Maastrichtian calcareous nannofossils of Kronsmoor-Section

The latest Campanian-earliest Maastrichtian interval is well known as a period of intense climate cooling. This cooling caused a distinctive bipolar biogeographic distribution of calcareous nannofossil assemblages: High latitude settings were dominated by newly evolving endemic taxa, former cosmopolitan species disappeared at the same time and equatorial communities experienced an invasion of cool water taxa. The impact of this cooling on northern mid-latitude assemblages is, however, less well known. In order to overcome this gap we studied the Kronsmoor section (northwest Germany). This section provides a continuous upper Campanian - lower Maastrichtian succession with moderately to well preserved nannofossils. Uppermost Campanian assemblages are dominated by Prediscosphaera cretacea; other common taxa include Prediscosphaera stoveri, Watznaueria barnesiae and Micula staurophora. The lower Maastrichtian is characterized by lower numbers of P. cretacea and frequent Kamptnerius magnificus, Arkhangelskiella cymbiformis and Cribrosphaerella ehrenbergii. These changes reflect, in part, the Campanian-Maastrichtian boundary cooling since some successful taxa (e.g. K. magnificus) are related to cool surface waters. Other shifts in the nannofossil communities were perhaps the result of a changing nutrient regime. Stronger latitudinal gradients may have increased wind velocities and thus the eolian input of ferruginous dust required by N-fixing bacteria. The enhanced high latitude deep-water formation probably changed the bottom-water environment in disfavor of denitrificating organisms. A decline of chemical weathering and fluviatile transport may have reduced the amount of bioavailable phosphate. These processes led to an increased nitrate and a decreased phosphate content shifting the nutrient regime from nitrate towards phosphate limitation.

Seawater carbonate chemistry and processes during experiments with amphipod Gammarus locusta, 2009

We report an investigation of the effects of increases in pCO2 on the survival, growth and molecular physiology of the neritic amphipod Gammarus locusta which has a cosmopolitan distribution in estuaries. Amphipods were reared from juvenile to mature adult in laboratory microcosms at three different levels of pH in nominal range 8.1-7.6. Growth rate was estimated from weekly measures of body length. At sexual maturity the amphipods were sacrificed and assayed for changes in the expression of genes coding for a heat shock protein (hsp70 gene) and the metabolic enzyme glyceraldehyde-3-phosphate dehydrogenase (gapdh gene). The data show that the growth and survival of this species is not significantly impacted by a decrease in sea water pH of up to 0.5 units. Quantitative real-time PCR analysis indicated that there was no significant effect of growth in acidified sea water on the sustained expression of the hsp70 gene. There was a consistent and significant increase in the expression of the gapdh gene at a pH of ~7.5 which, when combined with observations from other workers, suggests that metabolic changes may occur in response to acidification. It is concluded that sensitive assays of tissue physiology and molecular biology should be routinely employed in future studies of the impacts of sea water acidification as subtle effects on the physiology and metabolism of coastal marine species may be overlooked in conventional gross "end-point" studies of organism growth or mortality.

Benthic foraminifers in Mesozoic and Cenozoic sediments of the southwestern Atlantic

