses.
2. The image analysis method gave fast, non-destructive and reliable individual counts, even in cultures with high density and a large fraction of juveniles.
3. In Daphnia , animal width changes with nutritional status and growth within instar, while length changes only at the moult. Thus, estimation of individual biomass using an ellipsoidal model based on animal width gave improved biomass calculations compared to manual counting, sizing, and length : weight regressions.
4. The power of the image analysis technique for assessing population growth and size structure was demonstrated in two 40-day experiments, with Daphnia magna feeding on the green algae Selenastrum capricornutum in a two-stage chemostat system.     

Growth and survival of Daphnia in epilimnetic and metalimnetic water from oligotrophic lakes: the effects of food and temperature

SUMMARY 1. In oligotrophic lakes, phytoplankton and bacteria growing in the deep chlorophyll maximum in the cool metalimnion of lakes often dominate biomass and production, but the importance of this source of food for zooplankton is unknown.
2. During much of the day, Daphnia rosea in two mountain lakes inhabited deep chlorophyll layers where food availability was at least equal to that in the epilimnion.
3. To determine the importance of the two strata (epilimnion and metalimnion) for Daphnia , we used a cross-classified factorial experiment to measure how epilimnetic and metalimnetic food and temperature (10 and 16 °C) influenced survival, growth and reproduction.
4. Daphnia survived and grew better when fed seston from the epilimnion of one lake, although chlorophyll, particulate nitrogen and particulate carbon were 2–2.5 times greater in the metalimnion.
5. Temperature had no significant influence on Daphnia survival or growth. Similar results were obtained with food from the second lake, with Daphnia surviving and reproducing better when provided with epilimnetic, rather than metalimnetic food, although the quantities of chlorophyll and carbon in the two strata were similar.
6. Food quality, rather than quantity or temperature, appeared to be the most important determinant influencing survival, growth and reproduction, and the greater food quantity in the metalimnia was not used effectively by the Daphnia .     

Structure and functioning of the microbial loop in a boreal reservoir

The abundance and biomass of the main components of the microbial plankton food web (“microbial loop”)—heterotrophic bacteria, phototrophic picoplankton and nanoplankton, heterotrophic nanoflagellates, ciliates and viruses, production of phytoplankton and bacterioplankton, bacterivory of nanoflagellates, bacterial lysis by viruses, and the species composition of protists—have been determined in summer time in the Sheksna Reservoir (the Upper Volga basin). A total of 34 species of heterotrophic nanoflagellates from 15 taxa and 15 species of ciliates from 4 classes are identified. In different parts of the reservoir, the biomass of the microbial community varies from 26.2 to 64.3% (on average 45.5%) of the total plankton biomass. Heterotrophic bacteria are the main component of the microbial community, averaging 63.9% of the total microbial biomass. They are the second (after the phytoplankton) component of the plankton and contribute on average 28.6% to the plankton biomass. The high ratio of the production of heterotrophic bacteria to the production of phytoplankton indicates the important role of bacteria, which transfer carbon of allochthonous dissolved organic substances to a food web of the reservoir.     

Zooplankton-phytoplankton interactions: the case for refining methods, measurements and models

Many of the classic experiments on the interactions between animals and their food supply were performed using laboratory cultures of Daphnia but comparable predator-prey cycles have seldom been recorded in the field. In this paper, we report the results of a field experiment designed to examine the effect of seasonal variations in the supply of edible algae on the population dynamics of Daphnia hyalina var lacustris. The experiment was performed in a large (45 m diameter) enclosure that was fertilised at regular intervals to sustain phytoplankton production. The results demonstrate that predator prey interactions of the kind recorded in culture can be detected in the field if large numbers of samples are collected and the results analysed using appropriate methods. Analyses using the numbers of Daphnia collected at a single station and chlorophyll a as a measure of food availability produced inconclusive results. Similarly, Daphnia birth rates calculated using the total number of individuals provided a poor measure of the animals reproductive response to changes in the supply of food. The clearest evidence of systematic plant-herbivore interactions was obtained when improved estimates of Daphnia numbers and adult birth rates were combined with estimates of edible rather than total algal biomass. Daphnia birth rates were then positively correlated with the estimated amount of ingestible carbon (r = 0.77) and the proportion of gravid adults decreased dramatically when the concentration of food fell below 0.10 mg C l^-1. Measurements and models of phytoplankton loss rates were then used to assess the impact of Daphnia grazing on the growth and decline of three species of edible algae. The results demonstrated that most species were able to sustain positive growth rates despite short-term increases in the population filtration rate. For much of the summer, the Daphnia appeared to behave as non-interactive grazers and had very little effect on the growth rate of their main phytoplankton food. Their grazing activity did, however, arrest the growth of a late summer population of Chlorella when the Daphnia were estimated to be filtering all the water contained in the enclosure in less than a day.     

