The development and reproductive output of three species of Cladocera (Crustacea,Branchiopoda) with different size spectra as the result of vertebrate and invertebrate predation impact

Seasonal changes in development and reproduction of three species of Cladocera, Bosmina longirostris, Bosmina coregoni and Diaphanosoma brachyurum, were studied in regard to impact of vertebrate (planktivorous fish) and invertebrate (cladoceran Leptodora kindtii and cyclopoid copepods Mesocyclops leuckarti and Thermocyclops oithonoides) predators. Each of the cladocerans represented a different prey type. The population of the largest body-sized D. brachyurum was not affected by predators suggesting unidentified mechanisms of predation-avoidance. Diaphanosoma displayed low reproductive output coinciding with high density. Medium-sized B. coregoni, influenced by planktivorous fish, invested in reproduction and carried large egg-clutches resulting in gradual increase in population density. The population of small body-sized B. longirostris, that was influenced by invertebrate predators, displayed high density and reproductive effort in spring, followed by extreme collapse in abundance and reproduction during summer months. The results suggest that invertebrate predators may suppress populations of some Cladocera more strongly than planktivorous fish.     

Contaminated sediments in the Elbe estuary: Ecological and economic problems for the Port of Hamburg

The effects of food density on competition between Bosmina longirostris and Bosmina fatalis from Lake Kasumigaura were examined in the laboratory. When the animals were reared with a high concentration of Chlorella, B. fatalis, which is slightly larger than B. longirostris in body length, had larger brood sizes and a higher rate of population growth than B. longirostris. When the species were reared together at high food density, B. fatalis overcame B. longirostris. But B. longirostris won the competition at low food density.The effects of Microcystis on the competition were also examined. When the two species of Bosmina were reared in a mixture of Chlorella and Microcystis, the populations of both species hardly increased, although individual B fatalis showed slightly better individual growth and reproduction than B. longirostris. Decomposed Microcystis, however, was utilized well by both species.In Lake Kasumigaura, the decomposition of abundant Microcystis seemed to raise the food level, favouring B. fatalis over B. longirostris.     

Differential feeding behaviour of the dominant cladocera as an explanation of zooplankton community structure in the Loosdrecht Lakes

The zooplankton of Loosdrecht Lakes is dominated byBosmina longirostris in the Spring period but is replaced by the largerB.coregoni in early summer at a time when phytoplankton populations, dominated by filamentous cyanophytes, are rapidly increasing. Throughout the summerB.coregoni coexists with the longer, but more slender,Daphnia cucullata.Different size-categories of¹⁴C labelled lake-phytoplankton were fed to natural assemblages of crustacean zooplankton to investigate effects of body length on feeding and the relative selectivities of theDaphnia andBosmina species on food particle-size.B.coregoni feeds more effectively, especially on larger sizes of food, thanB.longirostris and this may be a factor in the observed change of bosminid species dominance.B.coregoni is able to feed on a wide size range of particles and grazes and assimilates greater volumes of food thanD.cucullata. B.coregoni preferentially feeds on food-particles greater than 7 m whereasD.cucullata selects food below 7 m. The coexistence of these species appears to depend on a niche separation through different feeding behaviours and selectivities.     

Effect of competition on the demography of planktonic cladocerans — Daphnia and Diaphanosoma

