Description: Nitrogen fixation is a key source of nitrogen in the Baltic Sea which counteracts nitrogen loss processes in the deep anoxic basins. Laboratory and field studies have indicated that single-strain nitrogen-fixing (diazotrophic) cyanobacteria from the Baltic Sea are sensitive to ocean acidification and warming, two drivers of marked future change in the marine environment. Here, we enclosed a natural plankton community in twelve indoor mesocosms (volume ~1400 L) and manipulated pCO2 to yield six CO2 treatments with two different temperature treatments (16.6°C and 22.4°C, pCO2 range = 360 – 2030 μatm). We followed the filamentous, heterocystous diazotrophic cyanobacteria community (Nostocales, primarily Nodularia spumigena) over four weeks. Our results indicate that heterocystous diazotrophic cyanobacteria may become less competitive in natural plankton communities under ocean acidification. Elevated CO2 had a negative impact on Nodularia sp. biomass, which was exacerbated by warming. Our results imply that Nodularia sp. may contribute less to new nitrogen inputs in the Baltic Sea in future.

Description: Highlights: • Ocean acidification increases phytoplankton standing stock. • This increase is more pronounced in smaller-sized taxa. • Primary consumers reac differently depending on nutrient availability. • Bacteria and micro-heterotrophs benefited under limiting conditions. • In general, heterotrophs are negatively affected at nutrient replete periods. Abstract: In situ mesocosm experiments on the effect of ocean acidification (OA) are an important tool for investigating potential OA-induced changes in natural plankton communities. In this study we combined results from various in-situ mesocosm studies in two different ocean regions (Arctic and temperate waters) to reveal general patterns of plankton community shifts in response to OA and how these changes are modulated by inorganic nutrient availability. Overall, simulated OA caused an increase in phytoplankton standing stock, which was more pronounced in smaller-sized taxa. This effect on primary producers was channelled differently into heterotroph primary consumers depending on the inorganic nutrient availability. Under limiting conditions, bacteria and micro-heterotrophs benefited with inconsistent responses of larger heterotrophs. During nutrient replete periods, heterotrophs were in general negatively affected, although there was an increase of some mesozooplankton developmental stages (i.e. copepodites). We hypothesize that changes in phytoplankton size distribution and community composition could be responsible for these food web responses.

Description: Highlights • Increasing influence of multiple environmental drivers produces changes on the temporal variability of species. • The intensity and hierarchy of drivers acting upon organisms within alternative regimes of variability may differ. • We identified regimes of variability of phytoplankton and depicted cascading effects of multiple drivers in each period. • The number of factors driving the response of phytoplankton increased along time and produced the erosion of productivity patterns. • The hierarchy and interactions of drivers changed over time, revealing that management policies require constant update. Abstract Estuaries are among the most valuable aquatic systems by their services to human welfare. However, increasing human activities at the watershed along with the pressure of climate change are fostering the co-occurrence of multiple environmental drivers, and warn of potential negative impacts on estuaries resources. At present, no clear understanding of how coastal ecosystems will respond to the non-stationary effect of multiple drivers. Here we analysed the temporal interaction among multiple environmental drivers and their changing priority on shaping phytoplankton response in the Bahía Blanca Estuary, SW Atlantic Ocean. The interaction among environmental drivers and the number of significant direct and indirect effects on chlorophyll concentration increased over time in concurrence with enhanced anthropogenic stress, changing winter climate and wind patterns. Over the period 1978–1993, proximal variables such as nutrients, water temperature and salinity, showed a dominant effect on chlorophyll, whereas in more recent years (1993–2009) climate signals (SAM and ENSO) boosted indirect effects through its influence on precipitation, wind, water temperature and turbidity. Turbidity emerged as the dominant driver of chlorophyll while in recent years acted synergistically with the concentration of dissolved nitrogen. As a result, chlorophyll concentration showed a significant negative trend and a loss of seasonal peaks reflecting a pronounced reorganisation of the phytoplankton community. We stress the need to account for the changing priority of drivers to understand, and eventually forecast, biological responses under projected scenarios of global anthropogenic change.