Benthic foraminiferal assemblages in Mesozoic and Cenozoic sediments were studied at Sites 511, 512, 513, and 514 drilled during Leg 71 in the southwestern Atlantic on the Maurice Ewing Bank and in the Argentine Basin. Benthic foraminifers in almost all stratigraphic subdivisions of Sites 511 and 512 reflect the gradual subsidence of the Falkland Plateau from shelf depths in the Barremian-Albian, when a semiclosed basin with restricted circulation of water masses and anaerobic conditions existed, to lower bathyal depths in the Late Cretaceous and Cenozoic, with an abrupt acceleration at the boundary of Lower and Upper Cretaceous. The composition, distribution, and preservation of Late Cretaceous assemblages of benthic foraminifers suggest considerable fluctuations of the foraminiferal lysocline and the CCD. This is evidenced by dissolution facies and foraminiferal assemblages in which agglutinated and resistant calcareous forms predominated during high stands of the CCD and by calcareous facies in which rich assemblages of calcareous species predominated during low stands. The highest position of the CCD on the Plateau (less than 1500-2000 m) was in the late Cenomanian, Turonian, and Coniacian. In the Santonian and Campanian the CCD was at depths below 1500-2000 meters. At the end of the Campanian the CCD shifted again to depths comparable with those of Cenomanian and Turonian time. In the latest Campanian and the Maestrichtian the CCD was low and nanno-foraminiferal oozes with a rich assemblage of benthic foraminifers accumulated. Foraminiferal assemblages at Sites 513 and 514 in the Argentine Basin also testify to oceanic subsidence from lower bathyal depths in the Oligocene to abyssal ones at present. This process was complicated by the influence of geographical migrations of the Polar Front caused by extensions of the ice sheet in the Antarctic after the opening of the Drake Passage during the Oligocene. In Mesozoic and Cenozoic deposits of the Falkland Plateau and the Argentine Basin seven assemblages of benthic foraminifers were distinguished by age: early-middle Albian, middle-late Albian, Late Cretaceous (including four groups), middle Eocene, late Eocene-early Miocene, middle-late Miocene, and Pliocene-Quaternary. The Albian assemblages contain many species common to the foraminiferal fauna of the Austral Biogeographical Province. The Late Cretaceous assemblage contains, along with Austral species, species common to foraminifers of North America, Western Europe, the Russian platform, and the south of the U.S.S.R. Deep-sea cosmopolitan species prevail in Cenozoic assemblages.

Diatom biostratigraphy of the Kerguelen Plateau and Prydz Bay, Southern Ocean

Samples were examined for diatoms from 22 holes at 11 sites cored by ODP Leg 119 on the Kerguelen Plateau and in Prydz Bay, East Antarctica. Diatoms were observed in Oligocene through Holocene sediments recovered from the Kerguelen Plateau. The diatom flora from the Kerguelen Plateau is characterized by species such as Azpeitia oligocenica, Rocella gelida, Rocella vigilans, and Synedra jouseana in the Oligocene and Crucidenticula nicobarica, Denticulopsis hustedtii, Nitzschia miocenica, and Thalassiosira miocenica in the Miocene. This somewhat cosmopolitan assemblage gives way to a Pliocene and Holocene assemblage characterized by species such as Nitzschia kerguelensis, Thalassiosira inura, and Thalassiosira torokina, which are endemic to the Southern Ocean region. Samples examined from Prydz Bay are generally devoid of diatoms. The exception is Site 739, where diatoms occur sporadically in lower Oligocene and upper Miocene through Quaternary sediments. The Leg 119 diatom biostratigraphic results allow the development of a stratigraphic framework for the Indian sector of the Southern Ocean. This diatom zonation integrates diatom zonations developed previously for other sectors of the Southern Ocean. The zonation proposed here is based on biostratigraphic events of both geographically widespread and endemic species calibrated to the paleomagnetic stratigraphy. As such, this zonation has application throughout the Southern Ocean and allows correlation from the southern high latitudes to the low latitudes.

Using the critical salinity (Scrit) concept to predict invasion potential of the anemone Diadumene lineata in the Baltic Sea

It is widely assumed that the ability of an introduced species to acclimate to local environmental conditions determines its invasion success. The sea anemone Diadumene lineata is a cosmopolitan invader and shows extreme physiological tolerances. It was recently discovered in Kiel Fjord (Western Baltic Sea), although the brackish conditions in this area are physiologically challenging for most marine organisms. This study investigated salinity tolerance in D. lineata specimens from Kiel Fjord in order to assess potential geographical range expansion of the species in the Baltic Sea. In laboratory growth assays, we quantified biomass change and asexual reproduction rates under various salinity regimes (34: North Sea, 24: Kattegat, 14: Kiel Fjord, 7: Baltic Proper). Furthermore, we used 1H-NMR-based metabolomics to analyse intracellular osmolyte dynamics. Within 4 weeks D. lineata exhibited a 5-fold population growth through asexual reproduction at high salinities (34 and 24). Biomass increase under these conditions was significantly higher (69%) than at a salinity of 14. At a salinity of 7, anemones ceased to reproduce asexually, their biomass decreased and metabolic depression was observed. Five main intracellular osmolytes were identified to be regulated in response to salinity change, with osmolyte depletion at a salinity of 7. We postulate that depletion of intracellular osmolytes defines a critical salinity (Scrit) that determines loss of fitness. Our results indicate that D. lineata has the potential to invade the Kattegat and Skagerrak regions with salinity >10. However, salinities of the Baltic Proper (salinity <8) currently seem to constitute a physiological limit for the species.

