Down-core distribution of live and dead benthic foraminifera in deep sea sediments from the Weddell Sea and the Californian continental borderland

Five short cores sub-sampled from box cores from three sites in the eastern Weddell Sea off Antarctica and in the eastern Pacific off southern California, covering a range in water depth from 500 to 2000 m, were analysed for the down-core distribution of live (stained with Rose Bengal) and dead benthic foraminifera. In the California continental borderland, Planulina ariminensis, Rosalina columbiensis and Trochammina spp. live attached to agglutinated polychaetes tubes that rise above the sedimentwater interface. Bolivina spissa lives exclusively in or on the uppermost sediment. Stained specimens of Chilostomella ovoidea are found down to 6 cm within the sediment and specimens of Globobulimina pacifica down to a maximum of 8 cm. Delta13C values of live G. pacifica decrease with increasing depth from the sediment surface down to 7 cm core depth, indicating that this infaunal species utilizes13C-depleted carbon from pore waters. In the dead, predominantly calcareous benthic forminiferal assemblage, selective dissolution of small delicate tests in the upper sediment column causes a continuous variation in species proportions. In the eastern Weddell Sea, the calcareous Bulimina aculeata lives in a carbonate corrosive environment exclusively in or on the uppermost sediment. The arenaceous Cribrostomoides subglobosum, Recurvoides contortus and some Reophax species are frequently found within the top 4 cm of the sediment, whereas stained specimens of Haplophragmoides bradyi, Glomospira charoides and Cribrostomoides wiesneri occur in maximum abundance below the uppermost 1.5 cm. Species proportions in the dead, predominantly arenaceous, benthic foraminiferal assemblage change in three distinct steps. The first change is caused by calcite dissolution at the sediment-water interface, the second coincides with the lower boundary of intense bioturbation, and the third results from the geochemical shift from oxidizing to reducing conditions below a compacted ash layer.

Stable carbon and oxygen isotope composition of benthic foraminifera

Seventeen surface sediment samples from the North Atlantic Ocean off NE-Greenland between 76° and 81°N, and nine samples from the South Atlantic Ocean close to Bouvet Island between 48° and 55°S were taken with the aid of a Multiple Corer and investigated for their live (Rose Bengal stained) benthic foraminiferal content within the upper 15 cm of sediment. Preferentially endobenthic Melonis barleeanum, Melonis zaandami, and Bulimina aculeata as well as preferentially epibenthic Lobatula lobatula were counted from 1-cm-thick sediment slices each and analyzed for stable carbon and oxygen isotopic compositions of their calcareous tests. Live and dead specimens were counted and measured separately. The carbon isotopic composition of the foraminifera was compared to that of the dissolved inorganic carbon (DIC) of simultaneously sampled bottom water. During a period of one month, one station off NE-Greenland was replicately sampled once every week and samples were processed as above. Live specimens of Lobatula lobatula are confined to the uppermost two centimeters of sediment. Live specimens of Melonis spp. are found down to 8 cm within the sediment but with a distinct sub-surface maximum between 2 and 5 cm. The down-core distribution of live Bulimina aculeata shows a distinct surface maximum in the top centimeter and constant but low numbers down to 11-cm subbottom depth. The average stable carbon isotopic composition (d13C versus per mil PDB) of live Lobatula lobatula off NE-Greenland is by 0.4±0.1 per mil higher than the d13CDIC of the ambient bottom water at the time of sampling. There is evidence that this species calcify before the ice-free season, when bottom water d13CDIC is supposed to be higher. This would reconfirm the one-to-one relationship between d13C of ambient water DIC and cibicids, widely used by paleoceanographers. Live Melonis barleeanum show a negative offset from bottom water DIC of -1.7±0.6 per mil in the uppermost sediment and of -2.2±0.5 per mil in 3-4-cm subbottom depth. All d13C values of live Melonis spp. decrease within the upper four centimeters, regardless of the time of sampling and site investigated. The offset of live Bulimina aculeata from bottom water d13CDIC values of 8 stations rather constantly amounts to -0.6±0.1 per mil, no matter what subbottom depth the specimens are from. At one station however, where is strong indication of elevated organic carbon flux, the negative offset averaged over all sub-bottom depths increases to -1.5±0.2 per mil. Buliminids actively move within the sediment and by this either record an average isotope signal of the pore water or the signal of one specific calcification depth. The recorded signal, however, depends on the organic carbon flux and reflects general but site-specific pore water d13CDIC values. If compared with epibenthic d13C values from the same site, not influenced by pore water and related phytodetritus layer effects, Buliminad13C values bear some potential as a paleoproductivity proxy. Specimens of Melonis spp. seem to prefer a more static way of life and calcify at different but individually fix depths within the sediment. Although live specimens thus record a stratified pore water d13C signal, there is no means yet to correct for bioturbational and early diagenetic effects in fossil faunas.

