Age and oxygen isotope data for bulk carbonate from DSDP Sites 41-366 and 72-516 and ODP Sites 111-677A and 130-807

A numerical model which describes oxygen isotope exchange during burial and recrystallization of deep-sea carbonate is used to obtain information on how sea surface temperatures have varied in the past by correcting measured d18O values of bulk carbonate for diagenetic overprinting. Comparison of bulk carbonate and planktonic foraminiferal d18O records from ODP site 677A indicates that the oxygen isotopic composition of bulk carbonate does reflect changes in sea surface temperature and d18O. At ODP Site 690, we calculate that diagenetic effects are small, and that both bulk carbonate and planktonic foraminiferal d18O records accurately reflect Paleogene warming of high latitude surface oceans, biased from diagenesis by no more than 1°C. The same is likely to be true for other high latitude sites where sedimentation rates are low. At DSDP sites 516 and 525, the effects of diagenesis are more significant. Measured d18O values of Eocene bulk carbonates are more than 2? lower at deeply buried site 516 than at site 525, consistent with the model prediction that the effects of diagenesis should be proportional to sedimentation rate. Model-corrections reconcile the differences in the data between the two sites; the resulting paleotemperature reconstruction indicates a 4°C cooling of mid-latitude surface oceans since the Eocene. At low latitudes, the contrast in temperature between the ocean surface and bottom makes the carbonate d180 values particularly sensitive to diagenetic effects; most of the observed variations in measured d18O values are accounted for by diagenetic effects rather than by sea surface temperature variations. We show that the data are consistent with constant equatorial sea surface temperatures through most of the Cenozoic, with the possible exception of the early Eocene, when slightly higher temperatures are indicated. We suggest that the lower equatorial sea surface temperatures for the Eocene and Oligocene reported in other oxygen isotope studies are artifacts of diagenetic recrystallization, and that it is impossible to reconstruct accurately equatorial sea surface temperatures without explicitly accounting for diagenetic overprinting.

Mid-Pleistocene Mg/Ca, temperature and stable isotopes from Globigerinoides ruber white and Globorotalia inflata of sediment core GeoB3801-6 from the subtropical South Atlantic

The glacial marine isotope stage 14 (MIS 14) appears in many climate records as an unusually warm glacial. During this period an almost monospecific, up to 1.5 m thick, laminated layer of the giant diatom Ethmodiscus rex has been deposited below the South Atlantic Subtropical Gyre. This oligotrophic region is today less favorable for diatom growth with sediments typically consisting of calcareous nannofossil oozes. We have reconstructed temperatures and the stable oxygen isotopic compositions of sea surface and thermocline water (d18Ow) from planktonic foraminiferal (Globigerinoides ruber and Globorotalia inflata) Mg/Ca and stable oxygen isotopes to test whether perturbations in surface ocean conditions contributed to the deposition of the diatom layer at ~530 kyr B.P. Temperatures and d18Ow values reconstructed from this diatom ooze interval are highly variable, with maxima similar to interglacial values. Since the area of the Ethmodiscus oozes resembles the region where Agulhas rings are present, we interpret these hydrographic changes to reflect the varying influence of warm and saline water of Indian Ocean origin that entered the Subtropical Gyre trapped in Agulhas rings. The formation of the Ethmodiscus oozes is associated with a period of maximum Agulhas leakage and a maximum frequency of Agulhas ring formation caused by a termination-type position of the Subtropical Front during the unusual warm MIS 14. The input of silica through the Agulhas rings enabled the shift in primary production from calcareous nannoplankton to diatoms, leading to the deposition of the massive diatom oozes.