Summary Isolated and mixed continuous cultures of Daphnia hyalina and Diaphanosoma brachyurum in lake water were maintained under laboratory conditions to elucidate demographic effects of competition. Population dynamics curves were obtained. Interspecific competition was revealed by the decrease in the average density of animals in the mixed versus isolated cultures and by the extinction of one species in the presence of the other. Within the first 50 days either Diaphanosoma (4 cases) or Daphnia (1 case) was the superior competitor, depending on the initial conditions. Further cultivation resulted in the extinction of Diaphanosoma in the mixed cultures. There were no statistically significant differences between the maximum rates of population increase (r _m ) in Daphnia and Diaphanosoma at the concentration of edible algae about 2·10⁵ m³ml^-1(0.293 and 0.286 days^-1, respectively). Time lags for density-dependent parameters of the populations were evaluated by means of rank cross-correlations. Regardless of the species identity the time lags of fecundity, birth rate, and the rate of population growth were significantly higher in the superior competitor. The initial conditions of culturing affected the time lags which in their turn influenced the outcome of the interaction. Enhanced competitive ability due to the maximized time lags in Daphnia was not associated with the loss of population stability. Conversely, it brought about destabilization of Diaphanosoma populations which seemed to be the ultimate cause of its extinction observed in the end of the experiment. Time lag of the population growth rate was well predicted based on the half-sum of time lags in birth and death rates (r²=0.80, P<0.001). Daphnia responded to competition with a sharp shortening of the time lags of fecundity, birth rate, and the population growth rate. It increased clutch size and showed inverse relationship between the fecundity time lag and average fecundity even though it was strongly suppressed by Diaphanosoma. The competitive ability was not related to the percentage of adults in the populations. In contrast to the current belief the major result of interspecific competition in the experiment was not a decrease in the rate of population growth but was a reduction in population time lags.     

Self-maintaining plankton: pelagic Cladocera in small microcosms with lake water

Monospecific and mixed populations of pelagic cladocerans, Daphnia galeata, Bosmina coregoni, and Diaphanosoma brachyurum of Lake Glubokoe (Moscow Region) were isolated in 0.6 litre flasks with their natural medium — epilimnion water. In the laboratory they persisted for more than 100 days without external additions of food until the experiment was stopped. In three-species cultures Diaphanosoma was outcompeted, but in the presence of only Bosmina it persisted. Daphnia coexisted with Bosmina in two- and three-species cultures the whole period of observations in spite of competition. Birth and death rates of animals in microcosms depended on their density indicating that experimental conditions did not spoil natural population regulation. Demographic time lags underlay the observed oscillations in numbers. Because the time of Cladocera persistence in the cultures was comparable with lake vegetational period I conclude that summer zooplankton is capable of existing solely at the expense of nutrient recycling within epilimnion. Density regulation of cladocerans reinforces the stability of the system herbivore — algae, but it is weakened by the demographic time lags.     

Abrupt Changes in Bosmina (Cladocera,Crustacea) Assemblages During the History of the Ostrowite Lake (Northern Poland)

During the 10000-year history of the Ostrowite Lake, there have been several episodes of change in dominance amongst species of the genus Bosmina. The dominants were alternately B.longirostris and Eubosmina spp. Amongst the subgenus Eubosmina, two species prevailed in different periods:Bosmina coregoni and Bosmina reflexa. The first species, Bosmina coregoni, is characterized by long antennae and short carapace mucrones, while the second one, Bosmina reflexa, by short antennae and very long mucrones. Bosmina reflexa was dominant at the beginning of the lake's history (Preboreal, 10000--9000 BP) and during the early Subboreal Period (5000–3500 BP). Only a small number of remains of Bosmina longispina(Eubosmina spp.) were present along the entire profile. Such strong changes in the domination of Bosmina species were observed for the first time in the sediments of Polish lakes. A similar transition from one species to another has been described in a few German lakes and was linked to climate changes. However, the changes in the Ostrowite Lake do not correlate with climate changes, but most probably, with changes in the lake's trophic level. This is suggested by the simultaneous increase of Bosmina reflexa and a good indicator of eutrophy, Bosmina longirostris.     

Staphylococci in Competition: III. Influence of pH and Salt on Staphylococcal Growth in Mixed Populations