Heterotrophic organisms and viruses in the Oka River and Cheboksary Reservoir during the abnormally hot summer of 2010

In July 2010, abnormally high water temperature (25–29°C), as well as increased biomass and phytoplankton production caused intensive development of heterotrophic bacteria and heterotrophic nanoflagellates. It was found that the abundance, biomass, and production of heterotrophic bacterioplankton, as well as the abundance and biomass of heterotrophic nanoflagellates, and the number of planktonic viruses, which were calculated on average for the reservoir under study, turned out to be higher in the years with lower water temperature (20–23°C). The virus-induced mortality of bacterioplankton in the Oka River and the Cheboksary Reservoir averaged 25.4 ± 3.4 and 22.4 ± 2.7% of the daily bacterioplankton production.     

Life history variation within a parthenogenetic population of Daphnia parvula (Crustacea: Cladocera)

Summary Evidence for genetically determined life history variability within a population or a species is rare. In this three year experimental examination of a parthenogenetically reproducing population of the planktonic crustacean Daphnia parvula, we found evidence for a succession of clones or groups of clones that exhibited distinctive body size and reproductive differences that were maintained after numerous generations under standardized conditions in the laboratory. The D. parvula population reached maximum density in the fall and maintained relatively high densities through the winter and spring. Isolates from this fall-winter-spring period all had a larger body size at death and higher fecundity when compared with summer isolates under natural food and temperature conditions. These differences could not be accounted for by differences in temperature and food abundance among the seasons. An additional difference in these experiments was a shift in reproductive effort by the summer isolate which produced a higher proportion of its offspring in the first two broods. The shift in life history characteristics and a summer decline of the Daphnia parvula population was correlated with both an increase in inedible and perhaps toxic blue-green algae and an increase in a dipteran predator Chaoborus. Comparison of the survivorship curves for all of the seasonal life history experiments indicated that D. parvula survivorship was not lower during the summer discounting a toxic effect from blue-green algae. Positive population growth on natural food in the laboratory at this time indicated food was not limiting and that predation was the probable cause of the population decline.Laboratory life history experiments under standardized food and temperature conditions were run with D. parvula isolates from the spring and summer plankton. Genetically based differences as determined in these experiments were smaller body size, lower fecundity, smaller brood size, and shorter life span for the summer animals relative to spring animals. Thirty seven percent of the summer animals also reproduced at an earlier age under standardized conditions. The shift in reproductive effort to earlier broods by summer animals rnder natural conditions appeared to be a phenotypic response as the summer isolate did not produce a higher proportion of its offspring in early broods under standardized conditions.When estimates of predatory mortality were added to the life tables of the standardized experiments, the earlier reproduction of some of the summer animals allowed a population increase under a regime of intense predation. Life tables for the spring animals predicted a population decline under these circumstances. Predictable seasonal changes in biotic factors such as predation suggest a mechanism whereby diverse life history patterns with corresponding differences in r may be maintained within a population.     

Effects of algal food quality on fecundity and population growth rates of Daphnia

1. Food quality was at least as important as food quantity for both fecundity and population growth responses of the cladoceran Daphnia pulicaria fed the green alga Ankistrodesmus falcatus grown under N limitation, P limitation, or non-limited condition.
2. The fecundity of D. pulicaria was reduced under conditions of low food quality (low N or low P) compared with that for animals fed control non-limited algae regardless of ration size. The reduced fecundity of D. pulicaria fed P-limited food could be partially alleviated by increasing the ration (hence, compensation), but such was not the case for animals fed N-limited food.
3. Population growth rates of D. pulicaria ( r _max) were significantly reduced under conditions of low-quality food for both N-limited and P-limited algae. Population growth rates were unaffected by ration size, indicating no compensation.     

