Relative abundance of dinoflagellate cysts in surface sediments and dinoflagellate counts from three sediment cores from the Atlantic sector of the Southern Ocean

In this study we investigate the potential of organic-walled dinoflagellate cysts (dinocysts) as tools for quantifying past sea-surface temperatures (SST) in the Southern Ocean. For this purpose, a dinocyst reference dataset has been formed, based on 138 surface sediment samples from different circum-Antarctic environments. The dinocyst assemblages of these samples are composed of phototrophic (gonyaulacoid) and heterotrophic (protoperidinioid) species that provide a broad spectrum of palaeoenvironmental information. The relationship between the environmental parameters in the upper water column and the dinocyst distribution patterns of individual species has been established using the statistical method of Canonical Correspondence Analysis (CCA). Among the variables tested, summer SST appeared to correspond to the maximum variance represented in the dataset. To establish quantitative summer SST reconstructions, a Modern Analogue Technique (MAT) has been performed on data from three Late Quaternary dinocyst records recovered from locations adjacent to prominent oceanic fronts in the Atlantic sector of the Southern Ocean. These dinocyst time series exhibit periodic changes in the dinocyst assemblage during the last two glacial/interglacial-cycles. During glacial conditions the relative abundance of protoperidinioid cysts was highest, whereas interglacial conditions are characterised by generally lower cyst concentrations and increased relative abundance of gonyaulacoid cysts. The MAT palaeotemperature estimates show trends in summer SST changes following the global oxygen isotope signal and a strong correlation with past temperatures of the last 140,000 years based on other proxies. However, by comparing the dinocyst results to quantitative estimates of summer SSTs based on diatoms, radiolarians and foraminifer-derived stable isotope records it can be shown that in several core intervals the dinocyst-based summer SSTs appeared to be extremely high. In these intervals the dinocyst record seems to be highly influenced by selective degradation, leading to unusual temperature ranges and to unrealistic palaeotemperatures. We used the selective degradation index (kt-index) to determine those intervals that have been biased by selective degradation in order to correct the palaeotemperature estimates. We show that after correction the dinocyst based SSTs correspond reasonably well with other palaeotemperature estimates for this region, supporting the great potential of dinoflagellate cysts as a basis for quantitative palaeoenvironmental studies.

Dinoflagellate cysts in surface sediments

Surface sediments at 439 sites throughout the North Atlantic Ocean and adjacent seas have been analyzed for dinoflagellate cysts in order to establish a reference database from which paleoenvironmental transfer functions can be developed. Laboratory procedures and systematics were standardized in order to avoid bias introduced by the selective loss of taxa and to facilitate site to site comparison. 371 sites were retained to develop the database that includes 41 taxa, some of which were grouped using morphological and/or ecological criteria. 27 taxa were retained for statistical purposes. Distribution maps of these latter taxa have been plotted on the basis of their relative abundance. Principal component analyses were performed in order to describe the distribution of assemblages. The relation between the assemblages, as well as the relative abundance of individual taxa, and selected sea-surface parameters are illustrated. The parameters which were considered include temperature and salinity for winter (February) and summer (August) together with the duration of sea-ice cover. Transfer functions using the best analogue method have been tested with a view to reconstruct past sea-surface parameters. Validation procedures on this transfer function demonstrate that more than 95% of the reconstructions are included within the interannual variability of modern sea-surface conditions. Therefore, these transfer functions give accurate results and can be applied to reconstructing paleo-temperatures and -salinities from analogous assemblages in Quaternary sedimentary sequences. Protoperidinium stellatum (Wall in Wall & Dale, 1968) Head, comb. nov. (basionym = Peridinium stellatum) is proposed as new, and Algidasphaeridium? minutum var. cezare de Vernal et al., 1989 ex de Vernal et al. is newly validated.

