Benthic foraminifera extinctions in the Cenozoic record

In the late Pliocene-middle Pleistocene a group of 95 species of elongate, cylindrical, deep-sea (lower bathyal-abyssal) benthic foraminifera became extinct. This Extinction Group (Ext. Gp), belonging to three families (all the Stilostomellidae and Pleurostomellidae, some of the Nodosariidae), was a major component (20-70%) of deep-sea foraminiferal assemblages in the middle Cenozoic and subsequently declined in abundance and species richness before finally disappearing almost completely during the mid-Pleistocene Climatic Transition (MPT). So what caused these declines and extinction? In this study 127 Ext. Gp species are identified from eight Cenozoic bathyal and abyssal sequences in the North Atlantic and equatorial Pacific Oceans. Most species are long-ranging with 80% originating in the Eocene or earlier. The greatest abundance and diversity of the Ext. Gp was in the warm oceanic conditions of the middle Eocene-early Oligocene. The group was subjected to significant changes in the composition of the faunal dominants and slightly enhanced species turnover during and soon after the rapid Eocene-Oligocene cooling event. Declines in the relative abundance and flux of the Ext. Gp, together with enhanced species loss, occurred during middle-late Miocene cooling, particularly at abyssal sites. The overall number of Ext. Gp species present began declining earlier at mid abyssal depths (in middle Miocene) than at upper abyssal (in late Pliocene-early Pleistocene) and then lower bathyal depths (in MPT). By far the most significant Ext. Gp declines in abundance and species loss occurred during the more severe glacial stages of the late Pliocene-middle Pleistocene. Clearly, the decline and extinction of this group of deep-sea foraminifera was related to the function of their specialized apertures and the stepwise cooling of global climate and deep water. We infer that the apertural modifications may be related to the method of food collection or processing, and that the extinctions may have resulted from the decline or loss of their specific phytoplankton or prokaryote food source, that was more directly impacted than the foraminifera by the cooling temperatures.

Ocean temperature and salinity measured by CTD SRDLs deployed on southern elephant seals from King George Island with links to datasets

West Antarctic ice shelves have thinned dramatically over recent decades. Oceanographic measurements that explore connections between offshore warming and transport across a continental shelf with variable bathymetry toward ice shelves are needed to constrain future changes in melt rates. Six years of seal-acquired observations provide extensive hydrographic coverage in the Bellingshausen Sea, where ship-based measurements are scarce. Warm but modified Circumpolar Deep Water floods the shelf and establishes a cyclonic circulation within the Belgica Trough with flow extending toward the coast along the eastern boundaries and returning to the shelf break along western boundaries. These boundary currents are the primary water mass pathways that carry heat toward the coast and advect ice shelf meltwater offshore. The modified Circumpolar Deep Water and meltwater mixtures shoal and thin as they approach the continental slope before flowing westward at the shelf break, suggesting the presence of the Antarctic Slope Current. Constraining meltwater pathways is a key step in monitoring the stability of the West Antarctic Ice Sheet.

NADW Photodegradation FTICR-MS

Duplicate, filtered samples of North Atlantic Deep Water (NADW) were irradiated for 28 days in a solar simulator. Duplicate dark controls were placed alongside the irradiated samples. After 28 days, samples were extensively photo-degraded based upon colored dissolved organic matter (CDOM) photo-bleaching (> 95%). Samples were solid phase extracted using PPL resin to isolate, concentrate and desalt the dissolved organic matter (DOM) in the samples. Ultrahigh resolution electrospray ionization Fourier transform ion cyclotron resonance mass spectrometry (FTICR-MS) enabled 3024 molecular formulas to be identified in the dark control. Photo-degradation decreased molecular diversity, with 2402 formulas assigned post-irradiation. Molecular formulas were classified based upon their photo-lability as 1) photo-resistant; 2) photo-labile; and, 3) photo-produced. Photo-resistant DOM made up 73% of all formulas and was dominated by highly unsaturated molecular signatures consistent with carboxylic-rich alicyclic molecules, suggesting that these apparently bio-refractory compounds may also survive multiple passages through sunlit surface waters and thus accumulate for timeframes exceeding ocean ventilation. Photo-labile DOM was enriched in low molecular weight formulas, aromatic molecular formulas, and sulfur and phosphorous containing formulas. Compounds containing both sulfur and nitrogen were particularly photo-labile and may represent an underappreciated component of the photo-reactive CDOM pool. Photo-produced DOM was enriched in higher molecular weight formulas, as well as aliphatic and peptide formulas. Molecular formulas are indexed by their photo-lability classification in the supplementary information.

