Resource limitation and food quality for cladocerans in a tropical Brazilian lake

Food quality of the natural food (seston) from Lake Monte Alegre was evaluated throughout a series of life-table experiments with cladocerans from the same lake carried out in summer and autumn. Experiments were performed using cohorts of newborns originated from ovigerous females cultured for several generations in the lab or taken directly from the lake. For these tests of food limitation, juveniles of different species were submitted to one of the following treatments: (1) lake seston and (2) lake seston + green algae. The age at first reproduction, mean clutch size, total offspring and the intrinsic rate of natural increase (r) were estimated. Cladocerans responded to both seasonal changes in food resources and to additions of green algae. In summer, food levels were higher and cladocerans grew better than in autumn. The low fecundity and population growth rates of Daphnia gessneri, together with high C:P ratios (>500) in the seston in summer, suggested P limitation. The other cladocerans (Moina micrura, Ceriodaphnia cornuta and Simocephalus mixtus) were apparently less sensitive than D. gessneri to P-limitation in summer. However, energy limitation cannot be disregarded even in summer, although in a lower degree compared to autumn. In this season, the predominance of large diatoms and dinophyceans of low nutritional value and the low food concentration suggested both quantitative and qualitative food limitation. Phytoplankton composition, nutritional value and particle size appeared to be important factors in determining cladocerans reproduction rates with seston diets.     

Effects of food limitation and temperature on cladocerans from a tropical Brazilian lake

Food limitation was tested in the laboratory by individual growth and reproduction of two cladoceran species, Ceriodaphnia richardi and Daphnia gessneri, from the shallow tropical Brazilian Lake Monte Alegre. The cladocerans were fed cultivated green alga Scenedesmus spinosus in concentrations of 0.20, 0.10, 0.05, and 0.025 mg C l⁻¹. Higher biomass and growth rates occurred in the two highest-food concentrations; the two lowest ones negatively affected clutch size and first reproduction. The threshold food concentration is lower than 0.025 mg C l⁻¹ and the incipient limiting level is a value between 0.10 and 0.20 mg C l⁻¹. The largest species, D. gessneri, was more sensitive to low food concentrations. The effects of low and high temperatures (19 and 27°C) were evaluated by life table experiments with three cladocerans from the lake—Daphnia ambigua, D. gessneri, and Moina micrura—with no food limitation (1 mg C l⁻¹ of S. spinosus). Higher population growth rates for the three species were found at 27°C; better performance in most life table parameters was observed for the former two species at the highest temperature, D. gessneri being the most sensitive to the lowest temperature. There are indications that temperature is an important abiotic factor that constrains populations of cladocerans for a short period in winter in the lake, when temperature decreases to 18–19°C. However, its influence cannot be separated from a biotic factor such as food, whose effect is stronger in the cool season, when concentrations are lower and contribution of inedible algae is relatively higher.     

Influence of biochemical,mineral and morphological features of natural food on tropical cladocerans

Five laboratory experiments were performed to evaluate the effect of supplements of fatty acids and a green alga on the individual growth and reproduction of three species of tropical cladocerans Ceriodaphnia richardi, Daphnia ambigua, and D. gessneri feeding on natural seston from the Brazilian Lake Monte Alegre. Cohorts of newborns from cultivated females were submitted to one of the following treatments: (1) Natural seston, (2) Natural seston + microcapsules of EPA and DHA or linoleic and linolenic fatty acids, (3) Natural seston + oil-free microcapsules, and (4) Natural seston + green alga Scenedesmus spinosus (1 mg C l⁻¹). Particulate organic carbon, algal carbon, C:P ratios of seston and green alga, polyunsaturated fatty acids (PUFA) content of seston and cladocerans, as well as phytoplankton composition, size, and shape were measured. The addition of fatty acids to seston did not significantly enhance growth and reproduction of the cladocerans, suggesting that sestonic PUFA content is sufficient for promoting cladoceran development, even in the cool–dry season when the fatty acids used in the experiments were 5–10 times lower in the seston than in the warm–wet season. Despite high C:P molar ratios in most experiments, there was only one indication of growth limitation by P. Reproduction was more affected than individual growth on some occasions by food quantity (energy) caused apparently by algal size, morphology, and digestion resistance. Energy availability, which is affected by algae morphological characteristics, seems to prevail over PUFA and P in controlling growth and reproduction of cladocerans in tropical Lake Monte Alegre.     