Stable isotope ratios and organic carbon accumulation rates of late Pleistocene sediments of ODP Hole 117-724C

Stable isotope records of coexisting benthic foraminifers Uvigerina spp. and Cibicidoides spp. and planktonic G. ruber (white variety) from Site 724 are used to study the late Pleistocene evolution of surface and intermediate water hydrography (593 m water depth) at the Oman Margin. Glacial-interglacial d18O amplitudes recorded by the benthic foraminifers are reduced when compared to the estimated mean ocean changes of d18Oseawater . Epibenthic d13C remains at its modern level or is increased during glacial times. This implies that Red Sea outflow waters which are enriched in d18Oseawater and d13C (Sum CO2) have been replaced during glacial periods by intermediate waters still positive in d13C (Sum CO2) but more negative in d18Oseawater. Glacial-interglacial amplitudes of the planktonic d18O record exceed those of the mean ocean d18Oseawater variation and imply decreased surface water temperatures (SST) during glacial times. Throughout most of the records these cooling events correlate with enhanced rates of carbon accumulation. However, both negative (colder) SST and positive Corg accumulation rate anomalies do not correlate with potential physical upwelling maxima as inferred from the orbital monsoon index. This is in conflict with the established hypothesis that upwelling in the estern Arabia Sea should be strongest during maxima of the southwest monsoon.

Magnetobiostratigraphy and stable isotope record of Paleocene sediments from ODP Hole 122-761B in the subtropical Indian Ocean

Surface and deep water circulation patterns in the eastern Indian Ocean during the Paleocene Epoch are inferred based on an integrated magnetobiostratigraphic and stable isotope investigation of Ocean Drilling Program Hole 761B, drilled on the Wombat Plateau. A combination of magnetostratigraphy, biostratigraphy and isotope stratigraphy demonstrates that numerous deep sea sites that have been considered to show continuous, or nearly continuous sedimentation through the Paleocene are punctuated by a series of hiatuses, some of which exceeding a duration of 1 Myr. Therefore, our study is based on a detailed temporal interpretation of the stratigraphic successions we used for paleoceanographic reconstructions. We compare detailed planktonic and benthic foraminiferal carbon and oxygen isotope records from Hole 761B with several temporally correlative records published from different oceanic provinces in order to distinguish between local and global patterns within the eastern Indian Ocean. Although Site 761 was situated at low latitudes during the Paleocene, its surface waters were predominantly influenced by circulation originating from the Southern Ocean as indicated by inferred cool sea surface temperatures and reduced surface to deep water temperature gradients. We suggest that deep waters in the eastern Indian Ocean were not directly fed by the Southern or Tethys Oceans. Rather, the more negative delta13C composition of the bottom waters recorded by benthic foraminifera implies the presence and/or active contribution of aged deep waters from the Pacific during this time, at least prior to ~60.2 Ma and subsequent to ~59.0 Ma. The Indian continent, Ninetyeast Ridge, Kerguelen Plateau and Broken Ridge may have played a significant role as submarine barriers to deep water circulation during the Paleocene.

Stable isotope record and trace element ratios of Eocene and Oligocene foraminifers from ODP Site 120-748

Stable carbon and oxygen isotope analyses were conducted on well-preserved planktonic and benthic foraminifers from a continuous middle Eocene to Oligocene sequence at Ocean Drilling Program (ODP) Site 748 on the Kerguelen Plateau. Benthic foraminifer d18O values show a 1.0 per mil increase through the middle and upper Eocene, followed by a rapid 1.2 per mil increase in the lowermost Oligocene (35.5 Ma). Surface-dwelling planktonic foraminifer d18O values increase in the lowermost Oligocene, but only by 0.6 per mil whereas intermediate-depth planktonic foraminifers show an increase of about l.0 per mil. Benthic foraminifer d13C values increase by 0.9 per mil in the lowermost Oligocene at precisely the same time as the large d18O increase, whereas planktonic foraminifer d13C values show little or no change. Site 748 oxygen isotope and paleontological records suggest that southern Indian Ocean surface and intermediate waters underwent significant cooling from the early to late Eocene. The rapid 1.2 per mil oxygen isotope increase recorded by benthic foraminifers just above the Eocene/Oligocene boundary represents the ubiquitous early Oligocene d18O event. The shift here is unique, however, as it coincided with the sudden appearance of ice-rafted debris (IRD), providing the first direct link between Antarctic glacial activity and the earliest Oligocene d18O increase. The d18O increase caused by the ice-volume change in the early Oligocene is constrained by (1) related changes in the planktonic to benthic foraminifer d18O gradient at Site 748 and (2) comparisons of late Eocene and early Oligocene planktonic foraminifer d18Ovalues from various latitudes. Both of these records indicate that 0.3 per mil to 0.4 per mil of the early Oligocene d18O increase was ice-volume related.

