Holocene millenial-scale productivity record in the Mediterranean Sea

The calcareous nannofossil assemblages of Ocean Drilling Program Hole 963D from the central Mediterranean Sea have been investigated to document oceanographic changes in surface waters. The studied site is located in an area sensitive to large-scale atmospheric and climatic systems and to high- and low-latitude climate connection. It is characterized by a high sedimentation rate (the achieved mean sampling resolution is <70 years) that allowed the Sicily Channel environmental changes to be examined in great detail over the last 12 ka BP. We focused on the species Florisphaera profunda that lives in the lower photic zone. Its distribution pattern shows repeated abundance fluctuations of about 10-15%. Such variations could be related to different primary production levels, given that the study of the distribution of this species on the Sicily Channel seafloor demonstrates the significant correlation to productivity changes as provided by satellite imagery. Productivity variations were quantitatively estimated and were interpreted on the basis of the relocation of the nutricline within the photic zone, led by the dynamics of the summer thermocline. Productivity changes were compared with oceanographic, atmospheric, and cosmogenic nuclide proxies. The good match with Holocene master records, as with ice-rafted detritus in the subpolar North Atlantic, and the near-1500-year periodicity suggest that the Sicily Channel environment responded to worldwide climate anomalies. Enhanced Northern Hemisphere atmospheric circulation, which has been reported as one of the most important forcing mechanisms for Holocene coolings in previous Mediterranean studies, had a remarkable impact on the water column dynamics of the Sicily Channel.

Faunal analysis and Mg/Ca records of planktic foraminifera in sediment core SO189/2_039KL

Climate phenomena like the monsoon system, El Niño Southern Oscillation (ENSO) and the Indian Ocean Dipole (IOD) are interconnected via various feedback mechanisms and control the climate of the Indian Ocean and its surrounding continents on various timescales. The eastern tropical Indian Ocean is a key area for the interplay of these phenomena and for reconstructing their past changes and forcing mechanisms. Here we present records of upper ocean thermal gradient, thermocline temperatures (TT) and relative abundances of planktic foraminifera in core SO 189-39KL taken off western Sumatra (0°47.400' S, 99°54.510' E) for the last 8 ka that we use as proxies for changes in upper ocean structure. The records suggest a deeper thermocline between 8 ka and ca 3 ka compared to the late Holocene. We find a shoaling of the thermocline after 3 ka, most likely indicating an increased occurrence of upwelling during the late Holocene compared to the mid-Holocene which might represent changes in the IOD-like mean state of the Indian Ocean with a more negative IOD-like mean state during the mid-Holocene and a more positive IOD-like mean state during the past 3 ka. This interpretation is supported by a transient Holocene climate model simulation in which an IOD-like mode is identified that involves an insolation-forced long-term trend of increasing anomalous surface easterlies over the equatorial eastern Indian Ocean.

Mg/Ca and Sr/Ca ratios of Hoeglundina elegans from surface sediments

Core top samples from Atlantic (Little Bahama Banks (LBB)) and Pacific (Hawaii and Indonesia) depth transects have been analyzed in order to assess the influence of bottom water temperature (BWT) and aragonite saturation levels on Mg/Ca and Sr/Ca ratios in the aragonitic benthic foraminifer Hoeglundina elegans. Both the Mg/Ca and Sr/Ca ratios in H. elegans tests show a general decrease with increasing water depth. Although at each site the decreasing trends are consistent with the in situ temperature profile, Mg/Ca and Sr/Ca ratios in LBB are substantially higher than in Indonesia and Hawaii at comparable water depths with a greater difference observed with increasing water depth. Because we find no significant difference between results obtained on "live" and "dead" specimens, we propose that these differences are due to primary effects on the metal uptake during test formation. Evaluation of the water column properties at each site suggests that in situ CO3 ion concentrations play an important role in determining the H. elegans Mg/Ca and Sr/Ca ratios. The CO3 ion effect is limited, however, only to aragonite saturation levels ([DeltaCO3]aragonite) below 15 µmol/kg. Above this level, temperature exerts a dominant effect. Accordingly, we propose that Mg/Ca and Sr/Ca in H. elegans tests can be used to reconstruct thermocline temperatures only in waters oversaturated with respect to the mineral aragonite using the following relationships: Mg/Ca = (0.034 ± 0.002)BWT + (0.96 ± 0.03) and Sr/Ca = (0.060 ± 0.002)BWT + (1.53 ± 0.03) (for [DeltaCO3]aragonite > 15 µmol/kg). The standard error associated with these equations is about ±1.1°C. Reconstruction of deeper water temperatures is complicated because in undersaturated waters, changes in Mg/Ca and Sr/Ca ratios reflect a combination of changes in [CO3] and BWT. Overall, we find that Sr/Ca, rather than Mg/Ca, in H. elegans may be a more accurate proxy for reconstructing paleotemperatures.