Influence of phytoplankton fractions on growth and reproduction of tropical cladocerans

The influence of two seston fractions, < 20 µm (nanoplankton) and 20 µm (microplankton), on growth and reproduction of cladoceran species with different sizes, from Lake Monte Alegre, was evaluated through individual growth and life table experiments. Size, shape and other features of the algae in the fractions were described. The procedure of filtering lake water through a 20 µm net for seston fractionation caused mutual contamination. The smallest cladoceran species, Ceriodaphnia cornuta Sars and Moina micrura Kurz, produced larger clutch sizes and exhibited higher intrinsic rates of population growth (r) in the nanoplankton, despite contamination of their food by inedible algae. The largest species, Simocephalus mixtus Sars, produced larger clutch sizes in the microplankton. There were no differences in juvenile biomass growth between treatments for M. micrura and Daphnia gessneri Herbst, but lower value of the exponential growth rate (g) in the microplankton was significant for M. micrura. Fecundity (eggs/total female) of M. micrura was significantly lower in the microplankton, while D. gessneri did not reproduce in this fraction, at the end of growth experiments. Spines, colonies, cenobium, filaments, hard cell wall, and gelatinous sheaths, represented constraints to cladoceran reproductive performance, besides algae size. Microplankton contamination by nanoplanktonic algae, in the experiments, probably minimized the negative effect of inedible algae. Nanoplankton was more suitable for the smallest species and microplankton for the largest one.     

Responses of tropical cladocerans to a gradient of resource quality

1. The response of three tropical cladocerans to a gradient of resource quality was compared in a series of growth bioassays using seston collected from five lakes of different depth and trophic structure in Michigan, U.S.A. To assess the food quality in terms of digestibility, assimilation experiments were performed with ³²P‐labelled seston from the same lakes. Animals were also analysed for P‐content in their tissues at the end of these assays. 2. In general, assimilation efficiency was higher when animals fed on seston from shallow compared to deep lakes, and was significantly correlated with growth rates, suggesting that shallow lakes have the best food resources in terms of digestibility and P availability. 3. Results also showed that all cladoceran species responded similarly to the resource gradient, with lower growth rates in deep lakes and higher growth rates in shallow lakes, although the strength of response (sensitivity) was different among the species tested. 4. The cladoceran Moina micrura was the most sensitive species, and also displayed the highest P‐content and maximal growth rate, a pattern consistent with the growth rate hypothesis. 5. However, seston C : P ratio and growth rates in the different resources did not correlate with the animals’ P‐contents, showing an uncoupling between RNA‐phosphorus demands for growth and seston food quality. 6. In conclusion, our results support the idea that digestion resistance in algae is a major constraint to cladocerans in natural plankton communities.     

Influence of phytoplankton fractions on growth and reproduction of tropical cladocerans

The influence of two seston fractions, < 20 μm (nanoplankton) and ≥ 20 μm (microplankton), on growth and reproduction of cladoceran species with different sizes, from Lake Monte Alegre, was evaluated through individual growth and life table experiments. Size, shape and other features of the algae in the fractions were described. The procedure of filtering lake water through a 20 μm net for seston fractionation caused mutual contamination. The smallest cladoceran species, Ceriodaphnia cornuta Sars and Moina micrura Kurz, produced larger clutch sizes and exhibited higher intrinsic rates of population growth (r) in the nanoplankton, despite contamination of their food by inedible algae. The largest species, Simocephalus mixtus Sars, produced larger clutch sizes in the microplankton. There were no differences in juvenile biomass growth between treatments for M. micrura and Daphnia gessneri Herbst, but lower value of the exponential growth rate (g) in the microplankton was significant for M. micrura. Fecundity (eggs/total female) of M. micrura was significantly lower in the microplankton, while D. gessneri did not reproduce in this fraction, at the end of growth experiments. Spines, colonies, cenobium, filaments, hard cell wall, and gelatinous sheaths, represented constraints to cladoceran reproductive performance, besides algae size. Microplankton contamination by nanoplanktonic algae, in the experiments, probably minimized the negative effect of inedible algae. Nanoplankton was more suitable for the smallest species and microplankton for the largest one.     

