Effect of ocean acidification and pH fluctuations on the growth and development of coralline algal recruits, and an associated benthic algal assemblage

Coralline algae are susceptible to the changes in the seawater carbonate system associated with ocean acidification (OA). However, the coastal environments in which corallines grow are subject to large daily pH fluctuations which may affect their responses to OA. Here, we followed the growth and development of the juvenile coralline alga Arthrocardia corymbosa, which had recruited into experimental conditions during a prior experiment, using a novel OA laboratory culture system to simulate the pH fluctuations observed within a kelp forest. Microscopic life history stages are considered more susceptible to environmental stress than adult stages; we compared the responses of newly recruited A. corymbosa to static and fluctuating seawater pH with those of their field-collected parents. Recruits were cultivated for 16 weeks under static pH 8.05 and 7.65, representing ambient and 4*preindustrial pCO2 concentrations, respectively, and two fluctuating pH treatments of daily (daytime pH = 8.45, night-time pH = 7.65) and daily (daytime pH = 8.05, night-time pH = 7.25). Positive growth rates of new recruits were recorded in all treatments, and were highest under static pH 8.05 and lowest under fluctuating pH 7.65. This pattern was similar to the adults' response, except that adults had zero growth under fluctuating pH 7.65. The % dry weight of MgCO3 in calcite of the juveniles was reduced from 10% at pH 8.05 to 8% at pH 7.65, but there was no effect of pH fluctuation. A wide range of fleshy macroalgae and at least 6 species of benthic diatoms recruited across all experimental treatments, from cryptic spores associated with the adult A. corymbosa. There was no effect of experimental treatment on the growth of the benthic diatoms. On the community level, pH-sensitive species may survive lower pH in the presence of diatoms and fleshy macroalgae, whose high metabolic activity may raise the pH of the local microhabitat.

Benthic foraminifera stable isotope data from the South Pacific sediment cores PS75/059-2 and PS75/056-1

Investigating the inter-basin deep water exchange between the Pacific and Atlantic Oceans over glacial-interglacial climate cycles is important for understanding circum-Antarctic Southern Ocean circulation changes and their impact on the global Meridional Overturning Circulation. We use benthic foraminiferal d13C records from the southern East Pacific Rise to characterize the d13C composition of Circumpolar Deep Water in the South Pacific, prior to its transit through the Drake Passage into the South Atlantic. A comparison with published South Atlantic deep water records from the northern Cape Basin suggests a continuous water mass exchange throughout the past 500 ka. Almost identical glacial-interglacial d13C variations imply a common deep water evolution in both basins suggesting persistent Circumpolar Deep Water exchange and homogenization. By contrast, deeper abyssal waters occupying the more southern Cape Basin and the southernmost South Atlantic have lower d13C values during most, but not all, stadial periods. We conclude that these values represent the influence of a more southern water mass, perhaps AABW. During many interglacials and some glacial periods, the gradient between Circumpolar Deep Water and the deeper southern Cape Basin bottom water disappears suggesting either no presence of AABW or indistinguishable d13C values of both water masses.

Neogene benthic foraminiferal community and organic carbon at DSDP Holes 95-612 and 95-613

Quantitative study of benthic foraminifers from the upper Miocene to lower Pliocene section at Site 612 (1404 m present water depth) and the Pliocene section at Site 613 (2323 m present water depth) shows no evidence of widespread downslope transport of shallow-water biofacies or reworking of older material in the greater than 150 µm size fraction. In contrast, upper Miocene sediments from Site 604 (2364 m present water depth) show extensive reworking and downslope transport. At Site 612, benthic foraminifers show a succession from an upper Miocene Bolivina alata-Nonionella sp. biofacies, to an uppermost Miocene Bulimina alazanensis biofacies, to a lower Pliocene Cassidulina reflexa biofacies, to an upper Pliocene Melonis barleeanum-Islandiella laevigata biofacies. Evidence suggests that the Pliocene biofacies are in situ, although they could have been transported downslope from the upper-middle bathyal zone. At Site 613, Uvigerina peregrina dominated the "middle" Pliocene, while Globocassidulina subglobosa was dominant in the early and late Pliocene. High abundances of U. peregrina at Site 613 are associated with high values of sedimentary organic carbon.