Seawater carbonate chemistry 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 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.

Ocean Acidification Affects Redox-Balance and Ion-Homeostasis in the Life-Cycle Stages of Emiliania huxleyi

Ocean Acidification (OA) has been shown to affect photosynthesis and calcification in the coccolithophore Emiliania huxleyi, a cosmopolitan calcifier that significantly contributes to the regulation of the biological carbon pumps. Its non-calcifying, haploid life-cycle stage was found to be relatively unaffected by OA with respect to biomass production. Deeper insights into physiological key processes and their dependence on environmental factors are lacking, but are required to understand and possibly estimate the dynamics of carbon cycling in present and future oceans. Therefore, calcifying diploid and non-calcifying haploid cells were acclimated to present and future CO2 partial pressures (pCO2; 38.5 Pa vs. 101.3 Pa CO2) under low and high light (50 vs. 300 µmol photons/m**2 /s). Comparative microarray-based transcriptome profiling was used to screen for the underlying cellular processes and allowed to follow up interpretations derived from physiological data. In the diplont, the observed increases in biomass production under OA are likely caused by stimulated production of glycoconjugates and lipids. The observed lowered calcification under OA can be attributed to impaired signal-transduction and ion-transport. The haplont utilizes distinct genes and metabolic pathways, reflecting the stage-specific usage of certain portions of the genome. With respect to functionality and energy-dependence, however, the transcriptomic OA-responses resemble those of the diplont. In both life-cycle stages, OA affects the cellular redox-state as a master regulator and thereby causes a metabolic shift from oxidative towards reductive pathways, which involves a reconstellation of carbon flux networks within and across compartments. Whereas signal transduction and ion-homeostasis appear equally OA-sensitive under both light intensities, the effects on carbon metabolism and light physiology are clearly modulated by light availability. These interactive effects can be attributed to the influence of OA and light on the redox equilibria of NAD and NADP, which function as major sensors for energization and stress. This generic mode of action of OA may therefore provoke similar cell-physiological responses in other protists.

Environmental controls on the Emiliania huxleyi calcite mass

Although ocean acidification is expected to impact (bio)calcification by decreasing the seawater carbonate ion concentration, [CO3]2-, there exists evidence of non-uniform response of marine calcifying plankton to low seawater [CO3]2-. This raises questions on the role of environmental factors other than acidification and on the complex physiological responses behind calcification. Here we investigate the synergistic effect of multiple environmental parameters, including temperature, nutrient (nitrate and phosphate) availability, and seawater carbonate chemistry on the coccolith calcite mass of the cosmopolitan coccolithophore Emiliania huxleyi, the most abundant species in the world ocean. We use a suite of surface (late Holocene) sediment samples from the South Atlantic and southwestern Indian Ocean taken from depths lying well above the modern lysocline. The coccolith calcite mass in our results presents a latitudinal distribution pattern that mimics the main oceanographic features, thereby pointing to the potential importance of phosphorus and temperature in determining coccolith mass by affecting primary calcification and possibly driving the E. huxleyi morphotype distribution. This evidence does not necessarily argue against the potentially important role of the rapidly changing seawater carbonate chemistry in the future, when unabated fossil fuel burning will likely perturb ocean chemistry beyond a critical point. Rather our study highlights the importance of evaluating the combined effect of several environmental stressors on calcifying organisms to project their physiological response(s) in a high CO2 world and improve interpretation of paleorecords.

