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: 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: Highlights: • Incubation experiments showed an omnivore feeding of Calanus finmarchicus. • Ciliates were positively selected by C. finmarchicus at varying food concentrations. • The degree of omnivory depended on the proportion of ciliates available. Abstract: The feeding selectivity of Calanus finmarchicus was studied by carrying out three incubation experiments; two experiments with natural seawater sampled during spring bloom (Exp. 1) and post-bloom conditions (Exp. 2) and a third experiment with cultured dinoflagellates and ciliates (Exp. 3). In the first two experiments a gradient in ciliate concentration was created to investigate the potential for prey density dependent selective feeding of C. finmarchicus. Results of microplankton counts indicated C. finmarchicus to be omnivorous. Diatoms contributed chiefly to the diet during spring bloom conditions. Despite the high microphytoplankton biomass during the spring bloom (Exp. 1), ciliates were selected positively by C. finmarchicus when the ciliate biomass exceeded 6.5 μg C L− 1. A selection in favor of large conic ciliates such as Laboea sp. and Strombidium conicum was indicated by positive selectivity indices. Ciliates were throughout positively selected by C. finmarchicus during Exp. 2, and selectivity indices indicated a negative selection of diatoms. The results from Exp. 3 showed that C. finmarchicus has the ability to switch from dinoflagellates to ciliates as sole food source, even if the dinoflagellate was offered in surplus. This suggests that other factors, such as nutrition may be of significance for the feeding selectivity of C. finmarchicus.

Description: Highlights: • A full-scale 5 year experimental study of ecosystem responses to increased nutrients. • Concentrations of DIN and DIP did not respond positively to increased nutrient input. • Concentrations of PON and POP and phytoplankton biomass responded positively. • PON is suggested as credible indicator for chemical and ecological state. • A general scientific concept for managing nutrient input to coastal waters is presented. Abstract: The objective of this study was to quantify chemical and biological responses to an experimentally increased nutrient input to an open coastal planktonic ecosystem and to contribute to a scientific concept and credible indicators for managing nutrient supply to coastal waters. Data were derived in a 5 year fertilisation experiment of a tidal driven coastal lagoon at the outer coast off Central Norway (63°36’ N, 9°33’ E), with a surface area of 275.000 m2, volume of 5.5 mill m3, mean depth of 22 m and a water exchange rate of 0.19 day- 1. The lagoon was fertilised in the summer season 1998 and 1999, while summer seasons 1996-97 and 2000 and inflowing water were used as unfertilised references. Most measured chemical and biological variables showed linear responses with an increasing loading rate of inorganic N and P (LN and LP, respectively). PON, POP and POC (〈 200 μm) responded significantly (P 〈 0.05) as did chlorophyll a and phytoplankton C. DIN and DIP remained, however, constant and independent of LN and LP, respectively (P 〉 0.05) as did heterotrophic biomass (P 〉 0.05). We evaluate the response variables assuming a stepwise incorporation process of nutrients in the planktonic ecosystem and how that will interact with biological response times and water dilution rates. We suggest that PON is a credible indicator of both chemical and ecological states of the planktonic ecosystem and that natural background and upper critical concentrations are 46 and 88 mg PON m- 3, respectively. The study was supported by data from mesocosms. We discuss the scientific relevance of our suggestions, how results can be extrapolated to a broader geographical scale, and we propose a science-based concept for the management of nutrient emission to open coastal waters.