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.     

Effects of saxitoxin- and non-saxitoxin-producing strains of the cyanobacterium Cylindrospermopsis raciborskii on the fitness of temperate and tropical cladocerans

Life-table experiments were performed with two strains of Cylindrospermopsis raciborskii, a saxitoxin-producing strain (T3) and a non-saxitoxin-producing strain (NPLP-1) combined with the green algae Ankistrodesmus falcatus, aiming to detect effects on the reproduction of three cladocerans species: Daphnia pulex, Daphnia gessneri and Moina micrura. Survivorship, age at first reproduction, clutch size and population growth rate during 12–15 days were recorded. Cladocerans showed different susceptibility to saxitoxin-producing strain (T3), with D. pulex and M. micrura being negatively affected, followed by and D. gessneri, which did not show any decrease in fitness and performed even better than the controls with this strain. A reverse response was found in experiments with the non-saxitoxin-producing strain (NPLP-1): while D. pulex and M. micrura had an increase, D. gessneri showed a significant decrease in fitness, suggesting that this strain may produce some bioactive compound. The contrasting responses of the cladoceran species to saxitoxins and non-saxitoxin-producing strains of C. raciborskii suggest that the presence of this cyanobacterium may be a selective factor in determining the composition of zooplankton communities.     

A Far East Moina,M. weismanni Ishikawa, 1896 found in an Italian ricefield

The study of heleoplankton samples collected in an experimental ricefield in the province of Reggio Emilia (Northern Italy) evidenced a successional pattern of cladoceran species, characterized by the dominance of Moinidae in the early phase the rice cultivation (from May to July). In particular a species new for Italy,Monia weismanni Ishikawa, 1896, typical of Far East ricefields, was found; it was probably introduced into Italy with seeds and constitutes a stable element of the heleoplankton biocoenosis. The population co-occurs withMoina brachiata, Moina affinis, andMoina micrura in the ricefield. In this short noteM. weismanni is described, pointing out the morphological differences with other populations of the same species and with similar species; furthermore, life cycles in Moinidae are analysed in comparison with those of the other dominant cladocerans.     

A laboratory study on the demography and competition of three species of littoral cladocerans from Lake Huetzalin,Xochimilco, Mexico

Lake Huetzalin (Xochimilco) is a shallow waterbody in Mexico City with a dominance of the littoral cladocerans Simocephalus vetulus, Alona rectangula, and Pleuroxus aduncus. Here, we present data on the life table and population dynamics of these three cladoceran species on two types of diets (alga Scenedesmus acutus and detritus). We also analyzed the effect of competition on population growth using Scenedesmus, which proved to be the better diet than detritus. We hypothesize that feces of the larger cladoceran species (S. vetulus) could be a food source for the smaller taxa (A. rectangula and P. aduncus). Experiments were conducted in 100 ml beakers with 50 ml of medium in each at a temperature of 23 ± 1°C. We studied the demography of three test species using two food types with four replicates for each treatment in the life table experiments. Based on the data thus obtained we further studied the competition between the cladocerans on a diet of Scenedesmus. The maximal growth rates obtained for P. aduncus, S. vetulus, and A. rectangula fed Scenedesmus acutus were: 0.1, 0.3, and 0.5 d⁻¹, respectively. Among the test species, S. vetulus had the highest survivorship of 77 days when cultured on detritus and P. aduncus the lowest of 20 days on Scenedesmus. The gross reproductive rate (offspring female⁻¹ lifespan⁻¹) ranged from 6 neonates in Pleuroxus to 160 in Simocephalus per female. From the competition experiments conducted using alga as the diet, we observed that A. rectangula and, especially P. aduncus, had higher growth rates in the presence of S. vetulus than in the controls. This facilitation could be one of the reasons for higher species diversity in the littoral of water bodies.     

