Calcareous nannofossil bioevents of early-middle Pleistocene sediments from the Mediterranean Sea and the Noth Atlantic

Quantitative distributions of calcareous nannofossils are analysed in the early-middle Pleistocene at the small Gephyrocapsa and Pseudoemiliania lacunosa zone transition in deep-sea cores from the Mediterranean Sea and North Atlantic Ocean (Ocean Drilling Program [ODP] Sites 977, 964 and 967, Deep Sea Drilling Project [DSDP] Site 607). The temporal and spatial mode of occurrence of medium-sized gephyrocapsids and reticulofenestrids has been examined to refine biostratigraphic constraints and evaluate possible relationships of stratigraphic patterns to environmental changes during a period of global climatic deterioration. The timing of bioevents has been calibrated using high-resolution sampling and correlation to the delta18O record in chronologically well-constrained sections. Newly identified events and ecostratigraphical signals enhance the stratigraphic resolution at the early-middle Pleistocene. The first occurrence (FO) of intermediate morphotypes between Pseudoemiliania and Reticulofenestra (Reticulofenestra sp.) is proposed as a reliable event within marine isotope stage (MIS) 35 or at the MIS 35/34 transition. The distribution of Reticulofenestra asanoi is characterized by rare and scattered occurrences in its lowest range, but the first common occurrence (FCO) is consistently identified at MIS 32 or 32/31; the last common occurrence (LCO) of the species is a distinctive event at MIS 23. In the studied interval, Gephyrocapsa omega dominates among medium-sized Gephyrocapsa. The FO of G. omega and contemporaneous re-entry of medium-sized gephyrocapsids at the lower-middle Pleistocene transition are diachronous between the Atlantic Ocean and Mediterranean Sea and from the western to eastern Mediterranean. In the Mediterranean, the LO of G. omega falls at MIS 15, insolation cycle 54 and is isochronous among the sites. Abundance fluctuations of G. omega show notable relations to early-middle Pleistocene climate changes; they considerably increase in abundance at the interglacial stages, suggesting warm water preferences. Gephyrocapsa omega temporarily disappears during the glacial MIS 22 and MIS 20. Above MIS 20, an impoverishment in G. omega and in the total abundance of medium-sized gephyrocapsids occurs. A decrease in abundance of G. omega is observed between the western Site 977 and the easternmost Site 967 in the Mediterranean Sea, as a possible response to high salinity and/or low nutrient content. Possible environmental influences on the distribution of R. asanoi and of Reticulofenestra sp. are discussed.

Maximum and mean diameter record of Orbulina universa from DSDP Hole 47-397 of Cape Bojador, eastern North Atlantic

Measurements of the diameter of O. universa carried out on 30 specimens from 39 samples covering a sediment thickness of 78 m and going back in time to approximately 750 000 y resulted in the construction of a curve of the mean diameter and a curve of the maximum diameter. Both curves, as well as those calculated with the running-averages technique, display cyclic fluctuations with durations of the order of 100 000 y and downwards decreasing amplitudes. These curves are compared with a carbonate curve (on bulk sediment) and an isotopic curve (on benthic foraminifers) obtained from the same set of samples. Correlations are fair to good, but a timelag is noticed between the isotopic curve and the faunal (O. universa mean diameter) curve, with the isotopic signal coming first, in the middle part of the Brunhes Epoch. Biostratigraphic calibration to the paleomagnetic record is provided by four datum planes (two based on calcareous nannofossils, two on diatoms) identified in the succession. Changes recorded in test porosity seem to be less meaningful than changes in test size.

Middle Miocene to recent sedimentological record of ODP sites 177-1088 and 177-1092