Biomass and oxygen dynamics of the epiphyte community in a Danish lowland stream

SUMMARY. 1. We examined the abundance and oxygen metabolism of epiphytic organisms on the dominant macrophyte, Potamogeton pectinatus , in headwaters of the eutrophic River Suså. Microbenthic algae were abundant in the stream during spring and macrophytes during summer.
2. The low macrophyte biomass in spring supported a dense epiphyte cover whereas the high macrophyte biomass during summer had a thin epiphyte cover of 10–100-fold lower abundance per unit area of macrophyte surface. The epiphyte community was dominated by microalgae in spring and by heterotrophs, probably bacteria, during summer. This seasonal shift was shown by pronounced reductions of the chlorophyll a content (from 2–3% to 0.1–0.7% of organic DW), the gross photosynthetic rate (from 20–85 to 3–15 mg O₂, g-¹ organic DW h⁻¹) and the ratio of gross photosynthesis to dark respiration in the epiphyte community (from 5–18 to 1). The reduced contributions of epiphytic microalgae correlated with reduced light availability during summer.
3. Both the density and the photosynthetic activity of epiphytic algae were low on a stream area basis relative to those of microbenthic algae and macrophytes. Rapid variations in water velocity and extensive light attenuation in water and macrophyte stands probably constrained the development of epiphytic algae. The epiphyte community was more important in overall stream respiration, contributing c. 10% to total summer respiration and c. 20% to summer respiration within the predominantly heterotrophic microbial communities on sediments and macrophyte surfaces.     

Chlorella-bearing ciliates dominate in an oligotrophic North Patagonian lake (Lake Pirehueico, Chile): abundance, biomass and symbiotic photosynthesis

1. Large mixotrophic ciliates ( Stentor araucanus , S. amethystinus and Ophrydium naumanni ) were a characteristic component of a temperate, oligotrophic lake in North Patagonia. During a 1-year study, the abundance, biomass and primary production of these large Chlorella -bearing ciliates were compared with those of the total plankton community.
2. Mixotrophic ciliates peaked in spring and from late summer to autumn, accounting for 1.6–43% (annual average: 16.3%) and 67–99% (annual average: 92%) of total ciliate abundance and biomass, respectively. Their contribution to total zooplankton biomass, including flagellates, rotifers, ciliates and crustaceans, was 14–76%, or 47% as an annual average. Endosymbiotic algae accounted for up to 25% of total autotrophic biomass (annual mean: 3.9%).
3. Maximum cell-specific photosynthetic rates of S. araucanus and S. amethystinus at light saturation varied between 80 and 4400 pg C ciliate^–1 h^–1 with high values during autumn and winter, and low values during summer. The depth-integrated rates of photosynthesis (0–40 m) of algal endosymbionts contributed 1–25% to total photosynthesis (annual mean: 6.5%).
4. A comparison of calculated ingestion rates with photosynthetic rates of Stentor indicates that photosynthate produced by endosymbionts generally exceeded heterotrophic food supply of Stentor during autumn and winter, but was much lower during summer, when food supply was high.
5. The mixotrophic ciliates represent an important 'link' between nanoplankton and higher trophic levels within the plankton community because of their high heterotrophic biomass and considerable contribution to total photosynthesis.     

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.     

Seasonal dynamics of crustacean zooplankton, heterotrophic nanoflagellates and bacteria in a shallow, eutrophic lake

1. The seasonal development of crustacean zooplankton, heterotrophic nanoflagellates (HNF) and bacteria was examined in Grosser Binnensee, a shallow, eutrophic lake in northern Germany. The grazing impact of Daphnia on bacteria and nanoflagellates was estimated from field data on population abundances and from clearance rates obtained in laboratory experiments. 2. The seasonal succession of zooplankton showed distinct peaks of Daphnia magna, cyclopopid copepods, Bosmina longirostris and Daphnia galeata and D. hynlina. The population dynamics of Dapfinia had the strongest impact on all sestonic components. Daphnia maxima coincided with clearwater phases, and were negatively correlated with particulate organic carbon (POC), HNF and phytoplankton. Bacterial abundance was only slightly affected although daphnids were at times more important as bacterial consumers than HNF, as estimated from measured bacterial clearance rates. Other crustaceans (copepods, Bosmina) were probably of minor importance as grazers of bacteria and nanoplankton. 3. HNF abundance varied from 550 ml^?1 to more than 30000 ml^?1. HNF appeared to be suppressed by daphnids and reached highest densities when copepods dominated the metazooplankton. The variation in HNF abundance was not reflected in the concentration of heterotrophic bacteria, which fluctuated rather irregularly between 5 and 20 ± 10⁶ ml^?1. Long filamentous bacteria which were probably resistant to protozoan grazing, however, appeared parallel to the development of HNF. These bacterial cells, although small in number, could comprise more than 30% of the total bacterial biomass.     

