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Fin whales (Balaenoptera physalus) aggregate west of Iceland during summer, where they feed mostly on krill, but their winter distribution is poorly known. This study investigates the winter behavior of fin whales that summer off Iceland by analyzing amino acid‐specific nitrogen isotope ratios in the baleen plates. This technique is increasingly used in large marine vertebrates to discriminate between changes in trophic position and changes in isotopic baseline levels, while ruling out the effect of fasting on these patterns. Analyses were conducted on samples extracted from two baleen plate segments: one corresponding to winter, another to summer. The trophic position was higher and slightly more variable in winter (3.6 ± 0.7) than in summer (3.0 ± 0.4), which indicates that during winter fin whales tend to feed on fish and are more generalist in feeding habits. Interestingly, the δ15NPhe values were not lower in winter than in summer (winter: 5.2‰ ± 2.4‰; summer: 4.0‰ ± 2.1‰). This was unexpected because baseline δ15N values (e.g., zooplankton) tend to be lower at low latitudes, where fin whales are believed to spend the winter. Fin whales may remain in relatively high latitudes during winter or visit low‐latitude areas with locally higher than average δ15N values, such as upwelling areas. [ABSTRACT FROM AUTHOR]

Eutrophication, increased temperatures and stratification can lead to massive, filamentous, N2‐fixing cyanobacterial (FNC) blooms in coastal ecosystems with largely unresolved consequences for the mass and energy supply in food webs. Mesozooplankton adapt to not top‐down controlled FNC blooms by switching diets from phytoplankton to microzooplankton, resulting in a directly quantifiable increase in its trophic position (TP) from 2.0 to as high as 3.0. If this process in mesozooplankton, we call trophic lengthening, was transferred to higher trophic levels of a food web, a loss of energy could result in massive declines of fish biomass. We used compound‐specific nitrogen stable isotope data of amino acids (CSIA) to estimate and compare the nitrogen (N) sources and TPs of cod and flounder from FNC bloom influence areas (central Baltic Sea) and areas without it (western Baltic Sea). We tested if FNC‐triggered trophic lengthening in mesozooplankton is carried over to fish. The TP of cod from the western Baltic (4.1 ± 0.5), feeding mainly on decapods, was equal to reference values. Only cod from the central Baltic, mainly feeding on zooplanktivorous pelagics, had a significantly higher TP (4.6 ± 0.4), indicating a strong carry‐over effect trophic lengthening from mesozooplankton. In contrast, the TP of molluscivorous flounder, associated with the benthic food web, was unaffected by trophic lengthening and quite similar reference values of 3.2 ± 0.2 in both areas. This suggests that FNC blooms lead to a large loss of energy in zooplanktivorous but not in molluscivorous mesopredators. If FNC blooms continue to trigger the detour of energy at the base of the pelagic food web due to a massive heterotrophic microbial system, the TP of cod will not return to lower TP values and the fish stock not recover. Monitoring the TP of key species can identify fundamental changes in ecosystems and provide information for resource management. [ABSTRACT FROM AUTHOR]

The mean trophic position (TP) of mesozooplankton largely determines how much mass and energy is available for higher trophic levels like fish. Unfortunately, the ratio of herbivores to carnivores in mesozooplankton is difficult to identify in field samples. Here, we investigated changes in the mean TP of mesozooplankton in a highly dynamic environment encompassing four distinct habitats in the southern South China Sea: the Mekong River plume, coastal upwelling region, shelf waters, and offshore oceanic waters. We used a set of variables derived from bulk and amino acid nitrogen stable isotopes from particulate organic matter and four mesozooplankton size fractions to identify changes in the nitrogen source and TP of mesozooplankton across these habitats. We found clear indications of a shift in N sources for biological production from nitrate in near-coastal waters with shallow mixed layer depths toward an increase in diazotroph-N inputs in oceanic waters with deep mixed layer depths where diazotrophs shaped the phytoplankton community. The N source shift was accompanied by a lengthening of the food chain (increase in the TP). This may provide further support for the connection between diazotrophy and the indirect routing of N through the marine food web. Our combined bulk and amino acid δ15N approach also allowed us to estimate the trophic enrichment (TE) of mesozooplankton across the entire regional ecosystem. When put in the context of literature values, a high TE of 5.1‰ suggested a link between ecosystem heterogeneity and the less efficient transfer of mass and energy across trophic levels. [ABSTRACT FROM AUTHOR]

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ13C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ13C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ13C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ13C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs. [ABSTRACT FROM AUTHOR]

