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: Allelopathic species can alter biodiversity. Using simulated assemblages that are characterised by neutrality, lumpy coexistence and intransitivity, we explore relationships between within-assemblage competitive dissimilarities and resistance to allelopathic species. An emergent behaviour from our models is that assemblages are more resistant to allelopathy when members strongly compete exploitatively (high competitive power). We found that neutral assemblages were the most vulnerable to allelopathic species, followed by lumpy and then by intransitive assemblages. We find support for our modeling in real-world time-series data from eight lakes of varied morphometry and trophic state. Our analysis of this data shows that a lake’s history of allelopathic phytoplankton species biovolume density and dominance is related to the number of species clusters occurring in the plankton assemblages of those lakes, an emergent trend similar to that of our modeling. We suggest that an assemblage’s competitive power determines its allelopathy resistance.

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: Global warming is assumed to alter the trophic interactions and carbon flow patterns of aquatic food webs. The impact of temperature on phyto-bacterioplankton coupling and bacterial community composition (BCC) was the focus of the present study, in which an indoor mesocosm experiment with natural plankton communities from the western Baltic Sea was conducted. A 6°C increase in water temperature resulted, as predicted, in tighter coupling between the diatom-dominated phytoplankton and heterotrophic bacteria, accompanied by a strong increase in carbon flow into bacterioplankton during the phytoplankton bloom phase. Suppressed bacterial development at cold in situ temperatures probably reflected lowered bacterial production and grazing by protists, as the latter were less affected by low temperatures. BCC was strongly influenced by the phytoplankton bloom stage and to a lesser extent by temperature. Under both temperature regimes, Gammaproteobacteria clearly dominated during the phytoplankton peak, with Glaciecola sp. as the single most abundant taxon. However, warming induced the appearance of additional bacterial taxa belonging to Betaproteobacteria and Bacteroidetes. Our results show that warming during an early phytoplankton bloom causes a shift towards a more heterotrophic system, with the appearance of new bacterial taxa suggesting a potential for utilization of a broader substrate spectrum.

Description: An integral concept of ecological research is the constraint of biodiversity along latitudinal and environmental gradients. The Red Sea features a natural example of a latitudinal gradient of salinity, temperature and nutrient richness. Coral reefs along the Red Sea coasts are supported with allochthonous resources such as oceanic and neritic phytoplankton and zooplankton; however, relatively little is known about how the ecohydrography correlates with plankton biodiversity and abundance. In this article we present the biodiversity of phytoplankton and zooplankton in Red Sea coral reefs. Oceanographic data (temperature, salinity), water samples for nutrient analysis, particulate organic matter, phytoplankton and zooplankton, the latter with special reference to Copepoda (Crustacea), were collected at nine coral reefs over ~1500 km distance along the Red Sea coast of Saudi Arabia. The trophic state of ambient waters [as indicated by chlorophyll a (Chl a)] changed from strong oligotrophy in the north to mesotrophy in the south and was associated with increasing biomasses of Bacillariophyceae, picoeukaryotes and Synechococcus as indicated by pigment fingerprinting (CHEMTAX) and flow cytometry. Net-phytoplankton microscopy revealed a Trichodesmium erythraeum (Cyanobacteria) bloom north of the Farasan Islands. Several potentially harmful algae, including Dinophysis miles and Gonyaulax spinifera (Dinophyceae), were encountered in larger numbers in the vicinity of the aquaculture facilities at Al Lith. Changes in zooplankton abundance were mainly correlated to the phytoplankton biomass following the latitudinal gradient. The largest zooplankton abundance was observed at the Farasan Archipelago, despite high abundances of copepodites, veligers (Gastropoda larvae) and Chaetognatha at Al Lith. Although the community composition changed over latitude, biodiversity indices of phytoplankton and zooplankton did not exhibit a systematic pattern. As this study constitutes the first current account of the plankton biodiversity in Red Sea coral reefs at a large spatial scale, the results will be informative for ecosystem-based management along the coastline of Saudi Arabia.

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: 1. Different components of the climate system have been shown to affect temporal dynamics in natural plankton communities on scales varying from days to years. The seasonal dynamics in temperate lake plankton communities, with emphasis on both physical and biological forcing factors, were captured in the 1980s in a conceptual framework, the Plankton Ecology Group (PEG) model. 2. Taking the PEG model as our starting point, we discuss anticipated changes in seasonal and long-term plankton dynamics and extend this model to other climate regions, particularly polar and tropical latitudes. Based on our improved post-PEG understanding of plankton dynamics, we also evaluate the role of microbial plankton, parasites and fish in governing plankton dynamics and distribution. 3. In polar lakes, there is usually just a single peak in plankton biomass in summer. Lengthening of the growing season under warmer conditions may lead to higher and more prolonged phytoplankton productivity. Climate-induced increases in nutrient loading in these oligotrophic waters may contribute to higher phytoplankton biomass and subsequent higher zooplankton and fish productivity. 4. In temperate lakes, a seasonal pattern with two plankton biomass peaks – in spring and summer – can shift to one with a single but longer and larger biomass peak as nutrient loading increases, with associated higher populations of zooplanktivorous fish. Climate change will exacerbate these trends by increasing nutrient loading through increased internal nutrient inputs (due to warming) and increased catchment inputs (in the case of more precipitation). 5. In tropical systems, temporal variability in precipitation can be an important driver of the seasonal development of plankton. Increases in precipitation intensity may reset the seasonal dynamics of plankton communities and favour species adapted to highly variable environments. The existing intense predation by fish on larger zooplankters may increase further, resulting in a perennially low zooplankton biomass. 6. Bacteria were not included in the original PEG model. Seasonally, bacteria vary less than the phytoplankton but often follow its patterns, particularly in colder lakes. In warmer lakes, and with future warming, a greater influx of allochthonous carbon may obscure this pattern. 7. Our analyses indicate that the consequences of climate change for plankton dynamics are, to a large extent, system specific, depending on characteristics such as food-web structure and nutrient loading. Indirect effects through nutrient loading may be more important than direct effects of temperature increase, especially for phytoplankton. However, with warming a general picture emerges of increases in bacterivory, greater cyanobacterial dominance and smaller-bodied zooplankton that are more heavily impacted by fish predation.

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.