Selective feeding and its effect on polymorphism and sexuality in the rotifer Asplanchna sieboldi

SUMMARY. Campanulate females of A. sieboldi (clone 12C1) attack Asplanchna brightwelli and A. girodi much more readily than either the alga Volvox aureus or the rotifer Brachionus calyciflorus . Pre-feeding campanulates on B. calyciflorus prior to testing does not appreciably affect their response to this prey. Discrimination during feeding occurs immediately after the campanulate's corona contacts a potential food organism and is probably mediated by chemoreceptors. Once a food organism is attacked, the probabilities of it being captured and subsequently swallowed are high and similar for each of the four organisms tested.
Since food organisms other than Asplanchna . such as V. aureus and B. calyciflorus , induce campanulates in this clone to produce cruciform, and thus frequently sexual, females in succeeding generations, a tendency of these campanulates to select congeneric prey would favour the maintenance of the campanulate morphotype and thus the continuation of parthenogenetic reproduction in the ensuing population.     

Effect of starvation time on the prey capture behaviour, functional response and population growth of Asplanchna sieboldi (Rotifera)

1. We examined the effect of different periods of prior starvation(from 30 min to 16 h) on the prey capture behaviour, and functional and numerical responses of the predatory rotifer Asplanchna sieboldi using the rotifer Brachionus calyciflorus as prey.
2. Feeding activity (i.e. encounter, attack, capture and ingestion) by Asplanchna increased significantly with increasing prey densities from 2 to 16 mL⁻⁻¹ and with increasing prior starvation periods from 0.5 to 8 h.
3.  Asplanchna sieboldi showed a type II functional response at all the prior starvation periods tested. The asymptotic prey density was highest after 8 h of starvation.
4. The instantaneous population growth rate of A. sieboldi ranged from 0.089 ± 0.044 (when starved for 8 h in every 24 h and at a prey density of 2 individuals mL⁻⁻¹ for the other 16 h) to 1.015 ± 0.142 in the control (no starvation and at a prey density of 16 individuals mL⁻⁻¹). The effect of starvation time on the numerical response was evident only at the higher prey density.     

Demography and population growth of Asplanchna girodi (Rotifera) as a function of prey (Anuraeopsis fissa) density

Laboratory studies on population growth and life table demography of Asplanchna girodi were conducted at 25±1 °c using Anuraeopsis fissa as prey at four (250, 500, 1000 and 2000 ind ml^–1) densities. A prey density of 100 ind ml^–1 per predator per day did not support A. girodi, while at the highest prey concentration, A. girodi reached a peak of 115±7 ind ml^–1. The age distribution of A. girodi indicated that non-adults constituted about 2/3rd of the population at all prey concentrations. A decrease in prey availability resulted in increased mortality of non-adults. At the highest prey density, the rate of population increase (r) was 1.51 d^–1. The significance of estimating mortality in population growth studies is discussed.Life table demography of A. girodi was also studied using the same prey at the same concentrations. None of the survivorship parameters (e.g. mean lifespan and mean survivorship) showed a significant relation to prey density. Net reproductive rate and generation time (but not rate of population increase) were affected by prey abundance.     

Effect of three food types on the population growth of Brachionus calyciflorus and Brachionus patulus (Rotifera: Brachionidae).

We compared the population growth of B. calyciflorus and B. patulus using the green alga Chlorella vulgaris, baker's yeast Saccharomyces cerevisiae or their mixture in equal proportions as food. Food was offered once every 24 h in two concentrations (low: 1 x 10(6) and high: 3 x 10(6) cells ml-1) separately for each species. The experiments were terminated after 15 days. In general, at any food type or concentration, B. patulus reached a higher population density. A diet of Chlorella alone supported a higher population growth of both rotifer species than yeast alone. B. calyciflorus and B. patulus achieved highest population densities (103 +/- 8 ind. ml-1 and 296 +/- 20 ind. ml-1, respectively) on a diet of Chlorella at 3 x 10(6) cells ml-1. When cultured using the mixture of Chlorella and yeast, the maximal population densities of B. calyciflorus were lower than those grown on Chlorella. Under similar conditions, the maximal abundance values of B. patulus were comparable in both food types. Regardless of food type and density the rate of population increase per day (r) for B. calyciflorus varied from 0.13 +/- 0.03 to 0.63 +/- 0.04. These values for B. patulus ranged from 0.19 +/- 0.01 to 0.37 +/- 0.01. The results indicated that even though Chlorella was a superior food for the tested rotifers, yeast can be effectively used at low concentrations to supplement algal requirements in rotifer culture systems.     