Pleistocene stable isotope record of planktonic foraminifera of ODP Site 138-847

We present Pleistocene oxygen and carbon isotope records from two planktonic foraminifer species (Globigerinoides sacculifer and Neogloboquadrina dutertrei) from Ocean Drilling Program Site 847 (0°16'N, 95°19'W; 3334 m water depth). An average sample resolution of 4500 yr was obtained by sampling at an interval of 15 cm through a continuous 35-m section from 0 to 1.15 Ma. Our d18O-based chronology is similar to that derived independently by astronomically tuning the gamma-ray attenuation porosity evaluator (GRAPE) record (Shackleton et al., 1995), though offsets as large as ± 30 k.y. occur on occasion. The surface waters at eastern equatorial Pacific Site 847, 380 km west of the Galapagos, are characterized by strong and constant upwelling, elevated nutrient concentrations, and high productivity. The isotopic composition of G. sacculifer (300-355 µm) reflects conditions in the thin-surface mixed layer, and the composition of N. dutertrei (355-425 µm) monitors the subsurface waters of the permanent shallow (10-40 m) thermocline. The Pleistocene d18O difference (N. dutertrei minus G. sacculifer, Dd18Od-s) averages 0.9 per mil and ranges from 0 per mil to 1.7 per mil. Neglecting species effects and shell size, the average Pleistocene d13C difference (G. sacculifer minus N. dutertrei, Dd13Cs-d) is 0.0 per mil and ranges from -0.5 per mil to 0.5 per mil. The Dd18Od-s and Dd13Cs-d records are used to infer vertical contrasts in upper ocean water temperature and nutrient concentration, though d13C may also be influenced by other factors, such as CO2 gas exchange. Variations in the isotopic differences are often synchronous with glacial/interglacial climate change. Glacial periods are characterized by smaller vertical contrasts in both temperature and nutrient concentration, and by notably greater accumulation rates of N. dutertrei and CaCO3. We attribute these responses to greater upwelling at the equatorial divergence. Superimposed on the glacial/interglacial Dd18Od-s pattern is a long-term trend possibly associated with the advection of Peru Current waters. The temporal fluctuations in the isotopic contrasts are strikingly similar to those observed at Site 851 (Ravelo and Shackleton, this volume), suggesting that the inferred changes in thermal and chemical profiles occurred over a broad region in the equatorial Pacific.

Delta 66Zn ratios and age of carbonates from ODP Site 138-849 (Table 1)

