(Appendix) Analysis of benthic foraminifera from ODP Leg 181 and DSDP Leg 90

Canonical correspondence analysis indicates that the distribution of Neogene benthic foraminiferal faunas (>63 µm) in seven DSDP and ODP sites (500-4500 m water depth) east of New Zealand (38-51°S, 170°E-170°W) is most strongly influenced by depth (water mass stratification), and secondly by age (palaeoceanographic changes influencing faunal composition and biotic evolution). Stratigraphic faunal changes are interpretted in terms of the pulsed sequential development of southern, and later northern, polar glaciation and consequent cooling of bottom waters, increased vertical and lateral stratification of ocean water masses, and increased overall and seasonal surface water productivity. Oligocene initiation of the Antarctic Circumpolar Current and Deep Western Boundary Current (DWBC), flowing northwards past New Zealand, resulted in extensive hiatuses throughout the Southwest Pacific, some extending through into the Miocene. Planktic foraminiferal fragmentation index values indicate that carbonate dissolution was significant at abyssal depths throughout most of the Neogene, peaking at upper abyssal depths in the late Miocene (11-7 Ma), with the lysocline progressively deepened thereafter. Miocene abyssal faunas are dominated by Globocassidulina subglobosa and Oridorsalis umbonatus, with increasing Epistominella exigua after 16 Ma at upper abyssal depths. Peak abundances of Epistominella umbonifera indicate increased input of cold Southern Component Water to the DWBC at 7-6 Ma. Faunal association changes imply establishment of the modern Oxygen Minimum Zone (upper Circumpolar Deep Water) in the latest Miocene. Significant latitudinal differences between the benthic foraminiferal faunas at lower bathyal depths indicate the existence of an oceanic front along the Chatham Rise (location of present Subtropical Front), since the early late Miocene at least, with more pulsed productivity (higher E. exigua) along the south side. Modern Antarctic Intermediate Water faunal associations were established north of the Chatham Rise at 10-9 Ma, and south of it at 3-1.5 Ma. Middle-upper bathyal faunas on the Campbell Plateau are dominated by reticulate bolivinids during the early and middle Miocene, indicative of sustained productivity above relatively sluggish, suboxic bottom waters. Faunal changes and hiatuses indicate increased current vigour over the Campbell Plateau from the latest Miocene on. Surface water productivity (food supply) appears to have increased in three steps (at times of enhanced global cooling) marked by substantially increased relative abundance of: (1) Abditodentrix pseudothalmanni, Alabaminella weddellensis, Cassidulina norvangi (16-15 Ma, increased pulsed productivity); (2) Bulimina marginata f. aculeata, Nonionella auris, Trifarina angulosa, Uvigerina peregrina (3-1.5 Ma, increased overall productivity); and (3) Cassidulina carinata (1-0.5 Ma, increased overall productivity). Three intervals of deep-sea benthic foraminiferal taxonomic turnover are recognised (16-15, 11.5-10, 2-0.5 Ma) corresponding to intervals of enhanced global cooling and possible productivity changes. The late Pliocene-middle Pleistocene extinction, associated with increasing Northern Hemisphere glaciation, culminating in the middle Pleistocene climatic transition, was more significant in the study area than the earlier Neogene turnovers.

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.

