Benthic foraminiferal assemblages and paleoproductivity reconstructions for the last 550 kyr of the Ontong Java Plateau, western equatorial Pacific

We analyzed foraminiferal and nannofossil assemblages and stable isotopes in samples from ODP Hole 807A on the Ontong Java Plateau in order to evaluate productivity and carbonate dissolution cycles over the last 550 kyr (kilo year) in the western equatorial Pacific. Our results indicate that productivity was generally higher in glacials than during interglacials, and gradually increased since MIS 13. Carbonate dissolution was weak in deglacial intervals, but often reached a maximum during interglacial to glacial transitions. Carbonate cycles in the western equatorial Pacific were mainly influenced by changes of deep-water properties rather than by local primary productivity. Fluctuations of the estimated thermocline depth were not related to glacial to interglacial alternations, but changed distinctly at ~280 kyr. Before that time the thermocline was relatively shallow and its depth fluctuated at a comparatively high amplitude and low frequency. After 280 kyr, the thermocline was deeper, and its fluctuations were at lower amplitude and higher frequency. These different patterns in productivity and thermocline variability suggest that thermocline dynamics probably were not a controlling factor of biological productivity in the western equatorial Pacific Ocean. In this region, upwelling, the influx of cool, nutrient-rich waters from the eastern equatorial Pacific or of fresh waters from rivers have probably never been important, and their influence on productivity has been negligible over the studied period. Variations in the inferred productivity in general are well correlated with fluctuations in the eolian flux as recorded in the northwestern Pacific, a proxy for the late Quaternary history of the central East Asian dust flux into the Pacific. Therefore, we suggest that the dust flux from the central East Asian continent may have been an important driver of productivity in the western Pacific.

Analysis of foraminifera from the Tagus Estuary, Portugal

The distribution and microhabitat of living benthic fora- minifera (15 calcareous and 6 agglutinated) have been studied in two box cores from the Tagus Prodelta. Stable oxygen and carbon isotopes were analysed for eight different species from six surface samples from the Tagus Prodelta and Estuary. At the two box core stations, most of the living foraminifera were restricted to the oxygenated top cm of the sediment and generally show a shallow infaunal behavior. Those taxa are e.g. Rectuvigerina phlegeri, Stainforthia fusiformis and species of the genus Bolivina, which is the most abundant genus in the Tagus Prodelta. Infaunal species are found down to 10 cm depth, and some infaunal taxa, e.g. Bulimina marginata, Globobulimina auriculata and Nonionella turgida, inhabit the low oxic or anoxic sediments. The deep infaunal species are suggested to feed selectively, on refractory organic matter or on the bacterial stocks, while the opportunistic shallow infaunal species are believed to feed on fresh phytodetritus or labile organic matter. Our data show that there is a close connection between the concentration of foraminifera and the distribution of organic matter in the area. The highest abundance of living benthic foraminifera was found in sediments close to the Tagus river plume, where the sediments have relatively high organic carbon contents. The spatial distribution of the stable isotope values of different benthic foraminifera reflects the distribution of the low salinity and relatively high temperature water with high organic carbon fluxes within the Tagus Estuary.

Stable oxygen isotope records of different benthic foraminiferal species of core GeoB3004-1 from the western Arabian Sea

The stable isotope composition of one epifaunal and three infaunal benthic foraminiferal species of a sediment core from 1800 m water depth of the western Arabian Sea was determined to evaluate deepwater oxygenation, organic matter remineralization, and early diagenetic processes during the past 190,000 years. The d18O records reveal species-specific metabolic effects, susceptibility to changes in carbonate ion concentration, and supralysoclinal calcite dissolution. The foraminiferal d13C records reveal changes in the stable carbon isotope gradients of pore water dissolved inorganic carbon (d13CDIC) and in the microhabitat depth of infaunal species. Maximum d13CDIC offsets between bottom and pore waters ranged between mean values of 0.8 and 1.2% corresponding to estimates of deepwater oxygen concentration between approximately 1 and 2.7 ml/l. Intervals of improved deepwater oxygenation coincided with high benthic foraminiferal diversity and indicate the admixture of well-oxygenated deepwater masses during interglacials. During interglacial maxima the d13C difference between epifauna and shallow infauna indicates highest organic matter remineralization rates at times of maximum organic matter fluxes.

