Ethological analysis of the trace fossil Zoophycos: hints from the Arctic Ocean

Löwemark, L. 2011: Ethological analysis of the trace fossil Zoophycos: Hints from the Arctic Ocean. Lethaia, Vol. 45, pp. 290–298. The distribution of the trace fossil Zoophycos in Quaternary marine sediments from the Arctic Ocean was studied in twelve piston and gravity cores retrieved during the Swedish icebreaker expeditions YMER80, Arctic Ocean‐96 and LOMROG I & II. The sampled cores span an area from the Makarov Basin to the Fram Strait. Zoophycos was only found in two cores taken at more than 2 km water depth on the slope of the Lomonosov Ridge, but was absent in cores obtained at shallower depth, confirming earlier observations of the trace maker’s bathymetric preferences. The two cores containing Zoophycos are characterized by quiet sedimentation and slightly enhanced food flux compared with the general Arctic. The occurrence of Zoophycos in these cores in a setting that is characterized by extreme seasonal variations in food flux due to the total ice coverage during winters and high primary productivity during the long summer days, is interpreted to be a cache‐behaviour response to pulsed flux of food to the benthic realm. □Arctic Ocean, ethology, Quaternary, spreiten, trace fossils, Zoophycos.     

Paleoceanography of the mid-Pleistocene

The fossil record of the Pleistocene calcareous nannoplankton indicates that during the mid-Pleistocene (0.93–1.25 my) occurred an episode of overwhelming dominance of smallGephyrocapsa. During this episode normally abundant, large size specimens of this genus (mainlyGephyrocapsa oceanica) were virtually excluded from the phytoplankton of tropical and subtropical oceans. The best modern analog of this dominantly smallGephyrocapsa assemblage is the subpolarEmiliania huxleyi assemblage, which implies that nutrient content was significantly greater and water temperature was lower in the photic water column of the tropical oceans than they are today. Increased equatorial upwelling in the oceans, on a scale much greater than today, may explain the above pattern.To achieve such broad equatorial upwelling there must be a source and a drive for cold, dense water. The Arctic Ocean, which was probably seasonally free of ice during this interval of the mid-Pleistocene, is capable of providing the requisite source as well as a drive for the inferred equatorial upwelling. The energy balance of a predominantly ice-free Arctic Ocean requires an approximately three to seven fold increase of hydrospheric heat transport from the North Atlantic to the Arctic Ocean, which dictates a corresponding or even greater increase in the volume of warm water entering the Arctic Ocean at the surface and cold dense water exiting at depth to the North Atlantic. Such enhanced dense water formation in the Arctic Ocean could drive the intensified equatorial upwelling implied by the smallGephyrocapsa dominance interval.If the above scenario is correct then the climate of the earth's northern hemisphere during the mid-Pleistocene may have been very different from the younger Pleistocene climate. One manifestation of this difference may be the mid-Pleistocene shift in climatic cycle periodicity from 40 ky to 100 ky. Another important aspect is that the enhanced greenhouse effect expected during the next century because of an increase of atmospheric CO₂ is thought to lead directly to melting of the Arctic Ocean ice cover and of the Greenland ice sheet. Thus, the “greenhouse” Arctic Ocean and its attendant ocean circulation would resemble the inferred mid-Pleistocene conditions.     

The development of planktonic biogeography in the Southern Ocean during the Cenozoic