Description: Highlights: • The natural ecohydrographic gradient of the Red Sea translates into an isoscape. • The Red Sea isoscape features increasing zooplankton δ15 N values towards the South. • Isotopic baseline variations propagate through the pelagic food web. • Eddy-induced upwelling modifies the natural ecohydrographic North-South gradient. Abstract: Although zooplankton occupy key roles in aquatic biogeochemical cycles, little is known about the pelagic food web and trophodynamics of zooplankton in the Red Sea. Natural abundance stable isotope analysis (SIA) of carbon (δ13C) and N (δ15N) is one approach to elucidating pelagic food web structures and diet assimilation. Integrating the combined effects of ecological processes and hydrography, ecohydrographic features often translate into geographic patterns in δ13C and δ15N values at the base of food webs. This is due, for example, to divergent 15N abundances in source end-members (deep water sources: high δ15N, diazotrophs: low δ15N). Such patterns in the spatial distributions of stable isotope values were coined isoscapes. Empirical data of atmospheric, oceanographic, and biological processes, which drive the ecohydrographic gradients of the oligotrophic Red Sea, are under-explored and some rather anticipated than proven. Specifically, five processes underpin Red Sea gradients: (a) monsoon-related intrusions of nutrient-rich Indian Ocean water; (b) basin scale thermohaline circulation; (c) mesoscale eddy activity that causes up-welling of deep water nutrients into the upper layer; (d) the biological fixation of atmospheric nitrogen (N2) by diazotrophs; and (e) the deposition of dust and aerosol-derived N. This study assessed relationships between environmental samples (nutrients, chlorophyll a), oceanographic data (temperature, salinity, current velocity [ADCP]), particulate organic matter (POM), and net-phytoplankton, with the δ13C and δ15N values of zooplankton collected in spring 2012 from 16°28′ to 26°57′N along the central axis of the Red Sea. The δ15N of bulk POM and most zooplankton taxa increased from North (Duba) to South (Farasan). The potential contribution of deep water nutrient-fueled phytoplankton, POM, and diazotrophs varied among sites. Estimates suggested higher diazotroph contributions in the North, a greater contribution of POM in the South, and of small phytoplankton in the central Red Sea. Consistent variation across taxonomic and trophic groups at latitudinal scale, corresponding with patterns of nutrient stoichiometry and phytoplankton composition, indicates that the zooplankton ecology in the Red Sea is largely influenced by hydrographic features. It suggests that the primary ecohydrography of the Red Sea is driven not only by the thermohaline circulation, but also by mesoscale activities that transports nutrients to the upper water layers and interact with the general circulation pattern. Ecohydrographic features of the Red Sea, therefore, aid in explaining the observed configuration of its isoscape at the macroecological scale.

Description: Highlights: • We propose a new Phytoplankton Index for monitoring the ecological status of lakes. • We develop a community index integrating the response to anthropogenic pressure. • We link high frequency Secchi depth with low frequency phytoplankton monitoring. • We demonstrate the correlation of the index with land use on the watershed scale. • The index offers a cost-effective tool, practical for medium-qualified personnel. Abstract: We propose and test a new Phytoplankton Community Index (PhyCoI) for monitoring the ecological status of lakes and reservoirs. The design of our PhyCoI is based on the fact that phytoplankton biomass and community structure respond to changes in water quality (mainly eutrophication) and by themselves also influence water quality. In order to accommodate this double role of phytoplankton as indicator and impact, PhyCoI is based on phytoplankton community properties at different hierarchical levels combining both specific metrics (total biomass, taxonomic group biomass, cyanobacteria contribution, taxonomic group species richness) and new or modified sub-indices. It is calculated from the scores of the different metrics/sub-indices resulting in a final index value in the range from 0 to 5, to assess water quality on the basis of five ecological classes according to the Water Framework Directive (WFD). The test of PhyCoI was based on Carlson's Trophic State Index (TSISD) based on water transparency (Secchi depth) in 26 Greek lakes and reservoirs covering the entire spectrum from oligotrophic to hypertrophic. A highly significant relationship at p 〈 0.001 between the two variables was found, with the values of the PhyCoI declining with increasing TSISD. Furthermore, a significant relationship between the PhyCoI and land use types at the watershed of the studied freshwaters was found identifying permanent crops, pastures and shrubs and herbaceous vegetation associations as significant predictors of PhyCoI values. Because of the amount of labor involved in obtaining the PhyCoI we suggest to combine low frequency PhyCoI determinations with a high frequency Secchi depth measurements for practical monitoring purposes.