The Cenozoic Gonyaulacacean Dinoflagellate Genera Operculodinium Wall, 1967 and Protoceratium Bergh, 1881 and Their Phylogenetic Relationships

The palynology of Ocean Drilling Program Site 1007, leeward of the present Bahamas Bank, provides insights into upper Oligocene–lower Pleistocene dinoflagellate cyst associations in the tropical Americas. These associations are reviewed along with the sedimentary paleoenvironment to provide context for a morphological study of the cystdefined dinoflagellate Operculodinium bahamense and its comparison with the thecadefined dinoflagellate Protoceratium reticulatum which produces a cyst assignable to the cyst-defined genus Operculodinium. Detailed reconstructions of the tabulation in both species reveal strong similarities, having a sexiform hyposomal tabulation and L-type or modified L-type ventral organization. Protoceratium reticulatum has dextral torsion of the hypotheca, requiring assignation of the genus to the subfamily Cribroperidinioideae, whereas Operculodinium bahamense has neutral torsion requiring assignation to the subfamily Leptodinioideae. Results either imply polyphyletic origins for the genus Operculodinium or that combinations of ventral organization and torsion cannot always be applied rigidly to subdivide the family Gonyaulacaceae.

Mid-Pliocene palynomorph record of the North Atlantic

The mid-Pliocene was an episode of prolonged global warmth and strong North Atlantic thermohaline circulation, interrupted briefly at circa 3.30 Ma by a global cooling event corresponding to marine isotope stage (MIS) M2. Paleoceanographic changes in the eastern North Atlantic have been reconstructed between circa 3.35 and 3.24 Ma at Deep Sea Drilling Project Site 610 and Integrated Ocean Drilling Program Site 1308. Mg/Ca ratios and d18O from Globigerina bulloides are used to reconstruct the temperature and relative salinity of surface waters, and dinoflagellate cyst assemblages are used to assess variability in the North Atlantic Current (NAC). Our sea surface temperature data indicate warm waters at both sites before and after MIS M2 but a cooling of ~2-3°C during MIS M2. A dinoflagellate cyst assemblage overturn marked by a decline in Operculodinium centrocarpum reflects a southward shift or slowdown of the NAC between circa 3.330 and 3.283 Ma, reducing northward heat transport 23-35 ka before the global ice volume maximum of MIS M2. This will have established conditions that ultimately allowed the Greenland ice sheet to expand, leading to the global cooling event at MIS M2. Comparison with an ice-rafted debris record excludes fresh water input via icebergs in the northeast Atlantic as a cause of NAC decline. The mechanism causing the temporary disruption of the NAC may be related to a brief reopening of the Panamanian Gateway at about this time.

Dinoflagellate cyst assemblages, oxygen isotope records of foraminifera, and alkenone and Mg/Ca-based SST estimates for the time interval 3.40-3.18 Ma (Late Pliocene) from five North Atlantic and Caribbean sediment cores

The early Late Pliocene (3.6 to ~3.0 million years ago) is the last extended interval in Earth's history when atmospheric CO2 concentrations were comparable to today's and global climate was warmer. Yet a severe global glaciation during marine isotope stage (MIS) M2 interrupted this phase of global warmth ~3.30 million years ago, and is seen as a premature attempt of the climate system to establish an ice-age world. Our geochemical and palynological records from five marine sediment cores along a Caribbean to eastern North Atlantic transect show that increased Pacific-to-Atlantic flow via the Central American Seaway weakened the North Atlantic Current (NAC) and attendant northward heat transport prior to MIS M2. The consequent cooling of the northern high latitude oceans permitted expansion of the Greenland ice sheet during MIS M2, despite near-modern atmospheric CO2 concentrations. Before and after MIS M2, heat transport via the NAC was crucial in maintaining warm climates comparable to those predicted for the end of this century.

