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.

New dinoflagellate cyst species from the upper Callovian-lower Oxfordian Rigaudella aemula zone, Timor sea, northwestern Australia

Four new species of dinoflagellate cysts are described from Callovian to lower Oxfordian (Jurassic) sediments of the Timor Sea, northwestern Australia. These comprise Evansia? lacryma, Egmontodinium elongatum, Leptodinium? ancoralium, and Nannoceratopsis reticulata. They are rare to common constituents of the Rigaudella aemula dinoflagellate cyst Interval Zone, and may prove useful for regional biostratigraphic correlation. (c) 2005 Elsevier B.V. All rights reserved.

Diversity of Symbiodinium dinoflagellate symbionts from the Indo-Pacific sea slug Pteraeolidia ianthina (Gastropoda : Mollusca)

The aeolid nudibranch Pteraeolidia ianthina hosts symbiotic dinoflagellates in the same way as many reef-building corals. This widespread Indo-Pacific sea slug ranges from tropical to temperate waters, and offers a unique opportunity to examine a symbiosis that occurs over a large latitudinal gradient. We used partial 28S and 18S nuclear ribosomal (nr) DNA to examine the genetic diversity of the Symbiodinium dinoflagellates contained within F ianthina. We detected Symbiodinium from genetic clades A, B, C and D. P. ianthina from tropical regions (Singapore, Sulawesi) host Symbiodinium clade C or D or both; those from the subtropical eastern Australian coast (Heron Island, Mon Repo, Moreton Bay, Tweed Heads) host Symbiodinium clade C, but those from the temperate southeastern Australian coastline (Port Stephens, Bare Island) host clade A or B or both. The Symbiodinium populations within 1 individual nudibranch could be homogeneous or heterogeneous at inter- or intra-clade levels (or both). Our results suggested that the Pteraeolidia-Symbiodinium symbiosis is flexible and favours symbiont phylotypes best adapted for that environment. This flexibility probably reflects the function of the symbiont clade in relation to the changing environments experienced along the latitudinal range, and facilitates the large geographic range of P. ianthina.

Transcriptome analysis of a cnidarian-dinoflagellate mutualism reveals complex modulation of host gene expression

Background: Cnidarian - dinoflagellate intracellular symbioses are one of the most important mutualisms in the marine environment. They form the trophic and structural foundation of coral reef ecosystems, and have played a key role in the evolutionary radiation and biodiversity of cnidarian species. Despite the prevalence of these symbioses, we still know very little about the molecular modulators that initiate, regulate, and maintain the interaction between these two different biological entities. In this study, we conducted a comparative host anemone transcriptome analysis using a cDNA microarray platform to identify genes involved in cnidarian - algal symbiosis. Results: We detected statistically significant differences in host gene expression profiles between sea anemones ( Anthopleura elegantissima) in a symbiotic and non-symbiotic state. The group of genes, whose expression is altered, is diverse, suggesting that the molecular regulation of the symbiosis is governed by changes in multiple cellular processes. In the context of cnidarian dinoflagellate symbioses, we discuss pivotal host gene expression changes involved in lipid metabolism, cell adhesion, cell proliferation, apoptosis, and oxidative stress. Conclusion: Our data do not support the existence of symbiosis- specific genes involved in controlling and regulating the symbiosis. Instead, it appears that the symbiosis is maintained by altering expression of existing genes involved in vital cellular processes. Specifically, the finding of key genes involved in cell cycle progression and apoptosis have led us to hypothesize that a suppression of apoptosis, together with a deregulation of the host cell cycle, create a platform that might be necessary for symbiont and/or symbiont-containing host cell survival. This first comprehensive molecular examination of the cnidarian - dinoflagellate associations provides critical insights into the maintenance and regulation of the symbiosis.

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.