(Table 1) Dinoflagellate cysts and acritarchs in upper Pliocene and Pleistocene sediments from ODP Hole 151-911A

Pliocene and Pleistocene sediments from Ocean Drilling Program Leg 151, Hole 911 A, drilled on the innermost Yermak Plateau (Eastern Arctic Ocean), were studied for their dinoflagellate cyst content. Three assemblage zones were tentatively defined, characterized by the predominance of few species. The composition of the assemblages changed markedly, even within single assemblage zones, during the last 2.6 to 2.8 m.y., reflecting the variable influence of warmer water from the Norwegian Sea, fluctuations in the influence of cold polar water masses, and the extent of sea-ice cover. Polar to subpolar surface water masses prevailed on the Yermak Plateau during the late Pliocene, when the eastern Arctic Ocean was probably isolated from the Norwegian-Greenland Sea. Intrusions of warmer water are recorded since the latest Pliocene, alternating with colder periods and a prolonged seasonal sea-ice cover. The composition of the dinoflagellate cyst assemblages has also changed considerably since the middle Pleistocene, reflecting the establishment of stronger fluctuations in surface water mass conditions than before at Yermak Plateau.

Dinoflagellate cysts analyses from sediment cores of the Po River discharge plume over the last two centuries

To obtain insight into the natural and/or human-induced changes in the trophic state of the distal portion of the Po River discharge plume over the last two centuries, high temporal resolution dinoflagellate cyst records were established at three sites. Cyst production rates appear to reflect the natural variability in the river's discharge, whereas cyst associations reflect the trophic state of the upper waters, which in turn can be related to agricultural development. The increased abundances of Lingulodinium machaerophorum and Stelladinium stellatum found as early as 1890 and 1920 correspond to the beginning of the industrial revolution in Italy and the first chemical production and dispersion of ammonia throughout Europe. After 1955, the increased abundances of these species and of Polykrikos schwartzii, Brigantedinium spp. and Pentapharsodinium dalei correspond to agriculturally induced alterations of the hypertrophic conditions. A slight improvement in water quality can be observed from 1987 onward.

(Figure 1) Dinoflagellate cyst stratigraphy of DSDP Hole 80-548A

Cyst assemblages from Sites 548, 549, and 550 were examined and gave evidence of early Eocene to late Miocene age. These assemblages were compared with other North Atlantic DSDP sites and with onshore sections in Denmark, southern England, Spain, and Italy. Some environmental interpretation is attempted for the Miocene assemblages; pollen, spores, and dinoflagellate cyst species were used to interpret the proximity of the shoreline. Key species are illustrated, along with some forms that are not discussed.

Dinoflagellate cyst abundances at DSDP Site 94-607 from the North Atlantic (Appendix B)

The climatic deterioration related to the onset of Northern Hemisphere glaciations (circa 2.52 Ma BP) must have lead to reorganization and relocation of species associations and may have enhanced species turnover. The present study investigates how this deterioration affects the dinoflagellate cyst and acritarch assemblages from two locations, DSDP Site 607 (North Atlantic) and the Singa section (southern Italy). The records from these locations cover the interval from 2.8 to 2.2 Ma with at least a 5 ka resolution and they have been correlated to the Milankovitch periodicities on a cycle to cycle basis by means of integrated high resolution stable isotope, calcium carbonate, foraminiferal, palynological and magnetostratigraphical datasets. In the present study this high resolution stratigraphic framework is used for a detailed correlation of events occurring in each of the depositional sequences. It also enables further assessment of the palaeoenvironmental preferences of some dinoflagellate cyst forms. Comparison of the two palynological records reveals a close correspondence in the timing of major assemblage changes and extinction events, confirming their Milankovitch cycle based correlation. A close link between periods of Northern Hemisphere cooling (at oxygen isotope stages 110, 104 and 100-96) and increased dinoflagellate cyst turnover appears to be present for both DSDP Site 607 and the Singa section. The turnover events can also be recognized in the records of planktic foraminifera and calcamous nannoplankton. Comparison of the Singa section with Site 607 and with other time equivalent marine palynological data sets, shows that some oceanic taxa respond similarly over a large area. The biostratigraphical implications are discussed. Notably the last occurrence of Invertocystu lucrymosa appears to be a valuable marker for isotope stage 110 in the Mediterranean and North Atlantic.

