Foraminifer, epifauna and infauna abundance of ODP Hole 183-1138A and core ELT54.007-PC, Kerguelen Plateau

Late Campanian and Maastrichtian benthic foraminifers are recorded from 12 samples from Ocean Drilling Program (ODP) Leg 183, Cores 183-1138A-52R through 63R (487.3-602.4 meters below seafloor), Kerguelen Plateau, Indian Ocean, and Danian benthics from one sample in the same section. The entire late Maastrichtian foraminifer fauna is noted from a dredge sample 220 km to the north. The structure of the fauna is compared with the Cenomanian-Turonian of the nearby Eltanin core E54-7. Faunas are reviewed in terms of planktonic percentage, composition, epifaunal/infaunal ratios, and dominance/diversity indices. The region was in the cool Austral Faunal Province through the Campanian-Maastrichtian and was probably warmer in the Cenomanian-Turonian. The ODP section is now 1600 meters below sea level and has subsided several hundred meters since deposition. Its fauna is dominated by epifaunal species suggesting little influence of upwelling. The dredge location has subsided little. Its fauna has a high infaunal content consistent with significant influence of upwelling near the plateau edge. The dominant benthic species remain constant through the ODP Cretaceous section, but subdominance changes, and the section is divided into three informal zones based on dominance/subdominance characteristics of the benthic fauna. Brief taxonomic comments are made on several species and some are figured.

Benthic foraminifera of the central Indian Ocean

Late Oligocene to late Pliocene vertical water-mass stratification along depth traverses in the northern Indian Ocean is depicted in this paper by benthic foraminifer index faunas. During most of this time, benthic faunas indicate well-oxygenated, bottom-water conditions at all depths except under the southern Indian upwelling and in the Pliocene in the southern Arabian Sea. Faunas suggest the initiation of lower oxygen conditions at intermediate depths in the northern Indian Ocean beginning in Oligocene Zone P21a. Lower oxygen conditions intensified during primary productivity pulses, possibly related to increased upwelling vigor, in the latest Oligocene and throughout most of the late middle through late Miocene. During times of elevated primary production, there may be more oxygen flux into sedimentary pore waters and the shallow infaunal habitat may become more oxygenated. One criterion for locating the source of "new" water masses is vertical homogeneity of benthic foraminifer indexes for well-oxygenated water masses from intermediate through abyssal depths. In the northern Mascarene Basin, this type of faunal homogeneity with depth corroborates the proposal that the northern Indian Ocean was an area of sinking well-oxygenated waters through most of the Miocene before Zone N17. Oxygenated, possibly "new" intermediate-water masses in the low- to middle-latitude Mascarene and Central Indian basins first developed in the late Oligocene. These well-oxygenated waters were probably more fertile than the Antarctic Intermediate Waters (AAIW) that cover intermediate depths in these areas today. Production of intermediate waters more similar to modern AAIW is indicated by the sparse benthic population of epifaunal rotaloid species in the northern Mascarene Basin during middle Miocene Zone N9 and from early through late Pliocene time. Deep-water characteristics are more difficult to interpret because of the extensive redeposition at the deeper sites. Redeposited intermediate, rather than shallow, water fossils and erosion from north to south in the Mascarene Basin are incompatible with the sluggish circulation from south to north through the western Indian Ocean basins today. Such erosion could result from the vigorous sinking of an intermediate-depth water mass of northern origin. Before late Oligocene Zone P22, benthic faunas indicate a twofold subdivision of the troposphere, with the boundary between upper and lower well-oxygenated water masses located from 2500-3000 mbsl. No characteristic bottom-water fauna developed before the end of late Oligocene Zone P22. Deep and abyssal benthic indexes suggest the development of water masses similar to those of the present day in the latest Miocene. Faunas containing deep-water benthic indexes, including the uvigerinids, suggestive of a water mass similar to modern Indian Deep Water (IDW), appeared during the late Miocene in the northern Mascarene and Central Indian basins. In the early Pliocene, this deep-water fauna was found only in the Central Indian Basin, whereas a fauna typical of modern Antarctic Bottom Water (AABW) spread through deep waters at 2800 mbsl in the Mascarene Basin. By late Pliocene Zone N21, however, deep-water faunas similar to their modern analogs were developed in both the eastern and western basins. Abyssal faunas, studied only in the Mascarene Basin, show more or less similarity to those under modern AABW. Bottom-water faunas containing Nuttallides umbonifera or Epistominella exiguua were first differentiated at the end of Zone P22, then appeared episodically during the early Miocene. These AABW-type faunas reappeared and migrated updepth into deep waters during the glacial episodes at the end of the Miocene and at the beginning of the Pliocene. By late Pliocene Zone N21, however, a bottom-water fauna similar to that under eastern Indian Bottom Water (IBW) developed in the Mascarene Basin. Modern bottom-water characteristics of the Mascarene Basin must have developed after ZoneN21.