Resource-mediated demographic variation during the midsummer succession of a cladoceran community

SUMMARY. 1. The midsummer succession of cladoceran zooplankton in Lake Texoma was examined by demographic characterization of lake populations and a parallel set of life table experiments in which the dominant cladoceran species were raised on naturally occurring lake phytoplankton at lake temperatures. Diaphanosoma leuchtenbergianum occurred throughout the study period (mid June-mid August 1981) but Daphnia parvula, D. galeata mendotae and Ceriodaphnia lacustris were rarely detected after 15 July.
2. Analyses of size structure and size-specific fecundity of the cladoceran populations showed that (1) size structure shifted to smaller individuals in mid July, (2) size-specific fecundity was relatively constant in the smaller size classes and was constant or declined in larger females of each species, and (3) size at maturation did not vary significantly during the study of any population.
3. Individual growth rates of the two Daphnia species and Ceriodaphnia in the life table experiments were significantly lower in mid July when populations were declining and when size structure of each population was more dominated by smaller Individuals. The reduction in individual growth rates was short-lived in the life table experiments; growth rates returned to pre-decline levels by the end of July.
4. Two effects of temporary reductions in individual growth rates when size-specific fecundity and size at maturation are constant are (1) that size structure is temporarily shifted to smaller individuals, and (2) the potential rate of population growth is reduced because of delays in reproduction. Because data on size structure and size-specific fecundity are commonly used to evaluate the relative effects of predators and resource changes on populations it is noteworthy that erroneous conclusions about the mechanism of population decline may be reached when individual growth rates vary.     

Trophic interactions in rockpool food webs: regulation of zooplankton and phytoplankton by Notonecta and Daphnia

We studied trophic interactions in experimental rockpools with three different food web structures: phytoplankton and small-bodied zooplankton; phytoplankton, small-bodied zooplankton and Daphnia ; and phytoplankton, small-bodied zooplankton, Daphnia and Notonecta . Nutrients, primary productivity, chlorophyll a and zooplankton species composition and biomass were measured over eight weeks.
2. Daphnia had a negative impact on other zooplankton and reduced the phytoplankton biomass and primary productivity. In the absence of Daphnia , small-bodied zooplankton species were abundant, in particular cyclopoid copepods. Concentrations of dissolved nutrients were lower and the standing crop of primary producers was higher when Daphnia was absent.
3. The presence of the invertebrate predator Notonecta produced a top-down effect which was similar to that reported for planktivorous fish, i.e. a selective reduction of daphnids followed by an increase of small-bodied zooplankton species and phytoplankton biomass.
4. The study showed that consumer regulation of Daphnia by Notonecta and of algae by Daphnia are important, but also demonstrated that trophic level biomasses were controlled by a combination of predation and resource limitation.     

Seasonal variation of phototrophic picoplankton in Lake Biwa (1994–1998)

Seasonal abundances of phototrophic picoplankton (PP) and heterotrophic nanoflagellates in Lake Biwa were studied from 1994 to 1998. Seasonal variation in cell volume and biomass of the phototrophic picoplankton were also studied. PP were counted using disposable glass microscopic plates, which gave superior accuracy to sample filtration onto membrane filters. Phycoerythrin-rich rod-shaped cyanobacteria (PEC), one of the major components of the picoplankton community, were sparse (about 10⁴ cells ml ^–1) in winter and began to increase in April. Several PEC peaks were observed during the period of thermal stratification, and a rapid fall took place after October or November. In the northern basin, PEC peaked during late June and early July in 3 of the 5 years, and in late summer in the remaining years. Phycocyanin-rich rod-shaped cyanobacteria (PCC) were abundant in the southern basin and were present in smaller numbers in the eutrophic nearshore area of the northern basin; they peaked several times during the period from July to October. Seasonal variations of these two kinds of picoplankton were correlated with seasonal changes in water temperature. Phycoerythrin-rich cylinder-shaped cyanobacteria exhibited narrow peaks in July, their abundance declining as the year progressed. The density of heterotrophic nanoflagellates was greatest in early spring. Average cell volume of PEC was largest in winter, then decreased gradually to a minimum in late summer; after the fall, it recovered to the winter cell volume. This change can likely be attributed to the depletion of nitrogen in the warmer seasons.     