The carbonate fraction of sediment core ODP 849, leg 138, located in the eastern equatorial Pacific, mostly consisting of coccoliths, was separated and analyzed for its Zn isotopic composition. The overall variation in Zn isotopic composition, as determined by multiple-collector, magnetic-sector, inductively coupled plasma mass spectrometry, was found to be on the order of 1? (expressed in delta66Zn, where deltaxZn=[(xZn/64Zn)sample/(xZn/64Zn)standard -1]*10**3 and x=66, 67 or 68) over the last 175 ka. The analytical precision was 0.04 per mil and the overall reproducibility was usually better than 0.07 per mil. The Zn isotopic composition signal exhibits several marked peaks and a high-frequency variability. A periodogram of the delta66Zn signal showed two periodicities of 35.2 and 21.2 ka. We suggest that the latter is caused by the precession of the Earth's axis of rotation. The periodogram exhibits a minimum at 41.1 ka, thus showing that the Zn isotopic composition is independent of the obliquity in the eastern equatorial Pacific. The range of delta66Zn values observed for the carbonate fraction of ODP 849 overlaps with the range observed for Fe-Mn nodules in the world's oceans, which suggests that seawater/carbonate Zn isotope fractionation is weak. We therefore assume that most of the Zn isotope variability is a result of the selective entrainment of the light isotopes by organic matter in the surface ocean. The ODP 849 delta66Zn record seems to follow the changes in the insolation cycles. Changes in the late summer/fall equatorial insolation modulate the intensity of the equatorial upwelling, hence the mixing between deep and surface waters. We propose that during decreased summer/fall equatorial insolation, when a steep thermocline can develop (El Niño-like conditions), the surface waters cannot be replenished by deep waters and become depleted in the lighter Zn isotopes by biological activity, thus resulting in the progressive increase of the delta66Zn values of the carbonate shells presumably in equilibrium with surface seawater.

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.

Stable oxygen isotope. Mg/Ca and Temperature record of Caribbean and Pacific sediments

The final phase of the closure of the Panamanian Gateway and the intensification of Northern Hemisphere Glaciation (NHG) both occurred during the Late Pliocene. Glacial-interglacial (G-IG) variations in sea level might, therefore, have had a significant impact on the remaining connections between the East Pacific and the Caribbean. Here, we present combined foraminiferal Mg/Ca and d18O measurements from Ocean Drilling Program (ODP) Site 1241 from the East Pacific and ODP Site 999 from the Caribbean. The studied time interval covers the first three major G-IG Marine Isotope Stages (MIS 95-100, ~2.5 Ma) after the intensification of NHG. Analyses were performed on the planktonic foraminifera Neogloboquadrina dutertrei and Globigerinoides sacculifer, representing water mass properties in the thermocline and the mixed-layer, respectively. Changes in sea water temperature, relative salinity, and water column stratification strongly suggest that the Panamanian Gateway temporarily closed during glacial MIS 98 and 100, as a result of changes in ice volume equivalent to a drop in sea level of 60-90 m. Reconstructed sea surface temperatures (SST) from G. sacculifer show a glacial decrease of 2.5°C at Site 1241, but increases of up to 3°C at Site 999 during glacial MIS 98 and 100 suggesting that the Panamanian Gateway closed during these glacial periods. The Mg/Ca-temperatures of N. dutertrei remain relatively stable in the East Pacific, but do show a 3°C warming in the Caribbean at the onset of these glacial periods suggesting that the closing of the gateway also changed the water column stratification. We infer that the glacial closure of the gateway allowed the Western Atlantic Warm Pool to extend into the southern Caribbean, increasing SST (G. sacculifer) and deepening the thermocline (N. dutertrei). Additionally, ice volume appears to have become large enough during MIS 100 to survive the relatively short lasting interglacial MIS 99 so that the gateway remained closed. Towards the end of MIS 98, during MIS 97 and into MIS 96 temperatures on both sides are mostly similar suggesting water masses exchanged again. Additionally, Caribbean variations in SST and d18Owater follow a precession-like cyclicity rather than the obliquity-controlled variations characteristic of the East-Pacific and many other tropical areas, suggesting that regional atmospheric processes related to the trade winds and the Intertropical Convergence Zone (ITCZ) had a dominant impact in the Caribbean.