Late Pliocene and Early Pleistocene dinoflagellate cysts and sea surface temperture reconstruction for several IODP and DSDP sites in the North Atlantic

In an attempt to document the palaeoecological affinities of individual extant and extinct dinoflagellate cysts, Late Pliocene and Early Pleistocene dinoflagellate cyst assemblages have been compared with geochemical data from the same samples. Mg/Ca ratios of Globigerina bulloides were measured to estimate the spring-summer sea-surface temperatures from four North Atlantic IODP/DSDP sites. Currently, our Pliocene-Pleistocene database contains 204 dinoflagellate cyst samples calibrated to geochemical data. This palaeo-database is compared with modern North Atlantic and global datasets. The focus lies in the quantitative relationship between Mg/Ca-based (i.e. spring-summer) sea-surface temperature (SSTMg/Ca) and dinoflagellate cyst distributions. In general, extant species are shown to have comparable spring-summer SST ranges in the past and today, demonstrating that our new approach is valid for inferring spring-summer SST ranges for extinct species. For example, Habibacysta tectata represents SSTMg/Ca values between 10° and 15°C when it exceeds 30% of the assemblage, and Invertocysta lacrymosa exceeds 15% when SSTMg/Ca values are between 18.6° and 23.5°C. However, comparing Pliocene and Pleistocene SSTMg/Ca values with present day summer values for the extant Impagidinium pallidum suggests a greater tolerance of higher temperatures in the past. This species occupies more than 5% of the assemblage at SSTMg/Ca values of 11.6-17.9°C in the Pliocene and Pleistocene, whereas present day summer SSTs are around -1.7 to 6.9°C. This observation questions the value of Impagidinium pallidum as reliable indicator of cold waters in older deposits, and may explain its bipolar distribution.

Distribution of dinoflagellate cycst at ODP Site 123-765 (Table 1)

Site 765 contains a sequence of tropical, middle Miocene to Holocene dinoflagellate cysts. These diverse assemblages are characterized by abundant Polysphaeridium zoharyi and Spiniferites bulloideus. Abundances of Impagidinium spp. and Nematosphaeridium spp. reflect the shelf-to-slope origin of the assemblages. One new genus, Blysmatodinium, and two new species, Nematosphaeridium (?) wrennii sp. nov. and Blysmatodinium argoi, are described.

(Table 1) Species abundance at DSDP Hole 63-467

Neogene palynofloras of southern California have been all too infrequently studied. Previous investigations of Pacific Coast sediments have been largely restricted to Pacific Northwest locales. Some important studies include those by Gray (1964), Wolfe, Hopkins, and Leopold (1966), Wolfe and Leopold (1967), Hopkins (1968), Piel (1969, 1977), Ballog, Sparks, and Waloweek (1972), and Musich (1973). The only published study of southern California materials is that of Heusser (1978) on Holocene sediments of the Santa Barbara basin. Most of these studies are concerned with the microflora from a particular formation; thus they have limited stratigraphic value and in most cases involve nonmarine to marginal marine rocks where no planktonic zonation was available. Musich's (1973) study was the first attempt at tying pollen assemblages to a planktonic zonation over an extended stratigraphic interval (Miocene to Pleistocene).Its location in the southern California Borderland and the sedimentary sections sampled make Leg 63 extremely valuable in deciphering the palynologic history of the Pacific Coast Neogene. Site 467 was chosen for our initial detailed study, because the relatively slow sedimentation rate provides an almost complete Neogene sequence of mainly terrigenous sediments and reliable planktonic age control is available.The goals of this study were to: (1) establish a reference section of Neogene palynomorph assemblages; (2) develop biostratigraphic criteria for use in correlation with other localities; (3) correlate the palynologic assemblages with the planktonic zonations; and (4) study the paleoenvironmental history in the southern California Neogene.