During Leg 177 of the Ocean Drilling Program (ODP), well-preserved Middle Miocene to Pleistocene carbonate-rich sediment records were recovered on a north-south transect through the south-eastern Atlantic sector of the Southern Ocean at Site 1088 on the Agulhas Ridge and Site 1092 on Meteor Rise. Both sites were dominated by the deposition of calcareous nannofossil oozes through the Miocene, indicating low biological productivity in warm to temperate surface waters. A continuous increase in the proportions of foraminifera since the latest Miocene (6.5 Ma) points to enhanced nutrient supply, possibly related to the global 'biogenic bloom' event across the Miocene-Pliocene boundary. Since the Late Pliocene, different styles of biological productivity developed between the sites. Enhanced deposition of biosiliceous constituents at the southern Site 1092, particularly in the Early Pleistocene, is consistent with the formation of the Circum-Antarctic Opal Belt since 2.5 Ma in a setting near the Polar Front, whereas carbonate deposition still prevailed at the northern Site 1088 situated near the Subtropical Front. Clay-mineral tracers of water-mass advection together with the pattern of sedimentation rates and hiatuses reflect distinct pulses in the development of regional ocean circulation between 14 and 12 Ma, around 8 Ma and since 2.8 Ma. These pulses can be related to Antarctic ice-sheet extension that mediates the production and flow of southern source water, and stepwise increases in North Atlantic Deep Water production that drives global conveyor circulation. At Site 1088, illite chemistry and silt/clay ratios of the terrigenous sediment fraction reflect the history of terrestrial climate in southern Africa, with humid conditions prior to the Early Late Miocene (9.7 Ma), followed by a dry episode until 7.7 Ma. The latest Miocene and Early Pliocene were characterized by a humid episode until modern aridity was established in the Late Pliocene between 4.0 and 2.8 Ma. These climate changes were related to the latitudinal migration of climate belts in response to tectonically caused reorganizations in atmospheric and ocean circulation.

Coccoliths in phytoplankton of the Eastern Bering Sea in August 2001

Based on results of field observations in August 1998, July 2000, and August 2001 composition and quantitative distribution of coccolithophorids in the middle part of the Eastern Bering Sea shelf between 56°052'N and 59°019'N was characterized. Emiliania huxleyi abundance, biomass, and population structure as well as role of species in the coccolithophorid community and phytoplankton as a whole were evaluated. Abundance of the species in the upper mixed layer in bloom areas was 1-3 mln cells/l and biomass made up 30-75 mg C/m**3. E. huxleyi share in total phytoplankton numbers and biomass at that reached 98% and 84% respectively. Significant spatial heterogeneity of E. huxleyi, quantitative distribution and population size structure, as well as asynchronism in population development in neighboring parts of the bloom area were shown. The time period, during which population structure in certain part of the area shifts from domination of juvenile cells without coccoliths to a phase of active detritus formation with dying coccolithophorid cells involved, may be estimated as two weeks. A conclusion is made that after anomalous E. huxleyi bloom in 1997 mass development of coccolithophorids became a characteristic feature of phytoplankton community's seasonal succession in the middle part of the Eastern Bering Sea shelf.

Benthic foraminiferal assemblages of middle to late Pliocene sediments in the North Atlantic Ocean

Records of benthic foraminifera from North Atlantic DSDP Site 607 and Hole 610A indicate changes in deep water conditions through the middle to late Pliocene (3.15 to 2.85 Ma). Quantitative analyses of modem associations in the North Atlantic indicate that seven species, Fontbotia wuellerstorfi, Cibicidoides kullenbergi, Uvigerina peregrina, Nuttallides umboniferus, Melonis pompilioides, Globocassidulina subglobosa and Epistominella exigua are useful for paleoenvironmental interpretation. The western North Atlantic basin (Site 607) was occupied by North Atlantic Deep Water (NADW) until c. 2.88 Ma. At that time, N. umboniferus increased, indicating an influx of Southern Ocean Water (SOW). The eastern North Atlantic basin (Hole 610A) was occupied by a relatively warm water mass, possibly Northeastern Atlantic Deep Water (NEADW), through c. 2.94 Ma when SOW more strongly influenced the site. These interpretations are consistent with benthic delta18O and delta13C records from 607 and 610A (Raymo et al., 1992). The results presented in this paper suggest that the North Atlantic was strongly influenced by northern component deep water circulation until 2.90-2.95 Ma. After that there was a transition toward a glacially driven North Atlantic circulation more strongly influenced by SOW associated with the onset of Northern Hemisphere glaciation. The circulation change follows the last significant SST and atmospheric warming prior to c. 2.6 Ma.

