Stoichiometric differences in food quality: impacts on genetic diversity and the coexistence of aquatic herbivores in a <Emphasis Type="Italic">Daphnia</Emphasis> hybrid complex

The maintenance of genetic and species diversity in an assemblage of genotypes (clones) in the Daphnia pulex species complex (Cladocera: Anomopoda) in response to variation in the carbon:phosphorus ratio (quantity and quality) of the green alga, Scenedesmus acutus, was examined in a 90-day microcosm competition experiment. Results indicated that mixed assemblages of seven distinct genotypes (representing clonal lineages of D. pulex, D. pulicaria and interspecific hybrids) showed rapid loss of genetic diversity in all treatments (2 × 2 factorial design, high vs. low quantity, and high vs. low quality). However, the erosion of diversity (measured as the effective number of clones) was slowest under the poorest food conditions (i.e., low quantity, low quality) and by the conclusion of the experiment (90 days) had resulted in the (low, low) treatment having significantly greater genetic diversity than the other three treatments. In addition, significant genotype (clone) × (food) environment interactions were observed, with a different predominant species/clone found under low food quality versus high food quality (no significant differences were detected for the two food quantities). A clone of D. pulex displaced the other clones under low food quality conditions, while a clone of D. pulicaria displaced the other clones in the high food quality treatments. Subsequent life-history experiments were not sufficient to predict the outcome of competitive interactions among members of this clonal assemblage. Our results suggest that genetic diversity among herbivore species such as Daphnia may be impacted not only by differences in food quantity but also by those in food quality and could be important in the overall maintenance of genetic diversity in natural populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Can suspension feeding by bivalves regulate phytoplankton biomass in Lake Waccamaw, North Carolina?

Suspension feeding by bivalves has been hypothesized to control phytoplankton biomass in shallow aquatic ecosystems. Lake Waccamaw, North Carolina, USA is a shallow lake with a diverse bivalve assemblage and low to moderate phytoplankton biomass levels. Filtration and ingestion rates of two relatively abundant species in the lake, the endemic unionid, Elliptio waccamawensis, and an introduced species, Corbicula fluminea, were measured in experiments using natural phytoplankton for durations of 1 to 6 days. Measured filtration and ingestion rates averaged 1.78 and 1.121 ind.^–1 d^–1, much too low to control phytoplankton at the observed phytoplankton biomass levels and growth rates. Measured ingestion rates averaged 4.80 and 1.50 µg chlorophyll a ind.^–1 d^–1, too low to support individuals of either species. The abundance of benthic microalgae in Lake Waccamaw reaches 200 mg chlorophyll a m^–2 in the littoral zone and averages almost an order of magnitude higher than depth-integrated phytoplankton chlorophyll a. Total microalgal biomass in the lake is therefore not controlled by suspension feeding by bivalves.     

Egg size and clutch size in two Daphnia species grown at different food levels

The effect of three different Scenedesmus food concentrations(0.04, 0.2 and 1 mg C l^–1) on maternal investment wasstudied in two cladoceran species of similar size, Daphnia pulicariaand D.hyalina. It was observed that as food concentration decreased(between 1 and 0.2 mg C l^–1), there was an increase insize, protein content, lipid content, carbon and mass of theegg, while, at the same time, the clutch size of the femalesbecame smaller. Such an increase in ‘per offspring investment’was reflected in an increase in body length, body carbon andbody mass of neonates as the food available for females decreased.However, in D.pulicaria this tendency was not maintained downat 0.04 mg C l^–1 in which there was a decrease of theegg characteristics mentioned above. Although, there are notavailable all the egg and neonate parameters of D.hyalina at0.04 mg C l^–1, the body length of the neonates was largerthan at 0.2 and 1 mg C l^–1. These results show that, asfood diminishes, these two cladoceran species are able to respondby decreasing clutch size, but increasing the size of egg, therebyincreasing the probability of neonate survival. This tendencyis probably maintained until the food concentration is too lowand the females have to reduce the energy allocated for reproduction.     