Changes in the complexity of planktonic food webs may be expected in future aquatic systems due to increases in sea surface temperature and an enhanced stratification of the water column. Under these conditions, the growth of unpalatable, filamentous, N2‐fixing cyanobacterial blooms, and their effect on planktonic food webs will become increasingly important. The planktonic food web structure in aquatic ecosystems at times of filamentous cyanobacterial blooms is currently unresolved, with discordant lines of evidence suggesting that herbivores dominate the mesozooplankton or that mesozooplankton organisms are mainly carnivorous. Here, we use a set of proxies derived from amino acid nitrogen stable isotopes from two mesozooplankton size fractions to identify changes in the nitrogen source and the planktonic food web structure across different microplankton communities. A transition from herbivory to carnivory in mesozooplankton between more eutrophic, near‐coastal sites and more oligotrophic, offshore sites was accompanied by an increasing diversity of microplankton communities with aging filamentous cyanobacterial blooms. Our analyses of 124 biotic and abiotic variables using multivariate statistics confirmed salinity as a major driver for the biomass distribution of non‐N2‐fixing microplankton species such as dinoflagellates. However, we provide strong evidence that stratification, N2 fixation, and the stage of the cyanobacterial blooms regulated much of the microplankton diversity and the mean trophic position and size of the metabolic nitrogen pool in mesozooplankton. Our empirical, macroscale data set consistently unifies contrasting results of the dominant feeding mode in mesozooplankton during blooms of unpalatable, filamentous, N2‐fixing cyanobacteria by identifying the at times important role of heterotrophic microbial food webs. Thus, carnivory, rather than herbivory, dominates in mesozooplankton during aging and decaying cyanobacterial blooms with hitherto uncharacterized consequences for the biogeochemical functions of mesozooplankton. R/V Meteor passing a decaying bloom of unpalatable, filamentous, N2‐fixing cyanobacteria in the Baltic Sea in August 2015 that are expected to increase in large areas of the global ocean in the future. While herbivory dominated the trophic structure of mesozooplankton in earlier bloom stages, direct estimates of carnivory in field mesozooplankton identified the at times important role of heterotrophic microbial food webs namely in such aging and decaying cyanobacterial blooms with hitherto uncharacterized consequences for the biogeochemical functions of mesozooplankton. Photo: Andreas Raeke, DWD. [ABSTRACT FROM AUTHOR]

Abstract: Cyanobacteria are the main autotrophs and N2‐fixing (diazotrophic) organisms in large parts of the oligotrophic global ocean, where generally all heterotrophic production depends on their activity. Amino acids (AAs) from cyanobacteria are essential macronutrients for these heterotrophic food webs, yet little is known about the de novo synthesis of AAs during N2 fixation. Through a combination of bulk and amino acid nitrogen (AAN) specific analyses of field based N2 fixation experiments, we demonstrate that the de novo synthesis of 13 AAs accounted for the majority of bulk N2 fixation rates at four stations in the central Baltic Sea in July 2015. Slow AA turnover times of 87 ± 14 d coincided with low phosphate concentrations and high cell‐carbon biomasses of unicellular cyanobacteria. Very fast turnover times of 17 ± 3 d coincided with high phosphate concentrations and undecayed Nodularia spumigena cells, but unexpectedly also with phosphate depletion and decayed N. spumigena cells. In a decayed bloom, volumetric N2 fixation rates into AAN provided a much better estimate of the net incorporation of N2 into biomass than fixation into bulk nitrogen that rather reflected gross N2 fixation. In an undecayed bloom, the turnover times of 13 AAs can be predicted from a single bulk N2 fixation rate. This is the first direct evidence that the very late, decayed stage of a cyanobacteria bloom can be a flashpoint of very fast AA turnover during N2 fixation with hitherto uncharacterized consequences for heterotrophic food webs and diazotroph N inputs to the global ocean. [ABSTRACT FROM AUTHOR]

Knowledge of how zooplankton can utilize cyanobacteria to sustain their nitrogen (N) demand for essential compounds like amino acids (AAs) is a key to predicting responses of higher trophic levels in terms of production and food web structure to future enhanced water column stratification. We explored the natural abundances of bulk N and AA-specific nitrogen isotopes (δ15N) in particulate organic matter and mesozooplankton size-fraction samples from three vertically separated water bodies in the central Baltic Sea during two summertime cyanobacteria blooms. The combination of plankton community and isotope data together with environmental variables helped to identify a mechanism of diazotrophic AA supply (synthesized during N2 fixation) for mesozooplankton, that largely depended on the sea surface temperature which regulated the access to the diazotrophic N-based food web in the surface water (SW). We found that in the warmer summer, thermophilic cladocerans (e.g., Bosmina spp.) benefited most from diazotrophic AAs in the SW (19.8°C), while only in the colder summer, temperate copepods (e.g., Temora longicornis) ascended from the subjacent winter water into the SW (16.2°C) and incorporated diazotrophic AAs. Trophic position estimates based on the phenylalanine and glutamic acid δ15N signatures revealed that the diazotrophic AA supply into mesozooplankton was mainly indirect via feeding on mixo- and heterotrophic diets. Significantly enriched δ15N signatures in phenylalanine in the deep mesozooplankton (mainly copepods of Pseudo- and Paracalanus spp.) from the bottom water (BW) that was a region of the suboxic zone point to a reliance on a local food web. Mesozooplankton in the BW was feeding on diets of heterotrophic origin and probably profited from the heterotrophic re-synthesis of AAs originating from sinking organic matter, as well as from the indirect incorporation of de novo synthesized AAs that most likely originated from chemoautotrophic bacteria or archaea communities in the suboxic zone. Our findings suggest that indirect feeding on diazotrophs and chemoautotrophs will be principal ways of amino acid supply for zooplankton in future enhanced stratified aquatic systems. Only a relatively small increase in temperature may restrict temperate key species from diazotrophic N-based food webs in the mixed layer. [ABSTRACT FROM AUTHOR]

