Upper Maestrichtian to Eocene benthic foraminifera in ODP Leg 121 holes

Late Maestrichtian to late Eocene bathyal benthic foraminiferal faunas at Sites 752,753, and 754 on Broken Ridge in the eastern Indian Ocean were analyzed as to their stratigraphic distribution of species to clarify the relation between faunal turnovers and paleoceanographic changes. Based on Q-mode factor analysis, eight varimax assemblages were distinguished: the Stensioina beccariiformis assemblage in the upper Maestrichtian to upper Paleocene; the Cibicidoides hyphalus assemblage in the upper Maestrichtian; the Cibicidoides cf. pseudoperlucidus assemblage in the upper Paleocene; the Anomalinoides capitatusldanicus assemblage in the uppermost Paleocene to lower Eocene; the Cibicidoides subspiratus assemblage in the lower Eocene; the Nuttallides truempyi assemblage in the lower and middle Eocene; the Osangularia sp. 1 - Hanzawaia ammophila assemblage in the upper Eocene; and the Lenticulina spp. assemblage in the uppermost Eocene, Oligocene, and lower Miocene. The presence of the Osangularia sp. 1 - Hanzawaia ammophila assemblage is related to the shallowing episode on Broken Ridge (upper bathyal), as a result of the rifting event that occurred in the middle Eocene. The most distinct faunal change (the disappearance of about 37% of the species) occurred between the S. beccariiformis assemblage and the A. capitatusldanicus assemblage, at the end of the upper Paleocene. A. capitatusldanicus, Lenticulina spp., and varied forms of Cibicidoides replaced the Velasco-type fauna at this time. The timing of this event is well correlated with the known age at South Atlantic sites (Thomas, 1990 doi:10.2973/odp.proc.sr.113.123.1990; Kennett and Stott, 1990 doi:10.2973/odp.proc.sr.113.188.1990; Katz and Miller, 1990 doi:10.2973/odp.proc.sr.114.147.1991). The primary cause of the extinction of the Stensioina beccariiformis assemblage is elusive, but may have resulted from the cessation of deep-water formation in the Antarctic (Katz and Miller, 1990), and subsequent arrival of warm saline deep water (Thomas, 1990; Kennett and Stott, 1990). Another possibility may be a weakened influence of high-salinity water formed at the low latitudes such as the Tethys Sea. The extinction event corresponds to the change from higher delta13C values in benthic foraminifers to lower ones. An interpretation of delta13C values is that the eastern Indian deep water, characterized by young and nutrient-depleted water, became old water which was devoid of a supply of new water during the latest Paleocene to early Eocene. Prior to this benthic event, signals of related faunal change were detected in the following short periods: early and late Paleocene, near the boundary of nannofossil Zone CP4, and Zone CP5 of the late Paleocene at Site 752. Among common taxa in the upper Maestrichtian, only seven species disappeared or became extinct at the Cretaceous/ Tertiary boundary at Site 752. The benthic foraminiferal population did not change for up to 2 m above the boundary, in contrast to the rapid decrease of the plankt onic foraminiferal population at the boundary. A decrease in the number of benthic foraminifers occurs after that level, corresponding to an interval of decreased numbers of planktonic foraminifers and higher abundance of volcanic ash. Reduced species diversity (H') suggests a secondary effect attributable to the dissolution of foraminiferal tests. The different responses of planktonic and benthic foraminifers to the event just above the boundary suggest that the Cretaceous/Tertiary event was a surface event as also suggested by Thomas (1990). In addition, a positive shift of delta13C in benthic foraminifers after the event indicates nutrient-depleted bottom water at Site 752.