Effects of crowding and different food levels on growth and reproductive investment of Daphnia

The effects of daphniid crowding on juvenile growth rate, length at first reproduction, clutch size and egg size of four species of Daphnia were compared with the effects of food level. Juvenile Daphnia were grown to primipary in a flow-through system in water conditioned by different densities of the same, or another, species. At high ambient food levels, water from Daphnia that had been crowded at densities 150 l^–1 depressed growth rate and lowered body size and clutch size of D. hyalina and D. galeata; effects on the same traits of D. magna and D. pulicaria were variable (stimulation, depression, or no effect). D. hyalina and D. galeata responded to gradients of increasing daphniid density (0–300 l^–1) by altering egg mass, somatic mass and clutch size to maintain a relatively constant reproductive investment; egg mass increased with crowding and then decreased in a pattern consistent with Glazier's (1992) hypothetical model of changes in offspring size in relation to food quantity and maternal demand. Effects of crowding by conspecifics were indistinguishable from those of other species. This study, which uncouples the effect of crowding per se from ambient resource depletion, shows that chemical substances released by high densities of Daphnia can cause changes in life-history traits comparable to those that occur in response to low food levels.     

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.     

Trait sorting in Daphnia colonising man‐made lakes

1. We used a zooplankton metacommunity to ask how dispersal, genetic drift and selection act to determine the local and regional distributions of trait variation. Since the formation of the lakes 80 years ago, cladoceran species have sorted into local assemblages that cluster by lake depth. Given this species sorting, we ask whether intraspecific variation in an ecologically important phenotypic trait – body size – has sorted as well. 2. We quantified changes in body size through time by measuring ephippia from D. pulicaria, D. dentifera and D. ambigua recovered from sediment cores from two lakes. We then estimated mean body size of contemporary populations of two competing species, Daphnia pulicaria and D. dentifera, in a laboratory common garden experiment. Finally, we used microsatellite loci to characterise genetic diversity and gene flow among local sites in the metacommunity. 3. Body size was variable both within and among years for the three species of Daphnia examined using sediment cores. For two lakes where we examined body size distributions through time, we observed a significant shift in body size of the first species to arrive after colonisation by other Daphnia species, which suggests selection has occurred historically. 4. Despite heritable variation in body size in the laboratory, evidence for trait sorting was only found for D. pulicaria, which was larger in deeper lakes. Mean body size varied among lakes, but did not sort relative to depth for D. dentifera. 5. Microsatellite data indicated that neutral genetic diversity was low in the region; only 27% of the individuals assayed were unique multi‐locus genotypes. We also found significant patterns of isolation by distance for both species. However, population structure was stronger in D. dentifera than in D. pulicaria. Hence, we conclude that a limited number of colonists have successfully invaded this metacommunity, and those genotypes arriving in this new region have experienced significant dispersal limitation among local sites. 6. Overall, while dispersal and selection have clearly led to the development of predictable community assemblages related to depth in this metacommunity, the distribution of phenotypic traits within species can differ substantially even between two trophically similar species. Our results highlight the complex roles of colonisation history, dispersal, selection and stochasticity in determining inter‐ and intra‐specific patterns in metacommunities.     

Acidity status and phytoplankton species richness,standing crop,and community composition in Adirondack,New York,U.S.A. lakes