Influence of sediment acidification and water flow on sediment acceptance and dispersal of juvenile soft-shell clams (Mya arenaria L.)

Although ocean acidification is expected to reduce carbonate saturation and yield negative impacts on open-ocean calcifying organisms in the near future, acidification in coastal ecosystems may already be affecting these organisms. Few studies have addressed the effects of sedimentary saturation state on benthic invertebrates. Here, we investigate whether sedimentary aragonite saturation (Omega aragonite) and proton concentration ([H+]) affect burrowing and dispersal rates of juvenile soft-shell clams (Mya arenaria) in a laboratory flume experiment. Two size classes of juvenile clams (0.5-1.5 mm and 1.51-2.5 mm) were subjected to a range of sediment Omega aragonite and [H+] conditions within the range of typical estuarine sediments (Omega aragonite 0.21-1.87; pH 6.8-7.8; [H+] 1.58 × 10**-8-1.51 × 10**- 7) by the addition of varying amounts of CO2, while overlying water pH was kept constant ~ 7.8 (Omega aragonite ~ 1.97). There was a significant positive relationship between the percent of juvenile clams burrowed in still water and Omega aragonite and a significant negative relationship between burrowing and [H+]. Clams were subsequently exposed to one of two different flow conditions (flume; 11 cm/s and 23 cm/s) and there was a significant negative relationship between Omega aragonite and dispersal, regardless of clam size class and flow speed. No apparent relationship was evident between dispersal and [H+]. The results of this study suggest that sediment acidification may play an important role in soft-shell clam recruitment and dispersal. When assessing the impacts of open-ocean and coastal acidification on infaunal organisms, future studies should address the effects of sediment acidification to adequately understand how calcifying organisms may be affected by shifting pH conditions.

Seawater carbonate chemistry and nutrient fluxes during experiments with brittlestar Amphiura filiformis, 2009

Rising levels of atmospheric carbon dioxide and the concomitant increased uptake of this by the oceans is resulting in hypercapnia-related reduction of ocean pH. Research focussed on the direct effects of these physicochemical changes on marine invertebrates has begun to improve our understanding of impacts at the level of individual physiologies. However, CO2-related impairment of organisms' contribution to ecological or ecosystem processes has barely been addressed. The burrowing ophiuroid Amphiura filiformis, which has a physiology that makes it susceptible to reduced pH, plays a key role in sediment nutrient cycling by mixing and irrigating the sediment, a process known as bioturbation. Here we investigate the role of A. filiformis in modifying nutrient flux rates across the sediment-water boundary and the impact of CO2- related acidification on this process. A 40 day exposure study was conducted under predicted pH scenarios from the years 2100 (pH 7.7) and 2300 (pH 7.3), plus an additional treatment of pH 6.8. This study demonstrated strong relationships between A. filiformis density and cycling of some nutrients; activity increases the sediment uptake of phosphate and the release of nitrite and nitrate. No relationship between A. filiformis density and the flux of ammonium or silicate were observed. Results also indicated that, within the timescale of this experiment, effects at the individual bioturbator level appear not to translate into reduced ecosystem influence. However, long term survival of key bioturbating species is far from assured and changes in both bioturbation and microbial processes could alter key biogeochemical processes in future, more acidic oceans.