Dinoflagellate cyst species composition of recent surface samples (Appendix)

Dinoflagellate cysts and other organic-walled microfossils have been studied in recent surface sediments from the entire Norwegian-Greenland Sea. More than 30 taxa have been recognized, of which only few show a distinct distribution pattern, and allow description of four assemblages. The occurrence of most taxa is related to the relatively warmer waters of the Norwegian Sea. Algidaspaeridium? minutum s.1., Brigantedinium simplex and Impagidinium? pallidum are the only species showing a preference for colder water masses. Two species, I.? pallidum and Nematosphaeropsis labyrinthus are mainly restricted to the oceanic environment, whereas the other species have also been reported from neritic environments in previous studies. Due to the limited knowledge of the ecological and sedimentological factors influencing the occurrence of dinoflagellate cysts in oceanic environments, their distribution in recent sediments can be only related to surface water masses in a broad sense. Although the distribution of assemblages correlates with specific surface water masses, comparison with assemblages recovered from sediment traps deployed basinwide in the Norwegian-Greenland Sea (Dale and Dale, 1992) revealed some major discrepancies in species composition and percentage abundances. The differences cannot be explained with certainty at the moment, although there is some evidence that transport of dinoflagellate cysts and other fossilizable microplankton in water masses by currents, in sea-ice and sediments may modify the assemblages found in recent oceanic surface sediments from the Norwegian-Greenland Sea.

Calcareous dinoflagellates in the eastern equatorial Atlantic

Sediments of the Equatorial Atlantic (core GeoB 1105-4) have been investigated for both calcareous dinoflagellates and organic-walled dinoflagellate cysts. In order to determine the ecological affinity of calcareous dinoflagellates the statistical methods of Detrended Correspondence Analysis (DCA) and Redundancy Analysis (RDA) were used. Utilising DCA, distribution patterns of calcareous dinoflagellates have been compared with those of the ecologically much better known organic-walled dinoflagellate cysts. This method was also used to determine which environmental gradients have a major influence on the species composition. By using existing environmental information based on benthic and planktic foraminifera, such as Sea Surface Temperature (SST) and stable oxygen and carbon isotopes, as well as information on the amount of Calcium Carbonate and Total Organic Carbon (TOC) in bottom sediments, these gradients could be interpreted in terms of productivity and glacial-interglacial trends. Using RDA, the direct relationships between the distribution patterns of calcareous dinoflagellates with the above mentioned external variables could be determined. For the studied region and time interval (141-6.7 ka) the calcareous dinoflagellates show enhanced abundances in periods with reduced productivity most probably related to decreased divergence and relatively stratified, oligotrophic oceanic conditions.

Geographic distribution of dinoflagellate cysts in surface sediments

Dinoflagellate cysts are useful for reconstructing upper water conditions. For adequate reconstructions detailed information is required about the relationship between modern day environmental conditions and the geographic distribution of cysts in sediments. This Atlas summarises the modern global distribution of 71 organicwalled dinoflagellate cyst species. The synthesis is based on the integration of literature sources together with data of 2405 globally distributed surface sediment samples that have been preparedwith a comparable methodology and taxonomy. The distribution patterns of individual cyst species are being comparedwith environmental factors that are knownto influence dinoflagellate growth, gamete production, encystment, excystment and preservation of their organic-walled cysts: surface water temperature, salinity, nitrate, phosphate, chlorophyll-a concentrations and bottom water oxygen concentrations. Graphs are provided for every species depicting the relationship between seasonal and annual variations of these parameters and the relative abundance of the species. Results have been compared with previously published records; an overview of the ecological significance as well as information about the seasonal production of each individual species is presented. The relationship between the cyst distribution and variation in the aforementioned environmental parameters was analysed by performing a canonical correspondence analysis. All tested variables showed a positive relationship on the 99% confidence level. Sea-surface temperature represents the parameter corresponding to the largest amount of variance within the dataset (40%) followed by nitrate, salinity, phosphate and bottom-water oxygen concentration, which correspond to 34%, 33%, 25% and 24% of the variance, respectively. Characterisations of selected environments as well as a discussion about how these factors could have influenced the final cyst yield in sediments are included.

(Table 2) Dinoflagellate cysts in surface sediment samples of METEOR cruises M20/2, M23/1, M34/2 and M34/2

The organic walled cyst content of 41 surface sediment samples from the south-eastern South Atlantic Ocean have been studied to create a dataset that can be used for palaeoceanographic reconstructions. In order to obtain insight into which environmental factors influence the distribution of individual cyst species, the cyst associations have been compared with oceanographic characteristics of the overlying water masses, i.e. temperature, salinity, density and stratification gradients. The associations and relationships have been established by visual examination of the dataset and the multivariate ordination techniques, Detrended Correspondence Analysis and Canonical Correspondence Analysis. Special attention has been given to the factors of transport and preservation of the cysts. Five associations have been recognised as being characteristic of (1) areas influenced by coastal upwelling and/or river outflow, (2) open ocean, (3) Agulhas Current and southern Benguela Current, (4) Benguela Current and (5) Walvis Bay, shelf break area. The factors dominant in influencing either directly or indirectly the cyst distributions appear to be the stratification in the upper 50 m of the water column, nutrient concentration and seasonality. Variations in sea surface temperatures and salinities have only minor effect on cyst distribution.