Neogene benthic foraminifers from the Kerguelen Plateau

Benthic foraminifers were studied quantitatively in 120 lower Miocene through upper Pleistocene samples from Ocean Drilling Program Site 747 (Central Kerguelen Plateau) and Sites 748 and 751 (Southern Kerguelen Plateau). These sites are situated on an 450-km-long, north-south transect between 54°49'S and 58°26'S at present water depths between 1696 and 1288 m. Principal component analysis on the census data of the most abundant 92 taxa helped to identify 8 benthic foraminifer assemblages. These benthic foraminifer assemblages were compared with Holocene faunas from southern high latitudes to reconstruct paleoenvironmental conditions. Middle lower Miocene sediments are characterized by a Uvigerina hispidocostata assemblage, indicating high paleoproductivity and/or not well-ventilated bottom water. From late early to late middle Miocene time, the Southern Kerguelen Plateau was bathed by a young, well-oxygenated, and carbonate-aggressive water mass, as indicated by a Nuttallides umbonifer-dominated benthic foraminifer assemblage. During late middle Miocene time, an Astrononion pusillum assemblage took over for only about 1 m.y., probably indicating the first injection of an aged water mass, similar to the North Atlantic Deep Water (NADW), into a developing circumpolar current system. Around the middle to late Miocene boundary, the fauna again became dominated by N. umbonifer. After the last appearance of N. umbonifer, reestablishment of the A. pusillum assemblage from the early late through at least the late late Miocene, indicated the established influence of a NADW-like water mass. The latest Miocene through middle late Pliocene benthic foraminifer assemblage was characterized by Epistominella exigua and strong carbonate dissolution, indicating very high biosiliceous production, and this in turn may indicate the formation and paleoposition of an Antarctic Polar Frontal Zone. From the late late Pliocene, a Trifarina angulosa assemblage (indicative today of sandy substrate and vigorous bottom currents) strongly dominated the fauna up to the late Pleistocene, when Bulimina aculeata (indicative today of calm sedimentation with high organic matter fluxes) became an important and partly dominating constituent of the fauna. This is interpreted as the faunal response to the decreased winnowing force (bottom current velocities) of the Antarctic Circumpolar Current during periods of global climatic amelioration and raised sea level.

Experiment: Food availability outweighs ocean acidification effects in juvenile Mytilus edulis

Ocean acidification is expected to decrease calcification rates of bivalves. Nevertheless in many coastal areas high pCO2 variability is encountered already today. Kiel Fjord (Western Baltic Sea) is a brackish (12-20 g kg-1) and CO2 enriched habitat, but the blue mussel Mytilus edulis dominates the benthic community. In a coupled field and laboratory study we examined the annual pCO2 variability in this habitat and the combined effects of elevated pCO2 and food availability on juvenile M. edulis growth and calcification. In the laboratory experiment, mussel growth and calcification were found to chiefly depend on food supply, with only minor impacts of pCO2 up to 3350 µatm. Kiel Fjord was characterized by strong seasonal pCO2 variability. During summer, maximal pCO2 values of 2500 µatm were observed at the surface and >3000 µatm at the bottom. However, the field growth experiment revealed seven times higher growth and calcification rates of M. edulis at a high pCO2 inner fjord field station (mean pCO2 ca. 1000 µatm) in comparison to a low pCO2 outer fjord station (ca. 600 µatm). In addition, mussels were able to outcompete the barnacle Amphibalanus improvisus at the high pCO2 site. High mussel productivity at the inner fjord site was enabled by higher particulate organic carbon concentrations. Kiel Fjord is highly impacted by eutrophication, which causes bottom water hypoxia and consequently high seawater pCO2. At the same time, elevated nutrient concentrations increase the energy availability for filter feeding organisms such as mussels. Thus M. edulis can dominate over a seemingly more acidification resistant species such as A. improvisus. We conclude that benthic stages of M. edulis tolerate high ambient pCO2 when food supply is abundant and that important habitat characteristics such as species interactions and energy availability need to be considered to predict species vulnerability to ocean acidification.

(Appendix A) Relative abundance of Emiliania huxleyi in sediment cores of the North Atlantic

The coccolithophore species Emiliania huxleyi is characterized by a wide range of sizes, which can be easily distinguished in the light microscope. In this study we have quantified the abundance of large (coccoliths > 4 µm in maximum length) E. huxleyi specimens during the last 25 kyr in sedimentary records from eleven cores and drill sites in the NE Atlantic and W Mediterranean Sea, to prove its usefulness in the reconstruction of water mass dynamics and biostratigraphic potential. During the Last Glacial Maximum this large form, a cold-water indicator, was common in the NE Atlantic and Mediterranean, and its regional variation in abundance indicates a displacement of the climatic zones southwards in agreement with the development of ice sheets and sea ice in the Northern Hemisphere during this period. On the other hand, the gradient between northern and southern surface water masses in the Subtropical Gyre appears to have been more pronounced than at present, while the Portugal and Canary Currents were more intense. In the western Mediterranean basin temperatures were cooler than in the adjacent Atlantic, provoking a quasi-endemism of these specimens until the end of Heinrich Event 1. This may have been due to a restriction in the communication between the Atlantic and Mediterranean through the Strait of Gibraltar, the arrival of cold surface water and the amplification of cooling after the development of ice sheets in the Northern Hemisphere. During the deglaciation, large E. huxleyi specimens decreased in abundance at medium and low latitudes, but were still numerous close to the Subarctic region during the Holocene. In transitional waters this decrease to present day abundances occurred after Termination Ib. The abrupt change in abundance of this large E. huxleyi form is proposed as a new biostratigraphic event to characterize the Holocene in mid- to low-latitude water masses in the North Atlantic, although this horizon seems to be diachronous by 5 kyr from tropical to subarctic regions, in agreement with the gradual onset of warm conditions.

