Experiment on inter- and intra-specific phenotypic plasticity of three phytoplankton species in response to ocean acidification

Phenotypic plasticity describes the phenotypic adjustment of the same genotype to different environmental conditions and is best described by a reaction norm. We focus on the effect of ocean acidification (OA) on inter - and intraspecific reaction norms of three globally important phytoplankton species (Emiliania huxleyi, Gephyrocapsa oceanica, Chaetoceros affinis). Despite significant differences in growth rates between the species, they all showed a high potential for phenotypic buffering (no significant difference in growth rates between ambient and high CO2 condition). Only three coccolithophore genotypes showed a reduced growth in high CO2. Largely diverging responses to high CO2 of single coc-colithophore genotypes compared to the respective mean species responses, however, raise the question if an extrapolation to the population level is possible from single genotype experiments. We therefore compared the mean response of all tested genotypes to a total species response comprising the same genotypes, which was not significantly different in the coccolithophores. Assessing species reac-tion norm to different environmental conditions on short time scale in a genotype-mix could thus reduce sampling effort while increasing predictive power.

(Table 1) Chemical composition of volcanic rocks from the Mariisky sequence and extrusive vent rocks of the Mary Cape, Schmidt Peninsula (North Sakhalin)

Early Cretaceous volcanic rocks of the Mariisky sequence and Early Cenozoic extrusive-vent rocks of the Mary Cape are exposed at the most northwest of the Schmidt Peninsula, North Sakhalin. In chemical composition, all the rocks are subdivided into four groups. Three groups include volcanic rocks of the Mariisky sequence, which consists, from bottom to top, of calc-alkaline rocks, transitional calc-alkaline-tholeiite rocks, and incompatible element-depleted tholeiites. These rocks show subduction geochemical signatures and are considered as a fragment of the Moneron-Samarga island arc system. Trace-element modeling indicates their derivation through successive melting of a garnet-bearing mantle and garnet-free shallower mantle sources containing amphibole; pyroxene; and, possibly, spinel. The mixed subduction and intra-plate characteristics of the extrusive vent rocks of the Mary Cape attest to their formation in a transform continental margin setting.

Temperatures, salinity, and stable oxygen isotope values of bottom water and of the mollusc shell Arctica islandica from Nantucket Shoals

The carbonate shell of the bivalve Arctica islandica has been recognized, for more than a decade, as a potentially important marine geochemical biorecorder owing to this species' great longevity (200+ years) and wide geographic distribution throughout the northern North Atlantic Ocean, a region vital to global climate and ocean circulation. However, until now this potential has not been realized owing to the difficulty of precisely sampling the shell of this slow growing species. Using newly available automated microsampling techniques combined with micromass stable isotope mass spectrometry, a stable oxygen isotope record (1956-1957 and 1961-1970) has been obtained from a live-captured, 38-year-old A. islandica specimen collected near the former position of the Nantucket Shoals Lightship (41°N. 69°W). The shell's delta18O signal is compared with an expected signal derived from ambient bottom temperature and salinity data recorded at the lightship for the same period. The results show that A islandica's delta18O record (1) is in phase with its growth banding, confirming the annual periodicity of this species' growth bands, (2) is in oxygen isotopic equilibrium with the ambient seawater, (3) shows a consistent shell growth shutdown temperature of ~6°C. which translates into an ~8-month (May-December) shell growth period at this location, and (4) records the ambient bottom temperature with a precision of ~ +/-1.2°C. These results add important information on the life history of this commercially important shellfish species and demonstrate that A. islandica shells can be used to reconstruct inter- and intra-annual records of the continental shelf bottom temperature.