Previous results showed definite repressive effects on the growth of staphylococci in mixed cultures due to the competitive growth of psychrophilic saprophytes. This study was continued, and the influence of other environmental factors, pH and salt, on the competition between staphylococci and saprophytes was investigated. Initial pH values varied from 5 to 9. At the extremes of the pH range, staphylococci failed to grow, while the saprophytes grew under all of the conditions tested. At pH 5, the growth curves for the saprophytes were markedly altered from those obtained at neutral pH. The lag phases were greatly lengthened at and below 20 C, but normal numbers of saprophytes were reached in the stationary phase. At pH 6 and 8, staphylococcal growth showed the same inhibition observed at pH 7, at and below 20 C; normal multiplication was observed above this temperature, but with accelerated death phases. Thus, pH did not primarily effect staphylococcal growth through its influence on saprophyte growth and competition, but rather directly affected the growth of Staphylococcus cultures. Salt concentrations from 3.5 to 9.5% were investigated for influence on staphylococcal growth in mixed populations. Above 3.5% salt, staphylococcal inhibition at and above 20 C was not as marked as in the controls, although normal numbers were never reached. The saprophytes were increasingly inhibited, and their lag phases materially lengthened as salt concentration was increased. Salt acted directly on the Staphylococcus population and also, by repressing saprophyte growth, decreased competition, which allowed the staphylococci to grow.     

Application of individual growth and population models of Daphnia pulex to Daphnia magna, Daphnia galeata and Bosmina longirostris

Daphnia models for individual growth and population dynamics have been developed in the manner of models developed by Gurney, McCauley, Andersen and others. All or most of the earlier models were parameterized for Daphnia pulex; we have used the D. pulex model as a baseline model for other species of Daphnia such as magna, galeata and also Bosmina longirostris. Because of the lack of ample data for D. magna, D. galeata and B. longirostris, some of the physiological data had to be relied on the other species whose data were available and in some case calibrated. We were able to produce reasonable results for individual growth as well as population dynamics under the controlled laboratory conditions. Most of the results were compared with the available laboratory data for population as well as growth. All the simulations have been done under high and low food concentrations. The animals are assumed to be feeding on green algae (Chlamydomonas reinhardtti) under the laboratory conditions of 18–20°C. The continuous growth until the end of the life was observed in smaller B. longirostris, whereas rapid growth in the beginning and slower after the start of the reproduction was observed in Daphnia species. The smaller species matured earlier than larger species. B. longirostris population sustained better than Daphnia species in medium food concentrations.     

Population dynamics of Bosmina longirostris (Crustacea: Cladocera) in Lake Kariba,Zimbabwe

The population dynamics of Bosmina longirostris in a large tropical reservoir, Lake Kariba, Zimbabwe, are described. Population density showed two peaks associated with periods of high nutrient fluxes. Fecundity was always low. Two eggs per female was the maximum ever recorded. Instantaneous rates of birth, death and population growth calculated from field data using the egg ratio method are discussed. The rate of biomass change fluctuated in a regular pattern, probably linked to food availability or predation by a clupeid, Limnothrissa miodon.     

Combined Effects of Food Concentration and Temperature on Competition Among Four Species of <Emphasis Type="Italic">Brachionus </Emphasis>(Rotifera)

We evaluated the effect of algal food density (1.5 × 10⁶, 3.0 × 10⁶ and 4.5 × 10⁶ cells ml⁻¹ of Chlorella) and temperature (22° and 28 °C) on competition among the rotifers Brachionus calyciflorus, Brachionus havanaensis, Brachionus patulus and Brachionus rubens, based on population growth experiments for 24 days. The growth experiments were conducted seperately for each individual rotifer species (i.e., controls), and in mixtures of all four species in equal initial proportions (i.e., under competition). The population growth of B. calyciflorus, B. havanaensis, B. patulus and B. rubens grown separately at two temperatures and at three algal food densities showed typical patterns of lag, exponential and retardation phases in the controls. This pattern differed considerably under competition. In general, we observed that in all of the test species, the highest growth rates were observed at higher food levels and in the absence of congenerics. At 22 °C, under the lowest food level, the differences in the population abundances of B. havanaensis, B. patulus and B. rubens grown alone and in the presence of competition were large. However, these differences reduced as food density was increased from 0.5 × 10⁶ to 4.5 × 10⁶ cells ml⁻¹. At 28 °C and at the lowest food level, all of the other rotifer species eliminated B. havanaensis in mixed cultures. Each brachionid species had a higher rate when grown alone than when cultured with other species. The highest r (mean ± standard error: 0.54 ± 0.01 day⁻¹) was recorded for B. havanaensis at 28 °C under 4.5 × 10⁶ cells ml⁻¹ of algal food density. At 28 °C at low algal food density, the presence of competitors resulted in negative population growth rates for three of the four rotifer species tested.     

