Benthic foraminifera in surface sediments of the Arabian Sea

Assemblages of living deep-sea benthic foraminifera, their densities, vertical distribution pattern, and diversity, were investigated in the intermonsoon period after the northeast monsoon in the Arabian Sea in spring 1997. Foraminiferal numbers show a distinct gradient from north to south, with a maximum of 623 foraminifera in 50 cm**3 at the northern site. High percentages of small foraminifera were found in the western and northern part of the Arabian Sea. Most stations show a typical vertical distribution with a maximum in the first centimeter and decreasing numbers with increasing sediment depths. But at the central station, high densities can be found even in deeper sediment layers. Diversity is very high at the northern and western sites, but reduced at the central and southern stations. Data and faunal assemblages were compared with studies carried out in 1995. A principal component analysis of intermonsoon assemblages shows that the living benthic foraminifera can be characterized by five principal component communities. Dominant communities influencing each site differ strongly between the two years. In spring 1997, stations in the north, west and central Arabian Sea were dominated by opportunistic species, indicating the influence of fresh sedimentation pulses or enhanced organic carbon fluxes after the northeast monsoon.

Auxiliary material and basic geochemical and benthic foraminiferal stable isotope data for the interval bracketing Eocene Thermal Maximum 2 (ETM2) at DSDP Sites 401 and 550 in the northeastern Atlantic Ocean

Ever since its discovery, Eocene Thermal Maximum 2 (ETM2; ~53.7 Ma) has been considered as one of the "little brothers" of the Paleocene-Eocene Thermal Maximum (PETM; ~56 Ma) as it displays similar characteristics including abrupt warming, ocean acidification, and biotic shifts. One of the remaining key questions is what effect these lesser climate perturbations had on ocean circulation and ventilation and, ultimately, biotic disruptions. Here we characterize ETM2 sections of the NE Atlantic (Deep Sea Drilling Project Sites 401 and 550) using multispecies benthic foraminiferal stable isotopes, grain size analysis, XRF core scanning, and carbonate content. The magnitude of the carbon isotope excursion (0.85-1.10 per mil) and bottom water warming (2-2.5°C) during ETM2 seems slightly smaller than in South Atlantic records. The comparison of the lateral d13C gradient between the North and South Atlantic reveals that a transient circulation switch took place during ETM2, a similar pattern as observed for the PETM. New grain size and published faunal data support this hypothesis by indicating a reduction in deepwater current velocity. Following ETM2, we record a distinct intensification of bottom water currents influencing Atlantic carbonate accumulation and biotic communities, while a dramatic and persistent clay reduction hints at a weakening of the regional hydrological cycle. Our findings highlight the similarities and differences between the PETM and ETM2. Moreover, the heterogeneity of hyperthermal expression emphasizes the need to specifically characterize each hyperthermal event and its background conditions to minimalize artifacts in global climate and carbonate burial models for the early Paleogene.

Biogeochemical investigation of the benthic nitrogen cycle in the Arabian Sea off Pakistan

A pronounced deficit of nitrogen (N) in the oxygen minimum zone (OMZ) of the Arabian Sea suggests the occurrence of heavy N-loss that is commonly attributed to pelagic processes. However, the OMZ water is in direct contact with sediments on three sides of the basin. Contribution from benthic N-loss to the total N-loss in the Arabian Sea remains largely unassessed. In October 2007, we sampled the water column and surface sediments along a transect cross-cutting the Arabian Sea OMZ at the Pakistan continental margin, covering a range of station depths from 360 to 1430 m. Benthic denitrification and anammox rates were determined by using 15N-stable isotope pairing experiments. Intact core incubations showed declining rates of total benthic N-loss with water depth from 0.55 to 0.18 mmol N m**-2 day**-1. While denitrification rates measured in slurry incubations decreased from 2.73 to 1.46 mmol N m**-2 day**-1 with water depth, anammox rates increased from 0.21 to 0.89 mmol N m**-2 day**-1. Hence, the contribution from anammox to total benthic N-loss increased from 7% at 360 m to 40% at 1430 m. This trend is further supported by the quantification of cd1-containing nitrite reductase (nirS), the biomarker functional gene encoding for cytochrome cd1-Nir of microorganisms involved in both N-loss processes. Anammox-like nirS genes within the sediments increased in proportion to total nirS gene copies with water depth. Moreover, phylogenetic analyses of NirS revealed different communities of both denitrifying and anammox bacteria between shallow and deep stations. Together, rate measurement and nirS analyses showed that anammox, determined for the first time in the Arabian Sea sediments, is an important benthic N-loss process at the continental margin off Pakistan, especially in the sediments at deeper water depths. Extrapolation from the measured benthic N-loss to all shelf sediments within the basin suggests that benthic N-loss may be responsible for about half of the overall N-loss in the Arabian Sea.