Stable isotopic composition of bottom water and benthic foraminifera from Hakon Mosby Mud Vulcano

To investigate the use of benthic foraminifera as a means to document ancient methane release, we determined the stable isotopic composition of tests of live (Rose Bengal stained) and dead specimens of epibenthic Fontbotia wuellerstorfi, preferentially used in paleoceanographic reconstructions, and of endobenthic high-latitude Cassidulina neoteretis and Cassidulina reniforme from a cold methane-venting seep off northern Norway. We collected foraminiferal tests from three push cores and nine multiple cores obtained with a remotely operated vehicle and a video-guided multiple corer, respectively. All sampled sites except one control site are situated at the Håkon Mosby mud volcano (HMMV) on the Barents Sea continental slope in 1250 m water depth. At the HMMV in areas densely populated by pogonophoran tube worms, d13C values of cytoplasm-containing epibenthic F. wuellerstorfi are by up to 4.4 per mil lower than at control site, thus representing the lowest values hitherto reported for this species. Live C. neoteretis and C. reniforme reach d13C values of -7.5 and -5.5 per mil Vienna Pee Dee Belemnite (VPDB), respectively, whereas d13C values of their empty tests are higher by 4 per mil and 3 per mil. However, d13C values of empty tests are never lower than those of stained specimens, although they are still lower than empty tests from the control site. This indicates that authigenic calcite precipitates at or below the sediment surface are not significantly influencing the stable isotopic composition of foraminiferal shells. The comparatively high d13C results rather from upward convection of pore water and fluid mud during active methane venting phases at these sites. These processes mingle tests just recently calcified with older ones secreted at intermittent times of less or no methane discharge. Since cytoplasm-containing specimens of suspension feeder F. wuellerstorfi are almost exclusively found attached to pogonophores, which protrude up to 3 cm above the sediment, and d13C values of bottom-water-dissolved inorganic carbon (DIC) are not significantly depleted, we conclude that low test d13C values of F. wuellerstorfi are the result of incorporation of heavily 13C-depleted methanotrophic biomass that these specimens feed on rather than because of low bottom water d13CDIC. Alternatively, the pogonophores, which are rooted at depth in the upper sediment column, may serve as a conduit for depleted d13CDIC that ultimately influences the calcification process of F. wuellerstorfi attached to the pogonophoran tube well above the sediment/water interface. The lowest d13C of live specimens of the endobenthic C. neoteretis and C. reniforme are within the range of pore water d13CDIC values, which exceed those that could be due to organic matter decomposition, and thus, in fact, document active methane release in the sediment.

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.

Vertical distribution of benthic foraminifers in the Arctic Ocean

The vertical distribution of living (Rose Bengal stained) benthic foraminifers was determined in the upper 15 cm of sediment cores taken along transects extending from the continental shelf of Spitsbergen through the Eurasian Basin of the Arctic Ocean. Cores taken by a multiple corer were raised from 50 stations with water depths between 94 and 4427 m, from areas with moderate primary production values to areas that are among the least productive ones in the world. We believe, that in the Arctic Ocean the vertical distribution of living foraminifers is determined by the restricted availability of food. Live foraminiferal faunas are dominated by potentially infaunal species or epifaunal species. Species confined to the infaunal microhabitat are absent in Arctic sediments that we examined, and predominantly infaunal living species are nowhere dominant. In general, an infaunal mode of life is restricted to the seasonally ice-free areas and thus to areas with at least moderate primary production during the summer period. Under the permanent ice cover living species are usually restricted to the top centimeter of the sediment surface, even though some are able to dwell deeper in the sediment under ice-free conditions.

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.

Benthic foraminiferal assemblage counts from the Gulf of California

Extensive CO2 vents have been discovered in the Wagner Basin, northern Gulf of California, where they create large areas with lowered seawater pH. Such areas are suitable for investigations of long-term biological effects of ocean acidification and effects of CO2 leakage from subsea carbon capture storage. Here, we show responses of benthic foraminifera to seawater pH gradients at 74-207 m water depth. Living (rose Bengal stained) benthic foraminifera included Nonionella basispinata, Epistominella bradyana and Bulimina marginata. Studies on foraminifera at CO2 vents in the Mediterranean and off Papua New Guinea have shown dramatic long-term effects of acidified seawater. We found living calcareous benthic foraminifera in low pH conditions in the northern Gulf of California, although there was an impoverished species assemblage and evidence of post-mortem test dissolution.