Pollen abundance in Late Quaternary sediment cores off West Africa

The pollen record of three marine late Quaternary cores off Senegal shows a juxtaposition of Mediterranean, Northern Saharan, Central Saharan elements, which are considered transported by the trade winds from a winter-rainfall area, and Sahelian, Soudanese, Soudano-Guinean elements, considered transported both by winds and mostly by the Senegal River, and coming from the monsoonal, summer tropical rainfall area of southern West Africa. Littoral vegetation is either the edaphically dry and saline Chenopodiaceae from sebkhas at the time of the main regression, or the warm tropical humid mangrove with Rhizophora during the humid optimum period. Four stratigraphic zones reflect, from basis to top: Zone 4. A semi-arid period with a balanced pollen input. Zone 3. A very arid period with the disappearance of monsoonal pollen, probably from the disappearance of the Senegal River, a very saline littoral plain with Chenopodiaceae, a larger input of northern Saharan pollen from intensified trade winds. Zone 2. A quite humid period, much more so than today, very suddenly established, with a northward extension of the monsoonal areas, a rich littoral mangrove, and weakening of the trade winds. Zone l. A slow and steady evolution toward the present semi-humid conditions with regression of the mangrove, and of the monsoonal areas toward the south. Tentative datations and correlations with the Tchad area suggested: zone 4: 22,500 to 19,000 years BP; zone 3: 19,000 to 12,500 years BP; zone 2: 12,500 to 5,500 years BP; zone 1: 5,500 years BP to top of core. Dinoflagellate cysts display a tropical assemblage with mostly estuarine neritic elements and also a weak oceanic component, mostly in the lower slope core 47. Cosmopolitan taxa dominate the assemblage and only a few species point to more specialized environments. Quantitative variations of the assemblage are the basis of stratigraphy which is not similar to the pollen stratigraphy, and an inshore-outshore gradient has to be taken into account to correlate the three cores.

(Table 2) Stratigraphic marker taxa in ODP Hole 162-985A sediments

The rich and diverse dinocyst assemblages in Cores 162-985A-32X through 62X confirm the importance of these microfossils in unraveling the evolution of the Norwegian Sea. Cosmopolitan taxa, with well-documented stratigraphic ranges in northwest Europe, indicate the following ages: Sections 162-985A-62X-1 through 51X-2, Rupelian (early Oligocene); 50X-5, Oligocene, possibly Chattian; 48X-6, Aquitanian? (early Miocene); 48X-4 through 37X-5, Aquitanian (early Miocene); and 36X-5 through 32X-1, Burdigalian (early Miocene). This stratigraphic interpretation suggests that a major hiatus, which can be correlated with an apparently coeval hiatus at Site 643, occurs within the Chattian at Site 985. Several endemic dinocyst taxa with unusual morphology and restricted stratigraphic occurrences are present in Hole 985A and other Norwegian Sea sites, especially Site 643. By using Hole 985A data for control, the Oligocene-Miocene sediments can be correlated with some degree of confidence in the Norwegian Basin.

Late Eocene to Recent deep-sea benthic foraminifera from the Central equatorial Pacific Ocean

Benthic foraminifers were studied in upper Eocene to Recent core-catcher samples from DSDP Sites 573, 574, and 575. The sites are on a north-south transect from the equator to about 05°N at about 133°W, water depth 4300 to 4600 m. At Site 574 additional samples were used to study the Eocene/Oligocene boundary in detail. About 200 specimens were counted per sample. The fauna is highly diverse (about 50 to 70 species per sample) and is of low dominance. The diversity is not related to age or sub-bottom depth. Many species are cosmopolitan and probably have wide environmental tolerances. Fluctuations in frequency of some taxa (e.g., Nuttallides umbonifera, Epistominella exigua, and Uvigerina spp.) cannot be correlated from one site to another. Several common species (e.g. Oridorsalis umbonatus and Globocassidulina subglobosa) range from late Eocene to Recent. First and last appearances are generally difficult to define precisely because many species are rare. For some species these datums differ from one site to another, but several datum levels are within 1 m.y. at all sites. First and last appearances are most numerous in two intervals, the late Eocene to early Oligocene (about 32 to 37 Ma) and the early to middle Miocene (about 13 to 18.5 Ma). Isotopic events occur within each of these periods of benthic faunal change, but the isotopic events have a shorter duration and start after the initiation of the changes in the fauna. Changes in deep-sea benthic faunal composition are not directly related to short-term oceanographic changes as expressed in isotopic records.