Pollen and dinoflagellate cyst counts on sediment core M72/5_628-1 (MD72/5-25-GC1) from the Black Sea

High-resolution pollen and dinoflagellate cyst records from sediment core M72/5-25-GC1 were used to reconstruct vegetation dynamics in northern Anatolia and surface conditions of the Black Sea between 64 and 20 ka BP. During this period, the dominance of Artemisia in the pollen record indicates a steppe landscape and arid climate conditions. However, the concomitant presence of temperate arboreal pollen suggests the existence of glacial refugia in northern Anatolia. Long-term glacial vegetation dynamics reveal two major arid phases ~64-55 and 40-32 ka BP, and two major humid phases ~54-45 and 28-20 ka BP, correlating with higher and lower summer insolation, respectively. Dansgaard-Oeschger (D-O) cycles are clearly indicated by the 25-GC1 pollen record. Greenland interstadials are characterized by a marked increase in temperate tree pollen, indicating a spread of forests due to warm/wet conditions in northern Anatolia, whereas Greenland stadials reveal cold and arid conditions as indicated by spread of xerophytic biomes. There is evidence for a phase lag of ~500 to 1500 yr between initial warming and forest expansion, possibly due to successive changes in atmospheric circulation in the North Atlantic sector. The dominance of Pyxidinopsis psilata and Spiniferites cruciformis in the dinocyst record indicates brackish Black Sea conditions during the entire glacial period. The decrease of marine indicators (marine dinocysts, acritarchs) at ~54 ka BP and increase of freshwater algae (Pediastrum, Botryococcus) from 32 to 25 ka BP reveals freshening of the Black Sea surface water. This freshening is possibly related to humid phases in the region, to connection between Caspian Sea and Black Sea, to seasonal freshening by floating ice, and/or to closer position of river mouths due to low sea level. In the southern Black Sea, Greenland interstadials are clearly indicated by high dinocyst concentrations and calcium carbonate content, as a result of an increase in primary productivity. Heinrich events show a similar impact on the environment in the northern Anatolia/Black Sea region as Greenland stadials.

(Appendix B) Dinoflagellate cysts in Southeast Pacific Ocean surface sediments

orty-eight surface sediment samples from the southeast (SE) Pacific (25-53°S) are investigated for the determination of the spatial distribution of organic-walled dinoflagellate cysts along the western South American continental margin. Fifty-five different taxa are recorded and reflect oceanic or coastal assemblages. The oceanic assemblages are characterised by low cyst concentrations and the dominance of autotrophs, while the coastal assemblages generally contain a higher number of cysts, which are mainly produced by heterotrophic species. Highest cyst concentrations are observed in the active upwelling system offshore Concepción (35-37°S). Brigantedinium spp., Echinidinium aculeatum, Echinidinium granulatum/delicatum and cysts of Protoperidinium americanum dominate assemblages related to upwelling. Echinidinium aculeatum appears to be the best indicator for the presence of all year round active upwelling cells. Other protoperidinioid cysts may also occur in high relative abundances in coastal regions outside active upwelling systems, if the availability of nutrients, co-responsible for the presence/absence of their main food sources such as diatoms and other protists, is sufficient. The importance of nutrient availability as a determining environmental variable influencing cyst signals on a regional scale (SE Pacific) is demonstrated through statistical analyses of the data. Because of the importance of nutrients, uncertainties about the outcomes of quantitative sea-surface temperature (SST) reconstructions (Modern Analogue Technique) based on dinoflagellate cysts may arise, since no interaction between different hydrographical variables is considered in this approach. The combination of the SE Pacific surface sample dataset with other published cyst data from the Southern Hemisphere resulted in a database which includes 350 samples: the 'SH350 database'. This database is used to test the accuracy of the quantitative reconstructions by calculating and comparing the estimated versus observed values for each site. An attempt to perform quantitative SST reconstructions on the last 25 cal ka of site ODP1233 (41°S; 74°27'W) is made and again stresses the importance of other environmental variables such as nutrient availability in determining the dinoflagellate cyst assemblages.

Dinocyst assemblages of site HM52-43

Past sea-surface conditions over the northern North Atlantic during the last glacial maximum were examined from the study of 61 deep-sea cores. The last glacial maximum time slice studied here corresponds to an interval between Heinrich layers H2 and H1, and spanning about 20-16 ka on a 14C time scale. Transfer functions based on dinocyst assemblages were used to reconstruct sea-surface temperature, salinity, and sea-ice cover. The results illustrate extensive sea-ice cover along the eastern Canadian margins and sea-ice spreading, only during winter, over most of the northern North Atlantic. On the whole, much colder winter prevailed, despite relatively mild conditions in August (10-15°C at most offshore sites), thus suggesting a larger seasonal contrast of temperatures than today. Lower salinity than at present is reconstructed, especially along the eastern Canadian and Scandinavian margins, likely because of meltwater supply from the surrounding ice sheets. These reconstructions contrast with those established by CLIMAP on the basis of planktonic foraminifera. These differences are discussed with reference to the stratigraphical frame of the last glacial maximum, which was not the coldest phase of the last glacial stage. The respective significance of dinocyst and foraminifer records is also examined in terms of the thermohaline characteristics of surface waters and the vertical structure of upper water masses, which was apparently much more stratified than at present in the northern North Atlantic, thus preventing deep-water formation.