Stable carbon and oxygen isotope compositions of Eocene benthic foraminifera from ODP Site 207-1258

'Hyperthermals' are intervals of rapid, pronounced global warming known from six episodes within the Palaeocene and Eocene epochs (~65-34 million years (Myr) ago) (Zachos et al., 2005, doi:10.1126/science.1109004; 2008, doi:10.1038/nature06588; Roehl et al., 2007, doi:10.1029/2007GC001784; Thomas et al., 2000; Cramer et al., 2003, doi:10.1029/2003PA000909; Lourens et al., 2005, doi:10.1038/nature03814; Petrizzo, 2005, doi:10.2973/odp.proc.sr.198.102.2005; Sexton et al., 2006, doi:10.1029/2005PA001253; Westerhold et al., 2007, doi:10.1029/2006PA001322; Edgar et al., 2007, doi:10.1038/nature06053; Nicolo et al., 2007, doi:10.1130/G23648A.1; Quillévéré et al., 2008, doi:10.1016/j.epsl.2007.10.040; Stap et al., 2010, doi:10.1130/G30777.1). The most extreme hyperthermal was the 170 thousand year (kyr) interval (Roehl et al., 2007) of 5-7 °C global warming (Zachos et al., 2008) during the Palaeocene-Eocene Thermal Maximum (PETM, 56 Myr ago). The PETM is widely attributed to massive release of greenhouse gases from buried sedimentary carbon reservoirs (Zachos et al., 2005; 2008; Lourenbs et al., 2005; Nicolo et al., 2007; Dickens et al., 1995, doi:10.1029/95PA02087; Dickens, 2000; 2003, doi:10.1016/S0012-821X(03)00325-X; Panchuk et al., 2008, doi:10.1130/G24474A.1) and other, comparatively modest, hyperthermals have also been linked to the release of sedimentary carbon (Zachos et al., 2008, Lourens et al., 2005; Nicolo et al., 2007; Dickens, 2003; Panchuk et al., 2003). Here we show, using new 2.4-Myr-long Eocene deep ocean records, that the comparatively modest hyperthermals are much more numerous than previously documented, paced by the eccentricity of Earth's orbit and have shorter durations (~40 kyr) and more rapid recovery phases than the PETM. These findings point to the operation of fundamentally different forcing and feedback mechanisms than for the PETM, involving redistribution of carbon among Earth's readily exchangeable surface reservoirs rather than carbon exhumation from, and subsequent burial back into, the sedimentary reservoir. Specifically, we interpret our records to indicate repeated, large-scale releases of dissolved organic carbon (at least 1,600 gigatonnes) from the ocean by ventilation (strengthened oxidation) of the ocean interior. The rapid recovery of the carbon cycle following each Eocene hyperthermal strongly suggests that carbon was resequestered by the ocean, rather than the much slower process of silicate rock weathering proposed for the PETM (Zachos et al., 2005; 2003). Our findings suggest that these pronounced climate warming events were driven not by repeated releases of carbon from buried sedimentary sources (Zachos et al., 2008, Lourens et al., 2005; Nicolo et al., 2007; Dickens, 2003; Panchuk et al., 2003) but, rather, by patterns of surficial carbon redistribution familiar from younger intervals of Earth history.

Late Neogene benthic foraminifera and stable carbon isotope record of the blake Outer Ridge