Fatty acid and stable isotope composition of calanoid copepods in the open eastern Atlantic Ocean during POLARSTERN cruise ANT-XXIX/1

The majority of global ocean production and total export production is attributed to oligotrophic oceanic regions due to their vast regional expanse. However, energy transfers, food-web structures and trophic relationships in these areas remain largely unknown. Regional and vertical inter- and intra-specific differences in trophic interactions and dietary preferences of calanoid copepods were investigated in four different regions in the open eastern Atlantic Ocean (38°N to 21°S) in October/November 2012 using a combination of fatty acid (FA) and stable isotope (SI) analyses. Mean carnivory indices (CI) based on FA trophic markers generally agreed with trophic positions (TP) derived from d15N analysis. Most copepods were classified as omnivorous (CI ~0.5, TP 1.8 to ~2.5) or carnivorous (CI >=0.7, TP >=2.9). Herbivorous copepods showed typical CIs of <=0.3. Geographical differences in d15N values of epi- (200-0 m) to mesopelagic (1000-200 m) copepods reflected corresponding spatial differences in baseline d15N of particulate organic matter from the upper 100 m. In contrast, species restricted to lower meso- and bathypelagic (2000-1000 m) layers did not show this regional trend. FA compositions were species-specific without distinct intra-specific vertical or spatial variations. Differences were only observed in the southernmost region influenced by the highly productive Benguela Current. Apparently, food availability and dietary composition were widely homogeneous throughout the oligotrophic oceanic regions of the tropical and subtropical Atlantic. Four major species clusters were identified by principal component analysis based on FA compositions. Vertically migrating species clustered with epi- to mesopelagic, non-migrating species, of which only Neocalanus gracilis was moderately enriched in lipids with 16% of dry mass (DM) and stored wax esters (WE) with 37% of total lipid (TL). All other species of this cluster had low lipid contents (< 10% DM) without WE. Of these, the tropical epipelagic Undinula vulgaris showed highest portions of bacterial markers. Rhincalanus cornutus, R. nasutus and Calanoides carinatus formed three separate clusters with species-specific lipid profiles, high lipid contents (>=41% DM), mainly accumulated as WE (>=79% TL). C. carinatus and R. nasutus were primarily herbivorous with almost no bacterial input. Despite deviating feeding strategies, R. nasutus clustered with deep-dwelling, carnivorous species, which had high amounts of lipids (>=37% DM) and WE (>=54% TL). Tropical and subtropical calanoid copepods exhibited a wide variety of life strategies, characterized by specialized feeding. This allows them, together with vertical habitat partitioning, to maintain high abundance and diversity in tropical oligotrophic open oceans, where they play an essential role in the energy flux and carbon cycling.

Multi-temporal mapping of seagrass cover, species and biomass of the Eastern Banks, Moreton Bay, Australia, with links to shapefiles

The spatial and temporal dynamics of seagrasses have been studied from the leaf to patch (100 m**2) scales. However, landscape scale (> 100 km**2) seagrass population dynamics are unresolved in seagrass ecology. Previous remote sensing approaches have lacked the temporal or spatial resolution, or ecologically appropriate mapping, to fully address this issue. This paper presents a robust, semi-automated object-based image analysis approach for mapping dominant seagrass species, percentage cover and above ground biomass using a time series of field data and coincident high spatial resolution satellite imagery. The study area was a 142 km**2 shallow, clear water seagrass habitat (the Eastern Banks, Moreton Bay, Australia). Nine data sets acquired between 2004 and 2013 were used to create seagrass species and percentage cover maps through the integration of seagrass photo transect field data, and atmospherically and geometrically corrected high spatial resolution satellite image data (WorldView-2, IKONOS and Quickbird-2) using an object based image analysis approach. Biomass maps were derived using empirical models trained with in-situ above ground biomass data per seagrass species. Maps and summary plots identified inter- and intra-annual variation of seagrass species composition, percentage cover level and above ground biomass. The methods provide a rigorous approach for field and image data collection and pre-processing, a semi-automated approach to extract seagrass species and cover maps and assess accuracy, and the subsequent empirical modelling of seagrass biomass. The resultant maps provide a fundamental data set for understanding landscape scale seagrass dynamics in a shallow water environment. Our findings provide proof of concept for the use of time-series analysis of remotely sensed seagrass products for use in seagrass ecology and management.