The mid-summer phytoplankton communities of more than 100 Adirondack lakes ranging in pH from 4.0 to 7.2 were characterized in relation to 25 physical-chemical parameters. Phytoplankton species richness declined significantly with increasing acidity. Acidic lakes (pH < 5.0) averaged fewer than 20 species while more circumneutral waters (pH > 6.5) averaged more than 33 species. Phytoplankton abundance was not significantly correlated with any of the measured physical-chemical parameters, but standing crop parameters, i.e., chlorophyll a and phytoplankton biovolume, did correlate significantly with several parameters. Midsummer standing crop correlated best with total phosphorus concentration but acidity status affected the standing crop-phosphorus relationship. Circumneutral waters of low phosphorus content, i.e. < 10 µg·1^–1 TP, averaged 3.62 µg·1^–1 chlorophyll a whereas acidic lakes of the same phosphorus content averaged only 1.96 µg·1^–1 chlorophyll a. The midsummer chlorophyll content of lakes of high phosphorus content, i.e. > 10 µg·1^–1 TP, was not significantly affected by acidity status.Adirondack phytoplankton community composition changes with increasing acidity. The numbers of species in midsummer collections within all major taxonomic groups of algae are reduced with increasing acidity. The midsummer phytoplankton communities of acidic Adirondack lakes can generally be characterized into four broad types; 1) the depauperate clear water acid lake assemblage dominated by dinoflagellates, 2) the more diverse oligotrophic acid lake community dominated by cryptomonads, green algae, and chrysophytes, 3) the productive acid lake assemblage dominated by green algae, and 4) the chrysophyte dominated community. The major phytoplankton community types of acid lakes are associated with different levels of nutrients, aluminum concentrations, and humic influences.     

Long‐term dynamics of a cladoceran community from an early stage of lake formation in Lake Fukami‐ike,Japan

An increase in nutrient levels due to eutrophication has considerable effects on lake ecosystems. Cladocerans are intermediate consumers in lake ecosystems; thus, they are influenced by both the bottom‐up and top‐down effects that occur as eutrophication progresses. The long‐term community succession of cladocerans and the effects cladocerans experience through the various eutrophication stages have rarely been investigated from the perspective of the early‐stage cladoceran community assemblage during lake formation. In our research, long‐term cladoceran community succession was examined via paleolimnological analysis in the currently eutrophic Lake Fukami‐ike, Japan. We measured the concentration of total phosphorus and phytoplankton pigments and counted cladoceran and other invertebrate subfossils in all layers of collected sediment cores, and then assessed changes in the factors controlling the cladoceran community over a 354‐year period from lake formation to the present. The cladoceran community consisted only of benthic taxa at the time of lake formation. When rapid eutrophication occurred and phytoplankton increased, the benthic community was replaced by a pelagic community. After further eutrophication, large Daphnia and high‐order consumers became established. The statistical analysis suggested that bottom‐up effects mainly controlled the cladoceran community in the lake''s early stages, and the importance of top‐down effects increased after eutrophication occurred. Total phosphorus and phytoplankton pigments had positive effects on pelagic Bosmina, leading to the replacement of the benthic cladoceran community by the pelagic one. In contrast, the taxa established posteutrophication were affected more by predators than by nutrient levels. A decrease in planktivorous fish possibly allowed large Daphnia to establish, and the subsequent increase in planktivorous fish reduced the body size of the cladoceran community.     

Growth of dominant zooplankton species feeding on plankton microflora in Lake Shira

Batch cultures and continuous flow cultures were used to study the growth rates of zooplankton species from Shira lake, the rotifer Brachionus plicatilis Muller and calanoid copepod Arctodiaptomus salinus Daday, which were fed on phytoplankton and bacterioplankton from the lake. Analyses of the birth and survival rates were used to demonstrate that the lake phytoplankton, consisting mostly of cyanobacteria and diatomaceous algae, is inadequotes for optimal realisation of the reproductive potential of B. plicatilis when compared with the bacterial diet. The study revealed that the kinetic growth characteristics of the two zooplankters were similar: B. plicatilis r _max, 0.120 d^–1; S ₀, 0.253; and K _s, 0.114 mg dry mass l^–1; and for A. salinus r _max, 0.129 d^–1; S ₀, 0.240; and K _s, 0.171 mg dry mass l^–1. Fluctuations in natural food concentration reduced the growth rate of both species. Even though the threshold concentration of food for B. plicatilis and A. salinus were quite similar, the copepods were less sensitive to food limitation.     