Benthic oxygen and nitrogen fluxes at different time series sites in the southern North Sea, 2012 to 2014

Benthic oxygen and nitrogen fluxes were quantified within the years 2012 to 2014 at different time series sites in the southern North Sea with the benthic lander NuSObs (Nutrient and Suspension Observatory). In situ incubations of sediments, in situ bromide tracer studies, sampling of macrofauna and pore water investigations revealed considerable seasonal and spatial variations of oxygen and nitrogen fluxes. Seasonal and spatial variations of oxygen fluxes were observed between two different time series sites, covering different sediment types and/or different benthic macrofaunal communities. On a sediment type with a high content of fine grained particles (<63 µm) oxygen fluxes of -15.5 to -25.1 mmol/m**2/d (June 2012), -2.0 to -8.2 mmol/m**2/d (March 2013), -16.8 to -21.5 mmol/m**2/d (November 2013) and -6.1 mmol/m**2/d (March 2014) were measured. At the same site a highly diverse community of small species of benthic macrofauna was observed. On a sediment type with a low content of fine grained particles (<63 µm) high oxygen fluxes (-33.2 mmol/m**2/d August 2012; -47.2 to -55.1 mmol/m**2/d November 2013; -16.6 mmol/m**2/d March 2014) were observed. On this sediment type a less diverse benthic macrofaunal community, which was dominated by the large bodied suspension feeder Ensis directus, was observed. Average annual rain rates of organic carbon and organic nitrogen to the seafloor of 7.44 mol C/m**2/y and 1.34 mol N/m**2/y were estimated. On average 79% of the organic bound carbon and 95% of the organic bound nitrogen reaching the seafloor are recycled at the sediment-water interface.

Benthic foraminifera of surface sediments from the continental slope off Sierra Leone, North-West Africa

On "Meteor" cruise 30 (1973) 22 piston-cores were collected off Sierra Leone from water-depths between about 5000 m (Sierra Leone Basin) and 500 m (upper continental slope) with the objective to study the sediment composition and age as well as processes of sedimentation on the continental slope in a tropical humid region. Granulometric analysis and determinations of the carbonate contents of the sediment samples were carried out, as well as qualitative and quantitative analysis of the components of the grain size fractions > 63 µm and of the planktonic and benthonic foraminifera > 160 µm. Presently, the cold Canary Current influences the composition of the planktonic foraminifera within the northwestern area of investigation (profile A), whereas the planktonic fauna of the eastern area (profile C) seems to be truly tropical. In all Quaternary sediments from the continental slope off Sierra Leone, species of Globorotalia are less abundant than in truly pelagic sediments. For that reason, the zonation of the Pleistocene sediments based on the presence or absence of Globorotalia cultrata does not always agree with the climatic changes reflected in the sediments. Concerning past climates better results can be obtained by using the changes in percentage abundances of Globigerina sp. sp. and Globigerinoides sp. sp. as indicators for cool and warm temperatures. The Tertiary sediments contain a pelagic foraminiferal assemblage. In the Holocene sediments the benthonic foraminifera do not only serve as good paleodepth indicators, but their communities are also restricted to defined water masses, which change their positions in accordance with climatic changes. Thus, Cassidulina carinata in the area of investigation is an excellent indicator for sediments deposited during times, which were cooler than today; this is true for all cores from the continental slope off Sierra Leone independent of water-depth although this species presently abounds at water-depths around 600 m. The cores from the continental rise and from the Sierra Leone Basin (M30-261, M30-146, M30-147) were deposited below the calcium carbonate compensation depth. Only small sections of the cores consist of the original carbonate-free sediments, whereas the main part of the sediment column is redeposited material, rich in foraminifera, which normally live on the upper continental slope, or even on the shelf. From these cores only M30-261 can be subdivided into biostratigraphic zones ranging from zone V to zone Y. In all cores from the middle and upper continental slope of the eastern area of investigation (profile C; KL 230, 209-204) and in cores KL 183 and KL 184 from the northwestern area (profile A) we observed an undisturbed succession of sediments from the biostratigraphic zones X (partly), Y and Z. All cores from the central area (M30-181, M30-182, M30-262 to 264) and M30-187 from the upper slope of profile A show variable hiatuses in the sedimentary record. Locally, high velocity bottom currents were probably responsible for erosion, nondeposition or minimal sedimentation rates. These currents might have been initiated partly by the somewhat exposed position of this part of the continental slope, where the shelf edge bends from a northwest towards an eastern direction, and partly by very young tectonic movements. Fracture zones with vertically displaced fault blocs are frequent at Sierra Leone continental margin. According to seismic measurements by McMaster et al. (1975) the sites of the central area are located on an uplifted fault bloc explaining the reduced sediment rates or erosion. Unlike the central area, the eastern area (profile C) is situated on a downfaulted bloc with high sediment rates. The sediments from the cores of profile B as well as the turbiditic deep-sea sediments were deposited under a higher flow regime; therefore they are coarser than the extremely fine-grained sediments of the cores from profile C. Since the sand fraction (> 63 µm) is mainly composed of foraminifera, besides pteropods and light-colored fecal pellets, the carbonate content increases with the increasing percentage of the coarse grain fraction. Higher concentrations of quartz were only observed in core sections with considerable carbonate dissolution (mainly in the X-Zone), and, in general, in all sediments from the eastern area with higher terrigenous input including larger concentration of mica. Especially during times transitional between glacials and interglacials (or interstadials) the bottom currents were intensified. The percentages of coarse fraction and carbonate increase with increasing current velocities. Calcium carbonate dissolution becomes important in water depths > 3500 m. During cooler times the lysokline is depressed. Light-colored fecal pellets were redeposited from Late Neogene sediments (M30-187, M30-181). In the area of investigation they occur in the Holocene and mainly the Pleistocene sediments of the cores from the northwestern and central area because only here Tertiary sediments have been eroded at the uppermost continental slope. In the central area there are at least two periods of non-sedimentation and/or erosion which can be confined as being (1) not older than middle Pliocene and not younger than zone V and (2) younger than zone W. The local character of the erosion is documented by the fact that a complete Late Quaternary section is present in the cores of the northwestern and eastern area, each within less than 100 km from incomplete cores from the central area.