Effect of Aeolosoma sp. (Aphanoneura: Aeolosomatidae) on the Population Dynamics of Selected Cladoceran Species

Although oligochaete worms naturally coexist with cladocerans in many shallow freshwater ponds and lakes, their influence on the latter is not well established. In this work we studied the effect of Aeolosoma sp. on the population growth of Alona rectangula, Ceriodaphnia dubia, Daphnia pulex, Macrothrix triserialis and Moina macrocopa. Population growth studies were conducted at one algal food density (1 × 10⁶cells ml^–1 of Chlorella vulgaris). The experimental design was similar for all five cladoceran species, where we used 100 ml capacity transparent jars containing 50 ml of EPA medium with the desired algal density and three replicates for each treatment. The test medium was changed daily and fresh algal food was added. The initial density of each of the cladoceran species in the population growth studies was 0.4 ind ml^–1 while that of the worms 1.0 ind ml^–1. Following inoculation, we estimated daily the number of cladocerans and the worms for duration of 21 days. Regardless of the presence of worms, Moina macrocopa and Macrothrix triserialis showed rapid population growth while A. rectangula took more than 2 weeks to reach peak abundances. With the exception of M. triserialis, all the other our cladoceran species declined in the presence of Aeolosoma sp. The lowest peak population density (about 1 ind ml^–1) was observed for M. triserialisin controls. The remaining species had peak densities of about 3–5 ind ml^–1. The rates of population increase per day varied from 0.03 to 0.19 depending on the cladoceran taxa and the treatment. In general we found that pelagic taxa were more adversely affected by the presence of the worms than were the littoral cladocerans.     

Influence of seston quantity and quality on growth of tropical cladocerans.

The seston of the small, shallow, and tropical lake Monte Alegre was tested for quantity and quality for cladocerans by growth bioassays, which were carried out in spring (Daphnia gessneri and Moina micrura), summer (D. gessneri, M. micrura, Ceriodaphnia cornuta, and Simocephalus mixtus), and winter (D. gessneri and D. ambigua). Cohorts of newborns originating from ovigerous females collected in the lake or from laboratory cultures were submitted, at a room temperature of 23 degrees C to the following treatments: (1) the chlorophytes Ankistrodesmus falcatus and/or Scenedesmus spinosus; (2) lake seston; and (3) lake seston + chlorophytes. Growth rate, clutch size, and fecundity were evaluated. Seston alone was not the best food for promoting cladoceran growth. There were seasonal differences in food quantity and quality with spring and summer seston being better for growth than that of the winter. Adding chlorophytes to the seston increased clutch size and fecundity for most species in summer and winter, but not in spring. Energy limitation seems to be the most important factor influencing cladoceran growth in summer and especially in winter.     

Differential Impacts of Copepods and Cladocerans on Lake Seston,and Resulting Effects on Zooplankton Growth

In an enclosure study in Schöhsee, a small mesotrophic lake in Northern Germany, the impact of copepods and daphniids on the seston community was studied. In general, these two guilds differ in their feeding behaviour. Copepods actively select their food, with a preference for larger particles, whereas most cladocerans are unselective filter-feeders. In this study we investigate how the impact of the two different grazers affects zooplankton growth. We combine results obtained in the laboratory with results measured in situ in the enclosures. Copepods and cladocerans were cultured on seston from enclosures that were inhabited by density gradients of copepods or daphniids. We observed that Daphnia grew faster on seston that was pre-handled by copepods than on seston that was pre-handled by daphniids, and that somatic growth decreased with increasing densities of daphniids in the enclosures. In contrast, we observed no differences in development rates for copepods grown on the different media. The population growth rates of Daphnia in the Daphnia treatments were determined in the enclosures. Growth differences in both somatic- and population growth of Daphnia were correlated to food quality aspects of the seston. In the laboratory we found that Daphnia growth was correlated with several fatty acids. The strongest regression was with the concentration of 20:43 (r ²= 0.37). This particular fatty acid also showed the highest correlation with growth after normalisation of the fatty acids to the carbon content of the enclosures (r ²= 0.33). On the other hand, in the enclosure the population growth correlated most to the particulate nitrogen content (r ²= 0.78) and only to the N:C ratio, when normalised to carbon (r ²= 0.51).     