Age model for ODP Leg 175 sites

Upwelling along the western coast of Africa south of the equator may be partitioned into three major areas, each having its own dynamics and history: (1) the eastern equatorial region, comprising the Congo Fan and the area of Mid-Angola; (2) the Namibia upwelling system, extending from the Walvis Ridge to Lüderitz; and (3) the Cape Province region, where upwelling is subdued. The highest nutrient contents in thermocline waters are in the northern region, the lowest in the southern one. Wind effects are at a maximum near the southern end of the Namibia upwelling system, and maximum productivity occurs near Walvis Bay, where the product between upwelling rate and nutrient content of upwelled waters is at a maximum. In the Congo/Angola region, opal tends to follow organic carbon quite closely in the Quaternary record. However, organic carbon has a strong precessional component, while opal does not. Despite relatively low opal content, sediments off Angola show the same patterns as those off the Congo; thus, they are part of the same regime. The spectrum shows nonlinear interference patterns between high- and low-latitude forcing, presumably tied to thermocline fertility and wind. On Walvis Ridge, as in the Congo-Angola region, the organic matter record behaves normally; that is, supply is high during glacial periods. In contrast, interglacial periods are favorable for opal deposition. The pattern suggests reduction in silicate content of the thermocline during glacial periods. The reversed phase (opal abundant during interglacials) persists during the entire Pleistocene and can be demonstrated deep into the Pliocene, not just on Walvis Ridge but all the way to the Oranje River and off the Cape Province. From comparison with other regions, it appears that silicate is diminished in the global thermocline, on average, whenever winds become strong enough to substantially shorten the residence time of silicate in upper waters (Walvis Hypothesis, solving the Walvis Paradox of reversed phase in opal deposition). The central discovery during Leg 175 was the documentation of a late Pliocene opal maximum for the entire Namibia upwelling system (early Matuyama Diatom Maximum [MDM]). The maximum is centered on the period between the end of the Gauss Chron and the beginning of the Olduvai Chron. A rather sharp increase in both organic matter deposition and opal deposition occurs near 3 Ma in the middle of the Gauss Chron, in association with a series of major cooling steps. As concerns organic matter, high production persists at least to 1 Ma, when there are large changes in variability, heralding subsequent pulsed production periods. From 3 to 2 Ma, organic matter and opal deposition run more or less parallel, but after 2 Ma opal goes out of phase with organic matter. Apparently, this is the point when silicate becomes limiting to opal production. Thus, the MDM conundrum is solved by linking planetary cooling to increased mixing and upwelling (ramping up to the MDM) and a general removal of silicate from the upper ocean through excess precipitation over global supply (ramping down from the MDM). The hypothesis concerning the origin of the Namibia opal acme or MDM is fundamentally the same as the Walvis Hypothesis, stating that glacial conditions result in removal of silicate from the thermocline (and quite likely from the ocean as a whole, given enough time). The Namibia opal acme, and other opal maxima in the latest Neogene in other regions of the ocean, marks the interval when a cooling ocean selectively removes the abundant silicate inherited from a warm ocean. When the excess silicate is removed, the process ceases. According to the data gathered during Leg 175, major upwelling started in the late part of the late Miocene. Presumably, this process contributed to the drawing down of carbon dioxide from the atmosphere, helping to prepare the way for Northern Hemisphere glaciation.

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.