Dinoflagellate and other palynomorph abundance in sediments from ODP Leg 189 sites

At Ocean Drilling Program (ODP) Leg 189 Sites 1170-1172, the climatologically critical Eocene-Oligocene (E-O) transition is barren of any calcareous microfossils but contains rich marine organic walled dinoflagellate cyst (dinocyst) and diatom assemblages, suitable for detailed biostratigraphic and paleoenvironmental analysis. The resulting first-ever integrated dinocyst/diatom magnetostratigraphy allows confident correlation of the E-O interval between all Leg 189 sites, including Site 1168. Our correlations indicate that the (deep) opening of the Tasmanian Gateway occurred quasi-synchronously throughout the Tasmanian region, starting at ~35.5 Ma. At Sites 1170-1172, quantitatively, three distinct dinocyst assemblages may be distinguished that reflect the relatively rapid and pronounced stepwise environmental changes associated with the E-O transition in the Tasmanian region, from a pro-deltaic setting to a deep marine pelagic setting. Moreover, synchronous with the deepening of the gateway, at the southern and eastern Sites 1170-1172, typical endemic Antarctic assemblages were replaced by more cosmopolitan dinocyst communities. In marked contrast, at Site 1168 off western Tasmania, endemic Antarctic taxa are virtually absent during the E-O transition. At Sites 1170-1172, the endemic Antarctic dinocyst assemblage (Transantarctic Flora) drastically changes into a more cosmopolitan assemblage at ~35.5 Ma, with a more offshore character, reflecting the arrival of different oceanographic and environmental conditions associated with the deepening of the Tasmanian Gateway. In turn, this assemblage grades at ~34 Ma into one more typical for even more offshore and/or upwelling conditions at Site 1172. In slightly younger deposits at all sites, organic microfossils are virtually absent, reflecting winnowing and oxidation, indicative of a next step of oceanographic development. This phase may be dated as close to the Oceanic Anoxic (Oi)-1 18O (Antarctic glaciation) event (~33.3 Ma). In a single productive sample from the earliest Oligocene the northern Site 1172, a relatively warm-water cosmopolitan assemblage has been recovered. This aspect contrasts findings from coeval deposits from the Ross Sea, where endemic Antarctic species remain dominant. Somewhere between the paleogeographic positions of Site 1172 and the Ross Sea, a strong differentiation of surface waters occurred in the earliest Oligocene, possibly reflecting the onset of the Antarctic Circumpolar Current.

Radiocarbon dates and dinoflagellate cysts of sediment core DP30PC

To obtain insight into character and potential forcing of short-term climatic and oceanographic variability in the southern Italian region during the "Roman Classical Period" (60 BC-AD 200), climatic and environmental reconstructions based on a dinoflagelate cyst record from a well dated site in the Gulf of Taranto located at the distal end of the Po-river discharge plume have been established with high temporal resolution. Short-term fluctuations in accumulation rates of the Adriatic Surface Water species Lingulodinium machaerophorum, the freshwater algae Concentricystes and species resistant to aerobic degradation indicate that fluctuations in the trophic state of the upper waters are related to river discharge of northern and eastern Italian rivers which in turn are strongly related to precipitation in Italy. The dinoflagellate cyst association indicates that local sea surface temperatures which in this region are strongly linked to local air temperatures were slightly higher than today. We reconstruct that sea surface temperatures have been relatively high and stable between 60 BC-AD 90 and show a decreasing trend after AD 90. Fluctuations in temperature and river discharge rates have a strong cyclic character with main cyclicities of 7-8 and 11 years. We argue that these cycles are related to variations of the North Atlantic Oscillation climate mode. A strong correlation is observed with global variation in Delta14C anomalies suggesting that solar variability might be one of the major forcings of the regional climate. Apart from cyclic climate variability we observed a good correlation between non-cyclic temperature drops and global volcanic activity indicating that the latter forms an additional major forcing factor of the southern Italian climate during the Roman Classical Period.