Carbon isotope and benthic foraminiferal data from Blake Outer Ridge, a sediment drift in the western North Atlantic (Ocean Drilling Program Sites 994 and 997, water depth ~ 2800 m), document variability in the relative volume of Southern Component (SCW) and Northern Component Waters (NCW) over the last 7 Ma. SCW was dominant before ~5.0 Ma, at ~3.6-2.4 Ma, and 1.2-0.8 Ma, whereas NCW dominated in the warm early Pliocene (5.0-3.6 Ma), and at 2.4-1.2 Ma. The relative volume of NCW and SCW fluctuated strongly over the last 0.8 Ma, with strong glacial-interglacial variability. The intensity of the Western Boundary Undercurrent was positively correlated to the relative volume of NCW. Values of Total Organic Carbon (TOC) were > 1.5% in sediments older than ~ 3.8 Ma, and not correlated to high primary productivity indicators, thus may reflect lateral transport of organic matter. TOC values decreased during the intensification of the Northern Hemisphere Glaciation (NHG, 3.8-1.8 Ma). Benthic foraminiferal assemblages underwent major changes when the sites were dominantly under SCW (3.6-2.4 and 1.2-0.8 Ma), coeval with the 'Last Global Extinction' of elongate, cylindrical deep-sea benthic foraminifera, which has been linked to cooling, increased ventilation and changes in the efficiency of the biological pump. These benthic foraminiferal turnovers were neither directly associated with changes in dominant bottom water mass nor with changes in productivity, but occurred during global cooling and increased ventilation of deep waters associated with the intensification of the NHG.

Biometry, stable isotopes and stomach contents of A. glacialis and B. saida from Northeast Greenland

Two gadoid fishes, Arctogadus glacialis and Boreogadus saida, often coexist (i.e. sympatric) in the fjords and shelf areas of the Arctic seas, where they likely share the same food resources. Diet composition from stomach contents, i.e. frequency of occurrence (FO) and Schoener's index (SI), and stable isotope signatures (d13C and d15N) in muscle of these sympatric gadoids were examined from two fjords in NE Greenland-Tyrolerfjord (TF, ~74°N, sill present) and Dove Bugt (DB, ~76°N, open). Twenty-three prey taxa and categories were identified and both gadoids ate mostly crustaceans. The SI values of 0.64-0.70 indicated possible resource competition, whereas FO differed significantly. A. glacialis fed mainly on the mysid Mysis oculata and other benthic-associated prey, whereas B. saida ate the copepod Metridia longa and other pelagic prey. Both diet and stable isotopes strongly suggest a spatial segregation in feeding habitat, with A. glacialis being associated with the benthic food web (mean d13C = -20.81 per mil, d15N = 14.92 per mil) and B. saida with the pelagic food web (mean d13C = -21.25 per mil, d15N = 13.64 per mil). The dietary differences and isotopic signals were highly significant in the secluded TF and less clear in the open DB, where prey and predators may be readily advected from adjacent areas with other trophic conditions. This is the first study on the trophic position of A. glacialis inferred from analyses of stable isotopes. The subtle interaction between the Arctic gadoids should be carefully monitored in the light of ocean warming and ongoing invasions of boreal fishes into the Arctic seas.

Stable hydrogen and carbon isotope records and concentration data for four long chain n-alkanes from core GIK16160-3 derived from the Zambezi delta in the Mozambique Channel during the last 37,000 years

Hydrogen isotope values (dD) of sedimentary terrestrial leaf wax such as n-alkanes or n-acids have been used to map and understand past changes in rainfall amount in the tropics because dD of precipitation is commonly assumed as the first order controlling factor of leaf wax dD. Plant functional types and their photosynthetic pathways can also affect leaf wax dD but these biological effects are rarely taken into account in paleo studies relying on this rainfall proxy. To investigate how biological effects may influence dD values we here present a 37,000-year old record of dD and stable carbon isotopes (d13C) measured on four n-alkanes (n-C27, n-C29, n-C31, n-C33) from a marine sediment core collected off the Zambezi River mouth. Our paleo d13C records suggest that each individual n-alkanes had different C3/C4 proportional contributions. n-C29 was mostly derived from a C3 dicots (trees, shrubs and forbs) dominant vegetation throughout the entire record. In contrast, the longer chain n-C33 and n-C31 were mostly contributed by C4 grasses during the Glacial period but shifted to a mixture of C4 grasses and C3 dicots during the Holocene. Strong correlations between dD and d13C values of n-C33 (correlation coefficient R2 = 0.75, n = 58) and n-C31 (R2 = 0.48, n = 58) suggest that their dD values were strongly influenced by changes in the relative contributions of C3/C4 plant types in contrast to n-C29 (R2 = 0.07, n = 58). Within regions with variable C3/C4 input, we conclude that dD values of n-C29 are the most reliable and unbiased indicator for past changes in rainfall, and that dD and d13C values of n-C31 and n-C33 are sensitive to C3/C4 vegetation changes. Our results demonstrate that a robust interpretation of palaeohydrological data using n-alkane dD requires additional knowledge of regional vegetation changes from which nalkanes are synthesized, and that the combination of dD and d13C values of multiple n-alkanes can help to differentiate biological effects from those related to the hydrological cycle.