Benthic foraminifera extinction at the base of the Brunhes Chronozone at ODP Holes 108-658A and 108-659A

A faunal boundary found at the base of the Brunhes Chronozone at Sites 658 and 659 confirms previous observations from several locations in the Atlantic Ocean and may be classified as a supraregional "extinction event". Several benthic foraminifer species typical of the Pliocene disappear near the Brunhes/Matuyama boundary, thus marking the upper limit of a faunal zone (faunal unit). Improved chronological dating indicates that the disappearance of these species occurs over a period of about 100,000 yr.

Early Cenozoic Atlantic and Pacific benthic foraminiferal isotopes from 12 sites

Oxygen and carbon isotope records are important tools used to reconstruct past ocean and climate conditions, with those of benthic foraminifera providing information on the deep oceans. Reconstructions are complicated by interspecies isotopic offsets that result from microhabitat preferences (carbonate precipitation in isotopically distinct environments) and vital effects (species-specific metabolic variation in isotopic fractionation). We provide correction factors for early Cenozoic benthic foraminifera commonly used for isotopic measurements (Cibicidoides spp., Nuttallides truempyi, Oridorsalis spp., Stensioina beccariiformis, Hanzawaia ammophila, and Bulimina spp.), showing that most yield reliable isotopic proxies of environmental change. The statistical methods and larger data sets used in this study provide more robust correction factors than do previous studies. Interspecies isotopic offsets appear to have changed through the Cenozoic, either (1) as a result of evolutionary changes or (2) as an artifact of different statistical methods and data set sizes used to determine the offsets in different studies. Regardless of the reason, the assumption that isotopic offsets have remained constant through the Cenozoic has introduced an 1-2°C uncertainty into deep sea paleotemperature calculations. In addition, we compare multiple species isotopic data from a western North Atlantic section that includes the Paleocene-Eocene thermal maximum to determine the most reliable isotopic indicator for this event. We propose that Oridorsalis spp. was the most reliable deepwater isotopic recorder at this location because it was best able to withstand the harsh water conditions that existed at this time; it may be the best recorder at other locations and for other extreme events also.

Benthic foraminifers in Mesozoic and Cenozoic sediments of the southwestern Atlantic

Benthic foraminiferal assemblages in Mesozoic and Cenozoic sediments were studied at Sites 511, 512, 513, and 514 drilled during Leg 71 in the southwestern Atlantic on the Maurice Ewing Bank and in the Argentine Basin. Benthic foraminifers in almost all stratigraphic subdivisions of Sites 511 and 512 reflect the gradual subsidence of the Falkland Plateau from shelf depths in the Barremian-Albian, when a semiclosed basin with restricted circulation of water masses and anaerobic conditions existed, to lower bathyal depths in the Late Cretaceous and Cenozoic, with an abrupt acceleration at the boundary of Lower and Upper Cretaceous. The composition, distribution, and preservation of Late Cretaceous assemblages of benthic foraminifers suggest considerable fluctuations of the foraminiferal lysocline and the CCD. This is evidenced by dissolution facies and foraminiferal assemblages in which agglutinated and resistant calcareous forms predominated during high stands of the CCD and by calcareous facies in which rich assemblages of calcareous species predominated during low stands. The highest position of the CCD on the Plateau (less than 1500-2000 m) was in the late Cenomanian, Turonian, and Coniacian. In the Santonian and Campanian the CCD was at depths below 1500-2000 meters. At the end of the Campanian the CCD shifted again to depths comparable with those of Cenomanian and Turonian time. In the latest Campanian and the Maestrichtian the CCD was low and nanno-foraminiferal oozes with a rich assemblage of benthic foraminifers accumulated. Foraminiferal assemblages at Sites 513 and 514 in the Argentine Basin also testify to oceanic subsidence from lower bathyal depths in the Oligocene to abyssal ones at present. This process was complicated by the influence of geographical migrations of the Polar Front caused by extensions of the ice sheet in the Antarctic after the opening of the Drake Passage during the Oligocene. In Mesozoic and Cenozoic deposits of the Falkland Plateau and the Argentine Basin seven assemblages of benthic foraminifers were distinguished by age: early-middle Albian, middle-late Albian, Late Cretaceous (including four groups), middle Eocene, late Eocene-early Miocene, middle-late Miocene, and Pliocene-Quaternary. The Albian assemblages contain many species common to the foraminiferal fauna of the Austral Biogeographical Province. The Late Cretaceous assemblage contains, along with Austral species, species common to foraminifers of North America, Western Europe, the Russian platform, and the south of the U.S.S.R. Deep-sea cosmopolitan species prevail in Cenozoic assemblages.