Dinoflagellate cysts abundance and events of ODP Hole 189-1168A sediments

Palynomorphs were studied in samples from Ocean Drilling Program (ODP) Leg 189, Hole 1168A (slope of the western margin of Tasmania; 2463 m water depth). Besides organic-walled dinoflagellate cysts (dinocysts), broad categories of other palynomorphs were quantified in terms of relative abundance. In this contribution, we provide an overview of the early late Eocene-Quaternary dinocyst distribution and illustrate main trends in palynomorph distribution. Dinocyst species throughout Hole 1168A are largely cosmopolitan with important contributions of typical low-latitude taxa and virtual absence of endemic Antarctic taxa. Dinocyst stratigraphic distribution broadly matches that known from the Northern Hemisphere and equatorial regions, although significant differences are noted. Selected potentially biochronostratigraphically useful events are summarized. The distribution of dinocysts in the middle-upper Miocene interval is rather patchy, probably due to prolonged exposure to oxygen. An important general aspect in the dinocyst assemblages is the near absence of Antarctic endemic species and the apparent influence of relatively warm waters throughout the succession at Site 1168. General palynomorph distribution indicates continued deepening from an initial shallow, even restricted, marine setting from late Eocene-Quaternary times. A curious massive influx of small skolochorate acritarchs is recorded throughout the late early-early middle Miocene; the significance of this signal is not yet understood. A general long-term oligotrophic nature of the surface waters influencing Site 1168 is suggested from the low abundance of (proto) peridinioid, presumably heterotrophic, species. The overall dinocyst distribution pattern corresponds to the long-term existence of a Leeuwin-like current influencing the region, including Site 1168, confirming results of earlier studies on other microfossil groups. The occasional influence of colder surface water conditions is, however, also apparent, notably during the late Pliocene-Quaternary, indicating the potential of high-resolution dinocyst analysis for future paleoceanographic studies.

Upper Cenozoic dinoflagellate cysts from the continental slope and rise off New Jersey

This report contains the occurrence data for dinoflagellate cysts recorded from 163 samples taken from Sites 902 through 906, during Ocean Drilling Program (ODP) Leg 150. The dinoflagellate cyst (dinocyst) stratigraphy has been presented in Mountain, Miller, Blum, et al. (1994, doi:10.2973/odp.proc.ir.150.1994), and was based on these data. This report provides the full dinocyst data set supporting the dinocyst stratigraphic interpretations made in Mountain, Miller, Blum, et al. (1994). For Miocene shipboard dinocyst stratigraphy, I delineated 10 informal zones: pre-A, and A through I, in ascending stratigraphic order. These zones are defined in Shipboard Scientific Party (1994a, doi:10.2973/odp.proc.ir.150.103.1994), and are based on my studies of Miocene dinocyst stratigraphy in the Maryland and Virginia coastal plain (de Verteuil and Norris, 1991, 1992; de Verteuil, 1995). This zonation has been slightly revised (de Verteuil and Norris, 1996), and the new formal zone definitions are repeated below. Each new zone has an alpha-numeric abbreviation starting with "DN" (for Dinoflagellate Neogene). The equivalence between the informal zones reported in Mountain, Miller, Blum, et al. (1994), and the new DN zones is illustrated in Figure 1. For clarity, I delineated both zonations in the range charts that accompany this report (Tables 1-6). De Verteuil and Norris (1996a), using these and other data, correlated the DN zonation with the geological time scale of Berggren et al. (1995). Figure 2 summarizes these correlations and can be used to check the chronostratigraphic position of samples in this report, as determined by dinocyst stratigraphy. A thorough discussion of the basis for, and levels of uncertainty associated with, these correlations to the Cenozoic time scale can be found in de Verteuil and Norris (1996a). The Appendix lists all the dinocyst taxa recorded during shipboard analyses of Leg 150 samples. Open nomenclature is used for undescribed taxa. The range charts and Appendix also include reference to several new taxa that de Verteuil and Norris (1996b) described from Miocene coastal plain strata in Maryland and Virginia. Names of these taxa in Tables 1 through 6 and in the Appendix of this report are not intended for effective publication as defined in the International Code of Botanical Nomenclature (ICBN, Greuter et al., 1994). Therefore, taxonomic nomenclature contained in this report is not to be treated as meeting the conditions of effective and valid publication (ICBN; Article 29).