Reconstruction of sea-surface conditions at middle to high latitudes of the Northern Hemisphere

A new calibration database of census counts of organic-walled dinoflagellate cyst (dinocyst) assemblages has been developed from the analyses of surface sediment samples collected at middle to high latitudes of the Northern Hemisphere after standardisation of taxonomy and laboratory procedures. The database comprises 940 reference data points from the North Atlantic, Arctic and North Pacific oceans and their adjacent seas, including the Mediterranean Sea, as well as epicontinental environments such as the Estuary and Gulf of St. Lawrence, the Bering Sea and the Hudson Bay. The relative abundance of taxa was analysed to describe the distribution of assemblages. The best analogue technique was used for the reconstruction of Last Glacial Maximum (LGM) sea-surface temperature and salinity during summer and winter, in addition to sea-ice cover extent, at sites from the North Atlantic (n=63), Mediterranean Sea (n=1) and eastern North Pacific (n=1). Three of the North Atlantic cores, from the continental margin of eastern Canada, revealed a barren LGM interval, probably because of quasi-permanent sea ice. Six other cores from the Greenland and Norwegian seas were excluded from the compilation because of too sparse assemblages and poor analogue situation. At the remaining sites (n= 54), relatively close modern analogues were found for most LGM samples, which allowed reconstructions. The new LGM results are consistent with previous reconstructions based on dinocyst data, which show much cooler conditions than at present along the continental margins of Canada and Europe, but sharp gradients of increasing temperature offshore. The results also suggest low salinity and larger than present contrasts in seasonal temperatures with colder winters and more extensive sea-ice cover, whereas relatively warm conditions may have prevailed offshore in summer. From these data, we hypothesise low thermal inertia in a shallow and low-density surface water layer.

Stable isotope record of Cibicidoides spp. from early and middle Miocene sediments

The middle Miocene delta18O increase represents a fundamental change in the ocean-atmosphere system which, like late Pleistocene climates, may be related to deepwater circulation patterns. There has been some debate concerning the early to early middle Miocene deepwater circulation patterns. Specifically, recent discussions have focused on the relative roles of Northern Component Water (NCW) production and warm, saline deep water originating in the eastern Tethys. Our time series and time slice reconstructions indicate that NCW and Tethyan outflow water, two relatively warm deepwater masses, were produced from ~20 to 16 Ma. NCW was produced again from 12.5 to 10.5 Ma. Another feature of the early and middle Miocene oceans was the presence of a high delta13C intermediate water mass in the southern hemisphere, which apparently originated in the Southern Ocean. Miocene climates appear to be related directly to deepwater circulation changes. Deep-waters warmed in the early Miocene by ~3°C (?20 to 16 Ma) and cooled by a similar amount during the middle Miocene delta18O increase (14.8 to 12.6 Ma), corresponding to the increase (?20 Ma) and subsequent decrease (~16 Ma) in the production of NCW and Tethyan outflow water. Large (>0.6 per mil), relatively rapid (~0.5 m.y.) delta18O increases in both benthic and planktonic foraminifera (i.e., the Mi zones of Miller et al. (1991a) and Wright and Miller (1992a)) were superimposed in the long-term deepwater temperature changes; they are interpreted as reflecting continental ice growth events. Seven of these m.y. glacial/interglacial cycles have been recognized in the early to middle Miocene. Two of these glacial/interglacial cycles (Mi3 and Mi4) combined with a 2° to 3°C decrease in deepwater temperatures to produce the middle Miocene delta18O shift.

Revision of the early-middle Pleistocene calcareous nannofossil biochronology

The extant nannofossil biostratigraphic and biochronologic framework for the early-middle Pleistocene time interval has been tested through the micropaleontological analysis of globally distributed high-quality low- to mid-latitude deep-sea successions. The quantitative temporal distribution patterns of relative abundances of selected taxa were reconstructed in critical intervals, and the following biohorizons were defined: first occurrence of medium-sized Gephyrocapsa spp. (bmG); last occurrence of Calcidiscus macintyrei (tCm); first occurrence of large Gephyrocapsa spp. (blG); last occurrence of large Gephyrocapsa spp. (tlG); first occurrence of Reticulofenestra asanoi (bRa); re-entrance of medium-sized Gephyrocapsa spp. (reemG) and last occurrence of Reticulofenestra asanoi (tRa). The detailed patterns of abundance change at these biohorizons were used to generate a detailed biostratigraphy, and the biostratigraphic data were transformed into a precise biochronology by means of correlation to isotope stratigraphies and astronomical timescales. The degree of isochrony or diachrony of the biohorizons was evaluated. Biohorizons tlG and tRa are isochronous occurring close to marine isotope stages (MIS)55 and MIS 22, respectively, and bmG and blG are slightly diachronous on the order of 30-40 kyr, whereas biohorizons tCm, reemG and bRa are confirmed as diachronous on the order of 100, 80 and 60 kyr, respectively. Some of the events are clearly controlled by environmental conditions, e.g. the last occurrence of R. asanoi, related to significant environmental changes associated with the first large-amplitude glaciation of the late Quaternary, MIS 22.