Gut filling, gut content and fatty acid composition of demersal fish species sampled during POLARSTERN cruise ANT-XXIV/2

The gut contents and fatty acid composition of 49 fish belonging to five Antarctic demersal families (Nototheniidae, Macrouridae, Channichtyidae, Bathydraconidae and Artedidraconidae) sampled at two stations at the Southern Ocean shelf and deep sea (600 and 2150 m) were analysed in order to identify their main food resource by linking trophic biomarkers with the dietary items found in the fish guts. Main food items of most fish analysed were amphipod crustaceans (e.g. in 63% of Trematomus bernachii guts) and polychaetes (e.g. in 80% of Bathydraco sp. guts), but other food items including fish, other crustaceans and gastropods were also ingested. The most prominent fatty acids found were 20:5(n-3), 16:0, 22:6(n-3) and 18:1(n-9). The results of gut content and fatty acid analyses indicate that all fish except the Channichthyidae share similar food resources irrespective of their depth distribution, i.e. benthic amphipods and polychaetes. A difference of the dietary spectrum can be observed with ontogenetic phases rather than between species, as high values of typical calanoid copepod marker fatty acids as 22:1(n-11) indicate that younger (smaller) specimens include more zooplankton in their diet.

Abyssal community analysis from replicate box cores in the central North Pacific

A 0.25 m**2 United States Naval Electronics Laboratory box corer was used to take replicate samples from an oligotrophic bottom under the North Pacific Central Water Mass (~28°N, 155°W). The bottom is a red clay with manganese nodules at a depth of 5500-5800 m. Macrofaunal density ranges from 84 to 160 individuals per m**2 and is therefore much the same as in Northwest Atlantic Gyre waters. Of the macrofaunal taxa, polychaetes dominate (55 %), followed by tanaids (18 %), bivalves (7 %), and isopods (6 %). Meiofaunal taxa were only partially retained by the 297 µm screen used in washing. Even then, they are 1.5-3.9 times as abundant as the macrofaunal taxa, with nematodes being numerically dominant by far. Foraminifera seem to comprise an important portion of the community, but could not be assessed accurately because of the inability to discriminate living and dead tests. Remains of what are probably xenophyophoridans are also very important, but offer the same problem. Faunal diversity is extremely high, with deposit feeders comprising the overwhelming majority. Most species are rare, being encountered only once. The distributions of only three species show any significant deviation from randomness. The polychaete fauna from box cores collected from 90 miles to the north was not significantly different from that of the principal study locality. Concordance appeared at several taxonomic levels, from species through macrofaunal/meiofaunal relationships. As a result, the variation in total animal abundance shows aggregation among cores. We discuss Sokolova's concept of a deep-sea oligotrophic zone dominated by suspension feeders, and reconcile it with our present findings. The high diversity of the fauna combined with the low food level contradict theories that relate diversity directly with productivity.

Benthic foraminiferal fauna in the Ionian Sea

To reconstruct paleoceanographic changes in the eastern Mediterranean during the last 330,000 years, we studied benthic foraminifera in a piston core from the Ionian Sea. The fauna exhibits large fluctuations in foraminiferal number, diversity, and species composition. Interglacials are characterized by low foraminiferal number and diversity indicating oligotrophic conditions. Directly below or above interglacial sapropels, increased numbers of low-oxygen-tolerant species indicate a strong reduction of deep water circulation. Glacials are characterized by increased foraminiferal number and diversity and faunas that are dominated by shallow infaunal species indicating mesotrophic conditions. Around glacial sapropel S6 very high foraminiferal numbers and the dominance of shallow and deep infaunal species suggest enhanced organic matter fluxes. These faunal results provide information about changes in the African and North Atlantic climate systems (monsoon and westerlies) controlling the humidity and wind stress in the Mediterranean region.

(Table 2) Species composition of agglutinated foraminifers from the abyssal zone of the Pacific Ocean

The quantitative study of distribution and taxonomic composition of recent living and dead (without plasma) benthic foraminifers revealed three foraminiferal assemblages in bottom sediments of the Pacific Ocean at depths of 3350 to 4981 m. The assemblage dominated by epibenthic Lagenammina difflugiformis, Reophax dentaliniformis, and Saccorhiza ramose occupies slopes of underwater hills. The assemblage with a high share of infaunal Cribrostomoides subglobosum, C. nitidum, and Ammobaculites agglutinans is registered on an abyssal plateau. The assemblage with a significant proportion of large Astrorhiza and Reophax species, which are characterized by active way of life, populates gentle slopes and narrow depressions with potentially strong bottom currents.

