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.

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.

A rapidly evolving secretome builds and patterns a sea shell

Background: Instructions to fabricate mineralized structures with distinct nanoscale architectures, such as seashells and coral and vertebrate skeletons, are encoded in the genomes of a wide variety of animals. In mollusks, the mantle is responsible for the extracellular production of the shell, directing the ordered biomineralization of CaCO3 and the deposition of architectural and color patterns. The evolutionary origins of the ability to synthesize calcified structures across various metazoan taxa remain obscure, with only a small number of protein families identified from molluskan shells. The recent sequencing of a wide range of metazoan genomes coupled with the analysis of gene expression in non-model animals has allowed us to investigate the evolution and process of biomineralization in gastropod mollusks. Results: Here we show that over 25% of the genes expressed in the mantle of the vetigastropod Haliotis asinina encode secreted proteins, indicating that hundreds of proteins are likely to be contributing to shell fabrication and patterning. Almost 85% of the secretome encodes novel proteins; remarkably, only 19% of these have identifiable homologues in the full genome of the patellogastropod Lottia scutum. The spatial expression profiles of mantle genes that belong to the secretome is restricted to discrete mantle zones, with each zone responsible for the fabrication of one of the structural layers of the shell. Patterned expression of a subset of genes along the length of the mantle is indicative of roles in shell ornamentation. For example, Has-sometsuke maps precisely to pigmentation patterns in the shell, providing the first case of a gene product to be involved in molluskan shell pigmentation. We also describe the expression of two novel genes involved in nacre (mother of pearl) deposition. Conclusion: The unexpected complexity and evolvability of this secretome and the modular design of the molluskan mantle enables diversification of shell strength and design, and as such must contribute to the variety of adaptive architectures and colors found in mollusk shells. The composition of this novel mantle-specific secretome suggests that there are significant molecular differences in the ways in which gastropods synthesize their shells.

(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 foraminiferal record and age dating of sediment cores from the Gullmar Fjord at the west coast of Sweden

A high-resolution study of benthic foraminiferal assemblages was performed on a ca. eight metre long sediment core from Gullmar Fjord on the west coast of Sweden. The results of 210Pb- and AMS 14C-datings show that the record includes the two warmest climatic episodes of the last 1500 years: the Medieval Warm Period (MWP) and the recent warming of the 20th century. Both periods are known to be anomalously warm and associated with positive NAO winter indices. Benthic foraminiferal successions of both periods are compared in order to find faunal similarities and common denominators corresponding to past climate changes. During the MWP, Adercotryma glomerata, Cassidulina laevigata and Nonionella iridea dominated the assemblages. Judging from dominance of species sensitive to hypoxia and the highest faunal diversity for the last ca. 2400 years, the foraminiferal record of the MWP suggests an absence of severe low oxygen events. At the same time, faunas and d13C values both point to high primary productivity and/or increased input of terrestrial organic carbon into the fjord system during the Medieval Warm Period. Comparison of the MWP and recent warming revealed different trends in the faunal record. The thin-shelled foraminifer N. iridea was characteristic of the MWP, but became absent during the second half of the 20th century. The recent Skagerrak-Kattegat fauna was rare or absent during the MWP but established in Gullmar Fjord at the end of the Little Ice Age or in the early 1900s. Also, there are striking differences in the faunal diversity and absolute abundances of foraminifera between both periods. Changes in primary productivity, higher precipitation resulting in intensified land runoff, different oxygen regimes or even changes in the fjord's trophic status are discussed as possible causes of these faunal differences.