Evaluation of Microcystis as food for zooplankton in a eutrophic lake

Four experiments were conducted to evaluate Microcystis as food for zooplankton in Lake Kasumigaura, and the following results were obtained. (1) Moina micrura (Cladocera) showed little growth and no reproduction when the animal was reared with Microcystis cultured in the laboratory. The animal did not grow nor reproduce well when Chlorella was mixed with Microcystis as food. (2) Moina micrura assimilated Microcystis much less than Chlorella when the animal fed on single species of Microcystis or a mixture with Chlorella. (3) Microcystis collected from Lake Kasumigaura could not be utilized by Moina micrura even though the colonies were broken up into edible sizes. However, the alga turned into utilizable food when it was decomposed. (4) No inhibitors of Moina micrura population growth could be found in the non-filtered water of Lake Kasumigaura where Microcystis was blooming heavily. Decomposed Microcystis seemed to be utilized by zooplankton as an important food source in Lake Kasumigaura.     

The effect of different algal densities of Scenedesmus acuminatus on the population growth of Moina micrura Kurz (Crustacea: Anomopoda,Moinidae)

Six densities (0.5 × 10⁶, 1.0 × 10⁶, 1.5 × 10⁶, 2.0 × 10⁶, 3.0 × 10⁶, and 4.0 × 10⁶ cells ml^–1) of the micro-alga Scenedesmus acuminatus, were fed to the cladoceran, Moina micrura, in 40-litre glass aquaria. Moina population increased with increasing cell densities of Scenedesmus only up to treatment 3 (i.e. 1.5 × 10⁶ cells ml^–1) where a peak population of 11303 individuals per litre was obtained. Moinapopulation growth was inhibited at higher algal densities. The percentage of egg-bearing females and the number of eggs per egg-bearing females, followed a similar pattern. Comparatively, the peak production density of approximately 11000 Moina per litre, is interesting from a mass production point of view and indicates that S. acuminatus is a satisfactory micro-alga food for M. micrura.     

Comparison of the roles of Ostracods and Cladocerans in regulating community structure and metabolism in freshwater microcosms

Laboratory microcosms were used to compare the effects of the littoral ostracod Cypridopsis vidua and the planktonic cladoceran Daphnia magna on community structure and metabolism. Filter-feeding by cladocerans, both in the presence and absence of ostracods, greatly reduced the abundance of planktonic algae when D. magna reached peak density around day 50; rotifers and euglenids were then limited to flocculent matter on the container bottom. Both net production and community respiration rates decreased as community composition changed. Microcosms containing ostracods as the only microcrustacean showed little reduction in total algal numbers but the otherwise dominant alga, Scenedesmus spp., was replaced by Ankistrodesmus spp. when peak ostracod density was reached around day 100. Rotifers were completely eliminated but euglenids were able to coexist with ostracods. Ostracods impacted community metabolism less than cladocerans, but depressed respiration slightly more than net production.     

Food quality effects on life history traits and fitness in the generalist herbivore Daphnia

Summary Life table experiments were conducted on the generalist suspension feeder Daphnia galeata, using as food the two green algae (Chlorophyta) Scenedesmus acutus and Oocystis lacustris. Oocystis was hypothesized to be a lower quality food because it is convered with a thick sheath, believed to reduce digestibility. Results showed that Oocystis is a lower quality food for Daphnia, but only at relatively low food concentrations (0.15 mg C/L) and not at higher concentrations (1.0 mg C/L). At 0.15 mg C/L, Daphnia intrinsic rate of increase (r) when grown on Oocystis was only half that when grown on Scenedesmus. Daphnia r was similar at 0.15 mg C/L Oocystis and 0.075 mg C/L Scenedesmus, indicating that Daphnia requires twice as much Oocystis as Scenedesmus to achieve the same fitness. Intrinsic rate of increase was lower on Oocystis mainly because age at first reproduction was greatly delayed compared to that on Scenedesmus (13.6 vs 7.3 d). In addition, juvenile growth and survivorship were reduced on Oocystis compared with Scenedesmus. Clutch sizes were similar on the two foods, indicating that once individuals reached adulthood, the two foods were similar in quality. In contrast, at high food concentrations (1.0 mg C/L), the two algae were similar in quality for both juveniles and adults, and r was not significantly different on the two foods. Ingestion and assimilation rate experiments whowed that Daphnia consumes the two algae at identical rates, and that adults assimilate the two algae at similar rates. However, juveniles assimilate Oocystis at much lower rates than Scenedesmus, possibly accounting for reduced juvenile growth and delay in age at maturity at low concentrations. Thus, Daphnia exhibits an ontogenetic shift in its ability to utilize Oocystis, and this can result in juvenile bottlenecks in which survival and growth of young age classes are of critical importance in determining population dynamics. Because food quality effects were manifested primarily in juveniles and at low concentrations, food quality effects in nature will depend on phytoplankton abundance and age-structure of Daphnia populations.     

