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 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.

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.

Compilation of respiration rate data for marine pelagic organisms

The metabolic rate of organisms may either be viewed as a basic property from which other vital rates and many ecological patterns emerge and that follows a universal allometric mass scaling law; or it may be considered a property of the organism that emerges as a result of the organism's adaptation to the environment, with consequently less universal mass scaling properties. Data on body mass, maximum ingestion and clearance rates, respiration rates and maximum growth rates of animals living in the ocean epipelagic were compiled from the literature, mainly from original papers but also from previous compilations by other authors. Data were read from tables or digitized from graphs. Only measurements made on individuals of know size, or groups of individuals of similar and known size were included. We show that clearance and respiration rates have life-form-dependent allometries that have similar scaling but different elevations, such that the mass-specific rates converge on a rather narrow size-independent range. In contrast, ingestion and growth rates follow a near-universal taxa-independent ~3/4 mass scaling power law. We argue that the declining mass-specific clearance rates with size within taxa is related to the inherent decrease in feeding efficiency of any particular feeding mode. The transitions between feeding mode and simultaneous transitions in clearance and respiration rates may then represent adaptations to the food environment and be the result of the optimization of tradeoffs that allow sufficient feeding and growth rates to balance mortality.

Compilation of growth rate data for marine pelagic organisms

The metabolic rate of organisms may either be viewed as a basic property from which other vital rates and many ecological patterns emerge and that follows a universal allometric mass scaling law; or it may be considered a property of the organism that emerges as a result of the organism's adaptation to the environment, with consequently less universal mass scaling properties. Data on body mass, maximum ingestion and clearance rates, respiration rates and maximum growth rates of animals living in the ocean epipelagic were compiled from the literature, mainly from original papers but also from previous compilations by other authors. Data were read from tables or digitized from graphs. Only measurements made on individuals of know size, or groups of individuals of similar and known size were included. We show that clearance and respiration rates have life-form-dependent allometries that have similar scaling but different elevations, such that the mass-specific rates converge on a rather narrow size-independent range. In contrast, ingestion and growth rates follow a near-universal taxa-independent ~3/4 mass scaling power law. We argue that the declining mass-specific clearance rates with size within taxa is related to the inherent decrease in feeding efficiency of any particular feeding mode. The transitions between feeding mode and simultaneous transitions in clearance and respiration rates may then represent adaptations to the food environment and be the result of the optimization of tradeoffs that allow sufficient feeding and growth rates to balance mortality.

Distribution of Upper Cretaceous deep water agglutinated foraminifera in sediments of the North Atlantic and its marginal seas (Tab. 1)

The stratigraphic and biogeographic distribution of more than 170 species of deep-water agglutinated benthic foraminifers (DWAF) from the North Atlantic and adjacent marginal seas has been compared with paleoenvironmental data (e.g. paleobathymetry, oxygenation of the bottom waters, amount of terrigenous input and substrate disturbance). Six general types of assemblages, in which deep water agglutinated taxa occur, are defined from the Turonian to Maastrichtian times: 1. High latitude slope assemblages 2. Low to mid latitude slope assemblages 3. Flysch-type assemblages 4. Deep water limestone assemblages (,,Scaglia,,-type) 5. Abyssal mixed calcareous-agglutinated assemblages 6. Abyssal purely agglutinated assemblages Latitudinal differences in faunal composition are observed, the most important of which is the lack or extreme paucity of calcareous forms in high latitude assemblages. East-to-west differences appear to be of comparatively minor importance. Most DWAF species occur in all studied regions and are thus considered as cosmopolitan. Biostratigraphic turnovers in the taxonomic content of assemblages are observed in the lowermost Turonian, mid-Campanian and in the upper Maastrichtian to lowermost Paleocene. These datum levels correspond to inter-regional and time-constant paleooceanographic events, which probably also affected the deep-water benthic biota. This allows us to use deep-water agglutinated foraminifers for biostratigraphy in the North Atlantic sequences deposited below CCD and to geographically extend the currently used zonal schemes which have been established in the Carpathian and Alpine areas.

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.