Description: The Red Sea features a natural environmental gradient characterized by increasing water temperature, nutrient and chlorophyll a concentrations from North to South. The aim of this study was to assess the relationships between ecohydrography, particulate organic matter (POM) and coral reef biota that are poorly understood by means of carbon (δ13C) and nitrogen (δ15N) stable isotopes. Herbivorous, planktivorous and carnivorous fishes, zooplankton, soft corals (Alcyonidae), and bivalves (Tridacna squamosa)were a priori defined as biota guilds. Environmental samples (nutrients, chlorophyll a), oceanographic data (salinity, temperature), POMand biotawere collected at eight coral reefs between 28°31′ N and 16°31′ N. Isotopic niches of guilds separated in δ13C and δ15N isotopic niche spaces and were significantly correlated with environmental factors at latitudinal scale. Dietary end member contributionswere estimated using the Bayesian isotope mixingmodel SIAR. POMand zooplankton 15N enrichment suggested influences by urban run-off in the industrialized central region of the Red Sea. Both δ15N and their relative trophic positions (RTPs) tend to increase southwards, but urban runoff offsets the natural environmental gradient in the central region of the Red Sea toward higher δ15N and RTPs. The present study reveals that consumer δ13C and δ15N in Red Sea coral reefs are influenced primarily by the latitudinal environmental gradient and localized urban runoff. This study illustrates the importance of ecohydrography when interpreting trophic relationships from stable isotopes in Red Sea coral reefs.

Description: Highlights: • We used indoor mesocosms to test the impact of warming on plankton communities. • Different stages of phytoplankton bloom were analysed. • Increased temperature and zooplankton grazing had similar effects on phytoplankton. • Warming and increased zooplankton density decreased phytoplankton richness. • Warming and increased zooplankton density increased phytoplankton evenness. Recent climate warming is expected to affect phytoplankton biomass and diversity in marine ecosystems. Temperature can act directly on phytoplankton (e.g. rendering physiological processes) or indirectly due to changes in zooplankton grazing activity. We tested experimentally the impact of increased temperature on natural phytoplankton and zooplankton communities using indoor mesocosms and combined the results from different experimental years applying a meta-analytic approach. We divided our analysis into three bloom phases to define the strength of temperature and zooplankton impacts on phytoplankton in different stages of bloom development. Within the constraints of an experiment, our results suggest that increased temperature and zooplankton grazing have similar effects on phytoplankton diversity, which are most apparent in the post-bloom phase, when zooplankton abundances reach the highest values. Moreover, we observed changes in zooplankton composition in response to warming and initial conditions, which can additionally affect phytoplankton diversity, because changing feeding preferences of zooplankton can affect phytoplankton community structure. We conclude that phytoplankton diversity is indirectly affected by temperature in the post-bloom phase through changing zooplankton composition and grazing activities. Before and during the bloom, however, these effects seem to be overruled by temperature enhanced bottom-up processes such as phytoplankton nutrient uptake.

Description: Simultaneous analysis of carbon, nitrogen and sulphur stable isotope ratios was applied in this pilot study to examine the food web of a Zostera marina L. system in the western Baltic Sea. Samples of three potential food sources: eelgrass, epiphytic algae and seston, as well as 69 consumer species were collected during the growing season of Z. marina from March to September 2011. The measured δ13C values of epiphytes (-14.1‰ ± 1.8 SD) were close to δ13C values of eelgrass (-11.6‰ ± 1.8 SD), impeding a clear distinction of those two carbon sources, whereas seston δ13C values (-20.9‰ ± 3.5 SD) were clearly different. This frequently encountered problem was solved by the additional use of δ34S, which resulted in easily distinguishable values for sediment and seawater derived sulphur. Values of primary producer δ34S ranged from 5.6‰ (± 2.3 SD) for Z. marina leaves to 14.2‰ (± 1.6 SD) for epiphytes and 11.9‰ (± 3.3 SD) for seston. The combination of δ34S and δ13C values made a separation of carbon sources possible and enabled the allocation of potential food sources to consumers and a description of their trophic relationships. The data of stable isotope ratio analysis of this eelgrass community strongly indicate a food web based on epiphyte and seston production. δ15N values show a food web consisting of large numbers of generalists and a high degree of omnivory amongst the consumer species analysed. This implies an occupation of every trophic position possible, which is supported by a continuous distribution of δ15N values. Previously described eelgrass food webs may have to be re-evaluated to include sulfur in order to provide a clear picture on primary carbon sources.