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.

Paleogene and early Neogene stable isotope record of foraminifera from ODP Leg 119 samples

Oxygen and carbon isotopic records of monogeneric and monospecific benthic and planktonic foraminifer samples from Sites 744 and 738 drilled on the southern end of the Kerguelen Plateau during ODP Leg 119 reveal the evolution of polar Indian Ocean water masses from the early Paleocene to the middle Miocene. Results from Site 738 are from sediments of early Paleocene to late Eocene age and those from Site 744 are late Eocene to middle Miocene. They suggest that intermediate waters at this location did not originate in the high latitudes during the early Eocene. Surface and near-surface waters cooled gradually after the maximum warming at 56 Ma, when surface waters were about 18°C. Intermediate waters cooled after 52 Ma. The highest temperatures (lowest d18O values) of the Cenozoic occurred from 56 to 52 Ma. The records of equatorial Pacific Site 577 and Weddell Sea Site 690 resemble that of the polar Indian Ocean in this interval. The well-documented d13C excursions toward positive values in the late Paleocene and negative values in the early Eocene are represented by foraminifers increases of 1.5 per mil and following decreases of about 3 per mil. Most of the cooling in the Paleogene occurred in the middle and late Eocene. A 2°C decrease of surface water at about 38.4 Ma heralded the beginning of extensive glacial conditions in Antarctica in the early Oligocene. At Site 744, the global d18O shift just above the Eocene/Oligocene boundary is 1.15 per mil, and occurred gradually in sediments dated at 36.5-35.9 Ma. Ice-rafted debris was deposited beginning at 36.1 Ma for about the next 2 m.y. This simultaneous occurrence of the global d18O shift with ice-rafted debris is evidence for early Oligocene glaciation in East Antarctica. Moreover, early and late Oligocene Cibicidoides d18O values between 2 and 2.2 per mil indicate intermediate water cooling and a small ice-volume effect. Production of cold dense bottom water in Antarctica was intensified with continental cooling and glaciation in the early Oligocene. Comparison of Oligocene and early Miocene isotopic data from high-latitude and low-latitude deepsea sites indicates that there were probably at least two sources of bottom waters at this time.

Late Cretaceous planktonic foraminiferal assemblages from the Exmouth Plateau, ODP Sites 122-762 and 122-763, eastern Indian Ocean

The evolution of planktonic foraminifera during the Late Cretaceous is marked in the Santonian by the disappearance of complex morphotypes (the marginotruncanids), and the contemporary increasing importance and diversification of another group of complex taxa, the globotruncanids. Upper Turonian to lower Campanian planktonic foraminiferal assemblages from Holes 762C and 763B (Ocean Drilling Program, Leg 122, Exmouth Plateau, 47°S palaeolatitude) were studied in detail to evaluate the compositional variations at the genus and species level based on the assumption that, in the Cretaceous oceans as in the modern, any faunal change was associated with changes in the characteristics and the degree of stability of the oceanic surface waters. Three major groups were recognised based on gross morphology, and following the assumption that Cretaceous planktonic foraminifera, although extinct, had life-history strategies comparable to those of modern planktonics: 1 - r-selected opportunists; 2 - k-selected specialists; 3 - r/k intermediate morphotypes which include all genera that display a range of trophic strategies in-between opportunist and specialist taxa. Although planktonic foraminiferal assemblages are characterised by a progressive appearance of complex taxa, this trend is discontinuous. Variation in number of species and specimens within genera has allowed recognition of five discrete intervals each of them reflecting different oceanic conditions based on fluctuations in diversity and abundance of the major morphotypes. Planktonic forms show cyclical fluctuations in diversity and abundance of cold (r-strategists) and warm taxa (k-strategists), perhaps representing alternating phases of unstable conditions (suggesting a weakly stratified upper water column in a mesotrophic environment), and well-stratified surface and near-surface waters (indicating a more oligotrophic environment). Interval 1, middle Turonian to early Coniacian in age, is dominated by the r/k intermediate morphotypes which alternate with r-strategists. These cyclical alternations are used to identify three additional subintervals. Interval 2, aged middle to late Coniacian, is characterised by the increasing number of species and relative abundance of k-strategists. After this maximum diversification the k-strategists show a progressive decrease reaching a minimum value in Interval 3 (early to late Santonian), which corresponds to the extinction of the genus Marginotruncana. In the Interval 4, latest Santonian in age, the k-strategists, represented mainly by the genera Globotruncana, increase again in diversity and abundance. The last Interval 5 (early Campanian) is dominated by juvenile globotruncanids and r-strategists which fluctuate in opposite phase. The positive peak (Interval 2) related to the maximum diversification of warm taxa (k-strategists) in the Coniacian seems to correspond to a warmer episode. It is followed by a marked decrease in the relative abundance of warm taxa (k-strategists crisis) with a minimum in the late Santonian (Interval 3), reflecting a decrease in temperature. Detailed analysis of faunal variations allows the Santonian faunal turnover to be ascribed to a cooling event strong enough to cause the extinction of the marginotruncanids.