Deep-sea drilling at high latitudes of the Southern Hemispheres has provided almost the only available data to evaluate the biogeographic development of the planktonic biota in the Southern Ocean during the Cenozoic (65 m.y. to Present Day). Paleontological investigations on Deep Sea Drilling Project (DSDP) materials have shown that the development of Cenozoic planktonic biogeography of the Southern Ocean is intimately linked with the evolution of the Southern Ocean water masses themselves. During the Cenozoic, this has included the development of the Circum-Antarctic Current system as obstructing land masses moved apart, the refrigeration and later extensive glaciation of the continent, and the development of the Antarctic Convergence (Polar Front) with related oceanic upwelling.Almost all evolution of calcareous planktonic microfossils has occurred outside of the Antarctic—Subantarctic region followed by limited migration into these water masses. Virtually no endemism occurs amongst calcareous microfossil groups at these latitudes. In contrast, conspicuous and widespread evolution has occurred within the siliceous microfossil groups especially during the Neogene. Low diversity and differences in stratigraphic ranges of Antarctic calcareous microfossils makes them only broadly useful for correlation. Relatively higher diversities within the Subantarctic provide a firmer basis for more detailed correlation, although the ranges of fossils are often different than at lower latitudes because of different paleoceanographic and paleoclimatic controls. Within the Antarctic water mass south of the Antarctic Convergence, siliceous microfossilsbiostratigraphy, oxygen isotopic stratigraphy and magnetostratigraphy, provide the only firm basis for correlation with low-latitude sequences.Eocene (55-38 Ma) sediments contain abundant calcareous microfossils even closely adjacent to the continent. Antarctic calcareous planktonic microfossils of this age exhibit relative high diversity, although this is lower than assemblages of equivalent age at middle and low latitudes. Within the Subantarctic region, Eocene planktonic foraminifera exhibit strong affinities with those in the temperate regions. Biogeographic differences exist between various sectors of the Southern Ocean related to biogeographic isolation preceding the development of the Circum-Antarctic Current. Subantarctic calcareous nannofossil assemblages of Paleocene and Eocene age exhibit higher diversity than Oligocene and Neogene assemblages. Siliceous microfossils are poorly represented or at best poorly known.One of the most dramatic changes in Southern Ocean planktonic biogeography occurred near the Eocene/Oligocene boundary (38 Ma). Since then, Antarctic planktonic foraminiferal assemblages have exhibited distinct polar characteristics, marked in particular by low diversity, and this event thus reflects the initiation of the Antarctic faunal and floral provinces. Profound paleoceanographic changes at this time, which triggered the biogeographic crisis, appear to be related to the initiation of widespread Antarctic sea-ice formation, and rapid cooling of deep and intermediate waters, in turn associated with increased Antarctic glaciation. During the Oligocene, planktonic microfossil diversity was low in all groups throughout the world's oceans. In Antarctic waters, the early Oligocene foraminiferal fauna is monospecific (Subbotina angiporoides), while in the later Oligocene two species (S. angiporoides and Catapsydrax dissimilis) were recorded. Calcareous nannofossil assemblages are of low diversity compared with the Eocene. Subantarctic foraminiferal faunas of Oligocene age display much higher diversity than those in the Antarctic, but early and middle Oligoceae faunas still exhibit the lowest diversities for the entire Cenozoic. Siliceous assemblages remain relatively inconspicuous in most regions of the Southern Ocean.The Paleogene-Neogene transition (22 Ma) is marked by a major change in the global planktonic biogeography, i.e. modern patterns developed in which permanent, steep faunal and floral diversity gradients existed between tropical and polar regions; a gradient which has persisted even during the most severe glacial episodes. Oligocene assemblages of low diversity and almost cosmopolitan distribution were replaced by distinctive belts of planktonic assemblages arranged latitudinally from the tropics to the poles. The establishment of the steep planktonic diversity gradients and latitudinal provinces near the beginning of the Neogene almost certainly were linked to the development of the Circum-Antarctic Current in the late Oligocene which effectively separated high- and low-latitude planktonic assemblages. These fundamental global circulation and biogeographic patterns have persisted through the Neogene.During the Neogene (22 Ma to Present Day), Antarctic calcareous microfossil assemblages exhibit persistent low diversity and high dominance, while Subantarctic assemblages are of much greater diversity. The beginning of the Neogene (= beginning of Miocene) heralded the development of the high-latitude siliceous microfossil assemblages towards their present-day dominant role. Siliceous biogenec productivity began to increase. These changes were linked to the initial development and later intensification of circulation associated with the Antarctic Convergence and Antarctic Divergence. The Antarctic Convergence sharply separates dominantly siliceous assemblages to the south from calcareous assemblages to the north. Radiolarian assemblages became more endemic. Relatively warm early and middle Miocene conditions are reflected by slightly higher diversity of planktonic foraminifera and by the presence, in the northern Subantarctic, of conspicuous discoasters in early Miocene sediments. In Antarctic waters, calcareous nannofossils become unimportant as biogenic elements after the middle Miocene.The latest Miocene ( 5 m.y. ago) was marked by northward movement of the Antarctic Convergence, corresponding expansion of the Antarctic water mass, and low diversity of calcareous assemblages. Pliocene planktonic foraminifera seem to be largely monospecific in Antarctic and southern Subantarctic sequences. During the Quaternary, Antarctic waters reached a maximum northward expansion and exhibit highest siliceous biogenic productivity for the Cenozoic. In the Subantarctic, Quaternary foraminiferal diversities are much higher than in Pliocene sequences. Although calcareous nannofossil diversity may be high, only a few species are abundant. Large northward shifts of Antarctic and Subantarctic water masses have occurred during the Quaternary although no southward penetrations have occurred much beyond that of the present day. Several radiolarian and foraminiferal species disappeared or appeared at or close to a number of paleomagnetic reversals during the last 4 m.y. These faunal events, which provide valuable datums, do not seem to be associated with major climatic changes.     