Stable carbon and oxygen isotope record of IODP Site 306-U1313, MIS 23-19

Since the seminal work by Hays et al. (1976), a plethora of studies has demonstrated a correlation between orbital variations and climatic change. However, information on how changes in orbital boundary conditions affected the frequency and amplitude of millennial-scale climate variability is still fragmentary. The Marine Isotope Stage (MIS) 19, an interglacial centred at around 785 ka, provides an opportunity to pursue this question and test the hypothesis that the long-term processes set up the boundary conditions within which the short-term processes operate. Similarly to the current interglacial, MIS 19 is characterised by a minimum of the 400-kyr eccentricity cycle, subdued amplitude of precessional changes, and small amplitude variations in insolation. Here we examine the record of climatic conditions during MIS 19 using high-resolution stable isotope records from benthic and planktonic foraminifera from a sedimentary sequence in the North Atlantic (Integrated Ocean Drilling Program Expedition 306, Site U1313) in order to assess the stability and duration of this interglacial, and evaluate the climate system's response in the millennial band to known orbitally induced insolation changes. Benthic and planktonic foraminiferal d18O values indicate relatively stable conditions during the peak warmth of MIS 19, but sea-surface and deep-water reconstructions start diverging during the transition towards the glacial MIS 18, when large, cold excursions disrupt the surface waters whereas low amplitude millennial scale fluctuations persist in the deep waters as recorded by the oxygen isotope signal. The glacial inception occurred at ~779 ka, in agreement with an increased abundance of tetra-unsaturated alkenones, reflecting the influence of icebergs and associated meltwater pulses and high-latitude waters at the study site. After having combined the new results with previous data from the same site, and using a variety of time series analysis techniques, we evaluate the evolution of millennial climate variability in response to changing orbital boundary conditions during the Early-Middle Pleistocene. Suborbital variability in both surface- and deep-water records is mainly concentrated at a period of ~11 kyr and, additionally, at ~5.8 and ~3.9 kyr in the deep ocean; these periods are equal to harmonics of precession band oscillations. The fact that the response at the 11 kyr period increased over the same interval during which the amplitude of the response to the precessional cycle increased supports the notion that most of the variance in the 11 kyr band in the sedimentary record is nonlinearly transferred from precession band oscillations. Considering that these periodicities are important features in the equatorial and intertropical insolation, these observations are in line with the view that the low-latitude regions play an important role in the response of the climate system to the astronomical forcing. We conclude that the effect of the orbitally induced insolation is of fundamental importance in regulating the timing and amplitude of millennial scale climate variability.

Colimitation of the unicellular photosynthetic diazotroph Crocosphaera watsonii by phosphorus, light, and carbon dioxide

We describe interactive effects of total phosphorus (total P = 0.1-4.0 µmol/L; added as H2NaPO4), irradiance (40 and 150 µmol quanta/m**2/s), and the partial pressure of carbon dioxide (P-CO2; 19 and 81 Pa, i.e., 190 and 800 ppm) on growth and CO2- and dinitrogen (N-2)-fixation rates of the unicellular N-2-fixing cyanobacterium Crocosphaera watsonii (WH0003) isolated from the Pacific Ocean near Hawaii. In semicontinuous cultures of C. watsonii, elevated P-CO2 positively affected growth and CO2- and N-2-fixation rates under high light. Under low light, elevated P-CO2 positively affected growth rates at all concentrations of P, but CO2- and N-2-fixation rates were affected by elevated P-CO2 only when P was low. In both high-light and low-light cultures, the total P requirements for growth and CO2- and N-2-fixation declined as P-CO2 increased. The minimum concentration (C-min) of total P and half-saturation constant (K-1/2) for growth and CO2- and N-2-fixation rates with respect to total P were reduced by 0.05 µmol/L as a function of elevated P-CO2. We speculate that low P requirements under high P-CO2 resulted from a lower energy demand associated with carbon-concentrating mechanisms in comparison with low-P-CO2 cultures. There was also a 0.10 µmol/L increase in C-min and K-1/2 for growth and N-2 fixation with respect to total P as a function of increasing light regardless of P-CO2 concentration. We speculate that cellular P concentrations are responsible for this shift through biodilution of cellular P and possibly cellular P uptake systems as a function of increasing light. Changing concentrations of P, CO2, and light have both positive and negative interactive effects on growth and CO2-, and N-2-fixation rates of unicellular oxygenic diazotrophs like C. watsonii.