Responses of the predatory rotifer Asplanchna intermedia to prey species differing in vulnerability: laboratory and field studies

1. We have studied the numerical and functional responses of campanulate morphs of Asplanchna intermedia fed five species of rotifer ( Brachionus rubens , B. patulus , B. calyciflorus , Hexarthra mira and Filinia longiseta ). The vulnerability of the prey varied with their morphology and mode of swimming.
2. To test the numerical and functional responses, prey species differing in their morphology and mode of swimming were provided. Responses were also tested with mixtures of evasive and non-evasive prey provided in three different ratios.
3. A. intermedia showed a type II functional response to all the prey species provided.
4. The population growth rate of A. intermedia on the various prey species provided ranged from a minimum of –0.24 to a maximum of 0.68. There was a significant correlation between the capturability of a prey species and the population growth rate of the predator feeding on it. The capturability of a prey species also has a significant influence on the maximal predator density but not on the time taken to reach it.
5. Observations from a field study undertaken over a 10-month period to study the prey preferences of A. intermedia in nature were corroborated by the laboratory findings.     

Responses of the predatory rotifer Asplanchna intermedia to prey species differing in vulnerability: laboratory and field studies

1. We have studied the numerical and functional responses of campanulate morphs of Asplanchna intermedia fed five species of rotifer ( Brachionus rubens , B. patulus , B. calyciflorus , Hexarthra mira and Filinia longiseta ). The vulnerability of the prey varied with their morphology and mode of swimming.
2. To test the numerical and functional responses, prey species differing in their morphology and mode of swimming were provided. Responses were also tested with mixtures of evasive and non-evasive prey provided in three different ratios.
3. A. intermedia showed a type II functional response to all the prey species provided.
4. The population growth rate of A. intermedia on the various prey species provided ranged from a minimum of –0.24 to a maximum of 0.68. There was a significant correlation between the capturability of a prey species and the population growth rate of the predator feeding on it. The capturability of a prey species also has a significant influence on the maximal predator density but not on the time taken to reach it.
5. Observations from a field study undertaken over a 10-month period to study the prey preferences of A. intermedia in nature were corroborated by the laboratory findings.     

Comparative population growth and life table demography of the rotifer Asplanchna girodi at different prey (Brachionus calyciflorus and Brachionus havanaensis) (Rotifera) densities

Population growth and life table demography of the predatory rotifer A. girodi using spineless Brachionus calyciflorus and spined Brachionus havanaensis as prey at densities of 1, 2, 4 and 8 ind. ml^–1 at 25°C were studied. Regardless of the prey species, the population of A. girodi increased with increasing availability of Brachionus in the medium. At any given prey density, A. girodi fed B. calyciflorus showed consistently better growth than when fed B. havanaensis. The maximum population densities of A. girodi varied from 0.28 to 1.8 ind. ml^–1 depending on the prey species and the density. The rate of population increase observed in population growth studies varied from 0.17 to 0.43 day^–1 when fed B. calyciflorus and 0.09 to 0.27 day^–1 when fed B. havanaensis. Male population of A. girodi was closely related to female density. The lowest average lifespan was observed for A. girodi when fed B. havanaensis at 1 ind. ml^–1, while the converse was the case when fed B. calyciflorus at comparable prey concentration. Net reproductive rates varied from 16 to 26 offspring female^–1 lifespan^–1 depending on the prey species and concentration. Generation time of A. girodi decreased with increasing food concentrations for both the prey species. The rates of population increase obtained from life table demography were lower for A. girodi when fed B. havanaensis than when fed B. calyciflorus.     