Composition of bioclastic sands, carbonates and pyroclastic rocks of the Great Meteor and Josephine Seamounts, eastern North Atlantic

1. Great Meteor Seamount (GMS) is a very large (24,000 km**3) guyot with a flat summit plateau at 330-275 m; it has a volcanic core, capped by 150-600 m of post-Middle-Miocene carbonate and pyroclastic rocks, and is covered by bioclastic sands. The much smaller Josephine Seamount (JS, summit 170- 500 m w. d.) consists mainly of basalt which is only locally covered by limestones and bioclastic sands. 2. The bioclastic sands are almost free of terrigenous components, and are well sorted, unimodal medium sands. (1) "Recent pelagic sands" are typical of water depths > 600 m (JS) or > 1000 m (GMS). (2) "Sands of mixed relict-recent origin" (10-40% relict) and (3) "relict sands" (> 40% relict) are highly reworked, coarse lag deposits from the upper flanks and summit tops in which recent constituents are mixed with Pleistocene or older relict material. 3. From the carbonate rocks of both seamounts, 12 "microfacies" (MF-)types were distinguished. The 4 major types are: (1) Bio(pel)sparites (MF 1) occur on the summit plateaus and consist of magnesian calcite cementing small pellets and either redeposited planktonic bioclasts or mixed benthonic-planktonic skeletal debris ; (2) Porous biomicrites (MF 2) are typical of the marginal parts of the summit plateaus and contain mostly planktonic foraminifera (and pteropods), sometimes with redeposited bioclasts and/or coated grains; (3) Dense, ferruginous coralline-algal biomicrudites with Amphistegina sp. (MF 3.1), or with tuffaceous components (MF 3.2); (4) Dense, pelagic foraminiferal nannomicrite (MF 4) with scattered siderite rhombs. Corresponding to the proportion and mineralogical composition of the bioclasts and of the (Mgcalcitic) peloids, micrite, and cement, magnesian calcite (13-17 mol-% MgCO3) is much more abundant than low-Mg calcite and aragonite in rock types (1) and (2). Type (3) contains an "intermediate" Mg-calcite (7-9 mol-X), possibly due to an original Mg deficiency or to partial exsolution of Mg during diagenesis. The nannomicrite (4) consists of low-Mg calcite only. 4. Three textural types of volcanic and associated gyroclastic rocks were distinguished: (1) holohyaline, rapidly chilled and granulated lava flows and tuffs (palagonite tuff breccia and hyaloclastic top breccia); (2) tachylitic basalts (less rapidly chilled; with opaque glass); and (3) "slowly" crystallized, holocrystalline alkali olivine basalts. The carbonate in most mixed pyroclastic-carbonate sediments at the basalt contact is of "post-eruptive" origin (micritic crusts etc.); "pre-eruptive" limestone is recrystallized or altered at the basalt contact. A deuteric (?hydrothermal) "mineralX", filling vesicles in basalt and cementing pyroclastic breccias is described for the first time. 5. Origin and development of GMS andJS: From its origin, some 85 m. y. ago, the volcano of GMS remained active until about 10 m. y. B. P. with an average lava discharge of 320 km**3/m. y. The volcanic origin of JS is much younger (?Middle Tertiary), but the volcanic activity ended also about 9 m. y. ago. During L a t e Miocene to Pliocene times both volcanoes were eroded (wave-rounded cobbles). The oldest pyroclastics and carbonates (MF 3.1, 3.2) were originally deposited in shallow-water (?algal reef hardground). The Plio (-Pleisto) cene foraminiferal nannomicrites (MF 4) suggest a meso- to bathypelagic environment along the flanks of GMS. During the Quaternary (?Pleistocene) bioclastic sands were deposited in water depths beyond wave base on the summit tops, repeatedly reworked, and lithified into loosely consolidated biopelsparites and biomicrites (MF 1 and 2; Fig. 15). Intermediate steps were a first intragranular filling by micrite, reworking, oncoidal coating, weak consolidation with Mg-calcite cemented "peloids" in intergranular voids and local compaction of the peloids into cryptocrystalline micrite with interlocking Mg-calcite crystals up to 4p. The submarine lithification process was frequently interrupted by long intervals of nondeposition, dissolution, boring, and later infilling. The limestones were probably never subaerially exposed. Presently, the carbonate rocks undergo biogenic incrustation and partial dissolution into bioclastic sands. The irregular distribution pattern of the sands reflects (a) the patchy distribution of living benthonic organisms, (b) the steady rain of planktonic organism onto the seamount top, (c) the composition of disintegrating subrecent limestones, and (d) the intensity of winnowing and reworking bottom current