(Table 1) Stable oxygen isotope ratios of Globigerinoides sacculifer of ODP Hole 130-806B

A detailed Pliocene oxygen isotope record from the Ontong Java Plateau, based on measurements of the surface-dwelling planktonic foraminifer Globigerinoides sacculifer, was produced for the period from 5 to 2 Ma. The record documents major long- and short-term climate changes. The results show periods of enhanced ice volume at 4.6 to 4.3 Ma and after 2.85 Ma, a long-term warming trend from 4.1 to 3.7 Ma, and a distinct cooling trend that was initiated at 3.5 Ma and progressed through the initiation of large-scale Northern Hemisphere glaciation after 2.85 Ma (according to the time scale of Shackleton and others proposed in 1990). Periods of high average ice volumes also show the highest d18O amplitudes. The pattern of climate cyclicity changed markedly at about 2.85 Ma. Earlier times were marked by high-frequency variability at the precessional frequencies or even higher frequencies, pointing to low-latitude processes as a main controlling factor driving planktonic d18O variability in this period. The high-frequency variability is not coherent with insolation and points to strong nonlinearity in the way the climate system responded to orbital forcing before the onset of large scale Northern Hemisphere glaciation. After 3 Ma, stronger 41-k.y. cyclicity appears in the record. The shift in pattern is clearest around 2.85 Ma (according to the time scale proposed by Shackleton and others in 1990), 100-200 k.y. before the most dramatic spread of Northern Hemisphere ice sheets. This indicates that high-latitude processes from this point on began to take over and influence most strongly the d18O record, which now reflects ice-volume fluctuations related to the climatic effects of obliquity forcing on the seasonality of high-latitude areas, most probably in the Northern Hemisphere. The general Pliocene trend is that high-latitude climate sensitivity and instability was increasing, and the causal factors producing the intensified glacial cyclicity during the Pliocene must be factors that enhance cooling and climate sensitivity in the subarctic areas.

Age determination and stable isotope composition of sediment cores from the Arctic Ocean

On the basis of 52 sediment cores, analyzed and dated at high resolution, the paleoceanography and climate of the Last Glacial Maximum (LGM) were reconstructed in detail for the Fram Strait and the eastern and central Arctic Ocean. Sediment composition and stable isotope data suggest three distinct paleoenvironments: (1) a productive region in the eastern to central Fram Strait and along the northern Barents Sea continental margin characterized by Atlantic Water advection, frequent open water conditions, and occasional local meltwater supply and iceberg calving from the Barents Sea Ice Sheet; (2) an intermediate region in the southwestern Eurasian Basin (up to 84-85°N) and the western Fram Strait characterized by subsurface Atlantic Water advection and recirculation, a moderately high planktic productivity, and a perennial ice cover that breaks up only occasionally; and (3) a central Arctic region (north of 85°N in the Eurasian Basin) characterized by a low-salinity surface water layer and a thick ice cover that strongly reduces bioproduction and bulk sedimentation rates. Although the total inflow of Atlantic Water into the Arctic Ocean may have been reduced during the LGM, its impact on ice coverage and halocline structure in the Fram Strait and southwestern Eurasian Basin was strong.