Population dynamics and production of Daphnia hyalina and Bosmina longirostris in a shallow, eutrophic reservoir

SUMMARY. 1. The population densities, dynamics and production of Daphnia hyalina and Bosmina longirostris were studied over a 2-year period in a shallow eutrophic reservoir in eastern England. The diet of the two species was assessed and their ecology was compared in relation to environmental factors. 2. Daphnia hyalina was characterized by small overall body size, probably due to heavy size-selective predation by fish. The first generation produced from the overwintering population appeared in April and Depopulation peaked in late spring and early summer. Adult survival was poor. Peak egg production varied between years, while mean brood size fell in summer probably due to food limitation during the cyanobacterial bloom. At this time selection for individual algal species was high and many guts were empty. Reproductive ratios were inversely related to population density. Bosmina longirostris exhibited a similar annual cycle. 3. Patterns of production were very different in the 2 years in D. hyalina. In 1981 production was high from August to early October, but in 1982 the peak was in spring. Annual production was 32.4 g C m^?2 in 1981 and 13.3 g C m^?2 in 1982. In 1981 B. longirostris production peaked in spring and late summer; in 1982 there was a high production in summer only. Annual production was 3.2 g C m^?2 in 1981 and 2.2 g Cm^?2 in 1982. 4. The number of adult D. hyalina was highly correlated with water temperature and total phytoplankton biomass, but these were negatively correlated with the number of eggs, mean brood size and instantaneous birth rate. In contrast, clutch size of B. longirostris was positively correlated with water temperature while birth rate was positively correlated with both temperature and phytoplankton biomass. The biomass of both species was positively related to total phytoplankton biomass, but neither relationship explained much of the variation in zooplankton biomass. 5. The significance of the observations on plankton in terms of managing the water quality in eutrophic reservoirs is discussed.     

The late Pleistocene/Holocene and recent Bosmina (Eubosmina) fauna (Crustacea: Cladocera) of the pre-alpine Starnberger See (FRG)

An analysis of the remains of Bosmina (Eubosmina) in a sedimentcore from the pre-alpine, mesotrophic Starnberger See showedthat the lake was inhabited by B. longispina since the OldestDryas. In the Subatlantic- -1900 years ago — the kesslerimorph of B. coregoni invaded the lake and both taxa have sinceco-existed. Although there was considerable morphological variation,B. longispina clearly differed from Circumbaltic populationsthrough most of the late Pleistocene and Holocene periods. Inthe recent Bosmina fauna of the lake two morphologically wellseparated taxa (B. longispina ruehei and a morph of B. coregoni)occur which differ clearly from the Bosmina remains found inthe core up to the uppermost sample from 7 cm sediment depth.B. coregoni f. kessleri appeared in the lake in a period whenit was supposedly still oligotrophic. The Bosmina successionobserved in the Starnberger See resembles the pattern foundin deep north German lakes.     

Effects of acidic pH and phytoplankton on survival and condition of Bosmina longirostris and Daphnia pulex

The effects of pH, algal composition and algal biomass on abundance, size, reproduction and condition of Daphnia pulex and Bosmina longirostris were tested in a field experiment using water and natural phytoplankton assemblages from a circumneutral (pH 6.43) and a moderately acidic (pH 5.75) lake in south-central Ontario. Both species were affected by pH and phytoplankton composition, with decreased egg production, lipid reserves, body size or abundance in treatments containing algae and/or water from the more acidic lake compared to treatments containing water and phytoplankton from the circumneutral lake. This result was unexpected for Bosmina, which often increases in relative and/or absolute abundance in acidified lakes. The negative effect of acidic conditions on Bosmina suggests that the population increase observed in most acidified lakes is not due to a positive response to low pH or ambient phytoplankton, but to altered biotic interactions possibly involving reduced competition.     