Oxygen and Carbon stable isotope ratios for Site U1334

The Oligocene-Miocene transition (OMT) (~23 Ma) is interpreted as a transient global cooling event, associated with a large-scale Antarctic ice sheet expansion. Here we present a 2.23 Myr long high-resolution (~3 kyr) benthic foraminiferal oxygen and carbon isotope (d18O and d13C) record from Integrated Ocean Drilling Program Site U1334 (eastern equatorial Pacific Ocean), covering the interval from 21.91 to 24.14 Ma. To date, five other high-resolution benthic foraminiferal stable isotope stratigraphies across this time interval have been published, showing a ~1 per mil increase in benthic foraminiferal d18O across the OMT. However, these records are still few and spatially limited and no clear understanding exists of the global versus local imprints. We show that trends and the amplitudes of change are similar at Site U1334 as in other high-resolution stable isotope records, suggesting that these represent global deep water signals. We create a benthic foraminiferal stable isotope stack across the OMT by combining Site U1334 with records from ODP Sites 926, 929, 1090, 1264, and 1218 to best approximate the global signal. We find that isotopic gradients between sites indicate interbasinal and intrabasinal variabilities in deep water masses and, in particular, note an offset between the equatorial Atlantic and the equatorial Pacific, suggesting that a distinct temperature gradient was present during the OMT between these deep water masses at low latitudes. A convergence in the d18O values between infaunal and epifaunal species occurs between 22.8 and 23.2 Ma, associated with the maximum d18O excursion at the OMT, suggesting climatic changes associated with the OMT had an effect on interspecies offsets of benthic foraminifera. Our data indicate a maximum glacioeustatic sea level change of ~50 m across the OMT.

Species abundance and water and sediment characteristics for three ANDEEP stations on Maud Rise and the Weddell Sea

The benthic fauna was investigated during the expedition ANT-XXIV/2 (2007/08) in relation to oceanographic features, biogeochemical properties and sediment characteristics, as well as the benthic, pelagic and air-breathing fauna. The results document that Maud Rise (MR) differs distinctly from surrounding deep-sea basins investigated during previous Southern Ocean expeditions (ANDEEP 2002, 2005). Considering all taxa, the overall similarity between MR and adjacent stations was low (~20% Bray-Curtis-Similarity), and analyses of single taxa show obvious differences in species composition, abundances and densities. The composition and diversity of bivalves of MR are characterised by extremely high abundances of three species, especially the small sized Vesicomya spp. Exceptionally high gastropod abundance at MR is due to the single species Onoba subantarctica wilkesiana, a small brooder that may prey upon abundant benthic foraminiferas. The abundance and diversity of isopods also show that one family, Haplomunnidae, occurs with a surprisingly high number of individuals at MR while this family was not found at any of the 40 bathyal and abyssal ANDEEP stations. Similarly, polychaetes, especially the tube-dwelling, suspension-feeder fraction, are represented by species not found at the comparison stations. Sponges comprise almost exclusively small specimens in relatively high numbers, especially a few species of Polymastiidae. Water-column sampling from the surface to the seafloor, including observations of top predators, indicate the existence of a prospering pelagic food web. Local concentrations of top predators and zooplankton are associated with a rich ice-edge bloom located over the northern slope of MR. There the sea ice melts, which is probably accelerated by the advection of warm water at intermediate depth. Over the southern slope, high concentrations of Antarctic krill (Euphausia superba) occur under dense sea ice and attract Antarctic Minke Whales (Balaenoptera bonaerensis) and several seabird species. These findings suggest that biological prosperity over MR is related to both oceanographic and sea-ice processes. Downward transport of the organic matter produced in the pelagic realm may be more constant than elsewhere due to low lateral drift over MR.