Dive depth and dive duration data of male southern elephant seals from Marion Island between 2004 and 2008 with links to datasets

We describe the habitat use of 22 male southern elephant seals (Mirounga leonina) satellite tagged at Marion Island between 2004 and 2008. While a few areas of increased utilization appeared to be associated with areas of shallower bathymetry (such as sea-floor ridges and fracture zones), seals in our study did not target other areas of shallow bathymetry within close proximity to Marion Island. Rather, most elephant seals foraged pelagically over very deep water where much variation was evident in diel vertical migration strategies. These strategies resulted in generally deeper and longer dives than what has been reported for male elephant seals from other colonies. No significant differences were recorded for dive durations or dive depths between adults and sub-adults. However, younger animals displayed a positive relationship between dive durations and age, as well as between dive depths and age, while these relationships became negative for older animals. Mixed model outputs suggested that seals increased their aerobic fitness as migrations progressed, enabling them to undertake longer dives. We conclude that Marion Island male elephant seals exhibit much variability in dive strategy and are seemingly capable of exploiting a range of different prey types occurring in various depth layers.

Depth tie points and ages of IODP Site 306-U1314

Integrated Ocean Drilling Program (IODP) Site U1314 of the North Atlantic is a critical sedimentary archive record of subpolar deep water from the southern Gardar Drift for which we derived an age model of orbital resolution for the last 1.8 Ma. This chronology combined with high-resolution (cm scale) X-ray fluorescence core scanning measurements of major elements allows tracking changes in terrigenous provenance during the last 1.1 Ma. Low Potassium to Titanium (K/Ti) ratios reflect enhanced transport of basalt-derived titanomagnetites during warm climate intervals, while high K/Ti ratios indicate a dominance of acidic sediment sources typical for glacial and stadial events. Changes in K/Ti and magnetic concentration at Site 1314 are coeval with fluctuations in smectite content and grain size data from nearby piston cores, suggesting that the provenance changes are mainly controlled by variable flow of the Iceland-Scotland Overflow Water, an important branch of North Atlantic Deep Water. Furthermore, K/Ti variations on orbital time scales show a striking similarity to the deep sea d13C record from ODP Site 607. Pervasive features of the K/Ti time series during and after the Mid-Pleistocene Transition are suborbital changes similar to Dansgaard/Oeschger and Bond oscillations that appear to be strongly amplified during ice growth phases when global benthic d18O was within the range of ~4.1-4.6 per mil. The strong increase in variability of sediment provenance and subsequently deep hydrography at benthic d18O values below ~4.1 suggests that the extent of glaciations and, therefore, sea level corresponding to this value constitutes an important physical threshold that was persistent at least for the last 1.1 Ma.

Planktic foraminifera at the Paleocene-Eocene transition in the Antarctic Indian Ocean

Isotopic depth stratification and relative abundance studies of planktic foraminifera at ODP Site 738 reveal three major faunal turnovers during the latest Paleocene and early Eocene, reflecting the climatic and structural changes in the Antarctic surface ocean. Faunal Event 1 occurred near the Paleocene/Eocene boundary and is characterized by a faunal turnover in deep dwellers, decreased relative abundance in intermediate dwellers and increased relative abundance in surface dwellers. This event marks a temporary elimination of the vertical structure in the surface ocean over a period of more than 63,000 years that is apparently associated with the sudden shutdown of the "Antarctic Intermediate Water" production. The appearance of morozovellids before this event suggests that polar warming is the cause for the shutdown in the production of this water mass. At this time warm saline deep water may have formed at low latitudes. Faunal Event 2 occurred near the AP5a/AP5b Subzonal boundary and is characterized by a faunal turnover in deep dwellers with no apparent change in surface and intermediate dwellers. Increased individual size, wall-thickness and relative abundance in deep dwelling chiloguembelinids suggests the formation of a deep oxygen minima in the Antarctic Oceans during the maximum polar warming possibly as a result of upwelling of nutrient-rich deep water. Faunal Event 3 occurred in Subzone AP6 and is characterized by a faunal turnover in surface dwellers and a delayed diversification in deep dwellers. This event marks the onset of Antarctic cooling. A drastic decrease in the delta13C/delta18O values of the deep assemblage in Zone AP7 suggests an intensified thermocline and reduced upwelling following the polar cooling.