Late Quateranry dinoflagellate cysts of ODP Site 202-1233

The late Quaternary organic-walled dinoflagellate cyst record of Site 1233 (41°S, offshore Chile) was studied with a ?200 year resolution spanning the last 25,000 years. The study provides the first continuous record of sub-recent and recent dinoflagellate cysts in the Southeast (SE) Pacific. Major changes in the composition of the cyst association, cyst concentration and morphology of Operculodinium centrocarpum reflect changes in sea surface temperature (SST), sea surface salinity (SSS), palaeoproductivity and upwelling intensity. These changes can be associated with latitudinal shifts of the circumpolar frontal systems. The high cyst concentration, high Brigantedinium spp. abundances, low species diversity and the occurrence of certain cold water species are supportive for a 7-10° equatorward shift of the Antarctic Circumpolar Current (ACC) during the coldest phase of the Last Glacial Maximum (LGM) between 25 and 21.1 cal ka BP. Deglacial warming initiated at ~18.6 cal ka BP. Termination I (18.6-11.1 cal ka BP) is interrupted by an unstable period of extreme seasonality, rather than a cooling event, between 14.4 and 13.2 cal ka BP, synchronous with the Antarctic Cold Reversal (ACR). The Holocene Maximum is observed between 11.6 and 9.8 cal ka BP and is typified by the most southward position of the northern margin of the ACC. A cooling phase occurred during the early Holocene (until ~7 cal ka BP) and during the last ~0.8 ka. Our data indicates that the SE Pacific (41°S) climate has been influenced over the whole record by changes in the Southern Hemisphere (SH) high-latitudes, while during the mid to late Holocene, also a tropical forcing mechanism was involved, including the El Niño Southern Oscillation and the variable Hadley cell intensity. Furthermore, this study showed a relationship between the variable morphology of the spines/processes of O. centrocarpum and the combined variation of sea surface salinity and temperature (SSS/SST-ratio).

An early Eocene carbon cycle perturbation at in the Southern Alps

At least two transient events of extreme global warming occurred superimposed on the long-term latest Paleocene and early Eocene warming trend in the Paleocene-Eocene thermal maximum (PETM) (or ETM1 ~55.5 Ma) and the Elmo (or ETM2 ?53.6 Ma). Other than warmth, the best known PETM is characterized by (1) significant injection of 13C-depleted carbon into the ocean-atmosphere system, (2) deep-sea carbonate dissolution, (3) strong biotic responses, and (4) perturbations of the hydrological cycle. Documentation of the other documented and suspected "hyperthermals" is, as yet, insufficient to assess whether they are similar in nature to the PETM. Here we present and discuss biomagnetostratigraphic data and geochemical records across two lower Eocene successions deposited on a continental margin of the western Tethys: the Farra and Possagno sections in the Venetian pre-Alps. We recognize four negative carbon isotope excursions within chron C24. Three of these shifts correlate to known or suspected hyperthermals: the PETM, the Eocene thermal maximum 2 (~53.6 Ma), and the informally named "X event" (~52.5 Ma). The fourth excursion lies within a reverse subchron and occurred between the latter two. In the Farra section, the X event is marked by a ~0.6 per mil negative carbon isotope excursion and carbonate dissolution. Furthermore, the event exhibits responses among calcareous nannofossils, planktic foraminifera, and dinoflagellates that are similar to, though less intense than, those observed across the PETM. Sedimentological and quantitative micropaleontological data from the Farra section also suggest increased weathering and runoff as well as sea surface eutrophication during this event.

Organic-walled dinoflagellate cysts analysis and statistical analysis of 116 surface sediment samples from the eastern Indian Ocean

Assemblages of organic-walled dinoflagellate cysts (dinocysts) from 116 marine surface samples have been analysed to assess the relationship between the spatial distribution of dinocysts and modern local environmental conditions [e.g. sea surface temperature (SST), sea surface salinity (SSS), productivity] in the eastern Indian Ocean. Results from the percentage analysis and statistical methods such as multivariate ordination analysis and end-member modelling, indicate the existence of three distinct environmental and oceanographic regions in the study area. Region 1 is located in western and eastern Indonesia and controlled by high SSTs and a low nutrient content of the surface waters. The Indonesian Throughflow (ITF) region (Region 2) is dominated by heterotrophic dinocyst species reflecting the region's high productivity. Region 3 is encompassing the area offshore north-west and west Australia which is characterised by the water masses of the Leeuwin Current, a saline and nutrient depleted southward current featuring energetic eddies.