Living benthic foraminiferal assemblages in surface samples of the Arabian Sea

The distribution, biomass, and diversity of living (Rose Bengal stained) deep-sea benthic foraminifera (>30 µm) were investigated with multicorer samples from seven stations in the Arabian Sea during the intermonsoonal periods in March and in September/October, 1995. Water depths of the stations ranged between 1916 and 4425 m. The distribution of benthic foraminifera was compared with dissolved oxygen, % organic carbon, % calcium carbonate, ammonium, % silica, chloroplastic pigment equivalents, sand content, pore water content of the sediment, and organic carbon flux to explain the foraminiferal patterns and depositional environments. A total of six species-communities comprising 178 living species were identified by principal component analysis. The seasonal comparison shows that at the western stations foraminiferal abundance and biomass were higher during the Spring Intermonsoon than during the Fall Intermonsoon. The regional comparison indicates a distinct gradient in abundance, biomass, and diversity from west to east, and for biomass from north to south. Highest values are recorded in the western part of the Arabian Sea, where the influence of coastal and offshore upwelling are responsible for high carbon fluxes. Estimated total biomass of living benthic foraminifera integrated for the upper 5 cm of the sediment ranged between 11 mg Corg m**-2 at the southern station and 420 mg Corg m**-2 at the western station. Foraminifera in the size range from 30 to 125 ?m, the so-called microforaminifera, contributed between 20 and 65% to the abundance, but only 3% to 28% to the biomass of the fauna. Highest values were found in the central and southern Arabian Sea, indicating their importance in oligotrophic deep-sea areas. The overall abundance of benthic foraminifera is positively correlated with oxygen content and pore volume, and partly with carbon content and chloroplastic pigment equivalents of the sediment. The distributional patterns of the communities seem to be controlled by sand fraction, dissolved oxygen, calcium carbonate and organic carbon content of the sediment, but the critical variables are of different significance for each community.