Daphnia population growth but not moulting is a substantial phosphorus drain for phytoplankton

SUMMARY 1. Negative effects of zooplankton on the availability of phosphorus (P) for phytoplankton as a result of the retention of nutrients in zooplankton biomass and the sedimentation of exoskeletal remains after moulting, have been recently proposed. 2. In a mesocosm study, the relative importance of these mechanisms was tested for the freshwater cladoceran Daphnia hyalina×galeata. A total of 13 mesocosm bags was suspended in a mesotrophic German lake during summer 2000 and fertilised with inorganic P in order to obtain a total nitrogen to total P ratio closer to the Redfield ratio. D. hyalina×galeata was then added at a logarithmically scaled density gradient of up to 40 ind. L^?1. Zooplankton densities, dissolved inorganic, particulate organic (seston <100 μm), as well as total nutrient concentrations were monitored. Additionally, nutrient concentrations of sediment water removed from the bottom of the mesocosm bags via a manual pump were determined. 3. Seston carbon (C), seston P and total P were significantly negatively correlated with Daphnia densities. The amount of particulate P (～5–6 μg P L^?1) sequestered from the seston compartment by Daphnia corresponded roughly to the increase of zooplankton biomass (population growth). Soluble reactive phosphorous (SRP) was at all times high (～25–35 μg P L^?1) and possibly unavailable to phytoplankton as a result of P adsorption to calcite during a calcite precipitation event (whiting). P concentrations determined in sediment water were generally <60 μg P m^?2 and thus never exceeded 1% of the total amount of P bound in particulate matter of the overlying water column. 4. Seston C : P ratios followed a polynomial second‐order function: At Daphnia densities <40 ind. L^?1 a positive linear relationship was evident, which is explained by the stronger reduction of P compared with C in seston, and transfer of seston P to zooplankton. Highest seston C : P ratios of ～300 : 1 were observed at Daphnia densities of ～30–50 ind. L^?1, which is in agreement with proposed threshold values limiting Daphnia reproductive growth. At Daphnia densities >40–50 ind. L^?1 C : P ratios were decreased because of the strong reduction of seston C at close to constantly low seston P‐values of ～3–4 μg P L^?1. 5. At least for Daphnia, it may be concluded that – unlike population growth – the sedimentation of faecal pellets and carapaces after moulting seem negligible processes in pelagic phosphorus dynamics.     

Egg production, development and growth in Moina micrura Kurz (1874) (Cladocera: Moinidae)

Laboratory observations on the egg production, development and growth in the Cladoceran Moina micrura show that this species has a life span of 13 days during which it produces 6M 8 eggs. Two preadult and eleven adult instars were recorded at a temperature range of 28°–30°C. The rate of egg production calculated on a cumulative basis was found to be lower (a = 1.0368) than that of other tropical species. The embryonic stages in Moina micrura closely resembled other cladoceran species, although with a short development period of 24 h. The total life span, growth rate and the total number of eggs produced in Moina micrura have been compared with those of allied tropical, temperate and arctic daphnids. The significance of this relationship to secondary productivity in tropical ecosystems has been discussed.     

A note about Daphnia in Venezuelan waterbodies

After examining numerous water samples of 19 lakes and reservoirs in Venezuela, only two species of Daphnia were found: D. laevis (in colder, high altitude lakes) and D. gessneri. This observation supports the theory of the scarcity of Daphnia species in tropical inland waters.