Abundance of organic-walled dinoflagellate cysts in surface sediments from the Atlantic sector of the Southern Ocean (Appendix 2)

Thirty-two surface sediment samples from the Southern Ocean (eastern Atlantic sector), between the Subtropical Front and the Weddell Gyre, were investigated to provide information on the distribution of modern organic-walled dinoflagellate cysts in relation to the oceanic fronts of the Antarctic Circumpolar Current (ACC). A clearly distinguishable distribution pattern was observed in relation to the water masses and fronts of the ACC. The dinoflagellate cysts of species characteristic of open oceanic environments, such as Impagidinium species, are highly abundant around the Subtropical Front, whereas south of this front, cosmopolitan species such as Nematosphaeropsis labyrinthus and the cysts of Protoceratium reticulatum characterise the transition from subtropical to subantarctic surface waters. The subantarctic surface waters are dominated by the cysts of heterotrophic dinoflagellates, such as Protoperidinium spp. and Selenopemphix antarctica. The cysts of Protoperidinium spp. form the dominant part of the assemblages around the Antarctic Polar Front, whereas S. antarctica concentrations increase further to the south. The presence of S. antarctica in sediments of the Maud Rise, a region of seasonal sea-ice cover, reflects its tolerance for low temperatures and sea-ice cover. A previously undescribed species, Cryodinium meridianum gen. nov. sp. nov., has a restricted distribution pattern between the Antarctic Polar Front and the ACC-Weddell Gyre Boundary.

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.

Dinoflagellate cysts counts of sediment core MD95-2042

Dinoflagellate cysts were analysed from IMAGES core MD952042 (37°48?N; 10°01?W) retrieved from the Tagus Abyssal Plain. Previous results of stable isotope and magnetic susceptibility measurements as well as of planktonic foraminiferal temperature reconstruction from this core, suggest the occurrence of "Heinrich-like events" (i.e. large ice-sheet decay) during Marine Isotopic Stage 5 (MIS 5). Dinoflagellate assemblages of this time period have revealed six dinocyst events that are characterised by peaks in Bitectatodinium tepikiense percentages. These events occur synchronously with "Heinrich-like events" previously identified. They are coeval with major retreats of the forest on land, indicating, therefore, drastic changes in the regional climate. However, results from the Ice-Rafted Detritus (IRD) analysis of the >150 ?m lithic fraction shows that MIS 5 of MD952042 has only recorded one significant input of iceberg discharge, located at the MIS 6/MIS 5 transition. It seems therefore that it is the only event that could be called a "true Heinrich event".

Organic-walled dinoflagellate cysts and environmental analysis of surface sediments from the Northwest African upwelling area

In order to examine the spatial distribution of organic-walled dinoflagellate cysts (dinocysts) in recent sediments related to environmental conditions in the water column, thirty-two surface sediment samples from the NW African upwelling region (20-32°N) were investigated. Relative abundances of the dinocyst species show distinct regional differences allowing the separation of four hydrographic regimes. (1) In the area off Cape Ghir, which is characterized by most seasonal upwelling and river discharge, Lingulodinium machaerophorum strongly dominates the associations which are additionally characterized by cysts of Gymnodinium nolleri, cysts of Polykrikos kofoidii and cysts of Polykrikos schwartzii. (2) Off Cape Yubi, a region with increasing perennial upwelling, L. machaerophorum, Brigantedinium spp., species of the genus Impagidinium and cysts of Protoperidinium stellatum occur in highest relative abundances. (3) In coastal samples between Cape Ghir and Cape Yubi, Gymnodinium catenatum, species of the genus Impagidinium, Nematosphaeropsis labyrinthus, Operculodinium centrocarpum, cysts of P. stellatum and Selenopemphix nephroides determine the species composition. (4) Off Cape Blanc, where upwelling prevails perennially, and at offshore sites, heterotrophic dinocyst species show highest relative abundances. A Redundancy Analysis reveals fluvial mud, sea surface temperature and the depth of the mixed layer in boreal spring (spring) as the most important parameters relating to the dinocyst species association. Dinocyst accumulation rates were calculated for a subset of samples using well-constrained sedimentation rates. Highest accumulation rates with up to almost 80.000 cysts cm**-2 ky**-1 were found off Cape Ghir and Cape Yubi reflecting their eutrophic upwelling filaments. A Redundancy Analysis gives evidence that primary productivity and the input of fluvial mud are mostly related to the dinocyst association. By means of accumulation rate data, quantitative cyst production of individual species can be considered independently from the rest of the association, allowing autecological interpretations. We show that a combined interpretation of relative abundances and accumulation rates of dinocysts can lead to a better understanding of the productivity conditions off NW Africa.