Calcareous dinoflagellate cysts in South and equatorial Atlantic surface sediments: diversity,distribution, ecology and potential for palaeoenvironmental reconstruction

Several marine, peridiniphycidean dinoflagellate species produce calcareous cysts during their life cycle, which are relatively resistant to chemical and physical degradation and are therefore often found in large quantities in oceanic bottom sediments. Although the use of these calcareous cysts as proxies for palaeoenvironmental and palaeoclimatic reconstructions has seen many advances over the last decade, until now only relatively patchy and regional information was available on their recent distribution patterns and ecology, especially at the species level. In this paper, comprehensive calcareous cyst diversity and distribution data have been compiled from published and unpublished work for 167 South and equatorial Atlantic Ocean surface sediments, ranging from 20°N to 50°S, and 30°E to 65°W. The main aim has been to focus on the complex, often non-linear, relationships between individual species’ distributions and the physicochemical and trophic conditions of the overlying (sub)surface waters through the use of x–y graphs of cyst abundance vs. (sub)surface water environmental parameters, and detrended correspondence analyses. Ten cyst species and the calcareous vegetative coccoid species Thoracosphaera heimii were observed in the bottom sediments, each species showing its own characteristic distribution pattern in relation to the environmental conditions of the upper water masses above them (e.g. sea surface temperature, productivity, stratification). The sensitive reactions of various species to unique combinations of environmental parameters shows that each species has its own specific ecological traits, thus rejecting the bundled use of ‘calcareous cyst accumulation’ as a general proxy for oligotrophy or stratification in future palaeoenvironmental analyses. The acquired ‘reference’ data set of this study is large and diverse enough to allow its future application in quantitative palaeoenvironmental reconstruction models, and shows that there is still an enormous reconstruction potential concealed in many fossil calcareous dinoflagellate cyst assemblages.     

Distributions of calcareous dinoflagellate cysts in surface sediments of the western equatorial Atlantic Ocean,and their potential use in palaeoceanography

Only very few studies focus on recent calcareous dinoflagellate cyst diversity, geographic distribution and ecology, so that information on the distribution patterns and environmental affinities of individual cyst species is extremely limited. This information is, however, essential if we want to use calcareous dinoflagellate cysts for palaeoenvironmental reconstruction. Surface sediment samples from the generally oligotrophic western equatorial Atlantic Ocean, offshore northeast Brazil, were therefore quantitatively analysed for their calcareous dinoflagellate cyst content, including the calcareous vegetative coccoid Thoracosphaera heimii. Seven calcareous dinoflagellate cyst species/morphotypes and T. heimii were encountered in high concentrations throughout the area. Substantial differences in the distribution patterns were observed. The highest concentrations of cysts are found in sediments of the more oligotrophic, oceanic regions, beyond the influence of Amazon River discharge waters. Dinoflagellates producing calcareous cysts thus appear to be capable of surviving low nutrient concentrations and produce large numbers of cysts in relatively stable and predictable environments affected by minimal seasonality. To test for the environmental affinities of individual species, distribution patterns in surface sediments were compared with temperature, salinity, density and stratification gradients within the upper water column (0–100 m) over different times of the year, using principal components analysis and redundancy analysis. T. heimii and four of the seven encountered cyst species (Sphaerodinella? albatrosiana, two morphotypes of Sphaerodinella? tuberosa and Scrippsiella regalis) relate to these parameters significantly and the variations in the cyst associations appear to be associated with the different surface water currents characterising the area. The results imply that calcareous dinoflagellate cyst distributions can potentially be used to distinguish between different open oceanic environments and they could, therefore, be useful in tracing water mass movements throughout the late Quaternary.     