Competition between Brachionus calyciflorus Pallas and Brachionus patulus (Müller) (Rotifera) in relation to algal food concentration and initial population density

Competitive laboratory experiments between Brachionus calyciflorus and B. patulus were conducted at low (1×10⁶ cells ml^–1) and high (3×10⁶ cells ml^–1) densities of Chlorella vulgaris and four initial inoculation densities (numerically, 100% B. calyciflorus; 75% B. calyciflorus + 25% B. patulus; 50% each of the two species; 25% B. calyciflorus + 75% B. patulus and 100% B. patulus). Population densities were enumerated and the medium was changed daily for 20 days. B. patulus was a superior competitor in low food density regardless of inoculation density. At high food density, B. calyciflorus showed higher population growth in the first week but thereafter was outcompeted by B. patulus regardless of initial density. When grown alone, B. calyciflorus reached peak abundances (mean ± standard error) of 31±3 and 81±7 individuals ml^–1 at low and high food densities, respectively. The corresponding values for B. patulus were 130±2 and 306±13. The adverse effects of B. patulus on the peak abundances of B. calyciflorus were higher at low food concentration. Data on egg ratios (eggs female^–1) revealed an inverse relation with population abundance of both tested rotifer species. Our results indicated that the rate of population increase of a species was not a good indicator of its competitive ability. Instead, the ability to reproduce under continuously diminishing food resources (until a threshold level) was responsible for the competitive edge of B. patulus over B. calyciflorus. This was further influenced by the relative inoculation densities of the tested rotifer species and the offered food densities.     

Epizoic mode of life in Brachionus rubens Ehrenberg as a deterrent against predation by Asplanchna intermedia Hudson

The planktonic rotifer Brachionus rubens has a propensity for an epizoic mode of life, and in nature is often found attached to cladocerans. In this way the rotifer avoids to a certain extent the adverse effects of interference competition with cladocerans. We test the hypothesis that the epizoic habit of B. rubens acts also as a deterrent against invertebrate predation. Using Asplanchna intermedia as predator, we followed the population growth patterns of B. rubens alone and in the presence of the host species Daphnia carinata and Ceriodaphnia rigaudi. In the absence of cladocerans, the prey was eliminated within three days, followed by extinction of the predator due to starvation. With D. carinata in the medium, the prey-predator system persisted much longer, with B. rubens reaching high population densities. With the smaller-sized C. rigaudi, allowing a significantly smaller fraction of B. rubens population to be epizoic, the system persisted longer than in the controls, but both the prey and predator eventually became extinct. We conclude that the epizoic habit of B. rubens, by acting as a prey refugium, helps a portion of the population to escape from predation, and facilitates its coexistence with Asplanchna intermedia.     

Behavioural and population responses to changing availability of <Emphasis Type="Italic">Artemia</Emphasis> prey by moulting black-necked grebes, <Emphasis Type="Italic">Podiceps nigricollis</Emphasis>

Chemical communication may inform about the location of prey, predators, co-specifics, and mate partners in zooplankton. In this study, we evaluated several life-history traits of the rotifer, Brachionus calyciflorus, exposed to conditioned media by a rotifer predator (Asplanchna brightwelli) and a cladocera competitor (Daphnia similis), quantifying population growth and life-table demography at two algal food levels (2.0 and 0.5 × 10⁶ cells ml⁻¹ of Chlorella pyrenoidosa). At both food levels, B. calyciflorus grown in predator-conditioned media had lower population abundance and slower population growth rate than controls. Conversely, the competitor-conditioned media treatments produced both higher rotifer population abundance and faster population growth rate than controls. Life-history parameters varied significantly depending on the presence of predator and competitor-conditioned media. The Asplanchna-conditioned media significantly decreased gross reproductive rate (GRR): 8–9 offsprings per female; net reproductive rate (R ₀): 6–7 offsprings per female; population growth rate (r): 0.34–0.37 day⁻¹; and increased generation time (T): 5.5–5.6 days. On the other hand, The Daphnia-conditioned media significantly increased the GRR (13–14 offsprings per female); net reproductive rate (8–9 offsprings per female); population growth rate (0.42–0.43 day⁻¹); and decreased generation time (4.9–5.0 days). However, the effects of food level on the life-history characteristic were not significant in both treatments. Maximum values of the population abundance and the population growth rate are significantly influenced by the predator densities and pre-culture time. This study suggests that rotifers use variable life-history strategies (low reproduction and high survivorship versus high reproduction and low survivorship) based on the presence of predators and competitors.     