Stable isotopes and Mg/Ca ratios on foraminifera and Ba/Ca ratios of sediment core SK168/GC-1

The past variability of the South Asian Monsoon is mostly known from records of wind strength over the Arabian Sea while high-resolution paleorecords from regions of strong monsoon precipitation are still lacking. Here, we present records of past monsoon variability obtained from sediment core SK 168/GC-1, which was collected at the Alcock Seamount complex in the Andaman Sea. We utilize the ecological habitats of different planktic foraminiferal species to reconstruct freshwater-induced stratification based on paired Mg/Ca and d18O analyses and to estimate seawater d18O (d18Osw). The difference between surface and thermocline temperatures (delta T) and d18Osw (delta d18Osw) is used to investigate changes in upper ocean stratification. Additionally, Ba/Ca in G. sacculifer tests is used as a direct proxy for riverine runoff and sea surface salinity (SSS) changes related to monsoon precipitation on land. Our delta d18Osw time series reveals that upper ocean salinity stratification did not change significantly throughout the last glacial suggesting little influence of NH insolation changes. The strongest increase in temperature gradients between the mixed layer and the thermocline is recorded for the mid-Holocene and indicate the presence of a significantly shallower thermocline. In line with previous work, the d18Osw and Ba/Ca records demonstrate that monsoon climate during the LGM was characterized by a significantly weaker southwest monsoon circulation and strongly reduced runoff. Based on our data the South Asian Summer Monsoon (SAM) over the Irrawaddyy strengthened gradually after the LGM beginning at ~18 ka. This is some 3 kyrs before an increase of the Ba/Ca record from the Arabian Sea and indicates that South Asian Monsoon climate dynamics are more complex than the simple N-S displacement of the ITCZ as generally described for other regions. Minimum d18Osw values recorded during the mid-Holocene are in phase with Ba/Ca marking a stronger monsoon precipitation, which is consistent with model simulations.

Comparison of the stable carbon and nitrogen isotopic values of gill and white muscle tissue of fish

The potential use of stable carbon and nitrogen isotope ratios (d13C, d15N) of fish gills for studies on fish feeding ecology was evaluated by comparing the d13C and d15N of gill tissue with the more commonly used white muscle tissue. To account for the effect of lipid content on the d13C signatures, a study-specific lipid correction model based on C:N ratios was developed and applied to the bulk d13C data. For the majority of species in the study, we found no significant difference in d13C values between gill and muscle tissue after correction, but several species showed a small (0.3-1.4 per mil) depletion in 13C in white muscle compared to gill tissue. The average species difference in d15N between muscle and gill tissue ranged from -0.2 to 1.6 per mil for the different fish species with muscle tissue generally more enriched in 15N. The d13C values of muscle and gill were strongly linearly correlated (R**2 = 0.85) over a large isotopic range (13 per mil), suggesting that both tissues can be used to determine long-term feeding or migratory habits of fish. Muscle and gill tissue bulk d15N values were also strongly positively correlated (R**2= 0.76) but with a small difference between muscle and gill tissue. This difference indicates that the bulk d15N of the two tissue types may be influenced by different isotopic turnover rates or a different composition of amino acids.

Stable isotope ratios and paleoceanaographic reconstructions from sediment cores 80-548 and 161-978A

Largely continuous millennial-scale records of benthic d18O, Mg/Ca-based temperature, and salinity variations in bottom waters were obtained from Deep Sea Drilling Project Site 548 (eastern Atlantic continental margin south of Ireland, 1250 m water depth) for the period between 3.7 and 3.0 million years ago. This site monitored mid-Pliocene changes in Mediterranean Outflow Water (MOW) documented by continuously high Nd values between -10.7 and -9. Site 978 (Alboran Sea, 1930 m water depth) provides a complementary record of bottom water variability in the westernmost Mediterranean Sea, which is taken to represent MOW composition at its source. Both sites are marked by a singular and persistent rise in bottom water salinities by 0.7-1.4 psu and in densities by ~1 kg m-3 from 3.5 to 3.3 Ma, which is matched by an average 3 °C increase in bottom water temperatures at Site 548. This event suggests the onset of strongly enhanced deep-water convection in the Mediterranean Sea and a related strengthened MOW flow, which implies a major aridification of the Mediterranean source region. In harmony with model suggestions, the enhanced MOW flow has possibly intensified Upper North Atlantic Deep Water formation.