Nd-Sr isotopic composition of foraminifera and bulk sediments

We present new isotopic data for sedimentary planktonic foraminifera, as well as for potential water column and sedimentary sources of neodymium (Nd), which confirm that the isotopic composition of the foraminifera is the same as surface seawater and very different from deep water and sedimentary Nd. The faithfulness with which sedimentary foraminifera record the isotopic signature of surface seawater Nd is difficult to explain given their variable and high Nd/Ca ratios, ratios that are often sedimentary foraminifera, ratios that are often much higher than is plausible for direct incorporation within the calcite structure. We present further data that demonstrate a similarly large range in Nd/Ca ratios in plankton tow foraminifera, a range that may be controlled by redox conditions in the water column. Cleaning experiments reveal, in common with earlier work, that large amounts of Nd are released by cleaning with both hydrazine and diethylene triamine penta-acetic acid, but that the Nd released at each step is of surface origin. While further detailed studies are required to verify the exact location of the surface isotopic signature and the key controls on foraminiferal Nd isotope systematics, these new data place the use of planktonic foraminifera as recorders of surface water Nd isotope ratios, and thus of variations in the past supply of Nd to the oceans from the continents via weathering and erosion, on a reasonably sure footing.

Amphipods in sediment traps of the eastern Fram Strait

Life-cycle characteristics of the free-swimming lysianassoid amphipod Cyclocaris guilelmi were investigated and compared to those of other regularly appearing amphipods in the Arctic deep-sea community. In this context we analysed time-series data of meso- and bathypelagic amphipods collected as swimmers in moored sediment traps from 2004 to 2008 at the deep-sea long-term observatory HAUSGARTEN (79°N/4°E) in the eastern Fram Strait, Arctic Ocean. Six mesopelagic and three bathypelagic deep-sea amphipod species regularly occurred in the traps. The lysianassoid C. guilelmi showed a stable interannual population size and seasonal peaks in its occurrence from August to February during the five-year sampling period. The investigation of its population structure and reproduction ecology indicated year-round breeding behavior of this species. Up to 4 cohorts consisting mainly of juvenile and female C. guilelmi were observed. We conclude that C. guilelmi plays an important role within the Arctic amphipod deep water community.

North Atlantic late Miocene stable-isotope stratigraphy, biostratigraphy, and magnetostratigraphy

