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.

Cretaceous dinoflagellate cysts of ODP Leg 103 holes

The dinoflagellate cyst assemblages in 42 samples collected from Sites 638 and 639 were analyzed. All samples from Site 639 are barren; relatively poor assemblages occur in samples from Site 638. The distribution of 61 dinoflagellate cyst taxa identified in samples from Holes 638B and 638C are tabulated. The assemblages from Site 638 are comparable to those of Tethyian stratotypes and from neighboring areas, which permits age determinations and correlations between Holes 638B and 638C. The interval from Cores 103-638C-14R to 103-638C-1R is late Berriasian through Valanginian in age. In Hole 638B, the interval from Core 103-638B-43R to Section 103-638B-23R-2 is dated as early Valanginian through middle Barremian. Sections 103-638B-21R-2 and 103-638B-21R-1 are late Aptian in age. Taxonomic remarks are made about some species; a new dinoflagellate cyst Heterosphaeridiuml galiciae is described.

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.

Dinoflagellate cyst accumulation rates of ODP Hole 146-893A from Santa Barbara Basin

The dinoflagellate cyst record from Ocean Drilling Program Hole 893A, Santa Barbara Basin, southern California, is examined at millennial-scale resolution for the past 40 kyr. Changes in cyst abundance, composition of cyst assemblages, and their diversity reflect major shifts in climate and ocean circulation in the region over this time interval. Throughout the sequence, dinoflagellate cyst assemblages are dominated by heterotrophic dinoflagellates. Brigantedinium spp. and other upwelling-related taxa such as Echinidinium and Protoperidinium americanum are abundant, indicating the continued influence of coastal upwelling on the basin during the late Quaternary. A significant increase in cyst accumulation rates is seen during the Holocene and, to a lesser extent, during shorter warming events such as Bolling/Allerod and Dansgaard-Oeschger interstadials, implying enhanced marine productivity during these periods. Cyst diversity is high during the Holocene. An increase in abundance of cysts produced by autotrophic dinoflagellates in the late Holocene suggests enhanced input of warm, nutrient-rich waters. In contrast, cyst assemblages from the Last Glacial Maximum exhibit a relatively low diversity and an increase in the cysts of heterotrophic dinoflagellates, in particular Selenopemphix nephroides. The presence of this taxon in association with Brigantedinium spp. implies substantial cooling of surface waters in the Santa Barbara Basin at that time.

Abundance of palynomorphs and dinoflagellate stratigraphy of ODP Hole 184-1148A, South China Sea

Dinoflagellate stratigraphy is described for the section from 364.75 to 843.85 meters below seafloor (mbsf) at Site 1148 (Sections 184-1148A-40X-1 through 76X-6 and 184-1148B-39X-CC through 56X-1) in the South China Sea. Two assemblage zones and two subzones are defined, based on characteristics of the assemblages and lowest/highest occurrences of some key species. These are the Cleistosphaeridium diversispinosum Assemblage Zone (Zone A; Oligocene), with the Enneadocysta pectiniformis Subzone (Subzone A-1) and the Cordosphaeridium gracile Subzone (Subzone A-2), and the Polysphaeridium zoharyi Assemblage Zone (Zone B; early Miocene). The highest concurrent occurrence of Enneadocysta arcuata, Eneadocysta multicornuta, Homotryblium plectilum, and Homotryblium tenuispinosum delineates the upper boundary of Zone A, which appears to mark a hiatus. Subzone A-1 is of early Oligocene age, as evidenced by the highest occurrences of E. pectiniformis and Phthanoperidinium amoenum at the upper boundary of the subzone. Subzone A-2 is of late Oligocene age based on the highest occurrences of C. gracile and Wetzeliella gochtii close to the upper boundary of the subzone and the occurrence of Distatodinium ellipticum and Membranophoridium aspinatum within the subzone. Zone B is dated as early Miocene based on the lowest occurrences of Cerebrocysta satchelliae, Hystrichosphaeropsis obscura, Melitasphaeridium choanophorum, Membranilarnacia? picena, and Tuberculodinium vancampoae within the zone. The present assemblage zones/subzones are correlative to various degrees with coeval zones/assemblages from areas of high to low latitudes in terms of common key species. We have compared the species content of the assemblage Zones A and B, and the subzones A-1 and A-2, with coeval assemblage(s)/zone(s) described from many, often widely distant, high- and low-latitude regions of the world. These comparisons show that, to various degrees and aside from a number of key species, the coordinated presence of certain important species may also help to assign an age to a given assemblage.

(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.