(Table 1) Stable nitrogen isotopes of ammonium in interstitial waters from ODP Site 164-997

Ammonium (NH4+) concentration profiles in piston-core sediments of the Carolina Rise and Blake Ridge generally have linear concentration profiles within the sulfate reduction zone (Borowski, 1998). Deep Sea Drilling Project (DSDP) Site 533, located on the Blake Ridge, also displayed a linear ammonium concentration profile through the sulfate reduction zone and the profile linearity continues into the upper methanogenic zone to a depth of ~200 meters below seafloor (mbsf), where the first methane gas hydrates probably occur (Jenden and Gieskes, 1983, doi:10.2973/dsdp.proc.76.114.1983; Kvenvolden and Barnard, 1983, doi:10.2973/dsdp.proc.76.106.1983). Sediments from the Ocean Drilling Program (ODP) Leg 164 deep holes (Sites 994, 995, and 997) also exhibit linear ammonium profiles above the top of the gas hydrate zone (~200 mbsf) (Paull, Matsumoto, Wallace, et al., 1996, doi:10.2973/odp.proc.ir.164.1996). We hypothesized that a possible cause of linear ammonium profiles was diffusion of ammonium from a concentrated ammonium source at depth. We further reasoned that if this ammonium were produced by microbial fermentation reactions at depth, that a comparison of the nitrogen isotopic composition of sedimentary organic nitrogen and the nitrogen with pore-water ammonium would test this hypothesis. Convergence with depth of d15N values of the nitrogen source (sedimentary organic matter) and the nitrogen product (dissolved NH4+) would strongly suggest that ammonium was produced within a particular depth zone by microbial fermentation reactions. Here, we report d15N values of pore-water ammonium from selected interstitial water (IW) samples from Site 997, sampled during ODP Leg 164.

Results from a stable isotope tracer experiment in Potter Cove, King George Island, Antarctica

Antarctic meiofauna is still strongly understud- ied, and so is its trophic position in the food web. Primary producers, such as phytoplankton, and bacteria may repre- sent important food sources for shallow water metazoans, and the role of meiobenthos in the benthic-pelagic coupling represents an important brick for food web understanding. In a laboratory, feeding experiment 13C-labeled freeze- dried diatoms (Thalassiosira weissflogii) and bacteria were added to retrieved cores from Potter Cove (15-m depth, November 2007) in order to investigate the uptake of 3 main meiofauna taxa: nematodes, copepods and cumaceans. In the surface sediment layers, nematodes showed no real difference in uptake of both food sources. This outcome was supported by the natural delta 13C values and the community genus composition. In the first centimeter layer, the dominant genus was Daptonema which is known to be opportunistic, feeding on both bacteria and diatoms. Copepods and cumaceans on the other hand appeared to feed more on diatoms than on bacteria. This may point at a better adaptation to input of primary production from the water column. On the other hand, the overall carbon uptake of the given food sources was quite low for all taxa, indicating that likely other food sources might be of relevance for these meiobenthic organisms. Further studies are needed in order to better quantify the carbon requirements of these organisms.

Stable carbon and oxygen isotope ratios of foraminifera from Cretaceous and Paleocene sediments