Benthic foraminifera analysed in Skagerrak and Kattegat sediments

a) In der horizontalen Verbreitung sind die vorwiegend kalkschaligen Benthos-Foraminiferen im Untersuchungsgebiet auf zwei Faciesbereiche verteilt: 1. Eine sandige Facies mit stärkeren Temperatur- und Salzgehaltschwankungen; Wasseroberfläche t = 2O-17°C, Salzgehalt nie über 32 per mil, Meerestiefe 30 bis 92 m. 2. Schlick-Facies mit zum Teil feinsandigen Beimengungen. Temperatur- und Salzgehaltschwankungen sind geringer; Wasseroberfläche t = ca. 4O-15° C, Salzgehalt bis 34 per mil, Meerestiefe 135-548 m. b) Einige Stoßröhren-Proben (Station 18, 21, 27, 28) zeigen in ihrer vertikalen Verbreitung auffallende Faunenunterschiede. c) Im Profil des Lotkerns wechseln in der Foraminiferenfauna Bolivinen- und Cassidulinen-Nonioninen-Provinzen miteinander ab. Die Profile der beiden tiefsten Stoßröhren-Kerne (Station 23, 26; s. Tab. I) stimmen in ihrer Mikrofauna mit der des oberen Teils des Lotkerns (s. Tab. 4) überein. d) Die unter b und C angefuührten Faunenwechsel werden auf langperiodische Klimaerwärmungen im skandinavischen Raum und den damit verbundenen Anstieg des Meeresspiegels zurückgeführt. e) Der Lotkern kann mit Hilfe von Untersuchungsergebnissen aus seiner näheren Umgebung (Bohuslän, Oslofjord) nur bedingt in ein stratigraphisches, durch Megafossilien belegtes Schema eingefügt werden, da er nach unten durch die Mikrofauna keine echte Begrenzung aufweist. Durch die Einwanderung mehrerer Foraminiferenarten mit boreal-lusitanischer Verbreitung in die Untersuchungsgebiete wird der Lotkern in die Isocardia-Absätze (Atlanticum-oberes Subboreal) eingegliedert. f) Aus einer Tabelle von PRATJE(1940) kann entnommen werden, daß dieser Zeitabschnitt nach DE GEER etwa um 5000 v.Chr. beginnt. Danach beträgt die geringste Sedimentation, die in dem Kerngebiet nach dieser Zeitrechnung möglich ist, bei einer Eindringtiefe des Lots von 10 m ungefähr 1,40 m pro Jahrtausend. Wahrscheinlich wird dieses Maß etwas größer sein.

Distribution of planktonic and benthic foraminifers in bottom sediments from Core LV28-42-5, Sea of Okhotsk

Distribution of diatoms and planktonic and benthic foraminifers, as well as correlation of components of sandy grain size fraction were studied in the Quaternary sediment core LV28-42-5 (720 cm long) col¬lected on the southeastern slope (1045 m depth) of the Institute of Oceanology Rise, Sea of Okhotsk. This study allowed to reconstruct principle features of paleoceanographic evolution. In the course of penultimate and last continental glaciations (isotope stages 6 and 4-2) and during the later period of the last interglacial (substages 5.d-5.a) the following conditions were characteristic of this area: low temperatures of surface water, terrigenous sediment accumulation including coarse grained ice-rafted material, minimum bioproductivity and microfossil content in sediments, low sea level, reduced water exchange with the ocean, low position of old deep Pacific water. During the interglacial optimum (substage 5.e), as well as in the last deglaciation and Holocene (stage 1) water temperature and bioproductivity increased, sea level rose, and active surface water exchange between the Sea of Okhotsk and the Pacific Ocean and the Sea of Japan took place. This resulted in intensive inflow of the old deep Pacific water into the Sea of Okhotsk and elevation of its upper boundary by few hundred meters. During the later intervals of these warm periods a dichothermal structure of the upper water layer formed and diatom oozes accumulated.

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.

Micropaleontological investigation of sediment core Kronsmoor

Benthic foraminiferal and calcareous nannofossil assemblages, as well as stable isotope data from the Campanian/Maastrichtian boundary interval (~71.4 to ~70.7 Ma) of the Kronsmoor section (North German Basin), were investigated in order to characterize changes in surface-water productivity and oxygen content at the seafloor and their link to climatic and paleoceanographic changes. A nutrient index based on calcareous nannofossils is derived for the high-latitude, epicontinental North German Basin, reflecting changes in surface-water productivity. Oxygen isotopes of well-preserved planktic foraminiferal specimens of Heterohelix globulosa reflect warmer surface-water temperatures in the lower part of the studied succession and a cooling of up to 2°C (0.5 per mil) in the upper part (after 71.1 Ma). For the lower and warmer part of the investigated succession, benthic foraminiferal assemblages and the calcareous nannofossils indicate well-oxygenated bottom waters and low-surface water productivity. In contrast, the upper part of the succession is characterized by cooler conditions, lower oxygen content at the seafloor and increasing surface-water productivity. It is proposed that the cooling phase starting at 71.1 Ma was accompanied by increasing surface-water mixing caused by westerly winds. As a consequence of mixing, nutrients were advected from sub-surface waters into the mixed layer, resulting in increased surface-water productivity and enhanced organic matter flux to the seafloor. We hypothesize that global sea-level fall during the earliest Maastrichtian (~71.3 Ma), indicated by decreasing carbon isotope values, may have led to a weaker water mass exchange through narrower gateways between the Boreal Realm and the open North Atlantic and Tethys oceans. Both the weaker water mass exchange and enhanced surface-water productivity may have led to slightly less ventilated bottom waters of the upper part of the studied section. Our micro-paleontological and stable isotopic approach indicates short-term (<100 kyr) changes in oxygen consumption at the seafloor and surface-water productivity across the homogeneous Boreal White Chalk succession of the North German Basin.