Field and experimental studies on the combined impacts of cyanobacterial blooms and small algae on crustacean zooplankton in a large,eutrophic, subtropical,Chinese lake

Field and experimental studies were conducted to evaluate the combined impacts of cyanobacterial blooms and small algae on seasonal and long-term changes in the abundance and community structure of crustacean zooplankton in a large, eutrophic, Chinese lake, Lake Chaohu. Seasonal changes of the crustacean zooplankton from 22 sampling stations were investigated during September 2002 and August 2003, and 23 species belonging to 20 genera were recorded. Daphnia spp. dominated in spring but disappeared in mid-summer, while Bosmina coregoni and Ceriodaphnia cornuta dominated in summer and autumn. Both maximum cladoceran density (310 ind. l⁻¹) and biomass (5.2 mg l⁻¹) appeared in autumn. Limnoithona sinensis, Sinocalanus dorrii and Schmackeria inopinus were the main species of copepods. Microcystis spp. were the dominant phytoplankton species and formed dense blooms in the warm seasons. In the laboratory, inhibitory effects of small colonial Microcystis on growth and reproduction of Daphnia carinata were more remarkable than those of large ones, and population size of D. carinata was negatively correlated with density of fresh large colonial Microcystis within a density range of 0–100 mg l⁻¹ (r = −0.82, P< 0.05). Both field and experimental results suggested that seasonal and long-term changes in the community structure of crustacean zooplankton in the lake were shaped by cyanobacterial blooms and biomass of the small algae, respectively, i.e., colonial and filamentous cyanobacteria contributed to the summer replacement of dominant crustacean zooplankton from large Daphnia spp. to small B. coregoni and C. cornuta, while increased small algae might be responsible for the increased abundance of crustacean zooplankton during the past decades.     

Prey handling time and ingestion probability for Mesocyclops sp. predation on small cladoceran species Bosmina longirostris, Bosminopsis deitersi, and Scapholeberis mucronata

We compared the vulnerability of small cladoceran species (Bosminopsis deitersi, Bosmina longirostris, and Scapholeberis mucronata) to predation by Mesocyclops sp. in the laboratory based on prey handling time and ingestion probability. We also estimated the effects of ontogenetic changes on handling time by testing prey of various body sizes. All tested prey species showed an increasing pattern of handling time along with growth (increase of body size). For juveniles smaller than 0.3mm, the prey handling time was similar for B. deitersi and B. longirostris, but markedly longer for S. mucronata. Mesocyclops sp. rejected large (>0.35mm) B. longirostris and S. mucronata soon after capture and consequently showed a low probability of successful ingestion, whereas most large individuals of B. deitersi were consumed. The shorter handling time and higher ingestion probability of small juveniles indicate that the copepods ingest small juveniles more efficiently than larger individuals. Thus, it seems that the vulnerability of small cladocerans to copepod predation differs ontogenetically, with the greatest vulnerability being during the juvenile period. The ontogenetic reduction in the vulnerability of B. longirostris and S. mucronata was shown in the rapidly reduced ingestion probability as well as by the increase in prey handling time for large individuals. The results show that the vulnerability to predation by copepods of B. longirostris and S. mucronata reduces more quickly with increasing size than does that of B. deitersi, and it seems that the former two species are more resistant to copepod predation.     

The effect of acidification in enclosures on the biomass and population size structure of Bosmina longirostris

An in situ enclosure experiment was performed from June to September, 1982, in Lake O'Woods, West Virginia (pH7.0), to examine the effects of acidification on the Bosmina longirostris population. Treatments were two control enclosures and two enclosures acidified from an initial pH of 7.2 to pH 4.2 over a 44 day period, using H₂SO₄. B. longirostris was not adversely affected by the acidification. In fact, its abundance, biomass, and mean body size increased in the acid treatment, as compared withe control. This positive response was probably not a direct consequence of the altered water chemistry, but instead may have occured due to reduced competition for food, and also reduced predation pressure. Acidification resulted in a decreased abundance of other herbivores, and elimination of the dominant carnivore, Mesocyclops edax.     