Detailed analysis of over 200 samples of uppermost Cretaceous and Paleocene sediments from Atlantic Ocean DSDP Sites 384, 86, 95, 152, 144, 20C, 21, 356, 357, and 329 provides new information on the temperature stratification of Paleocene planktonic foraminifera, the temperature and carbon isotopic changes across the Cretaceous/Tertiary boundary, and the fluctuating temperature and carbon isotopic records through the Paleocene ~64.5-54 m.y.). There was a significant temperature rise across the Cretaceous/Tertiary boundary both at the surface and in deep waters of the Atlantic Ocean. This temperature rise occurred before the basal Tertiary 'Globigerina' eugubina Zone, so that in the oldest Paleocene sample yet analyzed from the deep sea (Site 356) temperatures are already three degrees higher at the bottom and at the surface than in the Cretaceous. The temperature rise across the boundaryis more pronounced on the bottom and in samples from higher latitudes. Accompanying the temperature rise across the boundary there is a significant shift in the carbon isotope profile. In the basal Paleocene the foraminifera of the surface zone demonstrate very negative carbon isotope values (unlike in the Cretaceous of today's ocean), while deeper dwelling species have more positive values which then decrease to the bottom. The unusual carbon isotope gradients persist through the first three million years of the Paleocene until towards the top of planktonic foraminiferal Zone P.1 (G. trinidadensis Zone) the foraminifera record a profile more positive at the surface and decreasing towards the bottom (as in today's ocean). During the Paleocene there are two noteworthy rises in surface water temperature; the first around 62-61 m.y. (G. trinidadensis Zone), and the second near the base of the Globorotalia angulata Zone, 60-59 m.y. At this time surface temperatures at low to mid latitudes reached values near 25°C, while at mid-latitude Site 384 temperature highs near 22°C were registered. At a sample spacing of around one per million years, we have only produced some of the detail of these temperature fluctuations. The later Paleocene is generally cooler and there do not seem to be any large variations either through time or latitude. Middle-latitude sites average temperatures near 15°C at the surface, while high lower latitude site temperatures range near 18°C. The most salient feature of the bottom temperature record (based on multispecific samples) through the Paleocene is its lack of fluctuations. There is an overall temperature range of 5°C at these intermediate depth sites (paleodepth estimates between 1500 and 3000 m). Higher values near 13°C accompany the surface temperature peaks around 62 and 60 m.y., while low values near 8°C occur in Zone P.2 (61-60 m.y.). We detected no change in bottom temperature across the paleocene/Eocene boundary in the few samples studied so far. While there are several fluctuations in the carbon isotope values through the early Paleocene, the general trend is one of increasingly positive values at the surface and at depth. This trend culminates in the late Paleocene (upper Zone P.4, about 56-57 m.y.) with a major excursion in the carbon isotope values. At low latitudes the range between the surface and the deepest planktonic foraminifera is a delta13C of 4 per mil as compared with a range of 2 per mil today. The carbon values drop off slightly, but remain strongly positive through the remainder of the Paleocene at most sites. Accompanying the carbon isotope excursion at Site 384 is a productivity increase and a proposed rise in the CCD.

Stable isotope record of Uvigerina peregrina from the Mediterranean Sea

Based on the glacial to postglacial delta13C differences between endobenthic Uvigerina peregrina species from the Alboran basin and from other mediterranean basins, changes in the fertility of the western part of this basin during the last deglaciation are reconstructed. As a result of particulate organic carbon (POC) rain from the highly productive upwelling cell along the northwestern margin of the Alboran basin, U. peregrina is presently depleted by about 1.6per mil with respect to the measured delta13C values of bottom water SumCO2 and by about 0.9per mil with respect to specimens from other areas of the western Mediterranean or from the Gulf of Cadiz within the Mediterranean Outflow Water. The Uvigerina delta13C difference between the Alboran Sea and the Gulf of Cadiz (Delta delta13C), was close to 0per mil at the beginning of the last deglaciation and during the late glacial time. This suggests that highly fertile systems set in the Alboran Sea near 16 kyr B.P. Two rapid increases in the Delta delta13C offset are recorded near 15 kyr and 11 kyr B.P. Fluctuations around 1.1 to 1.2per mil occurred during the early Holocene, and a maximum was reached near 9 kyr B.P. After 4 kyr the Delta delta13C offset decreased to its present-day average value of 0.9per mil. Changes in the intensity of surficial production cannot account for all the observed fluctuations, especially in the early Holocene time. A strong decrease in the intermediate and deep water ventilation of the Alboran basin may have occurred near 8-9 kyr, in phase with the last stagnant phase in the eastern Mediterranean and the deposition of Sapropel S1. As a result, the redistribution and remineralization at depth of the produced organic matter was incomplete. The POC rain reaching the sediment was locally intensified and caused the lowering of the delta13C values of endobenthic foraminifers such as U. peregrina. The benthic 13C signal suggests that the difference between the Alboran Sea and the Gulf of Cadiz was at its maximum. At the same time, an important modification in the water masses structure may have occurred near 9-8 kyr B.P. The deepening of the permanent pycnocline probably related to a thicker Atlantic jet at a stage of high sea level stand is recorded by the replacement of the right coiling N. pachyderma dominance (coincident with a shallow pycnocline) by the G. inflata dominance (coincident with a deep pycnocline). Diatom abundances were strongly reduced indicating an important modification of the productive system. The glacial-postglacial evolution of productivity within the Alboran basin was therefore more complex than in the adjacent Atlantic Ocean and opposite to the global one which displays a general increase in productivity during glacial time. Although it is the global budget of paleoproductivity that would drive the partitioning of carbon within the ocean, local or regional discrepancies with the global glacial-interglacial model must be addressed. Local winds and regional atmospheric pressure systems, which are the forcing factors for circulation and exchange with the Atlantic, control the fertile systems of the Alboran basin.