Upper Maestrichtian to Eocene benthic foraminifera in ODP Leg 121 holes

Late Maestrichtian to late Eocene bathyal benthic foraminiferal faunas at Sites 752,753, and 754 on Broken Ridge in the eastern Indian Ocean were analyzed as to their stratigraphic distribution of species to clarify the relation between faunal turnovers and paleoceanographic changes. Based on Q-mode factor analysis, eight varimax assemblages were distinguished: the Stensioina beccariiformis assemblage in the upper Maestrichtian to upper Paleocene; the Cibicidoides hyphalus assemblage in the upper Maestrichtian; the Cibicidoides cf. pseudoperlucidus assemblage in the upper Paleocene; the Anomalinoides capitatusldanicus assemblage in the uppermost Paleocene to lower Eocene; the Cibicidoides subspiratus assemblage in the lower Eocene; the Nuttallides truempyi assemblage in the lower and middle Eocene; the Osangularia sp. 1 - Hanzawaia ammophila assemblage in the upper Eocene; and the Lenticulina spp. assemblage in the uppermost Eocene, Oligocene, and lower Miocene. The presence of the Osangularia sp. 1 - Hanzawaia ammophila assemblage is related to the shallowing episode on Broken Ridge (upper bathyal), as a result of the rifting event that occurred in the middle Eocene. The most distinct faunal change (the disappearance of about 37% of the species) occurred between the S. beccariiformis assemblage and the A. capitatusldanicus assemblage, at the end of the upper Paleocene. A. capitatusldanicus, Lenticulina spp., and varied forms of Cibicidoides replaced the Velasco-type fauna at this time. The timing of this event is well correlated with the known age at South Atlantic sites (Thomas, 1990 doi:10.2973/odp.proc.sr.113.123.1990; Kennett and Stott, 1990 doi:10.2973/odp.proc.sr.113.188.1990; Katz and Miller, 1990 doi:10.2973/odp.proc.sr.114.147.1991). The primary cause of the extinction of the Stensioina beccariiformis assemblage is elusive, but may have resulted from the cessation of deep-water formation in the Antarctic (Katz and Miller, 1990), and subsequent arrival of warm saline deep water (Thomas, 1990; Kennett and Stott, 1990). Another possibility may be a weakened influence of high-salinity water formed at the low latitudes such as the Tethys Sea. The extinction event corresponds to the change from higher delta13C values in benthic foraminifers to lower ones. An interpretation of delta13C values is that the eastern Indian deep water, characterized by young and nutrient-depleted water, became old water which was devoid of a supply of new water during the latest Paleocene to early Eocene. Prior to this benthic event, signals of related faunal change were detected in the following short periods: early and late Paleocene, near the boundary of nannofossil Zone CP4, and Zone CP5 of the late Paleocene at Site 752. Among common taxa in the upper Maestrichtian, only seven species disappeared or became extinct at the Cretaceous/ Tertiary boundary at Site 752. The benthic foraminiferal population did not change for up to 2 m above the boundary, in contrast to the rapid decrease of the plankt onic foraminiferal population at the boundary. A decrease in the number of benthic foraminifers occurs after that level, corresponding to an interval of decreased numbers of planktonic foraminifers and higher abundance of volcanic ash. Reduced species diversity (H') suggests a secondary effect attributable to the dissolution of foraminiferal tests. The different responses of planktonic and benthic foraminifers to the event just above the boundary suggest that the Cretaceous/Tertiary event was a surface event as also suggested by Thomas (1990). In addition, a positive shift of delta13C in benthic foraminifers after the event indicates nutrient-depleted bottom water at Site 752.

Neogene benthic foraminifers from the Kerguelen Plateau

Benthic foraminifers were studied quantitatively in 120 lower Miocene through upper Pleistocene samples from Ocean Drilling Program Site 747 (Central Kerguelen Plateau) and Sites 748 and 751 (Southern Kerguelen Plateau). These sites are situated on an 450-km-long, north-south transect between 54°49'S and 58°26'S at present water depths between 1696 and 1288 m. Principal component analysis on the census data of the most abundant 92 taxa helped to identify 8 benthic foraminifer assemblages. These benthic foraminifer assemblages were compared with Holocene faunas from southern high latitudes to reconstruct paleoenvironmental conditions. Middle lower Miocene sediments are characterized by a Uvigerina hispidocostata assemblage, indicating high paleoproductivity and/or not well-ventilated bottom water. From late early to late middle Miocene time, the Southern Kerguelen Plateau was bathed by a young, well-oxygenated, and carbonate-aggressive water mass, as indicated by a Nuttallides umbonifer-dominated benthic foraminifer assemblage. During late middle Miocene time, an Astrononion pusillum assemblage took over for only about 1 m.y., probably indicating the first injection of an aged water mass, similar to the North Atlantic Deep Water (NADW), into a developing circumpolar current system. Around the middle to late Miocene boundary, the fauna again became dominated by N. umbonifer. After the last appearance of N. umbonifer, reestablishment of the A. pusillum assemblage from the early late through at least the late late Miocene, indicated the established influence of a NADW-like water mass. The latest Miocene through middle late Pliocene benthic foraminifer assemblage was characterized by Epistominella exigua and strong carbonate dissolution, indicating very high biosiliceous production, and this in turn may indicate the formation and paleoposition of an Antarctic Polar Frontal Zone. From the late late Pliocene, a Trifarina angulosa assemblage (indicative today of sandy substrate and vigorous bottom currents) strongly dominated the fauna up to the late Pleistocene, when Bulimina aculeata (indicative today of calm sedimentation with high organic matter fluxes) became an important and partly dominating constituent of the fauna. This is interpreted as the faunal response to the decreased winnowing force (bottom current velocities) of the Antarctic Circumpolar Current during periods of global climatic amelioration and raised sea level.