Upper Miocene foraminiferal nannofossil ooze and chalk from DSDP Hole 552A in the northeast Atlantic Ocean have been closely sampled for biostratigraphic, paleomagnetic, and stable-isotopic studies. Sampling at 10-cm intervals resulted in an uppermost Miocene isotope stratigraphy with a 1000- to 3000-yr. resolution. Covariance in benthic (Planulina wuellerstorfi) and planktonic (Globigerina bulloides) foraminiferal d18O records is taken as evidence for variability in continental ice volume. Our best estimate is that glacial maxima occurred at -5.0 and ~ 5.5 Ma and lasted no more than 20,000 yrs. These events probably lowered sea level by 60 m below the latest Miocene average. There is little oxygen-isotope evidence, however, for a prolonged glaciation during the last 2 m.y. of the late Miocene. High- and low-frequency variability in the d13C record of foraminifers is useful for correlation among North Atlantic DSDP Sites 408, 410, 522, 610, and 611, and for correlation with sites in other oceans. Similar d13C changes are seen in P. wuellerstorfi and G. bulloides, but the amplitude of the signal is always greater in G. bulloides. Variability in d13C common to both species probably reflects variability in the d13C of total CO2 in seawater. Major long-term features in the d13C record include a latest Miocene maximum (P. wuellerstorfi = 1.5 per mil ) in paleomagnetic Chron 7, an abrupt decrease in d13C at -6.2 Ma, and a slight increase at -5.5 Ma. The decrease in d13C at -6.2 Ma, which has been paleomagnetically dated only twice before, occurs in the upper reversed part of Chronozone 6 at Holes 552A and 611C, in excellent agreement with earlier studies. Cycles in d13C with a period of ~ 10 4 yrs. are interpreted as changes in seawater chemistry, which may have resulted from orbitally induced variability in continental biomass. Samples of P. wuellerstorfi younger than 6 Ma from throughout the North Atlantic have d13C near lo, on average ~ l per mil greater than samples of the same age in the Pacific Ocean. Thus, there is no evidence for cessation of North Atlantic Deep Water production resulting from the Messinian "salinity crisis." Biostratigraphic results indicate continuous sedimentation during the late Miocene after about -6.5 Ma at Hole 552A. Nannofossil biostratigraphy is complicated by the scarcity of low-latitude marker species, but middle and late Miocene Zones NN7 through NN11 are recognized. A hiatus is present at -6.5 Ma, on the basis of simultaneous first occurrences of Amaurolithusprimus, Amaurolithus delicatus, Amaurolithus amplificus, and Scyphosphaera globulata. The frequency and duration of older hiatuses increase downsection in Hole 552A, as suggested by calcareous nannofossil biostratigraphy and magnetostratigraphy. Paleomagnetic results at Hole 552A indicate a systematic pattern of inclination changes. Chronozone 6 was readily identified because of its characteristic nannoflora (sequential occurrences of species assigned to the genus Amaurolithus) and the d13C decrease in foraminifers, but its lower reversed interval is condensed. Only the lower normal interval of Chronozone 5 was recognized at Hole 552A; the upper normal interval and the lowest Gilbert sediment are not recognized, owing to low intensity of magnetization and to coring disturbance. Interpreting magnetic reversals below Chronozone 6 was difficult because of hiatuses, but a lower normally magnetized interval is probably Chronozone 7. Correlation between DSDP Hole 552A and other North Atlantic sites is demonstrated using coiling direction changes in the planktonic foraminifer Neogloboquadrina. At most sites this genus changed its coiling preference from dominantly right to dominantly left during the late Miocene. At Hole 552A this event probably occurred about 7 m.y. ago. At the same time, P. wuellerstorfi had maximum d13C values. A similar d13C maximum and coiling change occurred together in Chron 7 at Hole 611C, and at Hole 610E. In sediment younger than -5.5 Ma, the coiling of small Neogloboquadrina species is random, but the larger species N. atlantica retains preferential left coiling.

Pliocene/Pleistocene benthic foraminiferal abundances from six drilling cores

Twenty percent (19 genera, 95 species) of cosmopolitan, deep-sea (500-4000 m), benthic foraminiferal species became extinct during the late Pliocene-Middle Pleistocene (3-0.12 Ma), with the peak of extinctions (76 species) occurring during the mid-Pleistocene Climate Transition (MPT, 1.2-0.55 Ma). One whole family (Stilostomellidae, 30 species) was wiped out, and a second (Pleurostomellidae, 29 species) was decimated with just one species possibly surviving through to the present. Our studies at 21 deep-sea core sites show widespread pulsed declines in abundance and diversity of the extinction group species during more extreme glacials, with partial interglacial recoveries. These declines started in the late Pliocene in southern sourced deep water masses (Antarctic Bottom Water, Circumpolar Deep Water) and extending into intermediate waters (Antarctic Intermediate Water, North Atlantic Deep Water) in the MPT, with the youngest declines in sites farthest downstream from high-latitude source areas for intermediate waters. We infer that the unusual apertural types that were targeted by this extinction period were adaptations for a specific kind of food source and that it was probably the demise of this microbial food that resulted in the foraminiferal extinctions. We hypothesize that it may have been increased cold and oxygenation of the southern sourced deep water masses that impacted on this deep water microbial food source during major late Pliocene and Early Pleistocene glacials when Antarctic ice was substantially expanded. The food source in intermediate water was not impacted until major glacials in the MPT when there were significant expansion of polar sea ice in both hemispheres and major changes in the source areas, temperature, and oxygenation of global intermediate waters.