Bathymetric distribution of fossil foraminifera within marine sediment cores from the eastern part of Lützow-Holm Bay, East Antarctica, and its paleoceanographic implications

Faunal analysis of fossil foraminifera from marine gravity and piston cores collected by the Japanese Antarctic Research Expeditions (1981 and 1992) is used to estimate the impact of the latest Quaternary paleoceanography on coastal environments of the eastern part of Lützow-Holm Bay, East Antarctica.Accelerator Mass Spectrometry (AMS) carbon-14 ages produced from sedimentary organic carbon were less than 16 ka (non-corrected). Detailed correlation among submarine cores and Holocene elevated marine deposits exposed on the eastern shore of the embayment is difficult due to the indefinite reservoir correction value for marine organic matter and to upward-increasing abnormal ages for some cores.A local carbonate dissolution level can be delineated around the present depth of 300–400 m or shallower in the eastern part of Lützow-Holm Bay during the Holocene, based on distributional trends of arenaceous, calcareous benthic, and planktonic foraminifera recognized within a depth less than 600 m. Downcore recovery of calcareous foraminifera containing Bulimina aculeata from two cores obtained in a drowned glacial trough deeper than 600 m situated far beyond the dissolution depth of CaCO₃ indicates the incursion of warm, high-nutrient, and CaCO₃-saturated Circumpolar Deep Water (CDW) from the offshore area along the trough toward the southeastern coast of Lützow-Holm Bay during the Holocene. The intrusion of CDW impacted on the marine environments of the southeastern coast, thereby contributing to peripheral retreat of the ice sheet as well as increasing calcareous benthic foraminiferal productivity along the southeastern coast of Lützow-Holm Bay.     

Patterns and controlling factors of species diversity in the Arctic Ocean

Aim The Arctic Ocean is one of the last near‐pristine regions on Earth, and, although human activities are expected to impact on Arctic ecosystems, we know very little about baseline patterns of Arctic Ocean biodiversity. This paper aims to describe Arctic Ocean‐wide patterns of benthic biodiversity and to explore factors related to the large‐scale species diversity patterns. Location Arctic Ocean. Methods We used large ostracode and foraminiferal datasets to describe the biodiversity patterns and applied comprehensive ecological modelling to test the degree to which these patterns are potentially governed by environmental factors, such as temperature, productivity, seasonality, ice cover and others. To test environmental control of the observed diversity patterns, subsets of samples for which all environmental parameters were available were analysed with multiple regression and model averaging. Results Well‐known negative latitudinal species diversity gradients (LSDGs) were found in metazoan Ostracoda, but the LSDGs were unimodal with an intermediate maximum with respect to latitude in protozoan foraminifera. Depth species diversity gradients were unimodal, with peaks in diversity shallower than those in other oceans. Our modelling results showed that several factors are significant predictors of diversity, but the significant predictors were different among shallow marine ostracodes, deep‐sea ostracodes and deep‐sea foraminifera. Main conclusions On the basis of these Arctic Ocean‐wide comprehensive datasets, we document large‐scale diversity patterns with respect to latitude and depth. Our modelling results suggest that the underlying mechanisms causing these species diversity patterns are unexpectedly complex. The environmental parameters of temperature, surface productivity, seasonality of productivity, salinity and ice cover can all play a role in shaping large‐scale diversity patterns, but their relative importance may depend on the ecological preferences of taxa and the oceanographic context of regions. These results suggest that a multiplicity of variables appear to be related to community structure in this system.     