Abundance of organic-walled dinoflagellate cysts of sediment core GeoB3603-2

Here, we reconstruct the varying influence of the Agulhas Current (AgC), the South Atlantic Current and the Antarctic Circumpolar Current on the Agulhas Retroflection (AgR) in the eastern South Atlantic Ocean for the last 160,000 years on the basis of the dinoflagellate cysts, pollen and spores present in a sediment core (GeoB 3603-2) from the southeastern Cape Basin offshore South Africa, where the Agulhas Current enters the Atlantic Ocean. Our analyses reveal strong orbital forcing on the heat exchange between the Indian Ocean and the South Atlantic Ocean during the Late Quaternary. Maxima in local productivity appear to be primarily related to a strengthening of the ocean circulation as a result of the high seasonal contrast during precession maxima. During precession minima, seasonal contrast was low and stratified, oligo- to mesotrophic conditions prevailed, notably when these minima coincided with the glacial terminations. The clear presence of periodicities on a sub-Milankovitch scale as well as modulations of the primary frequencies demonstrate that the Agulhas Retroflection furthermore is modulated substantially by complex interactions between the subtropical, 'precession-driven', climate and ocean circulation systems, and the southern, 'obliquity-driven', high latitudes.

Dinoflagellate cysts in western equatorial Atlantic surface sediments

In contrast to the wide range of studies carried out in temperate and high-latitude oceanic regions, only a few studies have focused on recent and Holocene organic-walled dinoflagellate cyst assemblages from the tropics. This information is, however, essential for fully understanding the ability of species to adapt to different oceanographic regimes, and ultimately their potential application to local and regional palaeoenvironmental and palaeoceanographic reconstructions. Surface sediment samples of the western equatorial Atlantic Ocean north of Brazil, an area greatly influenced by Amazon River discharge waters, were therefore analysed in detail for their organic-walled dinoflagellate cyst content. A diverse association of 43 taxa was identified, and large differences in cyst distribution were observed. The cyst thanatocoenosis in bottom sediments reflects the seasonal advection of Amazon River discharge water through the Guyana Current and the North Equatorial Countercurrent well into the North Atlantic. To establish potential links between cyst distribution and the environmental conditions of the upper water column, distribution patterns were compared with mean temperature, salinity, density and stratification gradients within the upper water column (0-100 m) over different times of the year, using correspondence analysis and canonical correspondence analysis. The analyses show that differences in these parameters only play a subordinate role in determining species distribution. Instead, nutrient availability, or related factors, dominates the distribution pattern. The only possible indicators of slightly reduced salinities are Trinovantedinium applanatum and Lingulodinium machaerophorum. Four assemblage groups of cyst taxa with similar environmental affinities related to specific water masses/currents can be distinguished and have potential for palaeoenvironmental reconstruction.

(Appendix A) Radiolarian abundance in ODP Hole 202-1241A

A quantitative radiolarian study at Ocean Drilling Program Site 1241 in the eastern tropical Pacific enables us to reconstruct paleoceanographic changes that occurred since the latest middle Miocene. Today, this site is located just under the Eastern Pacific Warm Pool (EPWP). Based on the abundance variations of radiolarian characteristic species which are indicators of upwelling and thermocline changes, it is suggested that three notable changes occurred at 10.6, 9.8, and 4.2 Ma in the region. Four distinct periods of oceanographic conditions bounded by these notable changes were characterized on the basis of the following: (1) stratified seawater (12.0 to 10.6 Ma); (2) a shallowing of the thermocline and an increasing of upwelling (10.6 to 9.8 Ma); (3) significant inflow of warm water to the eastern tropical Pacific caused by an intensified Northern Equatorial Countercurrent (NECC), resulting in the formation of EPWP (9.8 to 4.2 Ma); and (4) the reduction of the EPWP and the NECC, and an increase in upwelling (4.2 to 0 Ma). The timing of these paleoceanographic events indicated the strong relations with the opening and closing of the Indonesian and Central American (Panama) Seaways. The reduction of the EPWP (this study) and the deepening of the thermocline in western Pacific at about 4.2 Ma (Cannariato and Ravelo, 1997; Chaisson and Ravelo, 2000) indicated a change from a state resembling El Niño in the late Miocene and the early Pliocene time to a state resembling La Niña by the late Pliocene