Sedimentation patters in the eastern equatorial Pacific Ocean

The post-middle Miocene evolution of sedimentary patterns in the eastern equatorial Pacific Ocean has been deduced from a compilation and synthesis of CaCO3, opal, and nannofossil assemblage data from 11 sites drilled during Leg 138. Improvements in stratigraphic correlation and time scale development enabled the construction of lithostratigraphic and chronostratigraphic frameworks of exceptional quality. These frameworks, and the high sedimentation rates (often exceeding 4 cm/k.y.) provided a detailed and synoptic paleoceanographic view of a large and highly productive region. The three highlights that emerge are: (1) a middle late Miocene "carbonate crash" (Lyle et al., this volume); (2) a late Miocene-early Pliocene "biogenic bloom"; and (3) an early Pliocene "opal shift". During the carbonate crash, an interval of dissolution extending from -11.2 to 7.5 Ma, CaCO3 accumulation rates declined to near zero over much of the eastern equatorial Pacific, whereas opal accumulation rates remained substantially unchanged. The crash nadir, near 9.5 Ma, was marked by a brief shoaling of the regional carbonate compensation depth by more than 1400 m. The carbonate crash has been correlated over the entire tropical Pacific Ocean, and has been attributed to tectonically-induced changes in abyssal flow through the Panamanian seaway. The biogenic bloom extended from 6.7 to 4.5 Ma, and was characterized by an overall increase in biogenic accumulation and by a steepening of the latitudinal accumulation gradient toward the equator. The bloom has been observed over a large portion of the global ocean and has been linked to increased productivity. The final highlight, is a distinct and permanent shift in the locus of maximum opal mass accumulation rate at 4.4 Ma. This shift was temporally, and perhaps causally, linked to the final closure of the Panamanian seaway. Before 4.4 Ma, opal accumulation was greatest in the eastern equatorial Pacific Basin (near 0°N, 107°W). Since then, the highest opal fluxes in the equatorial Pacific have occurred in the Galapagos region (near 3°S, 92°W).

Isotopic-geochemical features of spontaneous gases from mud volcanoes in Eastern Georgia

Isotopic-geochemical study revealed presence of mantle He (3He/4He up to 223x10**-8) in gases from mud volcanoes of Eastern Georgia. This fact confirms that the Middle Kura basin fill encloses an intrusive body previously distinguished from geophysical data. Wide variations of carbon isotopic composition d13C in CH4 and CO2 and chemical composition of gas and water at temporally constant 3He/4He ratio indicate their relation with crustal processes. Unusual direct correlations of 3He/4He ratio with concentrations of He and CH4 and 40Ar/36Ar ratio can be explained by generation of gas in the Cenozoic sequence of the Middle Kura basin.

Stable isotope record, opal accumulation rate and geochemistry of middle Miocene sediments of IODP Site 321-U1338

During the middle Miocene, Earth's climate transitioned from a relatively warm phase (Miocene climatic optimum) into a colder mode with re-establishment of permanent ice sheets on Antarctica, thus marking a fundamental step in Cenozoic cooling. Carbon sequestration and atmospheric CO2 drawdown through increased terrestrial and/or marine productivity have been proposed as the main drivers of this fundamental transition. We integrate high-resolution (1-3 k.y.) benthic stable isotope data with XRF-scanner derived biogenic silica and carbonate accumulation estimates in an exceptionally well-preserved sedimentary archive, recovered at Integrated Ocean Drilling Program Site U1338, to reconstruct eastern equatorial Pacific productivity variations and to investigate temporal linkages between high- and low-latitude climate change over the interval 16-13 Ma. Our records show that the climatic optimum (16.8-14.7 Ma) was characterized by high amplitude climate variations, marked by intense perturbations of the carbon cycle. Episodes of peak warmth at (southern hemisphere) insolation maxima coincided with transient shoaling of the carbonate compensation depth and enhanced carbonate dissolution in the deep ocean. A switch to obliquity-paced climate variability after 14.7 Ma concurred with a general improvement in carbonate preservation and the onset of stepwise global cooling, culminating with extensive ice growth over Antarctica at ~13.8 Ma. We find that two massive increases in opal accumulation at ~14.0 and ~13.8 Ma occurred just before and during the final and most prominent cooling step, supporting the hypothesis that enhanced siliceous productivity in the eastern equatorial Pacific contributed to CO2 drawdown.