Effectiveness of phytoplankton control by large-bodied and small-bodied zooplankton

Employingin situ enclosures containing inocula of the lake zooplankton (mainlyDaphnia galeata, Daphnia cucullata andBosmina spp.) from a moderately eutrophic Lake Ros (Northern Poland) or large-bodiedDaphina magna, the following observations on succession of phytoplankton were made: 1) whereasD. magna could control the density of all the photoplankton size classes, the lake zooplankton could not suppress the large-sized phytoplankters or net phytoplankton; 2) the lake zooplankton was able to control the density of small algae (< 50μm), but its effect on large algae may be opposite: a promotion of net phytoplankton growth by removing small-sized algae which can out-compete net phytoplankton for limited PO₄-P resources (<5μg P l⁻¹). Since efficiency of phytoplankton density control byD. magna decreased with an increase in net phytoplankton abundance, biomanipulation could not be successful without introducing or maintaining a high population of large-bodied cladoceran species before high densities of large algae would make the control of phytoplankton inefficient.     

Plankton dynamics in a high mountain lake (Las Yeguas,Sierra Nevada,Spain). Indirect evidence of ciliates as food source for zooplankton

A detailed sampling programme during the ice-free season (July–September) in the oligotrophic lake Las Yeguas (Southern Spain) has shown a well-defined time lag between phytoplankton and zooplankton maximum standing stocks, the former displaying a peak (23 μgC l^-1) just after the ice-melting, and the latter by the end of September (80 μgC l^-1). A ratio of autotrophs to heterotrophs lower than 1 which lasted more than two thirds of the study period may suggest a high algal productivity per unit of biomass. The estimated strong top-down regulation of phytoplankton by zooplankton indicates an efficient utilization of resources. A comparative analysis between the available food supply and the critical food concentration that is necessary to maintain the population of Daphnia pulicaria (which constitutes up to 98% of the heterotrophic biomass) proves this species to be food-limited in the lake under study. To explain the dominance (and development) of such large-bodied cladoceran population, we discuss the possibility of the utilization of naked protozoan ciliates (Oligotrichidae) as a complementary high quality food source, and the exploitation of benthic resources through a coupled daily migration behaviour.     

Competitive ability of Daphnia under dominance of non-toxic filamentous cyanobacteria

It is generally assumed that Daphnia is more susceptible to the inhibitory effects of filamentous cyanobacteria than small cladocerans since daphnids have a larger gape size and filtrate the filaments, whereas small cladocerans do not. This study addresses the question whether food limitation has the potential to modify this scenario of cladoceran response to dominance of non-toxic filamentous cyanobacteria. Daphnia galeatawas grown under limited (0.1 mg C l^–1) and unlimited concentrations (1.0 mg C l^–1) of high-quality food algae both in the absence/presence of non-toxic filamentous Aphanizomenon flexuosum. As the effects of these cyanobacteria on D. galeatawere positive under food limiting conditions and negative at the high food density, it was concluded that D. galeatawas mainly affected by nutritional quality due to its ability to ingest the filaments, while mechanical interference with food collection was not important. In competition experiments between D. galeataand Bosmina longirostris, D. galeatawas the dominant species at regular additions of food (1.0 mg C l^–1) in the absence of Aphanizomenon. In the presence of these cyanobacteria, D. galeatawas inhibited during the first days of the experimental period. However, the negative effect at the initially high food density was outweighed by nutrition at food limiting conditions and the outcome in competitive dominance was not changed. The results demonstrate that the ability of D. galeata to ingest large-sized non-toxic cyanobacteria can be considered as advantageous under food limiting conditions.     