Morphometric and demographic responses of brachionid prey (Brachionus calyciflorus Pallas and Plationus macracanthus (Daday)) in the presence of different densities of the predator Asplanchna brightwellii (Rotifera: Asplanchnidae)

We quantified the indirect effects of different densities (1, 4, 16 individuals per jar) of the predator Asplanchna brightwellii on the morphometry and demography of their prey Brachionus calyciflorus and Plationus macracanthus. Population growth and life table demography experiments were separately conducted for each of the two prey rotifer species while keeping A. brightwellii in indirect contact. The brachionids were fed the green alga Chlorella vulgaris at a density of 1 × 10⁶ cells ml⁻¹. As compared to those cultured in the absence of the predator, in the presence of A. brightwellii the postero-lateral spines of P. macracanthus increased by about 15 μm, while in B. calyciflorus the increase was more (>80 μm). For both the brachionid species, the peak population density significantly decreased in the presence of A. brightwellii. The reproductive variables viz., net reproductive rate, generation time, and the rate of population increase of B. calyciflorus and P. macracanthus were negatively affected in the presence of kairomones from A. brightwellii.     

Life table demography of <Emphasis Type="Italic">Asplanchna brightwellii</Emphasis> Gosse, 1850 fed with five different prey items

We conducted life table experiments on the freshwater rotifer Asplanchna brightwellii to analyze its demography when fed with prey items from several taxonomic groups (cladocerans, protozoans, and rotifers) and under two different temperature regimes (20 and 25°C); the aim of the study was to determine the preferred prey for A. brightwellii in terms of fitness (evaluated as reproductive success) among five cladoceran, protozoan, and rotifer preys, and to test which temperature (20 or 25°C) is better for life table parameters of Asplanchna. Our analysis identified Brachionus calyciflorus as the preferred prey for A. brightwellii based on life table statistics, ingestion rate and electivity indices. The greatest values for net reproductive rate and intrinsic growth rate were achieved when A. brightwellii was fed B. calyciflorus. Greater reproductive values (R _o and r) were found at 25°C than at 20°C for A. brightwellii across the five prey species. We found significant differences in the ingestion rate and electivity index among zooplanktonic and benthic preys. The influence of temperature, the cost of predation, and how prey selection by A. brightwellii is influenced by: biomass, size, and swimming speed; they are discussed hoping to gain a better understanding of trophic transfers in zooplankton communities.     

Density-dependent regulation of natural and laboratory rotifer populations

Density-dependent regulation of abundance is fundamentally important in the dynamics of most animal populations. Density effects, however, have rarely been quantified in natural populations, so population models typically have a large uncertainty in their predictions. We used models generated from time series analysis to explore the form and strength of density-dependence in several natural rotifer populations. Population growth rate (r) decreased linearly or non-linearly with increased population density, depending on the rotifer species. Density effects in natural populations reduced r to 0 at densities of 1–10 l^–1 for 8 of the 9 rotifer species investigated. The sensitivities of these species to density effects appeared normally distributed, with a mean r=0 density of 2.3 l^–1 and a standard deviation of 1.9. Brachionus rotundiformis was the outlier with 10–100× higher density tolerance. Density effects in laboratory rotifer populations reduced r to 0 at population densities of 10–100 ml^–1, which is 10⁴ higher than densities in natural populations. Density effects in laboratory populations are due to food limitation, autotoxicity or to their combined effects. Experiments with B. rotundiformis demonstrated the absence of autotoxicity at densities as high as 865 ml^–1, a much higher density than observed in natural populations. It is, therefore, likely that food limitation rather than autotoxicity plays a major role in regulating natural rotifer populations.     