Benthic foraminiferal number of specimens in Cretaceous samples of various DSDP and ODP Sites (Table 2)

Benthic foraminifera from 24 DSDP/ODP sites were investigated to assess their global horizontal and vertical distribution in the deep-sea environment at the end of the Cretaceous period. The samples analyzed are from the late Maastrichtian and within the planktic foraminiferal Abathomphus mayaroensis Zone from a wide range of oceans and paleolatitudes, including the low-latitude Sites 10 and 384 (Atlantic Ocean), 47, 171, 305, and 465 (Pacific Ocean), the mid-latitude Sites 20, 111, 356, 363, 516, 525, 527, 548, and 605 (Atlantic Ocean), 216, 217, and 758 (Indian Ocean), and the high-latitude Sites 208 (Pacific Ocean), 689,698,700,738 and 750 (Southern Ocean). Correspondence analysis, based on the 75 most common taxa, shows a clear biogeographic trend along the first correspondence axis by arranging the sites in paleolatitudinal order. The assemblages from the Tethyan Realm (i.e., low latitudes) are marked by abundant heavily calcified buliminids (such as Bulimina incisa, B. trinitatensis, B. velascoensis, and Reussella szajnochae) and Aragonia spp., whereas high-latitude faunas are characterized by abundant Alabamina creta, Gyroidinoides quadratus, and Pullenia coryelli. The results indicate that the faunas at low and high latitudes, respectively, were influenced by quite different environmental conditions. This is based on the much higher abundance of infaunal morphotypes at low and mid latitudes compared to high latitudes, suggesting that the biogeographic trend found in the data set coincides with the trophic regime at the various sites. The results also provide support for the hypothesis that postulates two simultaneous sources and mechanisms for deep-water formation during the Late Cretaceous, including warm, saline deep water produced by evaporation at low (equatorial) latitudes in contrast to the formation of cold deep waters at high (southern) latitudes.

Activity and biomass of small benthic biota in the central Arctic Ocean

Sediment samples collected during the expedition "Arctic Ocean '96" with the Swedish ice-breaker ODEN were investigated to estimate for the first time heterotrophic activity and total microbial biomass (size range from bacteria to small metazoans) from the perennially ice-covered central Arctic Ocean. Benthic activities and biomass were evaluated analysing a series of biogenic sediment compounds (i.e. bacterial exoenzymes, total adenylates, DNA, phospholipids, particulate proteins). In contrast to the very time-consuming sorting, enumeration and weight determination, analyses of biochemical sediment parameters may represent a useful method for obtaining rapid information on the ecological situation in a given benthic system. Bacterial cell numbers and biomass were estimated for comparison with biochemically determined biomass data, to evaluate the contribution of the bacterial biomass to the total microbial biomass. It appeared that bacterial biomass made up only 8-31% (average of all stations = 20%) of the total microbial biomass, suggesting a large fraction of other small infaunal organisms within the sediment samples (most probably fungi, yeasts, protozoans such as flagellates, ciliates or amoebae, as well as a fraction of small metazoans). Activity and biomass values determined within this study were generally extremely low, and often even slightly lower than those given for other deep oceanic regions, thus characterizing the seafloor of the central Arctic Ocean as a "benthic desert". Nevertheless, some clear trends in the data could be found, e.g. generally sharply decreasing values within the sediment column, a vague tendency for declining values with increasing water depth of sampling stations, and also differences between various Arctic deep-sea regions.

Depth profiles of dissolved sulfide and sulfate in the pore water of hydrocarbon seep sediments offshore Pakistan

The interaction between fluid seepage, bottom water redox, and chemosynthetic communities was studied at cold seeps across one of the world's largest oxygen minimum zones (OMZ) located at the Makran convergent continental margin. Push cores were obtained from seeps within and below the core-OMZ with a remotely operated vehicle. Extracted sediment pore water was analyzed for sulfide and sulfate concentrations. Depending on oxygen availability in the bottom water, seeps were either colonized by microbial mats or by mats and macrofauna. The latter, including ampharetid polychaetes and vesicomyid clams, occurred in distinct benthic habitats, which were arranged in a concentric fashion around gas orifices. At most sites colonized by microbial mats, hydrogen sulfide was exported into the bottom water. Where macrofauna was widely abundant, hydrogen sulfide was retained within the sediment. Numerical modeling of pore water profiles was performed in order to assess rates of fluid advection and bioirrigation. While the magnitude of upward fluid flow decreased from 11 cm yr**-1 to <1 cm yr**-1 and the sulfate/methane transition (SMT) deepened with increasing distance from the central gas orifice, the fluxes of sulfate into the SMT did not significantly differ (6.6-9.3 mol m**-2 yr**-1). Depth-integrated rates of bioirrigation increased from 120 cm yr**-1 in the central habitat, characterized by microbial mats and sparse macrofauna, to 297 cm yr**-1 in the habitat of large and few small vesicomyid clams. These results reveal that chemosynthetic macrofauna inhabiting the outer seep habitats below the core-OMZ efficiently bioirrigate and thus transport sulfate down into the upper 10 to 15 cm of the sediment. In this way the animals deal with the lower upward flux of methane in outer habitats by stimulating rates of anaerobic oxidation of methane (AOM) with sulfate high enough to provide hydrogen sulfide for chemosynthesis. Through bioirrigation, macrofauna engineer their geochemical environment and fuel upward sulfide flux via AOM. Furthermore, due to the introduction of oxygenated bottom water into the sediment via bioirrigation, the depth of the sulfide sink gradually deepens towards outer habitats. We therefore suggest that - in addition to the oxygen levels in the water column, which determine whether macrofaunal communities can develop or not - it is the depth of the SMT and thus of sulfide production that determines which chemosynthetic communities are able to exploit the sulfide at depth. We hypothesize that large vesicomyid clams, by efficiently expanding the sulfate zone down into the sediment, could cut off smaller or less mobile organisms, as e.g. small clams and sulfur bacteria, from the sulfide source.

