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: We used marine phytoplankton from mesocosms seeded with different zooplankton densities to study the impact of mesozooplankton on phytoplankton nutrient limitation. After 7 d of grazing (copepod mesocosms) or 9 d (appendicularian mesocosms) phytoplankton nutrient limitation was studied by enrichment bioassays. After removal of mesozooplankton, bioassay bottles received either no nutrients, phosphorus or nitrogen alone, or a combination of nitrogen and phosphorus and were incubated for 2 d. Phytoplankton reproductive rates in the bottles without nutrient addition were calculated after correction for grazing by ciliates and indicated increasing nitrogen limitation with increasing copepod abundance. No nutrient limitation was found in the appendicularian mesocosms. The increase of nutrient limitation with increasing copepod density seems to be mainly the result of a trophic cascade effect: Copepods released nanoplankton from ciliate grazing pressure, and thereby enhanced nitrogen exhaustion by nanophytoplankton and reduced nitrogen excretion by ciliates. Nitrogen sequestration in copepod biomass, the mechanism predicted by the ecological stoichiometry theory, seems to have been a weaker effect because there was only little copepod growth during the experiment.

Description: Zooplankton grazing on bacterio- and phytoplankton was studied in the Gulf of Aqaba and the Northern Red Sea during Meteor Cruise Me 44-2 in February-March 1999. Protozoan grazing on bacterioplankton and autotrophic ultraplankton was studied by the Landry dilution method. Microzooplankton grazing on phytoplankton 〉6 µm was studied by incubation experiments in the presence and absence of microzooplankton. Mesozooplankton grazing was studied by measuring per capita clearance rates of individual zooplankton with radioactively labelled food organisms and estimating in situ rates from abundance values. Protozoan grazing rates on heterotrophic bacteria and on algae 〈6 µm were high (bacteria: 0.7 to 1.1 d-1, ultraphytoplankton: 0.7 to 1.3 d-1), while grazing rates on Synechococcus spp. were surprisingly low and undetectable in some experiments. Mesozooplankton grazing was weak, cumulative grazing rates being ca. 2 orders of magnitude smaller than the grazing rates by protozoans. Among mesozooplankton, appendicularians specialised on smaller food items and calanoid copepods on larger ones.

Description: In a mesocosm study, the appendicularian Oikopleura dioica bloomed after the reduction of copepod abundance, and in a second treatment showed a significantly negative correlation with copepod densities. Calculations, together with field data from the Baltic Sea, suggest that common calanoid copepods may control appendicularian population dynamics.

Description: An indoor mesocosm system was set up to study the response of phytoplankton and zooplankton spring succession to winter and spring warming of sea surface temperatures. The experimental temperature regimes consisted of the decadal average of the Kiel Bight, Baltic Sea, and three elevated regimes with 2°C, 4°C, and 6°C temperature difference from that at baseline. While the peak of the phytoplankton spring bloom was accelerated only weakly by increasing temperatures (1.4 days per degree Celsius), the subsequent biomass minimum of phytoplankton was accelerated more strongly (4.25 days per degree Celsius). Phytoplankton size structure showed a pronounced response to warming, with large phytoplankton being more dominant in the cooler mesocosms. The first seasonal ciliate peak was accelerated by 2.1 days per degree Celsius and the second one by 2.0 days per degree Celsius. The over-wintering copepod populations declined faster in the warmer mesocosm, and the appearance of nauplii was strongly accelerated by temperature (9.2 days per degree Celsius). The strong difference between the acceleration of the phytoplankton peak and the acceleration of the nauplii could be one of the “Achilles heels” of pelagic systems subject to climate change, because nauplii are the most starvation-sensitive life cycle stage of copepods and the most important food item of first-feeding fish larvae.

Description: A common elemental analyzer system connected to a temperature-controlled gas chromatography (GC) column and coupled to an isotope ratio mass spectrometer was improved to decrease the determination limit for a simultaneous stable isotope ratio measurement of nitrogen and carbon dioxide. The additional use of a special ashtray system to collect the combustion residuals permitted more time-efficient work. These modifications to the elemental analyzer allowed precise measurements to be made down to 1.5 µg nitrogen and 10 µg carbon for stable isotope analysis. Low system background values and an acceptable signal-to-noise ratio have made an additional blank correction for these low sample measurements unnecessary. We provide a precision of this stable isotope analysis for lowest amounts of 1.2–2 µg nitrogen with a standard deviation of ±0.496‰ (n = 27) and for 8.2–15 µg carbon with a standard deviation of ±0.257‰ (n = 31) across different sample runs under stipulated conditions. This application can be established in an automatic mode without cryofocusing procedures.

Description: Based on existing knowledge about phytoplankton responses to nutrients and food size spectra of herbivorous zooplankton, three different configurations of pelagic food webs are proposed for three different types of marine nutrient regimes: (1) upwelling systems, (2) oligotrophic oceanic systems, (3) eutrophicated coastal systems. Up-welling systems are characterised by high levels of plant nutrients and high ratios of Si to N and R. Phytoplankton consists mainly of diatoms together with a subdominant contribution of flagellates. Most phytoplankton falls into the food spectrum of herbivorous, crustacean zooplankton. Therefore, herbivorous crustaceans occupy trophic level 2 and zooplanktivorous fish occupy trophic level 3. Phytoplankton in oligotrophic, oceanic systems is dominated by picoplankton, which are too small to be ingested by copepods. Most primary production is channelled through the ‘microbial loop’ (picoplankton — heterotrophic nanoflagellates — ciliates). Sporadically, pelagic tunicates also consume a substantial proportion of primary production. Herbivorous crustaceans feed on heterotrophic nanoflagellates and ciliates, thus occupying a food chain position between 3 and 4, which leads to a food chain position between 4 and 5 for zooplanktivorous fish. By cultural eutrophication, N and P availability are elevated while Si remains unaffected or even declines. Diatoms decrease in relative importance while summer blooms of inedible algae (Phaeocystis, toxic dinoflagellates, toxic prymnesiophyceae, etc.) prevail. The spring bloom may still contain a substantial contribution of diatoms. The production of the inedible algae enters the pelagic energy flow via the detritus food chain: DOC release by cell lysis — bacteria — heterotrophic nanoflagellates — ciliates. Accordingly, crustacean zooplankton occupy food chain position 4 to 5 during the non-diatom seasons. Ecological efficiency considerations lead to the conclusion that fish production:primary production ratios should be highest in upwelling systems and substantially lower in oligotrophic and in culturally eutrophicated systems. Further losses of fish production may occur when carnivorous, gelatinous zooplankton (jellyfish) replace fish.