Retention of ice-associated amphipods: possible consequences for an ice-free Arctic Ocean

Recent studies predict that the Arctic Ocean will have ice-free summers within the next 30 years. This poses a significant challenge for the marine organisms associated with the Arctic sea ice, such as marine mammals and, not least, the ice-associated crustaceans generally considered to spend their entire life on the underside of the Arctic sea ice. Based upon unique samples collected within the Arctic Ocean during the polar night, we provide a new conceptual understanding of an intimate connection between these under-ice crustaceans and the deep Arctic Ocean currents. We suggest that downwards vertical migrations, followed by polewards transport in deep ocean currents, are an adaptive trait of ice fauna that both increases survival during ice-free periods of the year and enables re-colonization of sea ice when they ascend within the Arctic Ocean. From an evolutionary perspective, this may have been an adaptation allowing success in a seasonally ice-covered Arctic. Our findings may ultimately change the perception of ice fauna as a biota imminently threatened by the predicted disappearance of perennial sea ice.     

The (palaeo) environmental significance of modern calcareous dinoflagellate cysts: a review

Increased interest in the environmental applicability of calcareous dinoflagellate cysts for palaeo-environmental studies arose in the last two decades, when it was discovered that they constitute a widespread and abundant group within the calcareous phytoplankton. As primary producers they are directly influenced by environmental Parameters of the surrounding water masses, and therefore extremely suitable for detailed palaeo-environmental and -oceanographical reconstructions. During the last years, detailed investigations on their distribution in surface sediments of the South Atlantic Ocean, the Mediterranean Sea and the Arabian Sea were carried out, in order to reveal which environmental parameters influence their distribution. This, in turn, formed the base for palaeo-environmental and -oceanographical reconstructions. The compilation of the available data on calcareous dinoflagellate ecology, morphology, taxonomy and palaeo-environmental application presented here demonstrates their ample application potential as palaeo-environmental tools.      

Ammonia-oxidizing Archaea in the Arctic Ocean and Antarctic coastal waters

We compared abundance, distributions and phylogenetic composition of Crenarchaeota and ammonia-oxidizing Archaea (AOA) in samples collected from coastal waters west of the Antarctic Peninsula during the summers of 2005 and 2006, with samples from the central Arctic Ocean collected during the summer of 1997. Ammonia-oxidizing Archaea and Crenarchaeota abundances were estimated from quantitative PCR measurements of amoA and 16S rRNA gene abundances. Crenarchaeota and AOA were approximately fivefold more abundant at comparable depths in the Antarctic versus the Arctic Ocean. Crenarchaeota and AOA were essentially absent from the Antarctic Summer Surface Water (SSW) water mass (0–45 m depth). The ratio of Crenarchaeota 16S rRNA to archaeal amoA gene abundance in the Winter Water (WW) water mass (45–105 m depth) of the Southern Ocean was much lower (0.15) than expected and in sharp contrast to the ratio (2.0) in the Circumpolar Deep Water (CDW) water mass (105–3500 m depth) immediately below it. We did not observe comparable segregation of this ratio by depth or water mass in Arctic Ocean samples. A ubiquitous, abundant and polar-specific crenarchaeote was the dominant ribotype in the WW and important in the upper halocline of the Arctic Ocean. Our data suggest that this organism does not contain an ammonia monooxygenase gene. In contrast to other studies where Crenarchaeota populations apparently lacking amoA genes are found in bathypelagic waters, this organism appears to dominate in well-defined, ammonium-rich, near-surface water masses in polar oceans.     