Miocene planktonic foraminifera of ODP Hole 130-806B

We document the waxing and waning of a "proto-warm pool" in the western equatorial Pacific (WEP) based on a study of multi-species planktic foraminiferal isotope ratios and census data spanning the 13.2-5.8 Ma interval at ODP Site 806. We hypothesize that the presence or absence of a proto-warm pool in the WEP, caused by the progressive tectonic constriction of the Indonesian Seaway and modulated by sea level fluctuations, created El Niño/La Niña-like alternations of hydrographic conditions across the equatorial Pacific during the late Miocene. This hypothesis is supported by the general antithetical relationship observed between carbonate productivity and preservation in the western and eastern equatorial Pacific, which we propose is caused by these alternating ocean-climate states. Warming of thermocline and surface waters, as well as a major change in planktic foraminferal assemblages record a two-step phase of proto-warm pool development ~11.6-10 Ma, which coincides with Miocene isotope events Mi5 and Mi6, and sea-level low stands. We suggest that these changes in the biota and structure of the upper water column in the WEP mark the initiation of a more modern equatorial current system, including the development of the Equatorial Undercurrent (EUC), as La Niña-like conditions became established across the tropical Pacific. This situation sustained carbonate and silica productivity in the eastern equatorial Pacific (EEP) at a time when carbonate preservation sharply declined in the Caribbean. Proto-warm pool weakening after ~10 Ma may have contributed to the nadir of a similar "carbonate crash" in the EEP. Cooling of the thermocline and increased abundances of thermocline taxa herald the decay of the proto-warm pool and higher productivity in the WEP, particularly ~ 9.0-8.8 Ma coincident with a major perturbation in tropical nannofossil assemblages. We suggest that this interval of increased productivity records El Niño-like conditions across the tropical Pacific and the initial phase of the widespread "biogenic bloom". Resurgence of a later proto-warm pool in the WEP ~6.5-6.1 Ma may have spurred renewed La Niña-like conditions, which contributed to a strong late phase of the "biogenic bloom" in the EEP.

Size variations of Discoaster multiradiatus across the Paleocene-Eocene thermal maximum

Size measurements of the calcareous nannofossil taxon Discoaster multiradiatus were carried out across the Paleocene-Eocene Thermal Maximum (PETM) in Ocean Drilling Program Holes 690B (Maud Rise, Weddell Sea) and 1209B (Shatsky Rise, Pacific Ocean). Morphometric investigations show that D. multiradiatus specimens are generally larger at ODP Site 1209 than at ODP Site 690. A limited increase in size of D. multiradiatus is recorded at ODP Site 1209, whereas significant enlargements characterize ODP Site 690. Preservation is comparable at both sites: nannofossils are moderately preserved with some evidence of etching/overgrowth in the PETM interval. Yet, D. multiradiatus variations do not correlate with preservation state and morphometric data most likely represent primary signals rather than diagenetic artifacts. There is a direct relationship between D. multiradiatus size and paleotemperatures: largest specimens are coeval with global warming associated with the PETM, inferred to result from excess atmospheric CO2 due to (partial) oxidation of massive quantities of methane. Size increases and largest specimens of D. multiradiatus occur at different stratigraphic levels within PETM at ODP Sites 690 and 1209. A marked shift in diameter size was observed at the onset and peak of the Carbon Isotopic Excursion (CIE) at ODP Site 690, but only at the end of CIE and initial recovery interval at ODP Site 1209. This diachroneity is puzzling, but indeed correlates well with reconstructed changes in surface and thermocline water masses temperature and salinity in the PETM interval at low and high latitudes. The presumed high concentrations of carbon dioxide seem to have not influenced the morphometry of D. multiradiatus. The major size increase of D. multiradiatus in the CIE of ODP Site 690 could represent the migration of larger-sized allochtonus specimens that moved from peri-equatorial/subtropical areas to higher latitudes during the warmest interval of the PETM, although no direct evidence of distinct populations/subpopulations has been obtained from the frequency diagrams. As a result, we infer that D. multiradiatus is a proxy of water masses stratification and might be used for deriving temperature-salinity-nutrient conditions in the mixed layer and thermocline and their dynamics.