X-ray fluorescence scannings, diatoms, and dinocyst of ODP Site 189-1172

Eocene sediments drilled at the East Tasman Plateau (ETP) exhibit well-defined cycles, high-resolution magnetic stratigraphy, and environmentally-controlled dinoflagellate and diatom distribution patterns. We derive a cyclostratigraphy from the spectral analysis of high-resolution elemental concentration records (Ca, Fe) for this shallow marine time series spanning the middle to early late Eocene (C16n.2n - C21). Changes in carbonate content, the ratio between Gonyaulacoid and Peridinioid dinocysts, and relative abundance of "oligotrophic" diatoms serve as proxies for a high-resolution climatic and sea-level history with high values representing high sea-level stands and decreased eutrophy of surface waters. Changing ratios between high latitude dinocysts versus cosmopolitan species provide clues on sea surface temperature trends and water mass exchange. Our results show that the relatively shallow-water middle Eocene environments of the ETP are influenced by orbitally-forced climatic cycles superimposed on third order relative sea-level changes. Changes in the dominance of Milankovitch frequency at ~38.6 Ma (late Eocene) is related to an initial deepening-step within the Tasmanian Gateway prior to the major deepening during the middle late Eocene (~35.5 Ma). Decreasing sedimentation rates at 38 Ma and 37.2 Ma reflect winnowing associated with sea-level fall. This episode is followed by renewed transgression. Dinocyst distribution patterns indicate high latitude, probably cool temperate surface water conditions throughout, with the exception of a sudden surge in cosmopolitan species near the base of subchron C18.2r, at ~41 Ma; this event is tentatively correlated to the Middle Eocene Climatic Optimum.

(Table 1) Elemental mean values of the upper Paleocene and lower Eocene in IODP Hole 302-M0004A

We reconstruct the latest Paleocene and early Eocene (~57-50 Ma) environmental trends in the Arctic Ocean and focus on the Paleocene-Eocene thermal maximum (PETM) (~55 Ma), using strata recovered from the Lomonosov Ridge by the Integrated Ocean Drilling Program Expedition 302. The Lomonosov Ridge was still partially subaerial during the latest Paleocene and earliest Eocene and gradually subsided during the early Eocene. Organic dinoflagellate cyst (dinocyst) assemblages point to brackish and productive surface waters throughout the latest Paleocene and early Eocene. Dinocyst assemblages are cosmopolitan during this time interval, suggesting warm conditions, which is corroborated by TEX86'-reconstructed temperatures of 15°-18°C. Inorganic geochemistry generally reflects reducing conditions within the sediment and euxinic conditions during the upper lower Eocene. Spectral analysis reveals that the cyclicity, recorded in X-ray fluorescence scanning Fe data from close to Eocene thermal maximum 2 (~53 Ma, presence confirmed by dinocyst stratigraphy), is related to precession. Within the lower part of the PETM, proxy records indicate enhanced weathering, runoff, anoxia, and productivity along with sea level rise. On the basis of total organic carbon content and variations in sediment accumulation rates, excess organic carbon burial in the Arctic Ocean appears to have contributed significantly to the sequestration of injected carbon during the PETM.

Distribution of Upper Cretaceous deep water agglutinated foraminifera in sediments of the North Atlantic and its marginal seas (Tab. 1)

The stratigraphic and biogeographic distribution of more than 170 species of deep-water agglutinated benthic foraminifers (DWAF) from the North Atlantic and adjacent marginal seas has been compared with paleoenvironmental data (e.g. paleobathymetry, oxygenation of the bottom waters, amount of terrigenous input and substrate disturbance). Six general types of assemblages, in which deep water agglutinated taxa occur, are defined from the Turonian to Maastrichtian times: 1. High latitude slope assemblages 2. Low to mid latitude slope assemblages 3. Flysch-type assemblages 4. Deep water limestone assemblages (,,Scaglia,,-type) 5. Abyssal mixed calcareous-agglutinated assemblages 6. Abyssal purely agglutinated assemblages Latitudinal differences in faunal composition are observed, the most important of which is the lack or extreme paucity of calcareous forms in high latitude assemblages. East-to-west differences appear to be of comparatively minor importance. Most DWAF species occur in all studied regions and are thus considered as cosmopolitan. Biostratigraphic turnovers in the taxonomic content of assemblages are observed in the lowermost Turonian, mid-Campanian and in the upper Maastrichtian to lowermost Paleocene. These datum levels correspond to inter-regional and time-constant paleooceanographic events, which probably also affected the deep-water benthic biota. This allows us to use deep-water agglutinated foraminifers for biostratigraphy in the North Atlantic sequences deposited below CCD and to geographically extend the currently used zonal schemes which have been established in the Carpathian and Alpine areas.