Late Oligocene to Recent benthic foraminifers from the northeastern North Atlantic

Deep-sea benthic foraminiferal faunas were studied from Sites 608 (depth 3534 m, 42°50'N, 23°05'W) and 610 (depth 2427 m, 53°13'N, 18°53'W). The sampling interval corresponded to 0.1 to 0.5 m.y. at Site 608 and in the sections of Site 610 from which core recovery was continuous. First and last appearances of benthic foraminiferal taxa are generally not coeval at the two sites, although the faunal patterns are similar and many species occur at both sites. Major periods of changes in the benthic faunas, as indicated by the numbers of first and last appearances and changes in relative abundances, occurred in the early Miocene (19.2-17 Ma), the middle Miocene (15.5-13.5 Ma), the late Miocene (7-5.5 Ma), and the Pliocene-Pleistocene (3.5-0.7 Ma). A period of minor changes in the middle to late Miocene (10-9 Ma) was recognized at Site 608 only. These periods of faunal changes can be correlated with periods of paleoceanographic changes: there was a period of sluggish circulation in the northeastern North Atlantic from 19.2 to 17 Ma, and the deep waters of the oceans probably cooled between 15.5 and 13.5 Ma, as indicated by an increase in delta18O values in benthic foraminiferal tests. The period between 10 and 9 Ma was probably characterized by relatively vigorous bottom-water circulation in the northeastern Atlantic, as indicated by the presence of a widespread reflector. The faunal change at 7 to 5.5 Ma corresponds in time with a worldwide change in delta13C values, and with the Messinian closing of the Mediterranean. The last and largest faunal changes correspond in time with the onset and intensification of Northern Hemisphere glaciation.

Benthic foraminifera in the Arctic Ocean

We examine the quantitative composition of benthic foraminiferal assemblages of Rose Bengal-stained surface samples from 37 stations in the Laptev Sea, and combine this data set with an existing data set along a transect from Spitsbergen to the central Arctic Ocean. Foraminiferal test accumulation rates, diversity, faunal composition and statistically defined foraminiferal associations are analysed for living (Rose Bengal-stained) and dead foraminifers. We compare the results of several benthic foraminiferal diversity indices and statistically defined foraminiferal associations, including Fisher's alpha and Shannon-Wiener diversity indices, Q-mode principal component analysis and correspondence analysis. Diversity and faunal density (standing stock) of living benthic foraminifers are positively correlated to trophic resources. In contrast, the accumulation rate of dead foraminifers (BFAR) shows fluctuating values depending on test disintegration processes. Foraminiferal associations defined by Q-mode principal component analysis and correspondence analysis are comparable. The factor values of the correspondence analysis allow a quantitative correlation between the foraminiferal fauna and the local carbon flux, which may be used as a tool to estimate changes in primary productivity.

Benthic foraminifera off West Africa (1°N to 32°S)

Recent benthic foraminifera (> 125 µm) were investigated from multicorer samples on a latitudinal transect of 20 stations between 1°N and 32°S along the upper slope off West Africa. Samples were selected from a narrow water depth interval, between 1200 and 1500 m, so that changes in water masses are minimized, but changes in surface productivity are important and the only significant environmental variable. Live (Rose Bengal stained) benthic foraminifera were counted from the surface sediment down to a maximum of 12 cm. Dead foraminifera were investigated in the top 5 cm of the sediment only. Five live and five dead benthic foraminiferal assemblages were identified using Q-mode principal component analysis, matching distinct primary productivity provinces, characterized by different systems of seasonal and permanent upwelling. Differences in seasonality, quantity, and quality of food supply are the main controlling parameters on species composition and distribution of the benthic foraminiferal faunas. To test the sensitivity of foraminiferal studies based on the uppermost centimeter of sediment only, a comparative Q-mode principal component analysis was conducted on live and dead foraminiferal data from the top 1 cm of sediment. It has been demonstrated that, on the upper slope off West Africa, most of the environmental signals as recorded by species composition and distribution of the 'total' live and dead assemblages, i.e., including live and dead foraminifera from the surface sediment down to 12 cm and 5 cm, respectively, can be extracted from the assemblages in the top centimeter of sediment only. On the contrary, subsurface abundance maxima of live foraminifera and dissolution of empty tests strongly bias quantitative approaches based on the calculation of standing stocks and foraminiferal numbers in the topmost centimeter.