Stable isotope composition of Holocene benthic foraminifers from the Eastern Mediterranean Sea

Temporal and regional changes in paleoproductivity and paleoceanography in the eastern Mediterranean Sea during the past 12 kyr were reconstructed on the basis of the stable oxygen and carbon isotope composition of the epibenthic Planulina ariminensis and the shallow endobenthic Uvigerina mediterranea from three sediment cores of the Aegean Sea and Levantine Basin. The Younger Dryas is characterized by high d18O values, indicating enhanced salinities and low temperatures of deep water masses at all investigated sites. With the onset of the Holocene, d18O records show a continuous decrease towards the onset of sapropel S1 formation, mainly caused by a freshening and warming of surface waters at deep water formation sites. In the middle and late Holocene, the similarity of d18O values from the southern Aegean Sea and Levantine Basin suggests the influence of isotopically identical deep water masses. By contrast, slightly higher d18O values are observed the northern Aegean Sea, which probably point to lower temperatures of North Aegean deep waters. The epifaunal d13C records reveal clear changes in sources and residence times of eastern Mediterranean deep waters associated with period of S1 formation. Available data for the early and late phase of sapropel S1 formation and for the interruption around 8.2 kyr display drops by 0.5 and 1.5 per mil, indicating the slow-down of deep water circulation and enhanced riverine input of isotopically light dissolved inorganic carbon from terrestrial sources into the eastern Mediterranean Sea. The decrease in epifaunal d13C signals is particularly expressed in the southern Aegean Sea and Levantine Basin, while it is less pronounced in the northern Aegean Sea. This points to a strong reduction in deep water exchange rates in the southern areas, but the persistence of local deep water formation in the northern Aegean Sea. The d13C values of U. mediterranea records reveal temporal and regional differences in paleoproductivity during the past 12 kyr, with rather eutrophic and mesotrophic conditions in the North Aegean Sea and southeast Levantine Basin, respectively, while the South Aegean Sea is characterized by rather oligotrophic conditions. After S1 formation, increasing d13C values reflect a progressive decrease in surface water productivity in the eastern Mediterranean Sea during the middle and late Holocene. In the northern Aegean Sea, this time interval is marked by repetitive changes in organic matter fluxes documented by significant fluctuations in the d13C signal of U. mediterranea on millennial- to multi-centennial time scales. These fluctuations can be linked to short-term changes in river runoff driven by northern hemisphere climatic variability.

Benthic foraminifera in sapropels of the eastern Mediterranean Sea

High-resolution benthic foraminiferal and geochemical investigations were carried out across sapropels S5 and S6 from two sediment cores in the Levantine Sea to evaluate the impact of climatic and environmental changes on benthic ecosystems during times of sapropel formation. The faunal successions indicate that eutrophication and/or oxygen reduction started several thousand years prior to the onset of sapropel formation, suggesting an early response of the bathyal ecosystems to climatic changes. Severest oxygen depletions appear in the early phases of sapropel formation. The initial reduction of deep-water ventilation is caused by a warming and fresh water-induced stratification of Eastern Mediterranean surface waters. During the late phase of S5 formation improved oxygenation is restricted to middle bathyal ecosystems, indicating that at least some formation of subsurface water took place. During S6 formation oxygen depletions and eutrophication were less severe and more variable than during S5 formation. Estimated oxygen contents were low dysoxic at middle bathyal to anoxic at lower bathyal depths during the early phase of S6 formation but never dropped to anoxic values in its late phase. The high benthic ecosystem variability during S6 formation suggests that water column stratification at deep-water formation sites was in a very unstable mode and susceptible to minor temperature fluctuations at a millennial time-scale.