Isopoda E(S100) and surface sediment characteristics of ANDEEP I-III stations in the Southern Ocean deep sea

Shallow marine benthic communities around Antarctica show high levels of endemism, gigantism, slow growth, longevity and late maturity, as well as adaptive radiations that have generated considerable biodiversity in some taxa1. The deeper parts of the Southern Ocean exhibit some unique environmental features, including a very deep continental shelf2 and a weakly stratified water column, and are the source for much of the deep water in the world ocean. These features suggest that deep-sea faunas around the Antarctic may be related both to adjacent shelf communities and to those in other oceans. Unlike shallow-water Antarctic benthic communities, however, little is known about life in this vast deep-sea region2, 3. Here, we report new data from recent sampling expeditions in the deep Weddell Sea and adjacent areas (748-6,348 m water depth) that reveal high levels of new biodiversity; for example, 674 isopods species, of which 585 were new to science. Bathymetric and biogeographic trends varied between taxa. In groups such as the isopods and polychaetes, slope assemblages included species that have invaded from the shelf. In other taxa, the shelf and slope assemblages were more distinct. Abyssal faunas tended to have stronger links to other oceans, particularly the Atlantic, but mainly in taxa with good dispersal capabilities, such as the Foraminifera. The isopods, ostracods and nematodes, which are poor dispersers, include many species currently known only from the Southern Ocean. Our findings challenge suggestions that deep-sea diversity is depressed in the Southern Ocean and provide a basis for exploring the evolutionary significance of the varied biogeographic patterns observed in this remote environment.

Benthic foraminifera accumulation rate and benthic foraminiferal productivity proxies at ODP Site 165-999 in the Caribbean (Appendix A)

This study tests the hypothesis that the late Miocene to early Pliocene constriction and closure of the Central American Seaway (CAS), connecting the tropical Atlantic and East quatorial Pacific (EEP), caused a decrease in productivity in the Caribbean, due to decreased coastal upwelling and an end to the connection with high-productivity tropical Pacific waters. The present study compared paleoceanographic proxies for the interval between 8.3 and 2.5 Ma in 47 samples from south Caribbean ODP Site 999 with published data on EEP DSDP Site 503. Proxies for Site 999 include the relative abundance of benthic foraminiferal species representing bottom current velocity and the flux of organic matter to the sea floor, the ratio of infaunal/epifaunal benthic foraminiferal species and benthic foraminifer accumulation rates (BFARs). In addition, we calculated % resistant planktic foraminifers species and used the previously published % sand fraction and benthic carbon isotope values from Site 999. During early shoaling of the Isthmus (8.3-7.9 Ma) the Caribbean was under mesotrophic conditions, with little ventilation of bottom waters and low current velocity. The pre-closure interval (7.6-4.2 Ma) saw enhanced seasonal input of phytodetritus with even more reduced ventilation, and enhanced dissolution between 6.8 and 4.8 Ma. During the post-closure interval (4.2-2.5 Ma) in the Caribbean, paleoproductivity decreased, current velocity was reduced, and ventilation improved, while the seasonality of phytodetrital input was reduced dramatically, coinciding with the establishment of the Atlantic-Pacific salinity contrast at 4.2 Ma. Our data support the hypothesis that late Miocene constriction of the CAS at 7.9 Ma and its closure at 4.2 Ma caused a gradual decrease in paleoproductivity in the Caribbean, consistent with decreased current velocity and seasonality of the phytodetrital input.