Dinoflagellate cyst counts of sediment core GIK23071-3

Marine sediments from the Vøring Plateau (Norwegian Sea) have been studied for their dinoflagellate cyst (dinocyst) and foraminiferal content in order to reconstruct sea-surface conditions in the eastern Norwegian Sea during Marine Isotope Stage (MIS) 5e. In combination with stable oxygen isotope and ice rafted detritus (IRD) data, the variations in foraminiferal and dinocyst assemblage composition reflect a stepwise transition from the final phase of deglaciation (Termination II) into typical interglacial conditions. This stepwise change is repeated subsequently during the cooling conditions of glacial inception towards MIS 5d. The interval studied is characterized by relatively high abundances of Bitectatodinium tepikiense, in comparison to present-day values in the area, indicating a larger seasonal temperature amplitude with enhanced surface water stratification during MIS 5e. The important occurrence of the warm-temperate dinocyst Spiniferites mirabilis s.l. concurrent with subpolar foraminifers Turborotalita quinqueloba, Globigerina bulloides, and Globigerinita glutinata reveals that most pronounced interglacial marine conditions prevailed in the area just prior to the transition towards MIS 5d. The late stratigraphic position of this phase in the interglacial is verified by comparison with dinocyst data from south of Iceland, manifesting its over-regional implication. Besides the good agreement in dinocyst and foraminiferal assemblage changes, the variations in and between both fossil assemblages also point to the existence of some significant surface water variability in the eastern Norwegian Sea during MIS 5e.

(Appendix B Table 2) Abundance of organic-walled dinoflagellate cysts of sediment core GeoB2204-2

A palaeoceanographic reconstruction of the Late Quaternary tropical Atlantic Ocean has been made on the basis of dinoflagellate cyst associations of two sediment cores: the first core was recovered from below the highly productive waters of the equatorial divergence and the second from the oligotrophic western tropical Atlantic Ocean. Palaeoenvironmental indicators for productivity, sea surface temperature (SST) and salinity (SSS) based on selected organic-walled dinoflagellate cyst species have been established. On the basis of these palaeoenvironmental indicators, a strengthened intensity of the equatorial divergence in the eastern region during glacials and cold periods of interglacials has been reconstructed. The highest SST probably occurred around substage 5.5 and might refer to weakest upwelling intensity. In comparison, SST and SSS appear to have been generally higher in the western tropical Atlantic Ocean, with probably enhanced values during glacial intervals. Pronounced differences in accumulation rates and relative abundances of cysts formed by congruentidiacean dinoflagellates and relative abundances of oligotrophic cyst species between the eastern and the western region can be related to differences in palaeoproductivity, suggesting much higher values in the eastern area. The coherence between variation in frequency of the indicators for productivity and the boreal summer insolation and monsoon intensity in the eastern tropical Atlantic Ocean suggests an oceanographic reflection of regional intertropical, rather than boreal, dynamics.

Phytoplanktonic microfossil groups of sediments from the South and Equatorial Atlantic

Individual planktonic microfossil species, or assemblage groups of different species, are often used to, qualitatively and/or quantitatively, reconstruct past (sub)surface-water conditions of the world's oceans and seas. Until now, little information has been available on the surface sediment distribution patterns and paleoenvironmental reconstruction potential of coccolith, calcareous dinoflagellate cyst and organic-walled dinoflagellate cyst assemblages of the South and equatorial Atlantic, especially at the species level. This paper (i) summarizes the distributions of these three phytoplanktonic microfossil groups in numerous Atlantic surface sediments from 20°N-50°S and 30°E-65°W and determines their relationship with the physicochemical and trophic conditions of the overlying (sub)surface-waters, and (ii) determines the synecology of the three phytoplankton groups by carrying out statistical analyses (i.e. detrended and canonical correspondence analyses) on all groups simultaneously. Ecological relationships are additionally strengthened by statistically comparing the distribution patterns of the phytoplankton groups with those of planktonic foraminifera (Pflaumann et al. 1996; Niebler et al. 1998), as the ecological preferences of the latter are much better known. Many of the analyzed phytoplanktonic microfossil species or groups of species in the surface sediments do show restricted distributions which primarily reflect the environmental conditions of the upper water masses above them (e.g. sea-surface temperature, productivity, stratification). The acquired 'reference' data sets are large and diverse enough to allow future development of transfer functions for the reconstruction of past surface-water conditions, and show that there is still an enormous paleoenvironmental reconstruction potential concealed in many fossil coccolith and dinoflagellate cyst assemblages.

(Table T3) Distribution of radiolarian taxa in sediments of ODP Leg 180 sites

Cores from the 11 sites drilled during Leg 180 showed radiolarian assemblages that appear only in the Quaternary sediments. The most diverse and well-preserved assemblages were found in hemipelagic sediments from Holes 1108A, 1110A, and 1115B.