A conflict of morphological and genetic patterns in the Australian anostracan Branchinella longirostris

Branchinella longirostris is a fairy shrimp endemic to ephemeral pools on granite outcrops in southwestern Australia. The patchy nature of its habitat is thought to result in a high degree of subdivision among populations, potentially promoting speciation. We combined traditional taxonomy with a molecular phylogeny of cytochrome c oxidase I (COI) to test whether B. longirostris could be a species complex and whether the frontal appendage on the second antennae of males is a suitable character to differentiate new species. We also used nested clade analysis to assess the population structure and demographic factors explaining the geographical distributions of the mt DNA haplotypes. The results show that shapes of frontal appendages are not congruent with the mitochondrial genetic structure; however, they are positively correlated with geography. We conclude that the frontal appendage in B. longirostris is either subject to selection or a result of morphological plasticity; thus, its use in taxonomy remains uncertain. The intraspecific divergence in B. longirostris (≤7.7%) was approximately one third as large as the divergence from the outgroup (17.8–20.8%) and well within the ranges of divergence found in other crustaceans with fragmented population structure. There is some association between genetic structure and geography, resulting in the inference of restricted gene flow with isolation by distance and allopatric fragmentation as the most suitable models of the historical population processes. It is likely that the current distribution of haplotypes resulted from the dispersal of resting eggs by stochastic events (wind, birds) rather than from a fragmentation of previously continuous habitat as the estimated evolutionary age of the species (≤6.1 my) is much younger than its habitat (50–100 my). We found some evidence on the genetic level to support the hypothesis that B. longirostris could be a complex of species; however, the lack of correlation between the genetic pattern and the reproductively important frontal appendage lends a support to a view that B. longirostris is a single species with an exceptionally high intraspecific diversity.     

Effect of the epizoic rotifer Brachionus rubens on the population growth of three cladoceran species

Using population densities and growth rates as criteria, we studied interactions between the epizoic rotifer Brachionus rubens and each of three cladoceran species differing in size and reproductive rates — Daphnia carinata, Moina macrocopa and Ceriodaphnia rigaudi. In all mixed — species experiments, B. rubens existed in both the epizoic mode, attached to the cladoceran host, and in the free-swimming mode. Rotifer population growth rates were significantly depressed in the presence of M. macrocopa, presumably as a consequence of exploitative and interference competition. The largest cladoceran, D. carinata probably did not suppress B. rubens, because the epizoic component of the rotifer population escaped from the deleterious effects of mechanical interference. Peak population numbers and initial population growth rates reached by all three cladocerans were lower in the presence of B. rubens, probably because of the adverse effects of the epizoic infestation, which was maximal on D. carinata and least on C. rigaudi. In mixed-species cultures of D. carinata and M. macrocopa, the presence of B. rubens helped D. carinata coexist with M. macrocopa, which otherwise would have suppressed the Daphnia.     

Benthic–pelagic coupling in the population dynamics of the harmful cyanobacterium Microcystis

1. In eutrophic lakes, large amounts of the cyanobacterium Microcystis may overwinter in the sediment and re‐inoculate the water column in spring. 2. We monitored changes in pelagic and benthic populations of Microcystis in Lake Volkerak, The Netherlands. In addition, sedimentation rates and the rate of recruitment from the sediment were measured using traps. These data were used to model the coupling between the benthic and pelagic populations and to calculate the contribution of overwintering benthic and pelagic populations to the magnitude of the pelagic summer bloom. 3. Changes in the benthic Microcystis population showed a time lag of 3–14 weeks compared with the pelagic population. This time lag increased with lake depth. The largest amount of benthic Microcystis was found in the deepest parts of the lake. These observations suggest horizontal transport of sedimented Microcystis from shallow to deep parts of the lake. 4. Recruitment from and sedimentation to the sediment occurred throughout the year, with highest recruitment and sedimentation rates during summer. Model simulations indicate that the absence of benthic recruitment would reduce the summer bloom by 50%. 5. In spring, the total pelagic population was three to six times smaller than the total benthic population. Yet, model simulations predict that the absence of this small overwintering pelagic population would reduce the summer bloom by more than 64%. 6. Reduction of the overwintering pelagic populations, for instance by flushing, may be a useful management strategy to suppress or at least delay summer blooms of Microcystis.     

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.