Utilization of cyanobacteria by Brachionus calyciflorus: Anabaena flos-aquae (NRC-44-1) as a sole or complementary food source

The rotifer Brachionus calyciflorus can utilize the cyanobacterium Anabaena flos-aquae as either a sole or supplementary food source in laboratory culture. Positive population growth rates accompany food densities of 10 or 100 µg dry weight ml^–1, but slightly negative rates are found at a lower density (1.0 µg ml^–1). These results are consistent for rotifers feeding on two strains of A. flos-aquae, UTEX-1444 and NRC-44-1, with slightly enhanced survivorship and reproduction with the latter food. A 1:1 mixture (by dry weight) of Euglena gracilis and A. flos-aquae (NRC-44-1) produces survivorship comparable to that of control rotifer cohorts fed E. gracilis alone, but elicits significantly greater fecundity and population growth rates than found with the control food suspension at the same biomass density.     

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.     

Prey selection by Asplanchna girodi (Rotifera): the importance of prey defence mechanisms

1. Predator (laboratory-grown Asplanchna girodi and prey (several field-collected non-carnivorous rotifer species) interactions were studied in the laboratory as a function of both predator and prey densities. The clearance rates of A, girodi decreased with increasing prey density. Predator density had no effect on the feeding of A. girodi. 2. Asplanchna girodi selectively fed on Keratella cochlearis. Its clearance rates for K. cochlearis were much higher than those for Polyarthra and Brachionus. Short-spined forms (spine length less than 25 μm) showed a significantly higher susceptibility to predation than either the non-spined or the long-spined forms. 3. Large Asplanchna individuals fed selectively on reproductive females of K. cochlearis, thereby reducing the fecundity of their prey.     

Reproductive success of the rotifer Brachionus calyciflorus feeding on ciliates and flagellates of different trophic modes

SUMMARY 1. The nutritional value of the bacterivorous ciliate Tetrahymena pyriformis and the algivorous ciliate Coleps sp., as well as the heterotrophic flagellate Chilomonas paramecium and the autotrophic flagellate Cryptomonas ovata , were investigated in population growth experiments using the rotifer B. calyciflorus . The two ciliates, both flagellates, which were of similar size, shape and mobility, were each offered as a sole diet and as a supplement to the alga Monoraphidium minutum , known to support reproduction of B. calyciflorus .
2. To further test nutritional differences between the prey organisms, prey selection experiments were conducted in which B. calyciflorus was able to select between the bacterivorous and algivorous ciliate, and between the heterotrophic and autotrophic flagellate.
3. The results demonstrated that both ciliates and the heterotrophic flagellate were not sufficient to support reproduction of B. calyciflorus when offered as a sole diet. They were, however, a good supplement to algal prey (except for the bacterivorous ciliate T. pyriformis ). In the prey selection experiments, B. calyciflorus positively selected for the algivorous Coleps sp. and the autotrophic C. ovata.
4. Overall, ciliates and heterotrophic flagellates may enhance survival of B. calyciflorus , but reproduction of the rotifer is likely to rely on algal prey. Both higher population growth of B. calyciflorus when fed the algivorous Coleps and the autotrophic Cryptomonas, along with their positive selection, give evidence for prey specific differences in nutrition, with algivorous or autotrophic prey species tending to be of higher nutritional value.     