Late Miocene to Pleistocene paleoceanographic records from the Feni and Gardar Drifts: Pliocene reduction in abyssal flow

Magnetic fabric analyses from two North Atlantic drift deposits provide proxies for determining relative variations in the strength of abyssal flow over the last 10 my. The data show a cessation of current-controlled sedimentation at the shallower Feni Drift (2417 m) at the time of onset of Northern Hemisphere glaciation (2.6 Ma). Drift formation ended nearly 2 my earlier (4.2 Ma) at the deeper Gardar Drift (3220 m), implying stepwise reduction in deep-water flow. Relatively light delta18O values at the deeper Gardar Drift indicate a warmer, thus also more salty, water mass site prior to 6 Ma. We interpret this as representing Mediterranean Sea water, which flowed north at depths greater than that of the Feni Drift Site. The supply of Mediterranean Water to the North Atlantic was shut off as the Gibraltar Straits closed, causing the Messinian salinity crisis, and never returned to that position in the water column after the Mediterranean opened again.

Radiocarbon ages and stable isotope measurements on sediment cores of the subarctic Pacific and its marginal seas

Under modern conditions only North Pacific Intermediate Water is formed in the northwest Pacific Ocean. This situation might have changed in the past. Recent studies with general circulation models indicate a switch to deep-water formation in the northwest Pacific during Heinrich Stadial 1 (17.5-15.0 ka) of the last glacial termination. Reconstructions of past ventilation changes based on paleoceanographic proxy records are still insufficient to test whether a deglacial mode of deep-water formation in the North Pacific Ocean existed. Here we present deglacial ventilation records based on radiocarbon-derived ventilation ages in combination with epibenthic stable carbon isotopes from the northwest Pacific including the Okhotsk Sea and Bering Sea, the two potential source regions for past North Pacific ventilation changes. Evidence for most rigorous ventilation of the intermediate-depth North Pacific occurred during Heinrich Stadial 1 and the Younger Dryas, simultaneous to significant reductions in Atlantic Meridional Overturning Circulation. Concurrent changes in d13C and ventilation ages point to the Okhotsk Sea as driver of millennial-scale changes in North Pacific Intermediate Water ventilation during the last deglaciation. Our records additionally indicate that changes in the d13C intermediate-water (700-1750 m water depth) signature and radiocarbon-derived ventilation ages are in antiphase to those of the deep North Pacific Ocean (>2100 m water depth) during the last glacial termination. Thus, intermediate- and deep-water masses of the northwest Pacific have a differing ventilation history during the last deglaciation.

Log-ratio of silica to aluminium counts (ln(Si/Al)) from ODP site 108-658

Growing evidence suggests that the low atmospheric CO2 concentration of the ice ages resulted from enhanced storage of CO2 in the ocean interior, largely as a result of changes in the Southern Ocean1. Early in the most recent deglaciation, a reduction in North Atlantic overturning circulation seems to have driven CO2 release from the Southern Ocean**2, 3, 4, 5, but the mechanism connecting the North Atlantic and the Southern Ocean remains unclear. Biogenic opal export in the low-latitude ocean relies on silicate from the underlying thermocline, the concentration of which is affected by the circulation of the ocean interior. Here we report a record of biogenic opal export from a coastal upwelling system off the coast of northwest Africa that shows pronounced opal maxima during each glacial termination over the past 550,000 years. These opal peaks are consistent with a strong deglacial reduction in the formation of silicate-poor glacial North Atlantic intermediate water**2 (GNAIW). The loss of GNAIW allowed mixing with underlying silicate-rich deep water to increase the silicate supply to the surface ocean. An increase in westerly-wind-driven upwelling in the Southern Ocean in response to the North Atlantic change has been proposed to drive the deglacial rise in atmospheric CO2 (refs 3, 4). However, such a circulation change would have accelerated the formation of Antarctic intermediate water and sub-Antarctic mode water, which today have as little silicate as North Atlantic Deep Water and would have thus maintained low silicate concentrations in the Atlantic thermocline. The deglacial opal maxima reported here suggest an alternative mechanism for the deglacial CO2 release**5, 6. Just as the reduction in GNAIW led to upward silicate transport, it should also have allowed the downward mixing of warm, low-density surface water to reach into the deep ocean. The resulting decrease in the density of the deep Atlantic relative to the Southern Ocean surface promoted Antarctic overturning, which released CO2 to the atmosphere.