Seawater carbonate chemistry and particulate organic particles during a semicontinuous batch culture experiment with Trichodesmium IMS101, 2007

Diazotrophic (N2-fixing) cyanobacteria provide the biological source of new nitrogen for large parts of the ocean. However, little is known about their sensitivity to global change. Here we show that the single most important nitrogen fixer in today's ocean, Trichodesmium, is strongly affected by changes in CO2 concentrations. Cell division rate doubled with rising CO2 (glacial to projected year 2100 levels) prompting lower carbon, nitrogen and phosphorus cellular contents, and reduced cell dimensions. N2 fixation rates per unit of phosphorus utilization as well as C:P and N:P ratios more than doubled at high CO2, with no change in C:N ratios. This could enhance the productivity of N-limited oligotrophic oceans, drive some of these areas into P limitation, and increase biological carbon sequestration in the ocean. The observed CO2 sensitivity of Trichodesmium could thereby provide a strong negative feedback to atmospheric CO2 increase.

Direct linkage between dimethyl sulfide production and microzooplankton grazing, resulting from prey composition change under high partial pressure of carbon dioxide conditions

Oceanic dimethyl sulfide (DMS) is the enzymatic cleavage product of the algal metabolite dimethylsulfoniopropionate (DMSP) and is the most abundant form of sulfur released into the atmosphere. To investigate the effects of two emerging environmental threats (ocean acidification and warming) on marine DMS production, we performed a large-scale perturbation experiment in a coastal environment. At both ambient temperature and 2 °C warmer, an increase in partial pressure of carbon dioxide (pCO2) in seawater (160-830 ppmv pCO2) favored the growth of large diatoms, which outcompeted other phytoplankton species in a natural phytoplankton assemblage and reduced the growth rate of smaller, DMSP-rich phototrophic dinoflagellates. This decreased the grazing rate of heterotrophic dinoflagellates (ubiquitous micrograzers), resulting in reduced DMS production via grazing activity. Both the magnitude and sign of the effect of pCO2 on possible future oceanic DMS production were strongly linked to pCO2-induced alterations to the phytoplankton community and the cellular DMSP content of the dominant species and its association with micrograzers.

(Table S1) Trace element and isotopic compositions of anhydrites sampled in the Manus Basin, Papua New Guinea

Microchemical analyses of rare earth element (REE) concentrations and Sr and S isotope ratios of anhydrite are used to identify sub-seafloor processes governing the formation of hydrothermal fluids in the convergent margin Manus Basin, Papua New Guinea. Samples comprise drill-core vein anhydrite and seafloor massive anhydrite from the PACMANUS (Roman Ruins, Snowcap and Fenway) and SuSu Knolls (North Su) active hydrothermal fields. Chondrite-normalized REE patterns in anhydrite show remarkable heterogeneity on the scale of individual grains, different from the near uniform REEN patterns measured in anhydrite from mid-ocean ridge deposits. The REEN patterns in anhydrite are correlated with REE distributions measured in hydrothermal fluids venting at the seafloor at these vent fields and are interpreted to record episodes of hydrothermal fluid formation affected by magmatic volatile degassing. 87Sr/86Sr ratios vary dramatically within individual grains between that of contemporary seawater and that of endmember hydrothermal fluid. Anhydrite was precipitated from a highly variable mixture of the two. The intra-grain heterogeneity implies that anhydrite preserves periods of contrasting hydrothermal versus seawater dominant near-seafloor fluid circulation. Most sulfate d34S values of anhydrite cluster around that of contemporary seawater, consistent with anhydrite precipitating from hydrothermal fluid mixed with locally entrained seawater. Sulfate d34S isotope ratios in some anhydrites are, however, lighter than that of seawater, which are interpreted as recording a source of sulfate derived from magmatic SO2 degassed from underlying felsic magmas in the Manus Basin. The range of elemental and isotopic signatures observed in anhydrite records a range of sub-seafloor processes including high-temperature hydrothermal fluid circulation, varying extents of magmatic volatile degassing, seawater entrainment and fluid mixing. The chemical and isotopic heterogeneity recorded in anhydrite at the inter- and intra-grain scale captures the dynamics of hydrothermal fluid formation and sub-seafloor circulation that is highly variable both spatially and temporally on timescales over which hydrothermal deposits are formed. Microchemical analysis of hydrothermal minerals can provide information about the temporal history of submarine hydrothermal systems that are variable over time and cannot necessarily be inferred only from the study of vent fluids.