Linking herbivore-induced defences to population dynamics

1. Theoretical studies have shown that inducible defences have the potential to affect population stability and persistence in bi‐ and tritrophic food chains. Experimental studies on such effects of prey defence strategies on the dynamics of predator–prey systems are still rare. We performed replicated population dynamics experiments using the herbivorous rotifer Brachionus calyciflorus and four strains of closely related algae that show different defence responses to this herbivore. 2. We observed herbivore populations to fluctuate at a higher frequency when feeding on small undefended algae. During these fluctuations minimum rotifer densities remained sufficiently high to ensure population persistence in all the replicates. The initial growth of rotifer populations in this treatment coincided with a sharp drop in algal density. Such a suppression of algae by herbivores was not observed in the other treatments, where algae were larger due to induced or permanent defences. In these treatments we observed rotifer population densities to first rise and then decline. The herbivore went extinct in all replicates with large permanently defended algae. The frequency of herbivore extinctions was intermediate when algae had inducible defences. 3. A variety of alternative mechanisms could explain differential herbivore persistence in the different defence treatments. Our analysis showed the density and fraction of highly edible algal particles to better explain herbivore persistence and extinctions than total algal density, the fraction of highly inedible food particles or the accumulation of herbivore waste products or autotoxins. 4. We argue that the rotifers require a minimum fraction and density of edible food particles for maintenance and reproduction. We conjecture that induced defences in algae may thus favour larger zooplankton species such as Daphnia spp. that are less sensitive to shifts in their food size spectrum, relative to smaller zooplankton species, such as rotifers and in this way contributes to the structuring of planktonic communities.     

Influence of ethanol and ethylene on the seed germination of three nymphaeid water plants

1. We offered Scenedesmus obliquus in five densities, from 0.5 to 8 ± 10⁶ cells ml^?1, to the rotifer Anuraeopsis fissa. Growth rates (r) during the exponential phase (first 7 days) were significantly and positively related to food density. The r values (mean ± SD) varied between 0.454 ± 0.067 and 0,856 ± 0.090, from the lowest to the highest food concentration, respectively. Population growth went through a phase of exponential increase which lasted from 7 to 10 days before a plateau and, in some cases, a decrease occurred. 2. There was a linear relation between food density and rotifer plateau density. At the highest food density, a peak abundance (mean ± SD) of 2312 ± 226 individuals ml^?1 was reached, while at the lowest food density there was no identifiable single peak but a plateau, at a density of 361 ± 62 ind. ml^?1. 3. The egg ratio decreased with increasing population density. The ratio of loose eggs to eggs attached to females indicated that more eggs became detached at higher food densities. 4. As in many other rotifer species studied so far, population density of A. fissa was less stable at higher algal food concentrations. Numerically, A. fissa could be grown at twice the density achieved in Brachionus.     

Ecological problems of Lake Ladoga: causes and solutions

We studied the outcome of competition between a large (Brachionus calyciflorus) and a small (Anuraeopsis fissa) rotifer species at five algal (Scenedesmus acutus) concentrations (0.5 × 10⁶ to 40.5 × 10⁶ cells ml^–1) and with varying initial densities in mixed populations (100 to 0% of B. calcyciflorus or A. fissa), the combined initial biomass being 0.2 µg ml^–1 in all test jars. Experiments were conducted at 28 ± 1 °C.Regardless of food concentration, B. calcyciflorus showed a greater increase in biomass than A. fissa, peak densities (mean ± standard error) at the lowest food concentration in the controls being 1.34 ± 0.31 µg dry weight ml^–1 and 0.82 ± 0.08 dry weight ml^–1, respectively. At the lower food concentrations, A. fissa displaced B. calyciflorus and vice versa at the higher food concentrations. At the intermediate food concentrations of 4.5 × 10⁶ cells ml^–1, B. calyciflorus outcompeted A. fissa only if its initial population density was three times higher. The rates of population growth in controls varied from 0.792 ± 0.06 d^–1 to 1.492 ± 0.13 d^–1 for B. calyciflorus and 0.445 ± 0.04 to 0.885 ± 0.01 for A. fissa depending on food level. When both species were introduced together, low food levels favoured higher abundance of A. fissa than B. calyciflorus, suggesting, in nature, it is likely that small Anuraeopsis colonize oligotrophic water bodies more successfully than larger Brachionus. The results also suggest that the outcome of competition depends not only on the size of the competing species and food availability but also on their colonizing density.