Cenozoic benthic foraminifers from ODP Leg 134 holes

This paper discusses the Paleobathymetric and paleoenvironmental history of the New Hebrides Island Arc and North d'Entrecasteaux Ridge during Cenozoic time based on benthic foraminiferal and sedimentological data. Oligocene and Pliocene to Pleistocene benthic foraminiferal assemblages from Sites 827, 828, 829, and 832 of Ocean Drilling Program (ODP) Leg 134 (Vanuatu) are examined by means of Q-mode factor analysis. The results of this analysis recognize the following bathymetrically significant benthic foraminiferal biofacies: (1) Globocassidulina subglobosa biofacies and Bulimina aculeata-Bolivinita quadrilatera biofacies representing the upper bathyal zone (600-1500 m); (2) Gavelinopsis praegeri-Cibicides wuellerstorfi biofacies, indicating the Pacific Intermediate Water (water depth between 1500 and 2400 m); (3) Tosaia hanzawai-Globocassidulina muloccensis biofacies, Valvulineria gunjii biofacies, and the Melonis barleeanus-Melonis sphaeroides biofacies, which characterize the lower bathyal zone; (4) the Nuttallides umbonifera biofacies, which characterizes the interval between the lysocline (approximately 3500 m) and the carbonate compensation depth (approximately 4500 m); and (5) the Rhabdammina abyssorum biofacies representing the abyssal zone below the carbonate compensation depth. Benthic foraminiferal patterns are used to construct Paleobathymetric and paleogeographic profiles of the New Hebrides Island Arc and North d'Entrecasteaux Ridge for the following age boundaries: late Miocene/Pliocene, early/late Pliocene, Pliocene/Pleistocene, and Pleistocene/Holocene.

Global compilation of last glacial maximum oxygen isotopic ratios for planktonic and benthic foraminifera

We use the fully coupled atmosphere-ocean three-dimensional model of intermediate complexity iLOVECLIM to simulate the climate and oxygen stable isotopic signal during the Last Glacial Maximum (LGM, 21 000 yr). By using a model that is able to explicitly simulate the sensor (d18O), results can be directly compared with data from climatic archives in the different realms. Our results indicate that iLOVECLIM reproduces well the main feature of the LGM climate in the atmospheric and oceanic components. The annual mean d18O in precipitation shows more depleted values in the northern and southern high latitudes during the LGM. The model reproduces very well the spatial gradient observed in ice core records over the Greenland ice-sheet. We observe a general pattern toward more enriched values for continental calcite d18O in the model at the LGM, in agreement with speleothem data. This can be explained by both a general atmospheric cooling in the tropical and subtropical regions and a reduction in precipitation as confirmed by reconstruction derived from pollens and plant macrofossils. Data-model comparison for sea surface temperature indicates that iLOVECLIM is capable to satisfyingly simulate the change in oceanic surface conditions between the LGM and present. Our data-model comparison for calcite d18O allows investigating the large discrepancies with respect to glacial temperatures recorded by different microfossil proxies in the North Atlantic region. The results argue for a trong mean annual cooling between the LGM and present (>6°C), supporting the foraminifera transfer function reconstruction but in disagreement with alkenones and dinocyst reconstructions. The data-model comparison also reveals that large positive calcite d18O anomaly in the Southern Ocean may be explained by an important cooling, although the driver of this pattern is unclear. We deduce a large positive d18Osw anomaly for the north Indian Ocean that contrasts with a large negative d18Osw anomaly in the China Sea between the LGM and present. This pattern may be linked to changes in the hydrological cycle over these regions. Our simulation of the deep ocean suggests that changes in d18Osw between the LGM and present are not spatially homogenous. This is supported by reconstructions derived from pore fluids in deep-sea sediments. The model underestimates the deep ocean cooling thus biasing the comparison with benthic calcite d18O data. Nonetheless, our data-model comparison support a heterogeneous cooling of few degrees (2-4°C) in the LGM Ocean.