Absolute abundances of dinoflagellate cysts in surface sediment samples from four sites: Lycopodium marker-grain method

Absolute abundances (concentrations) of dinoflagellate cysts are often determined through the addition of Lycopodium clavatum marker-grains as a spike to a sample before palynological processing. An inter-laboratory calibration exercise was set up in order to test the comparability of results obtained in different laboratories, each using its own preparation method. Each of the 23 laboratories received the same amount of homogenized splits of four Quaternary sediment samples. The samples originate from different localities and consisted of a variety of lithologies. Dinoflagellate cysts were extracted and counted, and relative and absolute abundances were calculated. The relative abundances proved to be fairly reproducible, notwithstanding a need for taxonomic calibration. By contrast, excessive loss of Lycopodium spores during sample preparation resulted in non-reproducibility of absolute abundances. Use of oxidation, KOH, warm acids, acetolysis, mesh sizes larger than 15 µm and long ultrasonication (> 1 min) must be avoided to determine reproducible absolute abundances. The results of this work therefore indicate that the dinoflagellate cyst worker should make a choice between using the proposed standard method which circumvents critical steps, adding Lycopodium tablets at the end of the preparation and using an alternative method.

(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.

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.

Degradation rates and productivity signals of organic-walled dinoflagellate cyst species in surface sediment samples

To understand the role of the ocean within the global carbon cycle, detailed information is required on key-processes within the marine carbon cycle; bio-production in the upper ocean, export of the produced material to the deep ocean and the storage of carbon in oceanic sediments. Quantification of these processes requires the separation of signals of net primary production and the rate of organic matter decay as reflected in fossil sediments. This study examines the large differences in degradation rates of organic-walled dinoflagellate cyst species to separate these degradation and productivity signals. For this, accumulation rates of cyst species known to be resistant (R-cysts) or sensitive (S-cysts) to aerobic degradation of 62 sites are compared to mean annual chlorophyll-a, sea-surface temperature, sea-surface salinity, nitrate and phosphate concentrations of the upper waters and deep-water oxygen concentrations. Furthermore, the degradation of sensitive cysts, as expressed by the degradation constant k and reaction time t, has been related to bottom water [O2]. The studied sediments were taken from the Arabian Sea, north-western African Margin (North Atlantic), western-equatorial Atlantic Ocean/Caraibic, south-western African margin (South Atlantic) and Southern Ocean (Atlantic sector). Significant relationships are observed between (a) accumulation rates of R-cysts and upper water chlorophyll-a concentrations, (b) accumulation rates of S-cysts and bottom water [O2] and (c) degradation rates of S-cysts (kt) and bottom water [O2]. Relationships that are extremely weak or are clearly insignificant on all confidence intervals are between (1) S-cyst accumulation rates and chlorophyll-a concentrations, sea-surface temperature (SST), sea-surface salinity (SSS), phosphate concentrations (P) and nitrate concentrations (N), (2) between R-cyst accumulation rates and bottom water [O2], SST, SSS, P and N, and between (3) kt and water depth. Co-variance is present between the parameters N and P, N, P and chlorophyll-a, oxygen and water depth. Correcting for this co-variance does not influence the significance of the relationship given above. The possible applicability of dinoflagellate cyst degradation to estimate past net primary production and deep ocean ventilation is discussed.

The impact of fluctuating light on the dinoflagellate Prorocentrum micans depends on NO3- and CO2 availability

Increasing atmospheric pCO2 and its dissolution into oceans leads to ocean acidification and warming, which reduces the thickness of upper mixing layer (UML) and upward nutrient supply from deeper layers. These events may alter the nutritional conditions and the light regime to which primary producers are exposed in the UML. In order to better understand the physiology behind the responses to the concomitant climate changes factors, we examined the impact of light fluctuation on the dinoflagellate Prorocentrum micans grown at low (1 µmol/L) or high (800 µmol/L) [NO3(-)] and at high (1000 µatm) or low (390 µatm, ambient) pCO2. The light regimes to which the algal cells were subjected were (1) constant light at a photon flux density (PFD) of either 100 (C100) or 500 (C500) µmol/m**2/s or (2) fluctuating light between 100 or 500 µmol photons/m**2/s with a frequency of either 15 (F15) or 60 (F60) min. Under continuous light, the initial portion of the light phase required the concomitant presence of high CO2 and NO3(-) concentrations for maximum growth. After exposure to light for 3h, high CO2 exerted a negative effect on growth and effective quantum yield of photosystem II (F'(v)/F'(m)). Fluctuating light ameliorated growth in the first period of illumination. In the second 3h of treatment, higher frequency (F15) of fluctuations afforded high growth rates, whereas the F60 treatment had detrimental consequences, especially when NO3(-) concentration was lower. F'(v)/F'(m) respondent differently from growth to fluctuating light: the fluorescence yield was always lower than at continuous light at 100 µmol/m**2/s, and always higher at 500 µmol/m**2/s. Our data show that the impact of atmospheric pCO2 increase on primary production of dinoflagellate depends on the availability of nitrate and the irradiance (intensity and the frequency of irradiance fluctuations) to which the cells are exposed. The impact of global change on oceanic primary producers would therefore be different in waters with different chemical and physical (mixing) properties.