Description: The combined effects of warming and overwintering copepod densities on the spring succession of Baltic Sea plankton were investigated using indoor mesocosms. Three zooplankton (1.5, 4 and 10 copepods L-1) and two temperature levels called ∆0°C and ∆6°C (0°C and 6°C above the present day temperature scenario for Kiel Bight) were chosen. Both, the timing and the duration of the protozooplankton (PZP) bloom were significantly affected by temperature, but not by copepod density. In contrast, the bloom intensity of PZP was highly affected by the factors temperature and copepod density and its interaction. This suggests that at elevated temperature conditions PZP grows faster but, at the same time, are subject to higher top-down control by copepods. At low temperatures and low copepod densities, PZP in turn fully escaped from copepod predation. Further changes in the overwintering copepod densities resulted in a strong ciliate suppression of which small-sized ciliates (〈30 µm) were especially vulnerable to copepod predation while other PZP size classes remained unaffected. In conclusion, the results presented point at a pivotal regulating role of overwintering copepods under future warming condition. Further, warming was shown to cause a distinct match between phytoplankton and PZP thus strengthening trophic pathways through PZP. Our findings are discussed in the context of the ‘trophic link-sink’ debate by considering potential alterations in the flux of matter and energy up the food web.

Description: Inorganic dissolved macronutrient (nitrogen, N, and phosphorus, P) supply to surface waters in the eastern tropical South Pacific is influenced by expanding oxygen minimum zones, since N loss occurs due to microbial processes under anoxic conditions while P is increasingly released from the shelf sediments. To investigate the impact of decreasing N:P supply ratios in the Peruvian Upwelling, we conducted nutrient manipulation experiments using a shipboard mesocosm setup with a natural phytoplankton community. In a first experiment, either N or P or no nutrients were added with mesozooplankton present or absent. In a second experiment, initial nutrient concentrations were adjusted to four N:P ratios ranging from 2.5 to 16 using two "high N" and two "high P" levels in combination (i.e., +N, +P, +N and P, no addition). Over six and seven days, respectively, microalgal biomass development as well as nutrient uptake was monitored. Phytoplankton biomass strongly responded to N addition, in both mesozooplankton-grazed and not grazed treatments. The developing diatom bloom in the "high N" exceeded that in the "low N" treatments by a factor of two. No modulation of the total biomass by P-addition was observed, however, species-specific responses were more variable. Notably, some organisms were able to benefit from low N:P fertilization ratios, especially Heterosigma sp. and Phaeocystis globosa which are notorious for forming blooms that are toxic or inadequate for mesozooplankton nutrition. After the decline of the diatom bloom, the relative contribution of unsaturated fatty acid to the lipid content of seston was positively correlated to diatom biomass in the peak bloom, indicating that positive effects of diatom blooms on food quality of the protist community to higher trophic levels remain even after the phytoplankton biomass was incorporated by grazers. Our results indicate an overall N-limitation of the system, especially in the case of dominating diatoms, which were able to immediately utilize the available nitrate (within two days) and develop a biomass maximum within three days of incubation. After the decline of diatom biomass, detection of the cyanobacterial marker pigment aphanizophyll indicated the occurrence of diazotrophs, especially in those enclosures initially provided with high N supply. This was surprising, as diazotrophs are thought to play a role in compensating to some extent the N deficit above OMZs in the succession of phytoplankton after an upwelling event