Sedimentology, age models and stable isotope ratios of sediment cores from the equatorial Pacific

Based on detailed reconstructions of global distribution patterns, both paleoproductivity and the benthic d13C record of CO2, which is dissolved in the deep ocean, strongly differed between the Last Glacial Maximum and the Holocene. With the onset of Termination I about 15,000 years ago, the new (export) production of low- and mid-latitude upwelling cells started to decline by more than 2-4 Gt carbon/year. This reduction is regarded as a main factor leading to both the simultaneous rise in atmospheric CO2 as recorded in ice cores and, with a slight delay of more than 1000 years, to a large-scale gradual CO2 depletion of the deep ocean by about 650 Gt C. This estimate is based on an average increase in benthic d13C by 0.4-0.5 per mil. The decrease in new production also matches a clear 13C depletion of organic matter, possibly recording an end of extreme nutrient utilization in upwelling cells. As shown by Sarnthein et al., [1987], the productivity reversal appears to be triggered by a rapid reduction in the strength of meridional trades, which in turn was linked via a shrinking extent of sea ice to a massive increase in high-latitude insolation, i.e., to orbital forcing as primary cause.

(Table S1) Stable isotope record and growth rates of cultured Mytilus edulis

To study the effects of temperature, salinity, and life processes (growth rates, size, metabolic effects, and physiological/genetic effects) on newly precipitated bivalve carbonate, we quantified shell isotopic chemistry of adult and juvenile animals of the intertidal bivalve Mytilus edulis (Blue mussel) collected alive from western Greenland and the central Gulf of Maine and cultured them under controlled conditions. Data for juvenile and adult M. edulis bivalves cultured in this study, and previously by Wanamaker et al. (2006, doi:10.1029/2005GC001189), yielded statistically identical paleotemperature relationships. On the basis of these experiments we have developed a species-specific paleotemperature equation for the bivalve M. edulis [T °C = 16.28 (±0.10) - 4.57 (±0.15) {d18Oc VPBD - d18Ow VSMOW} + 0.06 (±0.06) {d18Oc VPBD - d18Ow VSMOW}**2; r**2 = 0.99; N = 323; p < 0.0001]. Compared to the Kim and O'Neil (1997) inorganic calcite equation, M. edulis deposits its shell in isotope equilibrium (d18Ocalcite) with ambient water. Carbon isotopes (d13Ccalcite) from sampled shells were substantially more negative than predicted values, indicating an uptake of metabolic carbon into shell carbonate, and d13Ccalcite disequilibrium increased with increasing salinity. Sampled shells of M. edulis showed no significant trends in d18Ocalcite based on size, cultured growth rates, or geographic collection location, suggesting that vital effects do not affect d18Ocalcite in M. edulis. The broad modern and paleogeographic distribution of this bivalve, its abundance during the Holocene, and the lack of an intraspecies physiologic isotope effect demonstrated here make it an ideal nearshore paleoceanographic proxy throughout much of the North Atlantic Ocean.