The role of hybridization in the origin and spread of asexuality in Daphnia

The molecular mechanisms leading to asexuality remain little understood despite their substantial bearing on why sexual reproduction is dominant in nature. Here, we examine the role of hybridization in the origin and spread of obligate asexuality in Daphnia pulex, arguably the best‐documented case of contagious asexuality. Obligately parthenogenetic (OP) clones of D. pulex have traditionally been separated into ‘hybrid’ (Ldh SF) and ‘nonhybrid’ (Ldh SS) forms because the lactase dehydrogenase (Ldh) locus distinguishes the cyclically parthenogenetic (CP) lake dwelling Daphnia pulicaria (Ldh FF) from its ephemeral pond dwelling sister species D. pulex (Ldh SS). The results of our population genetic analyses based on microsatellite loci suggest that both Ldh SS and SF OP individuals can originate from the crossing of CP female F₁ (D. pulex × D. pulicaria) and backcross with males from OP lineages carrying genes that suppress meiosis specifically in female offspring. In previous studies, a suite of diagnostic markers was found to be associated with OP in Ldh SS D. pulex lineages. Our association mapping supports a similar genetic mechanism for the spread of obligate parthenogenesis in Ldh SF OP individuals. Interestingly, our study shows that CP D. pulicaria carry many of the diagnostic microsatellite alleles associated with obligate parthenogenesis. We argue that the assemblage of mutations that suppress meiosis and underlie obligate parthenogenesis in D. pulex originated due to a unique historical hybridization and introgression event between D. pulex and D. pulicaria.     

Trophic interactions and habitat segregation between competing Daphnia species

Summary Two commonly coexisting species of Daphnia segregate by habitat in many stratified lakes. Daphnia pulicaria is mostly found in the hypolimnion whereas D. galeata mendotae undergoes diel vertical migration between the hypolimnion and the epilimnion. I examined how habitat segregation between these two potentially competing species might be affected by trophic interactions with their resources and predators by performing a field experiment in deep enclosures in which I manipulated fish predation, nutrient levels, and the density of epilimnetic Daphnia. The results of the experiment indicate that habitat use by D. pulicaria can be jointly regulated by competition for food from epilimnetic Daphnia and predation by fishes. Patterns of habitat segregation between the two Daphnia species were determined by predation by fish but not by nutrient levels: The removal of epilimnetic fish predators resulted in higher zooplankton and lower epilimnetic phytoplankton densities and allowed D. pulicaria to expand its habitat distribution into the epilimnion. In contrast, increased resource productivity resulted in higher densities of both Daphnia species but did not affect phytoplankton levels or habitat use by Daphnia. The two species exhibit a trade-off in their ability to exploit resources and their susceptibility to predation by fish. D. g. mendotae (the less susceptible species) may thus restrict D. pulicaria (the better resource exploiter) from the epilimnion when fish are common due to lower minimum resource requirements than those needed by D. pulicaria to offset the higher mortality rate imposed by selective epilimnetic fish predators. D. g. mendotae does not appear to have this effect in the absence of fish.     

Daphnia lumholtzi and Daphnia ambigua: population comparisons of an exotica and a native cladoceran in Lake Okeechobee, Florida

The population dynamics of an exotic cladoceran (Daphnia lumholtzi Sars) and a native cladoceran (Daphnia ambigua) were studied over a 12 month period in subtropical Lake Okeechobee, Florida (USA), to quantify the extent of invasion of the exotic species and compare ecological niches. Daphnia lumholtzi accounted for up to 70% of the Daphnia assemblage during the summer months (June-August), while D.ambigua accounted for up to 97% of the Daphnia assemblage from fall to spring (October-April). The densities of the two species were inversely corelated. The exotic species was most concentrated in the shallower, warmer, north and south ends of the lake during the summer. It also was present, but at much lower densities, in the central lake region during the fall. The native species displayed a ubiquitous distribution throughout the lake during spring and winter, but was concentrated in the deeper, cooler, central region during the summer. Relationships of the two species with environmental conditions indicate that water column temperature might affect the seasonal and spatial distribution of the two Daphnia species. The results also indicate that D.lumholtzi may be filling a 'vacant' seasonal or spatial niche when conditions are unfavorable for D.ambigua.      