(Table 1) Dominant species and total biomass of macrozoobenthos communities in the southwestern Kara Sea

The structure and distribution of the macrobenthic communities were studied in the southwestern Kara Sea. The material was collected in Baidaratskaya Bay in July 2007 and in a section running westward of the Yamal Peninsula in September 2007. The depths of the sampling stations ranged from 5 to 25 m in the Baidaratskaya Bay area and between 16 and 46 m in the Yamal section. A total of 212 benthic invertebrate species were recorded. In both areas, Bivalvia was the group with the highest biomass (54.88 g/m**2 in the Yamal section and 59.71 g/m**2 in the Baidaratskaya Bay area), while polychaetes were the group with the highest number of species (45 in the Yamal section and 64 the Baidaratskaya Bay area). Three major macrozoobenthic communities were recognized: the Astarte borealis community (20-46 m, the deepest sampling stations in both areas); the 'medium-depth' community (10-20 m, extremely mosaic, usually dominated by Serripes groenlandicus); and the Nephtys longosetosa community (depth smaller than 10 m, characterized by low biomass and the absence of large bivalves and echinoderms). The western Yamal shallow-water communities were shown to be generally similar to those of Baidaratskaya Bay. The comparison of these results with those of the benthos censuses performed in 1927-1945, 1975, and 1993 showed that the benthic communities in the southwestern Kara Sea remained relatively stable during the second half of the 20th century and the early 21st century.

The distribution of macrobenthos in surface sediments at Fladen Ground stations, North Sea

A general study of structure, biomass estimates and dynamics on the macrofauna was carried out in August 1975 and March 1976 during PREFLEX (1975) and FLEX (1976), the Fladen Ground Experiment. On the basis of these data an attempt was made to estimate macrobenthic production expressed as minimum production (MP). The macrobenthic production is discussed together with meiobenthic annual production and with indirectly estimated microbenthic production in relation to an energy input from the water column of about 25 g C m**-2 year**-1. From the production estimates of the three benthic components a rough energy budget is proposed. Sampling was performed at five stations for endofauna twice during the time of investigation and for epifauna once. At each station two replicate box core samples (30 X 20 cm) were taken for endofauna. Epifauna was sampled with an Agassiz trawl once at each station. The total numbers of endofauna increased from station 1 to 5. This was valid as well for August 1975 (4,233-12,166 individuals per m**2 and 10 cm sediment depth) as for March 1976 (1,008-2,925 individuals). The polychaetes were the dominant organisms with a share of 33 to 62 %. The densities for the endofauna decreased from August 1975 to March 1976 by a mean factor of 2.8. Abundances of epifauna amounted to values between 11 and 102 individuals per 1000 m**2. The biomass dry weights (DWT) for macrobenthic endofauna varied between 0.97 g DWT m**-2 and 6.42 g DWT m**-2 in August 1975 and between 0.27 g DWT m**-2 and 2.64 g DWT m**-2 in March 1976. The mean amounted to 1.74 g DWT m**-2. Dry weights of epifauna biomass gave values between 4.9 and 83.1 g DWT * 1000 m**-2. The minimum production for the total macro-endofauna at Fladen Ground amounted to 1.43 g DWT m**-2 yr**-1 or 0.82 g C m**-2 yr**-1. This resulted in a minimum turnover rate (P/B) of 0.8. The share produced by the polychaetes amounted to 1.06g DWT m**-2 yr**-1 or 74 %.