Benthic foraminifera and phytodetritus species in sediments from DSDP Leg 85, and ODP Legs 113 and 143

From late middle Eocene through earliest Oligocene, high-latitude regions cooled, and by the end of the period, continental ice sheets existed in Antarctica. Diversity of planktonic microorganisms declined, and modern groups of terrestrial vertebrates originated. Coeval faunal changes in deep-sea benthic foraminifers have been related to cooling of deep waters and increased oxygenation. Cooling, however, occurred globally, whereas species richness declined at high latitudes and not in the tropics. The late Eocene and younger lower-diversity, high-latitude faunas typically contain common Epistominella exigua and Alabaminella weddellensis, opportunistic phytodetritus-exploiting species that indicate a seasonally fluctuating input of organic matter to the sea floor. We speculate that the species-richness gradient and increase in abundance of phytodetritus-exploiting species resulted largely from the onset of a more unpredictable and seasonally fluctuating food supply, especially at high latitudes.

Benthic foraminifera of ODP Leg 104 holes

Benthic foraminifers have been studied in about 900 samples from Sites 642, 643, and 644 (ODP Leg 104, Voring Plateau), ranging in age from Eocene to Holocene. This sequence has been subdivided into seven assemblage zones. The Eocene to middle Miocene deposits are characterized by an agglutinated fauna. This reflects an environment causing dissolution of calcareous tests rather than the original living fauna. The upper Miocene to middle Pliocene deposits contain a diverse benthic foraminiferal fauna dominated by calcareous forms. The uppermost part of the sediment record, deposited during late Pliocene to Holocene, is characterized by many barren intervals and samples containing shallow-water species as well as ice-rafted material indicating glacial periods. Interglacials are reflected in samples containing a true oceanic foraminifer assemblage and no coarse clastic material.

Late Eocene to Recent deep-sea benthic foraminifera from the Central equatorial Pacific Ocean

Benthic foraminifers were studied in upper Eocene to Recent core-catcher samples from DSDP Sites 573, 574, and 575. The sites are on a north-south transect from the equator to about 05°N at about 133°W, water depth 4300 to 4600 m. At Site 574 additional samples were used to study the Eocene/Oligocene boundary in detail. About 200 specimens were counted per sample. The fauna is highly diverse (about 50 to 70 species per sample) and is of low dominance. The diversity is not related to age or sub-bottom depth. Many species are cosmopolitan and probably have wide environmental tolerances. Fluctuations in frequency of some taxa (e.g., Nuttallides umbonifera, Epistominella exigua, and Uvigerina spp.) cannot be correlated from one site to another. Several common species (e.g. Oridorsalis umbonatus and Globocassidulina subglobosa) range from late Eocene to Recent. First and last appearances are generally difficult to define precisely because many species are rare. For some species these datums differ from one site to another, but several datum levels are within 1 m.y. at all sites. First and last appearances are most numerous in two intervals, the late Eocene to early Oligocene (about 32 to 37 Ma) and the early to middle Miocene (about 13 to 18.5 Ma). Isotopic events occur within each of these periods of benthic faunal change, but the isotopic events have a shorter duration and start after the initiation of the changes in the fauna. Changes in deep-sea benthic faunal composition are not directly related to short-term oceanographic changes as expressed in isotopic records.

Communities and microhabitats of living benthic foraminifera from the tropical East Atlantic

Living (Rose Bengal stained) benthic foraminifera were collected with a multicorer from six stations between 2°N and 12°S off West Africa. The foraminiferal communities in the investigated area reflect the direct influence of different productivity regimes, and are characterized by spatially and seasonally varying upwelling activity. At five stations, foraminiferal abundance coincides well with the gradient of surface productivity. However, at one station off the Congo River, the influence of strong fresh water discharge is documented. Although this station lies directly in the center of an upwelling area, foraminiferal standing stocks are surprisingly low. It is suggested that the Congo discharge may induce a fractionation of the organic matter into small and light particles of low nutritional content, by contrast to the relatively fast-sinking aggregates found in the centers of high productivity areas. Quality and quantity of the organic matter seem to influence the distribution of microhabitats as well. The flux of organic carbon to the sea-floor controls the sequence of degradation of organic matter in sediment and the position of different redox fronts. The vertical foraminiferal stratification within sediment closely parallels the distribution of oxygen and nitrate in porewater, and reflects different nutritive strategies and adaptation to different types of organic matter. The epifauna and shallow infauna colonize oxygenated sediments where labile organic matter is available. The intermediate infauna (M. barleeanum) is linked to the zone of nitrate reduction in sediments where epifaunal and shallow infaunal species are not competitive anymore, and must feed on bacterial biomass or on metabolizable nutritious particles produced by bacterial degradation of more refractory organic matter. The deep infauna shows its maximum distribution in anoxic sediments, where no easily metabolizable organic matter is available.

