Description: The differences in the impact of two major groups of herbivorous zooplankton (Cladocera and Copepoda) on summer phytoplankton in a mesotrophic lake were studied. Field experiments were performed in which phytoplankton were exposed to different densities of two major types of herbivorous zooplankton, cladocerans and copepods. Contrary to expectation, neither of the two zooplankton groups significantly reduced phytoplankton biomass. However, there were strong and contrasting impacts on phytoplankton size structure and on individual taxa. Cladocerans suppressed small phytoplankton, while copepods suppressed large phytoplankton. The unaffected size classes compensated for the loss of those affected by enhanced growth. After contamination of the copepod mesocosms with the cladoceran Daphnia, the combined impact of both zooplankton groups caused a decline in total phytoplankton biomass.

Description: The foraging modes of calanoid copepods differ in that stationary suspension-feeding is more easily detected by prey with strong escape responses (ciliates) than is ‘cruising’ or ‘ambushing’ feeding. Thus, the ability of a copepod to include heterotrophic prey in its diet may be associated with its foraging mode and, further, with its nitrogen stable isotope signature (δ15N). This is because a more carnivorous diet may be expected to result in a higher δ15N. We tested this hypothesis in a mesocosm study using a density gradient (0 to 80 ind. l-1) of calanoid copepods. We expected copepod δ15N to generally increase with decreasing copepod density because of increased food availability, and predicted stronger increases for cruising than for stationary suspension-feeding species. As an assemblage, copepods had a pronounced impact on the food web: diatoms and ciliates decreased, whereas nanoflagellates increased with increasing copepod density. As expected, Centropages hamatus, a cruising species, showed the strongest isotopic increase and also highest population growth at low copepod density, suggesting that it was the most efficient species in capturing ciliates. Temora longicornis, a stationary suspension-feeder, showed a uniform isotopic increase in all mesocosms, which we believe resulted from nutritional stress arising from poor feeding on both ciliates (too fast for ingestion by T. longicornis) and nanoflagellates (too small). However, Pseudocalanus elongatus, a species equally categorised as a stationary suspension-feeder, showed increases in its δ15N similar to those for C. hamatus. While this may indicate potential switching in its foraging mode, alternative explanations cannot be ruled out, partly because qualitative and quantitative aspects of trophic enrichment in our experiment could not be clearly separated. This study shows that consumer δ15N are difficult to interpret, even if potential food sources and aspects of the species’ biology are known, and thus emphasises the necessity for further laboratory studies to help better interpret zooplankton δ15N in the field.

Description: The variation in life-history patterns that allow closely related species to co-exist has been an important theme in ecology for decades. We examined intra- and interspecific variation in a key life-history trait - diapause - for two congeneric copepods (Eudiaptomus gracilis and Eudiaptomus graciloides) inhabiting three lakes in Northern Germany. Diapause has been hypothesized in both theoretical and empirical studies to be important in the coexistence of competing species. We found no diapause for E. gracilis whereas we documented two distinct periods of diapause in the life cycle of E. graciloides. In the latter species, diapausing eggs were produced in all three lakes during autumn, however, relative investment in diapausing eggs differed between lakes. Diapausing egg production was delayed or fewer females switched to making diapausing eggs in the more productive lakes relative to the less productive systems. In the sediments of all three lakes, viable diapausing eggs were found buried from the sediment-water interface down several centimeters in each lake, suggesting the presence of a long-lived egg bank. In addition, detailed population studies carried out on both species in one lake (Schöhsee) revealed a second period of diapause in E. graciloides. After the diapausing eggs are produced in autumn, females accumulate lipids, reduce feeding and apparently over-winter in a state of active diapause. The prevalence of diapause in E. graciloides and lack of it in E. gracilis raises questions regarding the factors that shape the life-histories of these congeners, and the role that the life-history differences observed may play in their coexistence.

Description: 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 20:4ω3 (r2= 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 (r2= 0.33). On the other hand, in the enclosure the population growth correlated most to the particulate nitrogen content (r2= 0.78) and only to the N:C ratio, when normalised to carbon (r2= 0.51).

Description: 1. We performed a mesocosm experiment to investigate the structuring and cascading effects of two predominant crustacean mesozooplankton groups on microbial food web components. The natural summer plankton community of a mesotrophic lake was exposed to density gradients of Daphnia and copepods. Regression analysis was used to reveal top-down impacts of mesozooplankton on protists and bacteria after days 9 and 15. 2. Selective grazing by copepods caused a clear trophic cascade via ciliates to nanoplankton. Medium-sized (20-40 mum) ciliates (mainly Oligotrichida) were particularly negatively affected by copepods whereas nanociliates (mainly Prostomatida) became more abundant. Phototrophic and heterotrophic nanoflagellates increased significantly with increasing copepod biomass, which we interpret as an indirect response to reduced grazing pressure from the medium-sized ciliates. 3. In Daphnia-treatments, ciliates of all size classes as well as nanoflagellates were reduced directly but the overall predation effect became most strongly visible after 15 days at higher Daphnia biomass. 4. The response of bacterioplankton involved only modest changes in bacterial biomass and cell-size distribution along the zooplankton gradients. Increasing zooplankton biomass resulted either in a reduction (with Daphnia) or in an increase (with copepods) of bacterial biovolume, activity and production. Patterns of bacterial diversity, as measured by polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE), showed no distinct grouping after 9 days, whereas a clear treatment-coupled similarity clustering occurred after 15 days. 5. The experiment demonstrated that zooplankton-mediated predatory interactions cascade down to the bacterial level, but also revealed that changes occurred rather slowly in this summer plankton community and were most pronounced with respect to bacterial activity and composition.

Description: 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.