Distribution of benthic foraminifera in surface sediments on the continental margin off North-West Africa

The benthic foraminiferal populations along three traverses across the Northwest African continental margin were analyzed on the base of ca. 60 surface sediment samples. Depth ranges of 213 species were established and the main trends of vertical distribution are compared with those known from adjacent regions. Main faunal breaks occure at 100/200 m and 1000/1500 m depth of water. Some species show latitudinal distribution boundaries and the same applies to population density (standing stock), reflecting the regional distribution of nutrients supply by river discharge and upwelling processes. - High proportions of Bolivina test at the lower slope indicate extended downslope transport.

Paleogene benthic foraminifera from the Kerguelen Plateau

Benthic foraminifers were studied from lower Paleocene through upper Oligocene sections from Sites 747 and 748. The composition of the benthic foraminifer species suggests a middle to lower bathyal (600-2000 m) paleodepth during the Neogene and a probable upper abyssal (2000-3000 m) paleodepth during the Paleocene at Site 747. Site 748 is thought to have remained at middle to lower bathyal paleodepths throughout the Cenozoic. Principal component analysis distinguished four major benthic foraminifer assemblages: (1) a Paleocene Stensioina beccariiformis assemblage at Sites 747 and 748, (2) an early Eocene Nuttallides truempyi assemblage at lower bathyal Site 747, (3) an early through middle Eocene Stilostomella-Lenticulina assemblage at middle bathyal Site 748, and (4) a latest Eocene through Oligocene Cibicidoides-Astrononion pusillum assemblage at both sites. Major benthic foraminifer changes, as indicated by the principal components and first and last appearances, occurred at or close to the Paleocene/Eocene boundary, and in the late Eocene close to the middle/late Eocene boundary.

Benthic foraminiferal response to the emergence of the Isthmus of Panama and coincident paleoceanographic changes

Late Cenozoic benthic foraminiferal faunas from the Caribbean Deep Sea Drilling Project (DSDP) Site 502 (3052 m) and East Pacific DSDP Site 503 (3572 m) were analyzed to interpret bottom-water masses and paleoceanographic changes occurring as the Isthmus of Panama emerged. Major changes during the past 7 Myr occur at 6.7-6.2, 3.4, 2.0, and 1.1 Ma in the Caribbean and 6.7-6.4, 4.0-3.2, 2.1, 1.4, and 0.7 Ma in the Pacific. Prior to 6.7 Ma, benthic foraminiferal faunas at both sites indicate the presence of Antarctic Bottom Water (AABW). After 6.7 Ma benthic foraminiferal faunas indicate a shift to warmer water masses: North Atlantic Deep Water (NADW) in the Caribbean and Pacific Deep Water (PDW) in the Pacific. Flow of NADW may have continued across the rising sill between the Caribbean and Pacific until 5.6 Ma when the Pacific benthic foraminiferal faunas suggest a decrease in bottom-water temperatures. After 5.6 Ma deep-water to intermediate-water flow across the sill appears to have stopped as the bottom-water masses on either side of the sill diverge. The second change recorded by benthic foraminiferal faunas occurs at 3.4 Ma in the Caribbean and 4.0-3.2 Ma in the Pacific. At this time the Caribbean is flooded with cold AABW, which is either gradually warmed or is replaced by Glacial Bottom Water (GBW) at 2.0 Ma and by NADW at 1.1 Ma. These changes are related to global climatic events and to the depth of the sill between the Caribbean and Atlantic rather than the rising Isthmus of Panama. Benthic foraminiferal faunas at East Pacific Site 503 indicate a gradual change from cold PDW to warmer PDW between 4.0 and 3.2 Ma. The PDW is replaced by the warmer, poorly oxygenated PIW at 2.1 Ma. Although the PDW affects the faunas during colder intervals between 1.4 and 0.7 Ma, the PIW remains the principal bottom-water mass in the Guatemala Basin of the East Pacific.