Benthic foraminifera of the Red Sea

The distribution of living (Rose Bengal-stained), dead and fossil benthic foraminifera was investigated in six short cores (multicores, 30-32 cm total length) recovered from the central Red Sea. The ecological preferences as well as the relationship between the live and dead/fossil assemblages (preserved down-core) were examined. The sites, located along a W-E profile and between the depth of 366 and 1782 m, extend from the center of the oxygen minimum zone (OMZ, ~200-650 m), through its margin at ~600 m, and down to the well-aerated deep-water environment. Live (Rose-Bengal stained) and coexisting dead foraminifera were studied in the upper 5 cm of each of the sites, and the fossil record was studied down to ~32 cm. Q-mode Principal Component Analysis was used and four distinct foraminiferal fossil assemblages were determined. These assemblages follow different water mass properties. In the center of the OMZ, where the organic carbon content is highest and the oxygen concentration is lowest (<=0.5 ml O2/l), the Bolivina persiensis-Bulimina marginata-Discorbinella rhodiensis assemblage dominates. The slightly more aerated and lower organic-carbon-content seafloor, at the margin of the OMZ, is characterized by the Neouvigerina porrecta-Gyroidinoides cf. G. soldanii assemblage. The transitional environment, between 900-1200 m, with its well-aerated and oligotrophic seafloor, is dominated by the Neouvigerina ampullacea-Cibicides mabahethi assemblage. The deeper water (>1500 m), characterized by the most oxygenated and oligotrophic seafloor conditions, is associated with the Astrononion sp. A-Hanzawaia sp. A assemblage. Throughout the Red Sea extremely high values of temperature and salinity are constant below ~200 m depth, but the flux of organic matter to the sea floor varies considerably with bathymetry and appears to be the main controlling factor governing the distribution pattern of the benthic foraminifera. Comparison between live and the dead/fossil assemblages reveals a large difference between the two. Processes that may control this difference include species-specific high turnover rates, and preferential predation and loss of fragile taxa (either by chemical or microbial processes). Significant variations in the degree of loss of the organic-cemented agglutinants were observed down core. This group is preserved down to 5-10 cm at the shallow OMZ sites and down to greater depths at well-aerated and oligotrophic sites. The lower rate of disintegration of these forms, in the deeper locations of the Red Sea, may be related to low microbial activity. This results in the preservation of increasing numbers of organic-cemented shells down-core.

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.

Benthic foraminifera in surface sediments, South Atlantic

Sixty surface sediment samples from the eastern South Atlantic Ocean including the Walvis Ridge, the Angola and Cape basins, and the Southwest African continental margin were analysed for their benthic foraminiferal content to unravel faunal distribution patterns and ecological preferences. Live (stained with Rose Bengal) and dead faunas were counted separately and then each grouped by Q-mode principal component analysis into seven principal faunal end-members. Then, multiple regression technique was used to correlate Recent assemblages with available environmental variables and to finally differentiate between four principal groups of environmental agents acting upon the generation of benthic foraminiferal assemblages: (1) seasonality of food supply and organic carbon flux rates, together with oxygen content in the pore and bottom waters; (2) lateral advection of deep-water masses; (3) bottom water carbonate corrosiveness; and (4) energetic state at the benthic boundary layer and grain size composition of the substrate. Food supply and corresponding dissolved oxygen contents in the pore and bottom waters turned out to be the most important factors which control the distribution pattern of the Recent benthic foraminifera. At the continental margin, in the zone of coastal upwelling and its mixing area, benthic foraminiferal assemblages are dominated by stenobathic high-productivity faunas, characterized by elevated standing stocks, low diversities and a large number of endobenthic living species. At the continental shelf and upper continental slope the live assemblages are characterized by Rectuvigerina cylindrica, Uvigerina peregrina s.1., Uvigerina auberiana and Rhizammina spp. while the dead assemblages are characterized by Cassidulina laevigata, Bolivina dilatata, Bulimina costata and B. mexicana. At the lower continental slope strong influence of high organic matter fluxes on the species composition is restricted to the area off the Cunene river mouth, where the live assemblage is dominated by Uvigerina peregrina s.1., the corresponding dead assemblage by Melonis barleeanum and M. zaandamae. In the adjacent areas of the lower continental slope the biocoenosis is characterized by Reophax bilocularis, and Epistominella exigua which becomes dominant in the corresponding dead assemblage. At the Walvis Ridge and in the abyssal Angola and Cape basins, where organic matter fluxes are low and highly seasonal, benthic foraminiferal assemblages reflect both the oligotrophic situation and the deep and bottom water mass configuration. The top and flanks of the Walvis Ridge are inhabited by the Rhizammina, Psammosphaera and R. bilocularis live assemblages, the corresponding dead assemblages are dominated by G. subglobosa on the ridge top and E. exigua on the flanks. Within the highly diverse E. exigua dead assemblage several associated epibenthic species coincide with the core of NADW between about 1600 and 3700 m water depth. These species include Osangularia culter, Cibicidoides kullenbergi, Melonis pompilioides, Bolivinita pseudothalmanni and Bulimina alazanensis. The assemblages of the abyssal Cape and Angola basins are characterized by Nuttallides umbonifer and a high proportion of agglutinated species. These species are adapted to very low organic matter fluxes and a carbonate corrosive environment.