Population growth of some genera of cladocerans (Cladocera) in relation to algal food (Chlorella vulgaris) levels

We studied the patterns of population growth of 7 cladoceran species (Alona rectangula, Ceriodaphnia dubia, Daphnia laevis, Diaphanosoma brachyurum, Moina macrocopa, Scapholeberis kingi and Simocephalus vetulus) using 6 algal densities, viz. 0.05×10⁶, 0.1×10⁶, 0.2×10⁶, 0.4×10⁶, 0.8×10⁶ and 1.6×10⁶ cells ml^–1, of Chlorella vulgaris for 18 – 30 days. In terms of carbon content these algal concentrations corresponded to 0.29, 0.58, 1.16, 2.33, 4.65 and 9.31 g ml^–1, respectively. Cladocerans in the tested range of algal levels responded similarly, in that increasing the food concentrations resulted in higher numerical abundance and population growth rates (r). The peak population densities were (mean±standard error) 71±5; 17.1±0.4, 3.6±0.3, 12.7±1.1, 18.2±2.7, 15.8±1.0 and 10.9±0.02 ind. ml^–1, respectively for A. rectangula, C. dubia, D. laevis, D. brachyurum, M. macrocopa, S. kingi and S. vetulus. In general, the lowest r values were obtained for D. laevis (0.01±0.001) at 0.05×10⁶ cells ml^–1 food level while the highest was 0.283±0.004 for A. rectangula at 1.6×10⁶ cells ml^–1 of Chlorella. When the data of peak population density for each cladoceran species were plotted against the body length, we found an inverse relation, broadly curvilinear in shape. From regression equations between the food level and rate of population increase, we calculated the theoretical food quantity (the threshold level) required to maintain a zero population growth (r = 0) for each cladoceran species, which varied from 0.107 to 0.289 g ml^–1 d^–1 depending on the body size. When we plotted the cladoceran body size against the corresponding threshold food levels, we obtained a normal distribution curve. From this it became evident that for up to 1300 m body size, the threshold food level increased with increasing body size; however, beyond this, the threshold level decreased supporting earlier observations on rotifers and large cladocerans.     

Larval fish-induced phenotypic plasticity of coexisting Daphnia: an enclosure experiment

1. The indirect effects of predators on lower trophic levels have been studied without much attention to phenotypically plastic traits of key food web components. Phenotypic plasticity among species creates phenotypic diversity over a changing environmental landscape. 2. We measured the indirect effects of planktivorous larval walleye (Stizostedion vitreum) on phytoplankton biomass through their effects on the dominant herbivore species, Daphnia pulicaria and D. mendotae. 3. Fish had no effect on phytoplankton biomass or overall Daphnia density. We observed a compensatory response to predation by functionally comparable species within a trophic level in the form of shifting dominance and coexistence of Daphnia species. We hypothesized that this phenotypically plastic response to predation decoupled a potential trophic cascade in this freshwater pelagic system. Daphnia pulicaria density decreased over time with fish predation, but D. mendotae density increased over time with fish predation. 4. Phenotypically plastic life history trait shifts and reproductive rates differed between species in fishless and fish enclosures, accounting for population trends. Daphnia pulicaria were also proportionally higher in walleye larvae stomachs than in the enclosures, indicating that walleye preferred to feed on D. pulcaria over D. mendotae. The resultant shift in dominance may partially explain the overall benign effect of fish on grazers and supports the hypothesis that trophic level diversity can decouple a trophic cascade.     