Palaeoenvironmental turnover across the Ypresian–Lutetian transition at the Agost section, Southastern Spain: In search of a marker event to define the Stratotype for the base of the Lutetian Stage

Marker events to define the stratotype for the base of the Lutetian Stage are poorly defined. To elucidate such markers and characterize palaeoenvironmental turnovers, we conducted an integrated study of the Ypresian–Lutetian (Y–L; early-middle Eocene) transition at the continuous Agost section (southeastern Spain). This 115-m-thick section, which consists of hemipelagic marls intercalated with hemipelagic limestones and turbidity sandstones, spans from planktic foraminiferal Zones P9 to P12 (E7 to E10) and calcareous nannofossil Zones CP11 to CP14a (NP13 to NP16). We report quantitative analyses of planktic and benthic foraminifera and characterization of trace fossil assemblages that are integrated with mineralogical analyses.Relative to benthic forms, planktic foraminifera constitute more than 80% of the foraminiferal assemblage. We found that the most abundant planktic species belong to the genera Acarinina, Morozovella, Subbotina, and Pseudohastigerina. Benthic foraminiferal assemblages are strongly dominated by calcareous taxa, with bolivinids being the most abundant group. Trace fossils showed the succession Nereites–Zoophycos–Cruziana ichnofacies throughout the Agost section. In addition to changes in palaeobathymetry, we deduced that quantity and quality of organic matter flux influenced by turbidity currents are the main factors controlling benthic assemblages. We distinguished several mineralogical boundaries at the Agost section, each associated with lithological facies changes suggesting a change in provenance rather than changes in weathering conditions. We made three observations that indicate an increase in sea water temperatures or a possible hyperthermal event related to the first occurrence (FO) of hantkeninids (i.e., the P9/P10 boundary): 1) a distinct peak in abundance of the benthic foraminifera Aragonia aragonensis; 2) the low-diversity of benthic foraminiferal assemblages; and 3) the occurrence of the planktic foraminifera Clavigerinella eocenica and Clavigerinella jarvisi. Benthic foraminiferal and trace fossil assemblages also suggest an associated relative fall of sea level from upper-middle bathyal to sublittoral depths. These characteristic indicators point to this boundary as a promising feature for defining the Global Stratotype Section and Point (GSSP) for the base of the Lutetian Stage. However, complementary magnetobiostratigraphic studies carried out at the Agost section point to the FO of calcareous nannofossil Blackites inflatus (base of CP12b), which occurred 3–5 Myr before the P9/P10 boundary, as the most suitable primary marker event. Whatever the marker event chosen, all the successive events recognized at the Agost section allow a complete characterization of the Y–L transition, and thus this section may be a suitable candidate to locate the GSSP for the Ypresian/Lutetian boundary.     

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea: distribution patterns and influence of main environmental gradients

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.     

Planktic foraminifera from the mid-Cretaceous (Barremian–Early Albian) of the North Sea Basin: Palaeoecological and palaeoceanographic implications

A marine Cretaceous succession (Barremian–Albian) of a cored borehole (BGS 81/40), located in the Central North Sea Basin, has been examined with respect to its planktic and benthic foraminiferal content, as well as for calcareous nannofossils. The distribution patterns of foraminifera and calcareous nannofossils allow for a two fold division of the investigated interval. (1) The Barremian–earliest Aptian interval, which reflects a marine, temporary restricted setting. This is indicated by sporadic occurrences of planktic foraminifera with very rare planispiral forms suggesting short-term connections of the Boreal and Tethyan Realms. The benthic foraminiferal assemblages indicate aerobic, sometimes dysaerobic bottom-water conditions. High abundances of nannoconids in the Barremian suggest enhanced stratification and/or warm, oligotrophic surface water. (2) The late Aptian–early Albian interval, which was characterised by an open-oceanic environment with cool and aerobic surface water conditions. Planktic foraminifera are more abundant and diverse than in the lower interval. Trochospiral hedbergellids dominate the foraminiferal assemblages. The episodic occurrences of planispiral, clavate and trochospiral-flattened planktic morphotypes indicate the existence of a seaway between the Boreal and the Tethyan Realms. Aerobic to dysaerobic bottom-water conditions are suggested by the composition of the benthic foraminiferal assemblages. High abundances of cool-water taxa within the calcareous nannofossil assemblages indicate a cooling trend across the latest Aptian and earliest Albian.     