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.

Recent benthic foraminifera of the Scotia Sea and Argentine Basin

We investigated 88 surface sediment samples taken with a multiple corer from the southwestern South Atlantic Ocean for their live (Rose Bengal stained) and dead benthic foraminiferal content. Using Q-Mode Principal Component Analysis six live and six dead associations are differentiated. Live and dead association distributions correspond fairly well; differences are mainly caused by downslope transport and selective test destruction. In addition, four potential fossil associations are calculated from the dead data set after removal of non-fossilizable species. These potential fossil associations are expected to be useful for paleoceanographic reconstructions. Environments are described in detail for the live and potential fossil associations and for selected species. Along the upper Argentine continental slope strong bottom currents control the occurrence of live, dead and potential fossil Angulogerina angulosa associations. Here, particles of a high organic carbon flux rate remain suspended. Below this high energy environment live, dead and potential fossil Uvigerina peregrina dominated associations correlate with enhanced sediment organic carbon content and still high organic carbon flux rates. The live A. angulosa and U. peregrina associations correlate with high standing crops. Furthermore, live and dead Epistominella exigua-Nuttallides umbonifer associations were separated. Dominance of a Nuttallides umbonifer potential fossil association relates to coverage by Antarctic Bottom Water (AABW) and Lower Circumpolar Deep Water (LCDW), above the Calcite Compensation Depth (CCD). Three associations of mainly agglutinated foraminifera occur in sediments bathed mainly by AABW or CDW. A Reophax difflugiformis association was found in mud-rich and diatomaceous sediments. Below the CCD, a Psammosphaera fusca association occurs in coarse sediments poor in organic carbon while a Cribrostomoides subglobosus-Ammobaculites agglutinans association covers a more variable environmental range with mud contents exceeding 30%. One single Eggerella bradyi-Martinottiella communis association poor in both species and individuals remains from the agglutinated associations below the CCD if only preservable species are considered for calculation.

Benthic foraminifera asemblages and isotopic composition of surface sediment samples off Morocco

The influence of different primary productivity regimes on live (Rose Bengal stained) and dead benthic foraminiferal distribution, as well as on the stable carbon isotopic composition of foraminiferal tests, was investigated in sediment surface samples (0-1 cm) from the upwelling region off Morocco between Cape Ghir (31°N) and Cape Yubi (27°N). A combination of factor analysis, detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) was applied to the benthic foraminiferal data sets. Five major assemblages for both the live and dead fauna were revealed by factor analysis. In the cape regions organic matter fluxes are enhanced by high chlorophyll-a concentrations in the overlying surface waters. Here, benthic foraminiferal faunas are characterized by identical live and dead assemblages, high standing stocks, and low species delta13C values, indicating constant year-round high productivity. Bulimina marginata dominates the unique fauna at the shallowest station off Cape Ghir indicating highest chlorophyll-a concentrations. Off both capes, the succession of the Bulimina aculeata/Uvigerina mediterranea assemblage, the Sphaeroidina bulloides/Gavelinopsis translucens assemblage, and the Hoeglundina elegans assemblage from the shelf to the deep sea reflects the decrease in chlorophyll-a concentrations, hence the export flux. In contrast, the area between the capes is characterized by differently composed live and dead assemblages, low standing stocks, and less depleted delta13C values, thus reflecting low primary productivity. High foraminiferal numbers of Epistominella exigua, Eponides pusillus, and Globocassidulina subglobosa in the dead fauna indicate a seasonally varying primary productivity signal. Significantly lower mean delta13C values were recorded in Bulimina mexicana, Cibicidoides kullenbergi, H. elegans, U. mediterranea and Uvigerina peregrina. Cibicidoides wuellerstorfi is a faithful recorder of bottom water delta13C in the Canary Islands regions. The mean delta13C signal of this species is not significantly influenced by constant high organic matter fluxes. The species-specific offset between live and dead specimens is the same.