Phytoplankton productivity and chlorophyll-A concentration of Oguta Lake in Southeastern Nigeria

The phytoplankton productivity and chlorphyll-a concentration of Oguta Lake, the largest natural lake in south-eastern Nigeria, are presented (Dec. 1983. Nov. 1984). The gross productivity ranged from 1.3 to 3.77 g C.m^–2.day^–1 for the water column, dropped during the period of heavy rainfall and varied with depth. The chlorophyll-a concentration had monthly means ranging from 2.31 to 4.00 mg.m^–3, with a drop during the rains, but little depth variation. Both productivity and chlorophyll-a showed non-significant correlation with the physico-chemical features of the water. The values of the biological parameters showed the lake as mesotrophic. The values are compared with those of other African lakes.     

Ecology of Dictyosphaerium pulchellum Wood (Chlorophyta, Chlorococcales) in a shallow, acid, forest lake

This study relates to the ecology of Dictyosphaerium pulchellum Wood in Delamere Lake in Cheshire, UK. Dictyosphaerium pulchellum is a cosmopolitan, green colonial phytoplankton species that occasionally forms dense, monospecific populations in lakes. Delamere Lake is a small, shallow, acid lake (mean pH, 4.5) with very high phytoplankton biomass (annual mean chlorophyll a, 290 μg l⁻¹) and devoid of any significant cladoceran population, the efficient grazers of phytoplankton. A predominantly unicellular form of D. pulchellum was the dominant species in Lake Delamere, and it comprised on average ca. 80% (maximum >99%) of the lake phytoplankton biovolume. Laboratory and lake experiments were conducted on this species showed that its pH tolerance varied between 2.4 and 10.7, and its optimum tolerance range between 3.3 and 8.5 depending on other environmental variables. Low pH was not responsible for the unicellular habit of this alga, but a very high nutrient regime could be an important factor. Bioassays revealed that in Delamere Lake this species was limited by nitrogen, but nitrogen did not hamper high growth in the lake. Dictyosphaerium pulchellum can persist at low light levels, tolerate CO₂-deficiency and can grow in polyhumic water with water colour around 300 mg Pt l⁻¹, but probably not in darker waters. The dominance of D. pulchellum in Delamere Lake is apparently due to a combination of several factors: its ability to tolerate both low pH and high turbidity, exploit high nutrient conditions, absence of effective grazing pressure by zooplankton and being a superior competitor.     

Species-specific population-density thresholds in cladocerans?

The population density of a Daphnia species seems more dependent on properties specific to the species than on those specific to the point in the season, location within a lake basin or the given lake itself. In spite of week-to-week fluctuations, the population density for each of two common European Daphniaspecies was found to be remarkably similar within single lakes (from station to station on a single date, and from date to date at a single station) as well as from lake to lake, regardless of trophic state. All lakes on all dates revealed densities in the range 10–50 ind l^–1 for the smaller-bodied D. cucullata and 1–5 ind l^–1 for the larger-bodied D. hyalina, in spite of different intensity of reproduction resulting from different food levels (chlorophyll a between 0.2 and 4.2 g/l). It can be asserted that the population density of each species remains far below the carrying capacity of the habitat K, and does not depend on food levels, which merely set the rate of population increase, while the population density reflects the species' vulnerability to predation by planktivorous fish. The reactive distance (the distance over which a foraging fish can see its prey) in 1+ roach, a dominant planktivore in the lakes studied, has been found to be twice as great for D. hyalina as for D. cucullata, whatever the light intensity. The relative reactive field volume was therefore an order of magnitude greater for the former than for the latter, showing that the densities of the two prey species would be assessed by a foraging roach as equal when, in reality, they differed by an order of magnitude, as they do in various lakes and in various seasons.The first of the two conclusions is that whatever the growth and reproduction in a population of a cladoceran such as Daphnia, its density would be fixed by mortality induced by fish predation. The second would be, that the difference between the bottom-up and top-down effects in freshwater is more than merely the upward or downward direction along the food web, since the bottom-up effects are about the flow control (the rate of net production, individual growth rate, rate of reproduction) and the top-down about the standing-crop control (biomass, individual body size, population density level).