Growth response of an early successional assemblage of coralline algae and benthic diatoms to ocean acidification

The sustained absorption of anthropogenically released atmospheric CO2 by the oceans is modifying seawater carbonate chemistry, a process termed ocean acidification (OA). By the year 2100, the worst case scenario is a decline in the average oceanic surface seawater pH by 0.3 units to 7.75. The changing seawater carbonate chemistry is predicted to negatively affect many marine species, particularly calcifying organisms such as coralline algae, while species such as diatoms and fleshy seaweed are predicted to be little affected or may even benefit from OA. It has been hypothesized in previous work that the direct negative effects imposed on coralline algae, and the direct positive effects on fleshy seaweeds and diatoms under a future high CO2 ocean could result in a reduced ability of corallines to compete with diatoms and fleshy seaweed for space in the future. In a 6-week laboratory experiment, we examined the effect of pH 7.60 (pH predicted to occur due to ocean acidification just beyond the year 2100) compared to pH 8.05 (present day) on the lateral growth rates of an early successional, cold-temperate species assemblage dominated by crustose coralline algae and benthic diatoms. Crustose coralline algae and benthic diatoms maintained positive growth rates in both pH treatments. The growth rates of coralline algae were three times lower at pH 7.60, and a non-significant decline in diatom growth meant that proportions of the two functional groups remained similar over the course of the experiment. Our results do not support our hypothesis that benthic diatoms will outcompete crustose coralline algae under future pH conditions. However, while crustose coralline algae were able to maintain their presence in this benthic rocky reef species assemblage, the reduced growth rates suggest that they will be less capable of recolonizing after disturbance events, which could result in reduced coralline cover under OA conditions.

Response of benthic foraminifera to ocean acidification in their natural sediment environment: a long-term culturing experiment

Calcifying foraminifera are expected to be endangered by ocean acidification; however, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 6 months of incubation, foraminiferal assemblages were kept and treated in natural sediment with pCO2-enriched seawater of 430, 907, 1865 and 3247 µatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months of incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Omega calc was much higher in the sediment than in the water column in nearly all pCO2 treatments and remained close to saturation. As a result, the life cycle (population density, growth and reproduction) of living assemblages varied markedly during the experimental period, but was largely unaffected by the pCO2 treatments applied. According to the size-frequency distribution, we conclude that foraminifera start reproduction at a diameter of 250 µm. Mortality of living Ammonia aomoriensis was unaffected, whereas size of large and dead tests decreased with elevated pCO2 from 285 µm (pCO2 from 430 to 1865 µatm) to 258 µm (pCO2 3247 µatm). The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of CaCO3 weight. Living individuals had a calcium carbonate production rate of 0.47 g/m**2/a, whereas dead empty tests accumulated a rate of 0.27 g /m**2/a. Although Omega calc was close to 1, approximately 30% of the empty tests of Ammonia aomoriensis showed dissolution features at high pCO2 of 3247 µatm during the last 2 months of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. Our results emphasize that the sensitivity to ocean acidification of the endobenthic foraminifera Ammonia aomoriensis in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

The O2, pH and Ca2+ Microenvironment of Benthic Foraminifera in a High CO2 World

Ocean acidification (OA) can have adverse effects on marine calcifiers. Yet, phototrophic marine calcifiers elevate their external oxygen and pH microenvironment in daylight, through the uptake of dissolved inorganic carbon (DIC) by photosynthesis. We studied to which extent pH elevation within their microenvironments in daylight can counteract ambient seawater pH reductions, i.e. OA conditions. We measured the O2 and pH microenvironment of four photosymbiotic and two symbiont-free benthic tropical foraminiferal species at three different OA treatments (~432, 1141 and 2151 µatm pCO2). The O2 concentration difference between the seawater and the test surface (delta O2) was taken as a measure for the photosynthetic rate. Our results showed that O2 and pH levels were significantly higher on photosymbiotic foraminiferal surfaces in light than in dark conditions, and than on surfaces of symbiont-free foraminifera. Rates of photosynthesis at saturated light conditions did not change significantly between OA treatments (except in individuals that exhibited symbiont loss, i.e. bleaching, at elevated pCO2). The pH at the cell surface decreased during incubations at elevated pCO2, also during light incubations. Photosynthesis increased the surface pH but this increase was insufficient to compensate for ambient seawater pH decreases. We thus conclude that photosynthesis does only partly protect symbiont bearing foraminifera against OA.