Benthic megafaunal composition at the Arctic deep-sea observatory HAUSGARTEN

The Long-Term Ecological Research (LTER) observatory HAUSGARTEN, in the eastern Fram Strait, provides us the valuable ability to study the composition of benthic megafaunal communities through the analysis of seafloor photographs. This, in combination with extensive sampling campaigns, which have yielded a unique data set on faunal, bacterial, biogeochemical and geological properties, as well as on hydrography and sedimentation patterns, allows us to address the question of why variations in megafaunal community structure and species distribution exist within regional (60-110 km) and local (<4 km) scales. Here, we present first results from the latitudinal HAUSGARTEN gradient, consisting of three different stations (N3, HG-IV, S3) between 78°30'N and 79°45'N (2351 - 2788 m depth), obtained via the analysis of images acquired by a towed camera (OFOS - Ocean Floor Observation System) in 2011. We assess variability in megafaunal densities, species composition and diversity as well as biotic and biogenic habitat features, which may cause the patterns observed. While there were significant regional-scale differences in megafaunal composition and densities between the stations (N3 = 26.74 ± 0.63; HG-IV = 11.21 ± 0.25; S3 = 18.34 ± 0.39 individuals/m**2), significant local differences were only found at HG-IV. Regional-scale variations may be due to the significant differences in ice coverage at each station as well as the different quantities of protein available, whereas local-scale differences at HG-IV may be a result of variation in bottom topography or factors not yet identified.

Seawater carbonate chemistry and encrusting algal communities during a mesocosm experiment, 2007

Owing to anthropogenic emissions, atmospheric concentrations of carbon dioxide could almost double between 2006 and 2100 according to business-as-usual carbon dioxide emission scenarios. Because the ocean absorbs carbon dioxide from the atmosphere, increasing atmospheric carbon dioxide concentrations will lead to increasing dissolved inorganic carbon and carbon dioxide in surface ocean waters, and hence acidification and lower carbonate saturation states. As a consequence, it has been suggested that marine calcifying organisms, for example corals, coralline algae, molluscs and foraminifera, will have difficulties producing their skeletons and shells at current rates, with potentially severe implications for marine ecosystems, including coral reefs. Here we report a seven-week experiment exploring the effects of ocean acidification on crustose coralline algae, a cosmopolitan group of calcifying algae that is ecologically important in most shallowwater habitats. Six outdoor mesocosms were continuously supplied with sea water from the adjacent reef and manipulated to simulate conditions of either ambient or elevated seawater carbon dioxide concentrations. The recruitment rate and growth of crustose coralline algae were severely inhibited in the elevated carbon dioxide mesocosms. Our findings suggest that ocean acidification due to human activities could cause significant change to benthic community structure in shallow-warm-water carbonate ecosystems.

Seawater carbonate chemistry and benthic marine community during experiments, 2011

Ocean acidification is predicted to impact all areas of the oceans and affect a diversity of marine organisms. However, the diversity of responses among species prevents clear predictions about the impact of acidification at the ecosystem level. Here, we used shallow water CO2 vents in the Mediterranean Sea as a model system to examine emergent ecosystem responses to ocean acidification in rocky reef communities. We assessed in situ benthic invertebrate communities in three distinct pH zones (ambient, low, and extreme low), which differed in both the mean and variability of seawater pH along a continuous gradient. We found fewer taxa, reduced taxonomic evenness, and lower biomass in the extreme low pH zones. However, the number of individuals did not differ among pH zones, suggesting that there is density compensation through population blooms of small acidification-tolerant taxa. Furthermore, the trophic structure of the invertebrate community shifted to fewer trophic groups and dominance by generalists in extreme low pH, suggesting that there may be a simplification of food webs with ocean acidification. Despite high variation in individual species' responses, our findings indicate that ocean acidification decreases the diversity, biomass, and trophic complexity of benthic marine communities. These results suggest that a loss of biodiversity and ecosystem function is expected under extreme acidification scenarios.

Seawater carbonate chemistry and encrusting algal communities duirng a mesocosm experiment, 2007

Owing to anthropogenic emissions, atmospheric concentrations of carbon dioxide could almost double between 2006 and 2100 according to business-as-usual carbon dioxide emission scenarios. Because the ocean absorbs carbon dioxide from the atmosphere, increasing atmospheric carbon dioxide concentrations will lead to increasing dissolved inorganic carbon and carbon dioxide in surface ocean waters, and hence acidification and lower carbonate saturation states. As a consequence, it has been suggested that marine calcifying organisms, for example corals, coralline algae, molluscs and foraminifera, will have difficulties producing their skeletons and shells at current rates, with potentially severe implications for marine ecosystems, including coral reefs. Here we report a seven-week experiment exploring the effects of ocean acidification on crustose coralline algae, a cosmopolitan group of calcifying algae that is ecologically important in most shallowwater habitats. Six outdoor mesocosms were continuously supplied with sea water from the adjacent reef and manipulated to simulate conditions of either ambient or elevated seawater carbon dioxide concentrations. The recruitment rate and growth of crustose coralline algae were severely inhibited in the elevated carbon dioxide mesocosms. Our findings suggest that ocean acidification due to human activities could cause significant change to benthic community structure in shallow-warm-water carbonate ecosystems.