Planktonic foraminifera of sediment core MD01-2390

Sea-surface temperature (SST) estimates in the sediment core MD01-2390 based on planktonic foraminiferal species abundances using five different transfer function techniques suggest nearly unchanged or unusually higher temperatures in the tropical southern South China Sea (SCS) during the Last Glacial Maximum (LGM) relative to modern temperatures. These results are in contrast to substantial cooling of 2-5 °C inferred by geochemical (Uk'37, Mg/Ca ratios) and terrestrial proxies from the western tropical Pacific region. Using multivariate statistics we show that the glacial southern SCS harboured unique planktonic foraminiferal assemblages that have no modern analogs. Analyses of faunal variation through the core reveal that planktonic foraminiferal assemblages responded to temperature changes inferred from Mg/Ca data but that this signal is subdued by superimposed variations in the relative abundance of Pulleniatina obliquiloculata and Neogloboquadrina pachyderma (dextral). These species occur in glacial samples at proportions that are not observed in the calibration data set. The glacial high abundance of N. pachyderma (dextral) are interpreted to reflect a seasonal (winter) inflow of cold surface water from the northeast via the Bashi Strait due to the combined effects of an intensified winter monsoon, a southward shift of the polar front and the eastward migration of the Kuroshio Current. In contrast, processes controlling the high relative abundances of P. obliquiloculata during the LGM may be unique to the southern SCS. We propose a scenario involving a stronger (winter) mixing or enhanced upwelling due to an intensified winter monsoon that prevented shallow-dwelling, warm indicators to establish larger populations during the LGM. Our results indicate that a no-analog behaviour of planktonic foraminifera faunas is responsible for the warm glacial conditions in this part of the western Pacific warm pool as implied by foraminiferal transfer functions and that a more significant surface cooling in the region as implied by terrestrial and geochemical (Mg/Ca ratios; alkenone unsaturation index) marine proxies is a more likely scenario.

Diatom and hydrocarbon analysis of sediment core NBP00-03-JPC38

In this study, we present a unique high-resolution Holocene record of oceanographic and climatic change based on analyses of diatom assemblages combined with biomarker data from a sediment core collected from the Vega Drift, eastern Antarctic Peninsula (EAP). These data add to the climate framework already established by high-resolution marine sedimentary records from the Palmer Deep, western Antarctic Peninsula (WAP). Heavy sea ice conditions and reduced primary productivity were observed prior to 7.4 ka B.P. in relation with the proximity of the glacial ice melt and calving. Subsequent Holocene oceanographic conditions were controlled by the interactions between the Westerlies-Antarctic Circumpolar Current (ACC)-Weddell Gyre dynamics. A warm period characterized by short seasonal sea ice duration associated with a southern shift of both ACC and Westerlies field persisted until 5 ka B.P. This warm episode was then followed by climate deterioration during the middle-to-late Holocene (5 to 1.9 ka B.P.) with a gradual increase in annual sea ice duration triggered by the expansion of the Weddell Gyre and a strong oceanic connection from the EAP to the WAP. Increase of benthic diatom species during this period was indicative of more summer/autumn storms, which was consistent with changes in synoptic atmospheric circulation and the establishment of low- to high-latitude teleconnections. Finally, the multicentennial scale variability of the Weddell Gyre intensity and storm frequency during the late Holocene appeared to be associated with the increased El Niño-Southern Oscillation frequency.

Temperature, water level and bathymetry of thermokarst lakes in the continuous permafrost zone of northern Siberia - Lena River Delta, Siberia