Modem organic-walled dinoflagellate cysts in arctic marine environments and their (paleo-) environmental significance

The Arctic Ocean is one of the least known marine regions of the world. Because of its major influence on global climate and its hostile environmental conditions it is a fascinating area for paleoecological, paleoclimatic and paleoceanographic research. The composition of planktic microfossil assemblages, and both the trace-element and stable isotope compositions of hard parts, provide us with valuable information about the physical and biochemical parameters of surface waters in the high northern latitudes. Calcareous and biosiliceous microfossils that are traditionally used in Quaternary paleoenvironmental studies are of limited value in the Arctic Ocean because of their low abundances, low diversity and/or low preservation potential. The past several decades have seen considerable progress in our knowledge of the ecology and biogeography of dinoflagellates and their organic-walled cysts in the high northern latitudes, and these dinoflagellate cysts are now important proxies for reconstructing surface water conditions in the Quaternary. This arcticle gives an overview of the ecology of dinoflagellates and their cysts, the processes that transform the living communities into sediment communities, and the environmental gradients that may be reconstructed from fossil dinoflagellate cysts assemblages in the high northern latitudes.      

Correlation between the Paleocene/Eocene boundary and the Ilerdian at Campo, Spain

The Ilerdian is a well-established Tethyan marine stage, which corresponds to an important phase in the evolution of larger foraminifera not represented in the type-area of the classical Northwest-European stages. This biostratigraphic restudy of its parastratotype in the Campo Section (northeastern Spain) based on planktic foraminifera, calcareous nannofossils, dinoflagellate cysts and the distribution of the stable isotopes ∂¹³C and ∂¹⁸O is an attempt to correlate the Paleocene/Eocene boundary based on a characteristic carbon isotope excursion (CIE) marking the onset of the Initial Eocene Thermal Maximum (IETM) and the Ilerdian stage. The base of this ∂¹³C excursion has been chosen as the criterion for the recent proposal of the Global Stratotype Section and Point (GSSP) of the base of the Eocene (= base of the Ypresian) in the Dababiya Section (Egypt) to which an age of 54.9 Ma has been attributed. This level is also characterized by a marked extinction among the deep-water benthic foraminifera (Benthic Foraminifera Extinction Event, BFEE), a flood of representatives of the planktic foraminiferal genus Acarinina and the acme of dinoflagellate cysts of the genus Apectodinium. In the Campo Section, detailed biozonations (planktic foraminifera, calcareous nannofossils, dinoflagellate cysts) are recognized in the Lower and Middle Ilerdian. The correlation with the Ypresian stratotype is based on dinoflagellate cysts and calcareous nannofossils. The base of the Ilerdian is poor in planktic microfossils and its precise correlation with the redefined Paleocene/Eocene boundary remains uncertain.     

Response of deep-sea benthic foraminifera to Miocene paleoclimatic events, DSDP Site 289

Changes in the Miocene deep-sea benthic foraminifera at DSDP Site 289 closely correlate to the climatically induced variations in deep and bottom waters in the Pacific Ocean. In early Miocene time, oxygen and carbon isotopes indicate that bottom waters were relatively warm and poorly oxygenated. Benthic foraminiferal assemblages are characterized by various species inherited from the Oligocene. Expansion of the Antarctic icecap in the early middle Miocene, 14–16 m.y. ago, increased oxygen isotope values, produced cold, more oxygenated bottom waters and lead to a turnover in the benthic foraminifera. An Oligocene—early Miocene assemblage was replaced by a cibicidoid-dominated assemblage. Some species became extinct and benthic faunas became more bathymetrically restricted with the increased stratification of deep waters in the ocean. In mid-Miocene time, Epistominella exigua and E. umbonifera, indicative of young, oxygenated bottom waters, are relatively common at DSDP Site 289. Further glacial expansion 5–9 m.y. ago lowered sealevel, increased oceanic upwelling and associated biological productivity and intensified the oxygen minima. Abundant hispid and costate uvigerines become a dominant faunal element at shallow depths above 2500 m as E. umbonifera becomes common to abundant below 2500 m. By late Miocene time, benthic faunas similar in species composition and proportion to modern faunas on the Ontong-Java plateau, had become established.     

