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: 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.

Description: Marine food webs are highly compartmentalized, and characterizing the trophic niches among consumers is important for predicting how impact from human activities affects the structuring and functioning of marine food webs. Biomarkers such as bulk stable isotopes have proven to be powerful tools to elucidate trophic niches, but they may lack in resolution, particularly when spatiotemporal variability in a system is high. To close this gap, we investigated whether carbon isotope (delta C-13) patterns of essential amino acids (EAAs), also termed delta(13)C(AA)fingerprints, can characterize niche differentiation in a highly dynamic marine system. Specifically, we tested the ability of delta(13)C(AA)fingerprints to differentiate trophic niches among six functional groups and ten individual species in the Baltic Sea. We also tested whether fingerprints of the common zooplanktivorous fishes, herring and sprat, differ among four Baltic Sea regions with different biochemical conditions and phytoplankton assemblages. Additionally, we investigated how these results compared to bulk C and N isotope data for the same sample set. We found significantly different delta(13)C(AA)fingerprints among all six functional groups. Species differentiation was in comparison less distinct, due to partial convergence of the species' fingerprints within functional groups. Herring and sprat displayed region-specific delta(13)C(AA)fingerprints indicating that this approach could be used as a migratory marker. Niche metrics analyses showed that bulk isotope data had a lower power to differentiate between trophic niches than delta(13)C(AA)fingerprinting. We conclude that delta(13)C(AA)fingerprinting has a strong potential to advance our understanding of ecological niches, and trophic linkages from producers to higher trophic levels in dynamic marine systems. Given how management practices of marine resources and habitats are reshaping the structure and function of marine food webs, implementing new and powerful tracer methods are urgently needed to improve the knowledge base for policy makers.

Description: Mixotrophic organisms are increasingly recognized as important components of ecosystems, but the factors controlling their nutrition pathways (in particular their autotrophy-heterotrophy balance) are little known. Both autotrophy and heterotrophy are expected to respond to density-dependent mechanisms but not necessarily in the same direction and/or strength. We hypothesize that the autotrophy-heterotrophy balance of mixotrophic organisms might therefore be a function of population densities. To investigate this relationship, we sampled mixotrophic jellyfish holobionts (host, Mastigias papua etpisoni; symbiont, Cladocopium sp.) in a marine lake (Palau, Micronesia) on six occasions (from 2010 to 2018). Over this period, population densities varied similar to 100 fold. We characterized the nutrition of the holobionts using the delta C-13 and delta N-15 signatures as well as C:N ratios. delta C-13 values increased and delta N-15 values decreased with increasing population densities (respectively, R-2 = 0.86 and 0.70, P 〈 0.05). Although less distinct, C:N ratios increased with increasing population densities (R-2 = 0.59, 0.1 〉 P 〉 0.05). This indicates that the autotrophy-heterotrophy balance tends toward autotrophy when population densities increase. We propose that the availability of zooplanktonic prey is the main driver of this pattern. These results demonstrate that the autotrophy-heterotrophy balance of mixotrophic jellyfishes can be tightly regulated by density-dependent mechanisms.

Description: We hypothesized that the trophic level of marine copepods should depend on the composition of the protist community. To test this hypothesis, we manipulated the phytoplankton composition in mesocosms and measured grazing rates of copepods and mesozooplankton in those mesocosms. Twelve mesocosms with Northeast Atlantic phytoplankton were fertilised with different Si:N ratios from 0:1 to 1:1. After 1 week, ten of the mesocosms were filled with natural densities of mesozooplankton, mainly calanoid copepods, while two remained as mesozooplankton-free controls. Both before and after the addition of copepods there was a positive correlation of diatom dominance with Si:N ratios. During the second phase of the experiment, copepod and microzooplankton grazing rates on different phytoplankton species were assessed by a modification of the Landry-Hassett dilution technique, where the bottles containing the different dilution treatments were replaced by dialysis bags incubated in situ. The results indicated no overlap in the food spectrum of microzooplankton (mainly ciliates) and copepods. Ciliates fed on nanoplankton, while copepods fed on large or chain-forming diatoms, naked dinoflagellates, and ciliates. The calculated trophic level of copepods showed a significantly negative but weak correlation with Si:N ratios. The strength of this response was strongly dependent on the trophic levels assumed for ciliates and mixotrophic dinoflagellates.

Description: Mesozooplankton communities in the mesooligotrophic Gulf of Aqaba, Northern Red Sea, were investigated over a 2 years period (2005-2007) with emphasis on the trophodynamic relations among different taxonomic groups ranging from primary consumers to carnivorous predators. Based on stable isotope analyses, we present evidence for a strong contribution of 'new' nitrogen mainly derived from the utilization of aerosol nitrate by unicellular cyanobacteria especially during summer stratification and the propagation of exceptionally low δ15N onto higher trophic levels. In contrast, N2-fixation by diazotrophs seemed to play a minor role, while the utilization of deep water nitrate by cyanobacteria and eukaryotic algae might be of importance during winter mixing. Based on 15N enrichment of consumers, clear differences between exclusively herbivorous organisms (doliolids, appendicularians, pteropods) and those with omnivorous feeding modes were detected. The category of omnivores comprised a large variety of taxons ranging from small meroplanktonic larvae to non-calanoid copepods (harpacticoids, cyclopoids and poecilostomatoids) that together form a diverse and complex community with overlapping feeding modes. In addition, distinct seasonality patterns in δ15N of copepods were found showing elevated trophic positions during periods of winter mixing, which were most pronounced for non-calanoid copepods. In general, feeding modes of omnivores appeared rather unselective, and relative contributions of heterotrophic protists and degraded material to the diets of non-calanoid copepods are discussed. At elevated trophic positions, four groups of carnivore predators were identified, while calanoid copepods and meroplanktonic predators showing lowest 15N enrichment within the carnivores. The direct link between 'new' nitrogen utilization by primary producers and the 15N enrichment of consumers in the planktonic food web of the Gulf of Aqaba emphasizes the significant contribution of 'new' nitrogen to the nitrogen budget and ecosystem functions in subtropical and tropical oligotrophic oceans.

Description: A recent meta-analysis indicates that trophic cascades (indirect effects of predators on plants via herbivores) are weak in marine plankton in striking contrast to freshwater plankton (Shurin et al. 2002, Ecol. Lett., 5, 785–791). Here we show that in a marine plankton community consisting of jellyfish, calanoid copepods and algae, jellyfish predation consistently reduced copepods but produced two distinct, opposite responses of algal biomass. Calanoid copepods act as a switch between alternative trophic cascades along food chains of different length and with counteracting effects on algal biomass. Copepods reduced large algae but simultaneously promoted small algae by feeding on ciliates. The net effect of jellyfish on total algal biomass was positive when large algae were initially abundant in the phytoplankton, negative when small algae were dominant, but zero when experiments were analysed in combination. In contrast to marine systems, major pathways of energy flow in Daphnia-dominated freshwater systems are of similar chain length. Thus, differences in the length of alternative, parallel food chains may explain the apparent discrepancy in trophic cascade strength between freshwater and marine planktonic systems.

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.