Thermokarst lakes are typical features of the northern permafrost ecosystems, and play an important role in the thermal exchange between atmosphere and subsurface. The objective of this study is to describe the main thermal processes of the lakes and to quantify the heat exchange with the underlying sediments. The thermal regimes of five lakes located within the continuous permafrost zone of northern Siberia (Lena River Delta) were investigated using hourly water temperature and water level records covering a 3-year period (2009-2012), together with bathymetric survey data. The lakes included thermokarst lakes located on Holocene river terraces that may be connected to Lena River water during spring flooding, and a thermokarst lake located on deposits of the Pleistocene Ice Complex. Lakes were covered by ice up to 2 m thick that persisted for more than 7 months of the year, from October until about mid-June. Lake-bottom temperatures increased at the start of the ice-covered period due to upward-directed heat flux from the underlying thawed sediment. Prior to ice break-up, solar radiation effectively warmed the water beneath the ice cover and induced convective mixing. Ice break-up started at the beginning of June and lasted until the middle or end of June. Mixing occurred within the entire water column from the start of ice break-up and continued during the ice-free periods, as confirmed by the Wedderburn numbers, a quantitative measure of the balance between wind mixing and stratification that is important for describing the biogeochemical cycles of lakes. The lake thermal regime was modeled numerically using the FLake model. The model demonstrated good agreement with observations with regard to the mean lake temperature, with a good reproduction of the summer stratification during the ice-free period, but poor agreement during the ice-covered period. Modeled sensitivity to lake depth demonstrated that lakes in this climatic zone with mean depths > 5 m develop continuous stratification in summer for at least 1 month. The modeled vertical heat flux across the bottom sediment tends towards an annual mean of zero, with maximum downward fluxes of about 5 W/m**2 in summer and with heat released back into the water column at a rate of less than 1 W/m**2 during the ice-covered period. The lakes are shown to be efficient heat absorbers and effectively distribute the heat through mixing. Monthly bottom water temperatures during the ice-free period range up to 15 °C and are therefore higher than the associated monthly air or ground temperatures in the surrounding frozen permafrost landscape. The investigated lakes remain unfrozen at depth, with mean annual lake-bottom temperatures of between 2.7 and 4 °C.

Mineralogical, geochemical, and lipid biomarker study of cabonate precipitates at station GeoB9908-1

Carbonate precipitates recovered from 2,000 m water depth at the Dolgovskoy Mound (Shatsky Ridge, north eastern Black Sea) were studied using mineralogical, geochemical and lipid biomarker analyses. The carbonates differ in shape from simple pavements to cavernous structures with thick microbial mats attached to their lower side and within cavities. Low d13C values measured on carbonates (-41 to -32 per mill V-PDB) and extracted lipid biomarkers indicate that anaerobic oxidation of methane (AOM) played a crucial role in precipitating these carbonates. The internal structure of the carbonates is dominated by finely laminated coccolith ooze and homogeneous clay layers, both cemented by micritic high-magnesium calcite (HMC), and pure, botryoidal, yellowish low-magnesium calcite (LMC) grown in direct contact to microbial mats. d18O measurements suggest that the authigenic HMC precipitated in equilibrium with the Black Sea bottom water while the yellowish LMC rims have been growing in slightly 18O-depleted interstitial water. Although precipitated under significantly different environmental conditions, especially with respect to methane availability, all analysed carbonate samples show lipid patterns that are typical for ANME-1 dominated AOM consortia, in the case of the HMC samples with significant contributions of allochthonous components of marine and terrestrial origin, reflecting the hemipelagic nature of the primary sediment.

DNA alignment and Bayesian trees of partial sequences of COI, COIII and the haemocyanin functional unit f-g of 28 octopod species

Background: Octopods have successfully colonised the world's oceans from the tropics to the poles. Yet, successful persistence in these habitats has required adaptations of their advanced physiological apparatus to compensate impaired oxygen supply. Their oxygen transporter haemocyanin plays a major role in cold tolerance and accordingly has undergone functional modifications to sustain oxygen release at sub-zero temperatures. However, it remains unknown how molecular properties evolved to explain the observed functional adaptations. We thus aimed to assess whether natural selection affected molecular and structural properties of haemocyanin that explains temperature adaptation in octopods. Results: Analysis of 239 partial sequences of the haemocyanin functional units (FU) f and g of 28 octopod species of polar, temperate, subtropical and tropical origin revealed natural selection was acting primarily on charge properties of surface residues. Polar octopods contained haemocyanins with higher net surface charge due to decreased glutamic acid content and higher numbers of basic amino acids. Within the analysed partial sequences, positive selection was present at site 2545, positioned between the active copper binding centre and the FU g surface. At this site, methionine was the dominant amino acid in polar octopods and leucine was dominant in tropical octopods. Sites directly involved in oxygen binding or quaternary interactions were highly conserved within the analysed sequence. Conclusions: This study has provided the first insight into molecular and structural mechanisms that have enabled octopods to sustain oxygen supply from polar to tropical conditions. Our findings imply modulation of oxygen binding via charge-charge interaction at the protein surface, which stabilize quaternary interactions among functional units to reduce detrimental effects of high pH on venous oxygen release. Of the observed partial haemocyanin sequence, residue 2545 formed a close link between the FU g surface and the active centre, suggesting a role as allosteric binding site. The prevalence of methionine at this site in polar octopods, implies regulation of oxygen affinity via increased sensitivity to allosteric metal binding. High sequence conservation of sites directly involved in oxygen binding indicates that functional modifications of octopod haemocyanin rather occur via more subtle mechanisms, as observed in this study.