(Appendix B) Benthic foraminifera in Indian Ocean surface sediments

Multivariate analysis was performed on percentages of 46 species of unstained deep-sea benthic foraminifera from 131 core-top to near-core-top samples (322-5013 m) from across the Indian Ocean. Faunal data are combined with GEOSECS geochemical data to investigate any relationship between benthic foraminifera (assemblages and species) and deep-sea properties. In general, benthic foraminifera show a good correlation to surface productivity, organic carbon flux to the sea floor, deep-sea oxygenation and, to a lesser extent, to bottom temperature, without correlation with the water depths. The foraminiferal census data combined with geochemical data has enabled the division of the Indian Ocean into two faunal provinces. Province A occupies the northwestern Indian Ocean (Arabian Sea region) where surface primary production has a major maximum during the summer monsoon season and a secondary maximum during winter monsoon season that leads to high organic flux to the seafloor, making the deep-sea one of the most oxygen-deficient regions in the world ocean, with a pronounced oxygen minimum zone (OMZ). This province is dominated by benthic foraminifera characteristic of low oxygen and high organic food flux including Uvigerina peregrina, Robulus nicobarensis, Bolivinita pseudopunctata, Bolivinita sp., Bulimina aculeata, Bulimina alazanensis, Ehrenbergina carinata and Cassidulina carinata. Province B covers southern, southeastern and eastern parts of the Indian Ocean and is dominated by Nuttallides umbonifera, Epistominella exigua, Globocassidulina subglobosa, Uvigerina proboscidea, Cibicides wuellerstorfi, Cassidulina laevigata, Pullenia bulloides, Pullenia osloensis, Pyrgo murrhina, Oridorsalis umbonatus, Gyroidinoides (= Gyroidina) soldanii and Gyroidinoides cf. gemma suggesting well-oxygenated, cold deep water with low (oligotrophic) and pulsed food supply.

Benthic foraminifera in eastern equatorial Indian Ocean surface sediments

Study of Recent abyssal benthic foraminifera from core-top samples in the eastern equatorial Indian Ocean has identified distinctive faunas whose distribution patterns reflect the major hydrographic features of the region. Above 3800 m, Indian Deep Water (IDW) is characterized by a diverse and evenly-distributed biofacies to which Globocassidulina subglobosa, Pyrgo spp., Uvigerina peregrina, and Eggerella bradyi are the major contributors. Nuttalides umbonifera and Epistominella exigua are associated with Indian Bottom Water (IBW) below 3800 m. Within the IBW fauna, N. umbonifera and E. exigua are characteristic of two biofacies with independent distribution patterns. Nuttalides umbonifera systematically increases in abundance with increasing water depth. The E. exigua biofacies reaches its greatest abundance in sediments on the eastern flank of the Ninetyeast Ridge and in the Wharton-Cocos Basin. The hydrographic transition between IDW and IBW coincides with the level of transition from waters supersaturated to waters undersaturated with respect to calcite and with the depth of the lysocline. Carbonate saturation levels, possibly combined with the effects of selective dissolution on the benthic foraminiferal populations, best explain the change in faunas across the IDW/IBW boundary and the bathymetric distribution pattern of N. umbonifera. The distribution of the E. exigua fauna cannot be explained with this model. Epistominella exigua is associated with the colder, more oxygenated IBW of the Wharton-Cocos Basin. The distribution of this biofacies on the eastern flank of the Ninetyeast Ridge agrees well with the calculated bathymetric position of the northward flowing deep boundary current which aerates the eastern basins of the Indian Ocean.