Diatom abundance in surface sediments of the northern North Pacific

In order to map the modern distribution of diatoms and to establish a reliable reference data set for paleoenvironmental reconstruction in the northern North Pacific, a new data set including the relative abundance of diatom species preserved in a total of 422 surface sediments was generated, which covers a broad range of environmental variables characteristic of the subarctic North Pacific, the Sea of Okhotsk and the Bering Sea between 30° and 70°N. The biogeographic distribution patterns as well as the preferences in sea surface temperature of 38 diatom species and species groups are documented. A Q-mode factor analysis yields a three-factor model representing assemblages associated with the Arctic, Subarctic and Subtropical water mass, indicating a close relationship between the diatom composition and the sea surface temperatures. The relative abundance pattern of 38 diatom species and species groups was statistically compared with nine environmental variables, i.e. the summer sea surface temperature and salinity, annual surface nutrient concentration (nitrate, phosphate, silicate), summer and winter mixed layer depth and summer and winter sea ice concentrations. Canonical Correspondence Analysis (CCA) indicates 32 species and species groups have strong correspondence with the pattern of summer sea surface temperature. In addition, the total diatom flux data compiled from ten sediment traps reveal that the seasonal signals preserved in the surface sediments are mostly from spring through autumn. This close relationship between diatom composition and the summer sea surface temperature will be useful in deriving a transfer function in the subarctic North Pacific for the quantitative paleoceanographic and paleoenvironmental studies. The relative abundance of the sea-ice indicator diatoms Fragilariopsis cylindrus and F. oceanica of >20% in the diatom composition is used to represent the winter sea ice edge in the Bering Sea. The northern boundary of the distribution of F. doliolus in the open ocean is suggested to be an indicator of the Subarctic Front, while the abundance of Chaetoceros resting spores may indicate iron input from nearby continents and shelves and induced productivity events in the study area.

Late Quaternary deep sea sedimentation off the Niger delta

The oxygen isotopes ratios of benthic foraminifera and detailed radiocarbon ages of the organic matter of an over 15 m long sediment core from the outer Niger delta allow us to date the oxygen isotope stage boundaries 1/2 to 11500 (+/- 650) years BP, 2/3 to approximately 23000 (+/- 2000) years BP. The composition of the predominantly terrigenous clays and accessory pelagic fossils reflects the evolution of the climate over the southwestern Sahel zone and the response of the Eastern Tropical Atlantic to these climatic fluctuations during the Late Quaternary. The dilution of the pelagic fossil concentrations by the terrigenous material and the oxygen isotopes ratios of planktonic foraminifera indicate large fluctuations in the freshwater discharge from the Niger, with high precipitations over the drainage area of this river from 4500 (+/- 300) to 11500 (+/- 650) years BP and from 11800 (+(- 600) to 13000 (+/- 600) years BP while the time intervals in between were as dry as today. Relative increase of kaolinite during wet phases and the association of smectite, chlorite and attapulgite during dry ones characterize the response of the weathering in the Niger drainage basins to the climatic fluctuations. The occurrence of 10-14 A mixed-layers prior to 26000 years BP is correlated with moderate alteration of the crystalline substratum outcrops from the middle-lower part of the Niger Basin. High quartz concentrations are particularly typical for the transition between oxygen isotope stages 1 and 2 at the inception of heavy precipitations in the southern Sahel zone. Sedimentation rates were quite constant, 30-35 cm/1000 years; they became unusually large at the beginning of the Holocene from 10900 (+/- 650) to 11500 (+/- 650) years BP where they reached more than 600 cm/1000 years. Bottom waters around 1100 m depth in the Gulf of Guinea responded to changes in paleo-oceanography of the entire Atlantic Ocean as well as to local influences. Abnormal carbon isotopes ratios and the drastic changes from a highly diversified fauna (during stages 2 and 3. and during the last part of stage 1 after approx. 7000 years BP) to a poorly diversified fauna in the intervenin time span point to the development of a local benthic environment which cannot easily be compared with the corresponding continental and slope environments of the entire Atlantic Ocean.