Abstract: We report on methane enrichments that were observed during summer in the upper water column of the Gotland Basin, central Baltic Sea. In the eastern part of the basin, methane concentrations just below the thermocline varied between 15 nM and 77 nM, in contrast to the western part where no methane enrichments could be detected. Stable carbon isotope ratios of methane (δ13C‐CH4 of −67.6‰) indicated its in situ biogenic origin from CO2 reduction, which was supported by clonal sequences that clustered with Methanomicrobiaceae, a family of methanogenic Archaea. Incubation experiments with a Temora longicornis dominated seston fraction obtained from the relevant depth showed a positive correlation between seston concentration and methane production rates. Our results, in combination with previous literature outcomes, suggest that the methane enrichment in the eastern basin might be sustained by a diet‐consumer relationship between the dinoflagellate Dinophysis norvegica and the copepod T. longicornis. However, our mass balance indicates that a local methane production of 110 pmol L−1 d−1 was needed to maintain the methane enrichment, and that the estimated production rate from our incubation experiments of 0.3 pmol CH4 d−1 per adult T. longicornis (about 1 pmol L−1 d−1) was too low to maintain the methane enrichment by zooplankton associated methane production only. These calculations also showed that methane was consumed below the thermocline and not transported into the upper‐ocean, suggesting that other sources in the mixed layer in the range of 95 pmol L−1 d−1 are needed to maintain the observed methane air–sea flux. [ABSTRACT FROM AUTHOR]

The plasticity of nitrogen specific net growth efficiency (NGE) in marine mesozooplankton is currently unresolved, with discordant lines of evidence suggesting that NGE is constant, or that it varies with nitrogen source, food availability, and food quality in marine ecosystems. Specifically, the fate of nitrogen from nitrogen fixation is poorly known. We use 15N : 14N ratios in plankton in combination with hydrological data, nutrient profiles, and nitrogen fixation rate measurements to investigate the relationship between new nitrogen sources and the nitrogen specific NGE in three plankton communities along the outer Amazon River plume. The NGE of small (200-500 μm) mesozooplankton was estimated from the δ15N differences between particulate nitrogen and zooplankton using an open system Rayleigh fractionation model. The transfer efficiency of nitrogen among larger (> 500 μm) mesozooplankton was estimated from the change in δ15N as a function of zooplankton size. The Amazon River was not a significant source of bioavailable nitrogen anywhere in our study region, and subsurface nitrate was the primary new nitrogen source for the outer shelf community, which was dominated by diatoms. N2 fixation was the principal new nitrogen source at sites of high diatom diazotroph association abundance and at oceanic sites dominated by Trichodesmium spp. and Synechococcus spp. Although we found clear spatial differences in food quantity, food quality, and diazotroph inputs into mesozooplankton, our data show no significant differences in mesozooplankton nitrogen transfer efficiency and NGE (for latter, mean ± SD: 59 ± 10%) among sites. [ABSTRACT FROM AUTHOR]

Eurytemora affinis (Copepoda) were fed 15N-labeled Rhodomonas salina (Cryptophyta) or 15N-labeled Nodularia spumigena (Cyanobacteria) in excess under controlled laboratory conditions. Zooplankton collected from the Baltic Sea were fed natural phytoplankton amended with 15N-labeled N. spumigena. We quantified the direct incorporation of 15N tracer from N2-fixing N. spumigena (diazotroph nitrogen) and ammonium-utilizing R. salina into the amino acid nitrogen (AA-N) of zooplankton using complementary gas chromatography-combustion-isotope ratio mass spectrometry, gas chromatography-mass spectrometry, and elemental analysis-isotope ratio mass spectrometry approaches. Specific and mass-specific TN and AA-N incorporation rates of the 15N tracers were calculated for zooplankton. Highest incorporation of 15N was found in field zooplankton relying on N. spumigena and in E. affinis relying on R. salina. Lowest incorporation was found in E. affinis relying on N. spumigena. Decreasing specific and mass-specific rates during field experiments possibly were due to food shortage, whereas decreasing rates in E. affinis grazing on R. salina were more likely due to satiation. Specific and mass-specific rates were consistently low in E. affinis when exposed to N. spumigena, suggesting that these animals were reluctant to feed on N. spumigena. Essential isoleucine received most of the diazotroph nitrogen in field zooplankton, while nonessential amino acids received most 15N tracer in E. affinis. N. spumigena was clearly an important amino acid nitrogen source for Baltic Sea zooplankton. [ABSTRACT FROM AUTHOR]