Oliogocene to Pleistocene benthic foraminifera in ODP Leg 121 sites

Oligocene to Pleistocene bathyal benthic foraminifers at Broken Ridge (Site 754) and Ninetyeast Ridge (Site 756), eastern Indian Ocean, were investigated for then- stratigraphic distribution and their response to paleoceanographic changes. Q-mode factor analysis was applied to relative abundance data of the most abundant benthic foraminifers. At Site 754, seven varimax assemblages were recognized from the upper Oligocene to the Pleistocene: the Gyroidina orbicularis-Rectuvigerina striata Assemblage in the uppermost Oligocene; the Lenticulina spp. Assemblage in the upper Oligocene to lower Miocene, and in lower Miocene to lowermost middle Miocene; the Burseolina cf. pacifica-Cibicidoides mundulus Assemblage in the lower Miocene; the Planulina wuellerstorfi Assemblage in the upper middle Miocene; the Globocassidulina spp. Assemblage in the upper Miocene; the Gavelinopsis lobatulus-Uvigerina proboscidea Assemblage in the Pliocene; and the Ehrenbergina spp. Assemblage in the Pleistocene. The major faunal changes are complex, but exist between the Lenticulina spp. Assemblage and the P. wuellerstorfi Assemblage at ~13.8 Ma, and between the Ehrenbergina spp. Assemblage and the G. lobatulus Assemblage at ~5 Ma. The development of the P. wuellerstorfi and Globocassidulina spp. Assemblages after 13.8 Ma is correlated with the decrease in temperature of the intermediate waters of the ocean, in turn related to Antarctic glacial expansion. The faunal changes at ~5 Ma are related to the development of low oxygen intermediate water, formed in the presence of a strong thermocline. At Site 756, six varimax assemblages are distributed as follows: the Cibicidoides cf. mundulus-Oridorsalis umbonatus Assemblage in the lower Oligocene; the Epistominella umbonifera-Cibicidoides mundulus Assemblage from the upper Oligocene to the lower Miocene; the Cibicidoides mundulus-Burseolinapacifica Assemblage from lower Miocene to the lower middle Miocene; the Globocassidulina spp. Assemblage from the upper lower Miocene to the Pliocene; the Uvigerina proboscidea Assemblage in the upper Miocene and the Pliocene; and the Globocassidulina sp. D Assemblage in the Pliocene. The main faunal change at this site is between the E. umbonifera Assemblage and the Globocassidulina spp. Assemblage, at ~17.1 Ma. The timing of this faunal change is coeval with faunal changes in the North Atlantic and the Pacific. The change is related to a change in bottom water characteristics caused by an increased influence of carbonate corrosive water from the Antarctic source region, and a change in surface productivity. A low oxygen event at Site 756, which started at about 7.3 Ma, occurred about 2.3 m.y. before that at Site 754. The different response to global paleoceanographic changes is not yet explained, but may be due to the difference of marine topography and the degree of upwelling