Benthic foraminiferal record from Santa Barbara basin

The oxygen minimum zone (OMZ) of the late Quaternary California margin experienced abrupt and dramatic changes in strength and depth in response to changes in intermediate water ventilation, ocean productivity, and climate at orbital through millennial time scales. Expansion and contraction of the OMZ is exhibited at high temporal resolution (107-126 year) by quantitative benthic foraminiferal assemblage changes in two piston cores forming a vertical profile in Santa Barbara Basin (569 m, basin floor; 481 m, near sill depth) to 34 and 24 ka, respectively. Variation in the OMZ is quantified by new benthic foraminiferal groupings and new dissolved oxygen index based on documented relations between species and water-mass oxygen concentrations. Foraminiferal-based paleoenvironmental assessments are integrated with principal component analysis, bioturbation, grain size, CaCO3, total organic carbon, and d13C to reconstruct basin oxygenation history. Fauna responded similarly between the two sites, although with somewhat different magnitude and taxonomic expression. During cool episodes (Younger Dryas and stadials), the water column was well oxygenated, most strongly near the end of the glacial episode (17-16 ka; Heinrich 1). In contrast, the OMZ was strong during warm episodes (Bølling/Allerød, interstadials, and Pre-Boreal). During the Bølling/Allerød, the OMZ shoaled to <360 m of contemporaneous sea level, its greatest vertical expansion of the last glacial cycle. Assemblages were then dominated by Bolivina tumida, reflecting high concentrations of dissolved methane in bottom waters. Short decadal intervals were so severely oxygen-depleted that no benthic foraminifera were present. The middle to late Holocene (6-0 ka) was less dysoxic than the early Holocene.

Standing stocks and carbon isotopes of live benthic foraminifera from the South Atlantic

Live (Rose Bengal stained) and dead benthic foraminifera of surface and subsurface sediments from 25 stations in the eastern South Atlantic Ocean and the Atlantic sector of the Southern Ocean were analyzed to decipher a potential influence of seasonally and spatially varying high primary productivity on the stable carbon isotopic composition of foraminiferal tests. Therefore, stations were chosen so that productivity strongly varied, whereas conservative water mass properties changed only little. To define the stable carbon isotopic composition of dissolved inorganic carbon (d13CDIC) in ambient water masses, we compiled new and previously published d13CDIC data in a section running from Antarctica through Agulhas, Cape and Angola Basins, via the Guinea Abyssal Plain to the Equator. We found that intraspecific d13C variability of all species at a single site is constantly low throughout their distribution within the sediments, i.e. species specific and site dependent mean values calculated from all subbottom depths on average only varied by +/-0.09 per mil. This is important because it makes the stable carbon isotopic signal of species independent of the particular microhabitat of each single specimen measured and thus more constant and reliable than has been previously assumed. So-called vital and/or microhabitat effects were further quantified: (1) d13C values of endobenthic Globobulimina affinis, Fursenkoina mexicana, and Bulimina mexicana consistently are by between -1.5 and -1.0 per mil VPDB more depleted than d13C values of preferentially epibenthic Fontbotia wuellerstorfi, Cibicidoides pachyderma, and Lobatula lobatula. (2) In contrast to the Antarctic Polar Front region, at all stations except one on the African continental slope Fontbotia wuellerstorfi records bottom water d13CDIC values without significant offset, whereas L. lobatula and C. pachyderma values deviate from bottom water values by about -0.4 per mil and -0.6 per mil, respectively. This adds to the growing amount of data on contrasting cibicid d13C values which on the one hand support the original 1:1-calibration of F. wuellerstorfi and bottom water d13CDIC, and on the other hand document severe depletions of taxonomically close relatives such as L. lobatula and C. pachyderma. At one station close to Bouvet Island at the western rim of Agulhas Basin, we interpret the offset of -1.5 per mil between bottom water d13CDIC and d13C values of infaunal living Bulimina aculeata in contrast to about -0.6 +/- 0.1 per mil measured at eight stations close-by, as a direct reflection of locally increased organic matter fluxes and sedimentation rates. Alternatively, we speculate that methane locally released from gas vents and related to hydrothermal venting at the mid-ocean ridge might have caused this strong depletion of 13C in the benthic foraminiferal carbon isotopic composition. Along the African continental margin, offsets between deep infaunal Globobulimina affinis and epibenthic Fontbotia wuellerstorfi as well as between shallow infaunal Uvigerina peregrina and F. wuellerstorfi, d13C values tend to increase with generally increasing organic matter decomposition rates. Although clearly more data are needed, these offsets between species might be used for quantification of biogeochemical paleogradients within the sediment and thus paleocarbon flux estimates. Furthermore, our data suggest that in high-productivity areas where sedimentary carbonate contents are lower than 15 weight %, epibenthic and endobenthic foraminiferal d13C values are strongly influenced by 13C enrichment probably due to carbonate-ion undersaturation, whereas above this sedimentary carbonate threshold endobenthic d13C values reflect depleted pore water d13CDIC values.