Depth profiles of iodide and iodate during POLARSTERN cruise ANT-XXIV/3 to the Atlantic sector of the Southern Ocean

The biogeochemistry of iodine in the waters of the Atlantic sector of the Southern Ocean was investigated during the Polarstern cruise ANTXXIV-3 ZERO&DRAKE. The speciation and distribution of iodine (iodate and iodide) in seawater was examined across gradients of iron concentrations and phytoplankton abundance, ranging from an open ocean region along the Zero Meridian to the Weddell Sea and Drake Passage. Iodine cycling in high latitudes differs from that in low latitudes due to differences in the plankton community composition and the physicochemical characteristics. Iodate concentrations ranged between 400 and 450 nmol/L from the surface to the bottom. Surface concentrations of iodide (17 to over 60 nmol/L) were about an order of magnitude higher than below the pycnocline. The peak values of iodide lay nearly always within the euphotic zone and showed a weak, positive correlation with nitrite concentrations in the upper 200 m. In all vertical profiles a pronounced sub-surface maximum in iodide appears between 50 and 200 m depth indicating an iodide drawdown at the near surface. Iodide distribution in the Weddell Sea showed elevated levels in Weddell Sea Bottom Water (WSBW) indicating slow oxidation kinetics and the potential for iodide as a tracer of WSBW formation.

Oxygen and hydrogen isotopes measured in the Northeast Atlantic

Only a few studies have examined the variation of oxygen and hydrogen isotopes of seawater in NE Atlantic water masses, and data are especially sparse for intermediate and deep-water masses. The current study greatly expands this record with 527 d18O values from 47 stations located throughout the mid- to low-latitude NE Atlantic. In addition, dD was analyzed in the 192 samples collected along the GEOTRACES North Atlantic Transect GA03 (GA03_e=KN199-4) and the 115 Iberia-Forams cruise samples from the western and southern Iberian margin. An intercomparison study between the two stable isotope measurement techniques (cavity ring-down laser spectroscopy and magnetic-sector isotope ratio mass spectrometry) used to analyze GA03_e samples reveals relatively good agreement for both hydrogen and oxygen isotope ratios. The surface (0-100 m) and central (100-500 m) water isotope data show the typical, evaporation related trend of increasing values equatorward with the exception for the zonal transect off Cape Blanc, NW Africa. Off Cape Blanc, surface water isotope signatures are modified by the upwelling of fresher Antarctic Intermediate Water (AAIW) that generally has isotopic values of 0.0 to 0.5 per mil for d18O and 0 to 2 per mil for dD. Along the Iberian margin the Mediterranean Outflow Water (MOW) is clearly distinguished by its high d18O (0.5-1.1 per mil) and dD (3-6 per mil) values that can be traced into the open Atlantic. Isotopic values in the NE Atlantic Deep Water (NEADW) are relatively low (d18O: -0.1 to 0.5 per mil; dD: -1 to 4 per mil) and show a broader range than observed previously in the northern and southern convection areas. The NEADW is best observed at GA03_e Stations 5 and 7 in the central NE Atlantic basin. Antarctic Bottom Water isotope values are relatively high indicating modification of the original Antarctic source water along the flow path. The reconstructed d18O-salinity relationship for the complete data set has a slope of 0.51, i.e., slightly steeper than the 0.46 described previously by Pierre et al. (1994, J. Mar. Syst. 5 (2), 159-170.) for the tropical to subtropical Northeast Atlantic. This slope decreases to 0.46 for the subtropical North Atlantic Central Water (NACW) and the MOW and to 0.32 for the surface waters of the upper 50 m. The dD-salinity mixing lines have estimated slopes of 3.01 for the complete data, 1.26 for the MOW, 3.47 for the NACW, and 2.63 for the surface waters. The slopes of the d18O-dD relationship are significantly lower than the one for the Global Meteoric Water Line with 5.6 for the complete data set, 2.30 for the MOW, 4.79 for the NACW, and 3.99 for the surface waters. The lower slopes in all the relationships clearly reflect the impact of the evaporation surplus in the subtropics.