Zooplankton distribution and migration in low-oxygen modewater eddies

The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300-600 m depth. Here, oxygen concentrations rarely fall below 40 µmol O2 kg-1, but are expected to decline under future projections of global warming. The recent discovery of mesoscale eddies that harbour a shallow suboxic (<5 µmol O2 kg-1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by on-going ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. At nighttime, when a large proportion of acoustic scatterers is ascending into the upper 150 m, a drastic reduction in mean volume backscattering (Sv, shipboard ADCP, 75kHz) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers were avoiding the depth range between about 85 to 120 m, where oxygen concentrations were lower than approximately 20 µmol O2 kg-1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time-series observations of a moored ADCP (upward looking, 300kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies have been identified to be followed by zooplankton in response to the eddy OMZ: i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids), ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods), iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes), and iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy i), ii) and iv), compression of the habitable volume in the surface may increase prey-predator encounter rates, rendering zooplankton and micronekton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 µmol O2 kg-1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton and micronekton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Mn/Ca and Fe/Ca systematics in benthic foraminifera from the Peruvian OMZ

In this study we present an initial dataset of Mn/Ca and Fe/Ca ratios in tests of benthic foraminifera from the Peruvian oxygen minimum zone (OMZ) determined with SIMS. These results are a contribution to a better understanding of the proxy potential of these elemental ratios for ambient redox conditions. Foraminiferal tests are often contaminated by diagenetic coatings, like Mn rich carbonate- or Fe and Mn rich (oxyhydr)oxide coatings. Thus, it is substantial to assure that the cleaning protocols are efficient or that spots chosen for microanalyses are free of contaminants. Prior to the determination of the element/Ca ratios, the distributions of several elements (Ca, Mn, Fe, Mg, Ba, Al, Si, P and S) in tests of the shallow infaunal species Uvigerina peregrina and Bolivina spissa were mapped with an electron microprobe (EMP). To visualize the effects of cleaning protocols uncleaned and cleaned specimens were compared. The cleaning protocol included an oxidative cleaning step. An Fe rich phase was found on the inner test surface of uncleaned U. peregrina specimens. This phase was also enriched in Al, Si, P and S. A similar Fe rich phase was found at the inner test surface of B. spissa. Specimens of both species treated with oxidative cleaning show the absence of this phase. Neither in B. spissa nor in U. peregrina were any hints found for diagenetic (oxyhydr)oxide or carbonate coatings. Mn/Ca and Fe/Ca ratios of single specimens of B. spissa from different locations have been determined by secondary ion mass spectrometry (SIMS). Bulk analyses using solution ICP-MS of several samples were compared to the SIMS data. The difference between SIMS analyses and ICP-MS bulk analyses from the same sampling sites was 14.0-134.8 µmol mol-1 for the Fe/Ca and 1.68(±0.41) µmol mol-1 for the Mn/Ca ratios. This is in the same order of magnitude as the variability inside single specimens determined with SIMS at these sampling sites (1sigma[Mn/Ca] = 0.35-2.07 µmol mol-1; 1sigma[Fe/Ca] = 93.9-188.4 µmol mol-1). The Mn/Ca ratios in the calcite were generally relatively low (2.21-9.93 µmol mol-1) but in the same magnitude and proportional to the surrounding pore waters (1.37-6.67 µmol mol-1). However, the Fe/Ca ratios in B. spissa show a negative correlation to the concentrations in the surrounding pore waters. Lowest foraminiferal Fe/Ca ratios (87.0-101.0 µmol mol-1) were found at 465 m water depth, a location with a strong sharp Fe peak in the pore water next to the sediment surface and respectively, high Fe concentrations in the surrounding pore waters. Previous studies found no living specimens of B. spissa at this location. All these facts hint that the analysed specimens already were dead before the Fe flux started and the sampling site just recently turned anoxic due to fluctuations of the lower boundary of the OMZ near the sampling site (465 m water depth). Summarized Mn/Ca and Fe/Ca ratios are potential proxies for redox conditions, if cleaning protocols are carefully applied. The data presented here may be rated as base for the still pending detailed calibration.