Middle Miocene stable carbon and oxygen isotope record of ODP Holes 130-805B and 130-806B

Bulk carbon isotope records are an effective chemostratigraphic tool for the middle Miocene because of the large and systematic variation in first-order d13C signals. Bulk d13C measurements support the presence of a hiatus at 305 mbsf in Hole 805B (latest middle Miocene), provisionally located while on board ship using biostratigraphic and magnetostratigraphic events. Records at Holes 805B and 806B show the middle Miocene Monterey carbon isotope excursion although the record at Hole 806B is apparently more stratigraphically continuous. Detailed analysis of multispecies foraminiferal carbon isotope records during the middle Miocene ("Monterey excursion") segment at Hole 806B support the assertion that this carbon isotope excursion comprises mainly between-reservoir effects. The benthic d18O data increase after 15.3 Ma, which we suggest corresponds to the mid-Miocene cooling step/ice volume increase of other authors. Planktonic foraminiferal d18O evidence exists for steepening of the thermocline at 17.4 Ma. A second-order d13C excursion superimposed at 13.8 Ma on the first-order Monterey excursion is associated with a second-order negative d18O excursion.

Part of the global DOC versus AOU (dissolved organic carbon/apparent oxygen utilization) data compilation, Azores-I

Temperate, transitional and subtropical waters of the remote Azores Front region east of Azores (24-40°N, 22-32°W) were sampled during three cruises conducted under increasing stratification conditions (April 1999, May 1997 and August 1998). Despite the temporal increase of surface temperature (by 5 °C) and stratification (by 2.1 1/min**2), as well as the thermocline shoaling (by ~15 m), dissolved organic carbon (DOC) and nitrogen (DON) in the surface layer were not significantly different for the early spring, late spring and summer periods, with average concentrations of 69±2 µM-C and 5.2±0.4 µM-N, respectively. The surface excess of semi-labile DOC, compared with the baseline DOC concentration in the deep ocean (47±2 µM-C), represents 33% of the bulk DOC concentration and as much as 85% of the TOC (=POC+DOC) excess. When compared with the winter baseline (56±2 µM-C), the seasonal surface DOC excess is 20% of the bulk DOC concentration and 87% of the seasonal TOC excess. These results confirm the major role played by DOC in the carbon cycle of surface waters of the Azores Front region. The total amount of bioreactive DOC transported from the temperate to the subtropical North Atlantic by the Ekman flux between March and December represents only ~15% of the average annual primary production, and ~15% and ~30% of the measured sinking POC flux+vertical DOC eddy diffusion during early spring and summer, respectively. Vertical eddy diffusion is 35% and 2% of the spring and summer sinking POC flux, respectively. On the other hand, DOC only contributes 13% to the local oxidation of organic matter in subsurface waters (between the pycnocline and 500 m) of the study region.

Stable carbon and oxygen isotope ratios of Miocene to Recent foraminifera from the Rio Grande Rise, South Atlantic

Miocene to Recent species of planktic foraminifera in the Globorotalia (Globoconella) lineage evolved entirely within the thermocline. All species are most abundant within subtropical-temperate watermasses throughout their history. The near stasis in distribution within the thermocline and the subtropical convergence suggests the major morphological changes in Globorotalia (Globoconella) may have occurred through habitat subdivision rather than by vicariant shifts into new watermasses. At the Rio Grande Rise, in the South Atlantic, modern G. inflata is 0.66-0.84? more positive for delta18O than the most enriched coexisting Globigerinoides sacculifer and probably grows in the mid thermocline deeper than 325 m. All extinct globoconellid species have mean delta18O ratios 0.5-0.8? more positive than Globigerinoides trilobus and G. sacculifer and probably lived within the thermocline as well. Major events in skeletal evolution are poorly correlated with changes in delta18O in this group. These include evolutionary transitions to compressed, smooth-walled tests and acquisition of keels. In addition, morphological reversals from the umbilically-inflated G. conomiozea to biconvex G. pliozea and to unkeeled G. puncticulata occur in the absence of changes in delta18O signature. Instead, the ranges of delta18O between different species almost completely overlap once corrected for temporal changes in delta18O of sea water. Foraminifera morphologies have been widely considered to evolve in response to changes in watermasses or depth habitats. However, the variety of skeletal shapes in the globoconellid lineage apparently are not adaptations to a progressive radiation from the surface mixed layer into deeper waters.