Paleoceanographic changes during the past 1.9 Myr at DSDP Site 238, Central Indian Ocean Basin: Benthic foraminiferal proxies

Deep-sea benthic foraminifera have been quantitatively analyzed in samples (> 125 μm size fraction) from Ocean Drilling Program (ODP) Site 238, to understand paleoceanographic changes in the Central Indian Basin over the past 1.9 Myr. Factor and cluster analyses of the 25 highest-ranked species made it possible to identify five biofacies, characterizing distinct deep-sea environmental settings. The environmental interpretation of each biofacies is based on the ecology of recent deep-sea benthic foraminifera. The benthic faunal record indicates fluctuating deep-sea conditions in environmental parameters including oxygenation, surface productivity and organic food supply. These changes appear to be linked to Indian summer monsoon variability, the main climatic feature of the Indian Ocean region. The benthic assemblages show a major shift at ∼ 0.7 to 0.6 Ma, marked by major turnovers in the relative abundances of species, coinciding with an increased amplitude of glacial cycles. These cycles appear to have influenced low latitude monsoonal climate as well as deep-sea conditions in the Central Indian Ocean Basin.     

Spatial distribution of calcareous dinoflagellate cysts in surface sediments of the Atlantic Ocean between 13 degrees N and 36 degrees S

To enhance the limited information available about the palaeo-ecological significance of calcareous dinoflagellates, we have studied their lateral distribution in surface sediments of the equatorial and south Atlantic between 13 degrees N and 36 degrees S. Calcareous dinoflagellate cysts appear to be widely distributed throughout the studied area. In the surface sediments, concentrations (cyst per gram dry sediment) of the vegetative stage Thoracosphaera heimii are generally higher than that of the (presumably) calcareous resting cysts. Distribution patterns in surface sediments of Orthopithonella granifera (Fütterer) Keupp and Versteegh, Rhabdothorax spp. Kamptner., Sphaerodinella albatrosiana (Kamptner) Keupp and Versteegh S. albatrosiana praratabulated, Sphaerodinella tuberosa var. 1 (Kamptner) Keupp and Versteegh and S. tuberosa var. 2 and the ratios between these species have been compared with temperature, salinity, density and stratification gradients in the upper water column. Rhabdothorax spp. is characteristically present in sediments of more temperate regions characterized by high seasonality. Dinoflagellates producing these cysts are able to tolerate high nutrient concentrations, and mixing of the water column. S. albatrosiana is abundant in regions characterized by high sea surface temperatures and oligotrophic surface water conditions. In contrast, the distribution of S. tuberosa var. 2 is negatively related to temperature. The other cyst species did not show a characteristic pattern in relation to the studied environmental gradients.The ratio of Sphaerodinella tuberosa var. 2 to Orthopithonella granifera can be used for reconstructing the presence of stratification in the upper 50m of the water column, whereas the ratios of S. tuberosa var. 2 to Sphaerodinella albatrosiana and of O. granifera to Rhabdothorax spp. might be used for palaeotemperature reconstructions. Calcareous dinoflagellate cysts are abundant in oligotrophic areas and may be useful for the reconstruction of palaeoenvironmental conditions.     

Microbial diversity in Cenozoic sediments recovered from the Lomonosov Ridge in the Central Arctic Basin

The current understanding of microbes inhabiting deeply buried marine sediments is based largely on samples collected from continental shelves in tropical and temperate latitudes. The geographical range of marine subsurface coring was expanded during the Integrated Ocean Drilling Program Arctic Coring Expedition (IODP ACEX). This expedition to the ice-covered central Arctic Ocean successfully cored the entire 428 m sediment stack on the Lomonosov Ridge during August and September 2004. The recovered cores vary from siliciclastic sediment low in organic carbon (< 0.2%) to organic rich (∼3%) black sediments that rapidly accumulated in the early middle Eocene. Three geochemical environments were characterized based on chemical analyses of porewater: an upper ammonium oxidation zone, a carbonate dissolution zone and a deep (> 200 m below sea floor) sulfate reduction zone. The diversity of microbes within each zone was assessed using 16S rRNA phylogenetic markers. Bacterial 16S rRNA genes were successfully amplified from each of the biogeochemical zones, while archaea was only amplified from the deep sulfate reduction zone. The microbial communities at each zone are phylogenetically different and are most closely related to those from other deep subsurface environments.