Dynamics and trophic roles of heterotrophic protists in the plankton of a freshwater tidal estuary

Freshwater tidal estuaries comprise the most upstream reaches of estuaries and are often characterised by the presence of dense bacterial and algal populations which provide a large food source for bacterivorous and algivorous protists. In 1996, the protistan community in the freshwater tidal reaches of the Schelde estuary was monitored to evaluate whether these high food levels are reflected in a similarly high heterotrophic protistan biomass. Protistan distribution patterns were compared to those of metazoan zooplankton to evaluate the possible role of top-down regulation of protists by metazoans. Apart from the algivorous sarcodine Asterocaelum, which reached high densities in summer, heterotrophic protistan biomass was dominated by ciliates and, second in importance, heterotrophic nanoflagellates (HNAN). HNAN abundance was low (annual average 2490 cells ml^–1) and did not display large seasonal variation. It is hypothesised that HNAN were top-down controlled by oligotrich ciliates throughout the year and by rotifers in summer. Ciliate abundance was generally relatively high (annual average 65 cells ml^–1) and peaked in winter (maximum 450 cells ml^–1). The decline of ciliate populations in summer was ascribed to grazing by rotifers, which developed dense populations in that season. In winter, ciliate populations were probably regulated `internally' by carnivorous ciliates (haptorids and Suctoria). Our observations suggest that, in this type of productive ecosystems, the microbial food web is mainly top-down controlled rather than regulated by food availability.     

The influence of predation on horizontal distribution of zooplankton species

SUMMARY. 1. The horizontal distributions of Daphnia longispina and Bosmina tongispina in Lake Kvernavatn (Norway) were investigated twice in 1982. In late spring, when populations were small, the two species inhabited the same areas, and they were evenly distributed from the littoral to the pelagic. At high population densities, during midsummer, the species were spatially segregated, D. longispina being pelagic and B. longispina littoral in distribution.
2. The distribution and feeding of three-spined sticklebacks (Gasteros-teus aculeatus) were also studied. The sticklebacks were apparently forced into littoral areas by larger piscivorous predators in the pelagic and they were consequently restricted to foraging primarily on B. longispina, which formed dense swarms during daytime in summer.
3. We suggest that predation and competition influence the spatial distribution of zooplankton species. The feeding efficiency of fish foraging on high-density zooplankton populations can be reduced by spatial segregation of zooplankton species. Where high local densities occur, due to swarm formation, predation is changed from size-selective feeding to consumption of spatially isolated individuals.     

Life history consequences for Daphnia pulex feeding on nutrient-limited phytoplankton

1. The growth and feeding of Daphnia pulex De Geer on different algal species was examined. The green algae Chlamydomonas reinhardtii Dangeard and Scenedesmus acutus Meyen, the diatom Synedra tenuissima Kützing, the cryptophyte Cryptomonas pyrenoidifera Geitler and the cyanobacterium Microcystis aeruginosa Kützing were cultured in non-limiting and in N- or P-limiting medium and used as food for D. pulex.
2. Growth limitations were reflected in the elemental and biochemical composition and the morphological characteristics of the algal resources.
3. The clearance rates of D. pulex feeding on nutrient-limited algae were reduced. This was not observed when nutrient-limited mutant Chlamydomonas cells without cell walls were used as food, indicating that the cell wall may play an important part.
4. In comparison with animals grown on nutrient-sufficient cells, nutrient-limited algae resulted in smaller body length, reduced brood sizes, reduced size at maturity, increased age at first reproduction and, consequently, in reduced Daphnia population growth rates.
5. Daphnia population growth rates ( r ) were negatively correlated with the C : P ratio and the carbohydrate content of the food. Moreover, significant correlations between r and clearance rates were found.
6. The observed differences in the grazing and the life history parameters of Daphnia feeding on non-limited and nutrient-limited algae may be the result of both reduced nutritional value and reduced digestibility of nutrient-limited algae.     

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.