Description: The Eastern Tropical North Atlantic (ETNA) is characterised by a strong east to west gradient in the vertical upward flux of dissolved inorganic nitrogen to the photic zone. We measured the stable nitrogen isotope (δ15N) signatures of various zooplankton taxa covering twelve stations in the ETNA (04°–14°N, 016–030°W) in fall 2009, and observed significant differences in δ15N values among stations. These spatial differences in δ15N within zooplankton taxa exceeded those between trophic levels and revealed an increasing atmospheric input of nitrogen by N2 fixation and Aeolian dust in the open ocean as opposed to remineralised NO3− close to the NW African upwelling. In order to investigate the spatial distribution of upwelling-fuelled versus atmospheric-derived nitrogen more closely, we examined the δ15N signatures in size-fractionated zooplankton as well as in three widely distributed epipelagic copepod species on a second cruise in fall 2010 in the ETNA (02-17°35′N, 015–028°W). Copepods were sampled for δ15N and RNA/DNA as a proxy for nutritional condition on 25 stations. At the same stations, vertical profiles of chlorophyll-a and dissolved nutrients were obtained. High standing stocks of chl-a were associated with shallow mixed layer depth and thickening of the nutricline. As the nitracline was generally deeper and less thick than the phosphacline, it appears that non-diazotroph primary production was limited by N rather than P throughout the study area, which is in line with enrichment experiments during these cruises. Estimated by the δ15N in zooplankton, atmospheric sources of new N contributed less than 20% close to the African coast and in the Guinea Dome area and up to 60% at the offshore stations, depending on the depth of the nitracline. δ15N of the three different copepod species investigated strongly correlated with each other, in spite of their distinct feeding ecology, which resulted in different spatial patterns of nutritional condition as indicated by RNA/DNA. Highlights: ► We studied δ15N and RNA/DNA of eastern tropical Atlantic zooplankton along with nutrients and Chl-α. ► Zooplankton −δ15N was decreasing from east (West African Shelf) to west (oligotrophic open ocean). ► Total integrated Chl-a depended mainly on nutricline depth and was N-limited throughout the area. ► Zooplankton δ15N and nutricline depth were used to estimate atmospheric N sources to the food web. ► Estimated atmospheric nitrogen sources were less than 20% at the shelf slope and up to 60% offshore.

Description: Changing seawater chemistry towards reduced pH as a result of increasing atmospheric carbon dioxide (CO2) is affecting oceanic organisms, particularly calcifying species. Responses of non-calcifying consumers are highly variable and mainly mediated through indirect ocean acidification effects induced by changing the biochemical content of their prey, as shown within single species and simple 2-trophic level systems. However, it can be expected that indirect CO2 impacts observed at the single species level are compensated at the ecosystem level by species richness and complex trophic interactions. A dampening of CO2-effects can be further expected for coastal communities adapted to strong natural fluctuations in pCO2, typical for productive coastal habitats. Here we show that a plankton community of the Kiel Fjord was tolerant to CO2 partial pressure (pCO2) levels projected for the end of this century (〈1400 µatm), and only subtle differences were observed at the extremely high value of 4000 µatm. We found similar phyto- and microzooplankton biomass and copepod abundance and egg production across all CO2 treatment levels. Stoichiometric phytoplankton food quality was minimally different at the highest pCO2 treatment, but was far from being potentially limiting for copepods. These results are in contrast to studies that include only a single species, which observe strong indirect CO2 effects for herbivores and suggest limitations of biological responses at the level of organism to community. Although this coastal plankton community was highly tolerant to high fluctuations in pCO2, increase in hypoxia and CO2 uptake by the ocean can aggravate acidification and may lead to pH changes outside the range presently experienced by coastal organisms.