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.

Zooplankton distribution and migration in low-oxygen modewater eddies

The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300-600 m depth. Here, oxygen concentrations rarely fall below 40 µmol O2 kg-1, but are expected to decline under future projections of global warming. The recent discovery of mesoscale eddies that harbour a shallow suboxic (<5 µmol O2 kg-1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by on-going ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. At nighttime, when a large proportion of acoustic scatterers is ascending into the upper 150 m, a drastic reduction in mean volume backscattering (Sv, shipboard ADCP, 75kHz) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers were avoiding the depth range between about 85 to 120 m, where oxygen concentrations were lower than approximately 20 µmol O2 kg-1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time-series observations of a moored ADCP (upward looking, 300kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies have been identified to be followed by zooplankton in response to the eddy OMZ: i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids), ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods), iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes), and iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy i), ii) and iv), compression of the habitable volume in the surface may increase prey-predator encounter rates, rendering zooplankton and micronekton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 µmol O2 kg-1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton and micronekton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Development of bacterial and archaeal communities in erupted subsurface muds at the Håkon Mosby mud volcano

The microbial oxidation of methane controls the emission of the greenhouse gas methane from the ocean floor. However, some seabed structures such as mud volcanoes have leaky microbial methane filters and can be important sources of methane. We investigated the disturbance and recovery of a methanotrophic mud volcano microbiome (Håkon Mosby mud volcano, 1250 m water depth), to assess time scales of community succession and function in the natural deep-sea environment. We analyzed 10 surface and 5 subsurface sediment samples across HMMV mud flows from most recently discharged subsurface muds towards old consolidated muds as well as one reference site (REF) located approximately 0.5 km outside of the HMMV. Surface samples were obtained in 2003, 2009 and 2010. The surface of the new mud flows at the geographical center was sampled in 2009 and 2010. Around 100 m south of the center, we sampled more consolidated aged muds in 2003 and 2010. Old mud flows were sampled around 300 m southeast and 100 m north of the geographical center in 2003, 2009 and 2010. Surface sediment samples (0-20 cm) were recovered either by TV-guided Multicorer or by push cores using the remotely operated vehicle Quest (Marum, University Bremen). Subsurface sediments of all zones (>2 m below sea floor) were obtained in 2003 by gravity corer. After recovery, sediments were immediately subsampled in a refrigerated container (0°C) and further processed for biogeochemical analyses or preserved at -20°C for later DNA analyses. Our study show that freshly erupted muds hosted heterotrophic deep subsurface communities, which were replaced by surface communities within a few years of exposure. Aerobic methanotrophy was established at the top surface layer within less than a year, followed by anaerobic methanotrophy, sulfate reduction and finally thiotrophy. Our data indicate that it takes decades in cold environments before efficient methanotrophic communities establish to control methane emission. The observed succession provides insights to the response time of complex deep-sea communities to seafloor disturbances.