43 resultados para Low diversity


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Indicators of surface-water productivity and bottom-water oxygenation have been studied for the age interval from the latest Pleistocene to the Holocene at three holes (679D, 680B, and 68IB) located in the center and at the edges of an upwelling cell at approximately 11°S on the Peruvian continental margin. Upwelling activity was maximal at this latitude during d18O Stages 1 (lower part), 3, the upper part of 5, the lower part of 6, and 7, as documented by high diatom abundance. During these time intervals, the bottom water was poorly oxygenated, as documented by low diversity benthic foraminiferal assemblages that are dominated by B. seminuda s.l. Both surface- and bottom-water-circulation patterns appear to have changed rapidly over short time intervals. Due to changes in surface circulation, the intensity of upwelling decreased, thereby decreasing the concentration of nutrients, and reducing the supply of organic matter to the bottom. Radiolarians became more abundant in the surface waters, and the bottom-water environment was less depleted in oxygen, allowing for the establishment of more diverse benthic foraminiferal assemblages. Surface-water productivity was probably minimal during the early part of d18O Stages 5 and 9, as indicated by the increased abundance of planktonic foraminifers and pteropods and their subsequent preservation.

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Sparse terrestrial palynomorphs (spores and pollen) were recovered from glacigene Lower Miocene and Oligocene core samples from the Cape Roberts Project (CRP) drillhole CRP-2/2A, Victoria Land Basin, Antarctica. Rarity of palynomorphs probably results from the spares periglacial vegetation in the surrounding landscape at the time of deposition, as well as dilution from rapid sediment accumulation. The Miocene and Late Oligocene vegetation is interpreted as including herb-moss tundra with low-growing woody plants (including Nothofagus and podocarp conifers) in more protected areas, similar to that encountered in the Miocene of CRP-1. Species richness and numbers of specimens increase downhole, a trend that begins very gradually below ~307 mbsf, and increases below ~443 mbsf through the Early Oligocene. These lower assemblages reflect low diversity woody vegetation dominated by several species of Nofhofagus and podocarps, growing in somewhat milder conditions, though still cold temperate to periglacial in the Early Oligocene. The CRP-2/2A core provides new biostratigraphical information, such as the First Appearance Datums (FADS) of Tricolpites sp. a near the Oligocene/Miocene boundary, and Marchantiaceae in the Early/Late Oligocene transition: these are taxa that along with N. lachlaniae, Coptospora spp. and Podocarpidites sp.b characterize assemblages recovered from outcrops of the Pliocene Sirius Group in the Transantarctic Mountains. Some elements of the extremely hardy periglacial tundra vegetation that survived in Antarctica into the Pliocene had their origin in the Oligocene during a time of deteriorating (colder, drier) climatic conditions. The CRP results highlight the long persistence of this tundra vegetation, through approximately 30 million years of dynamically changing climatic conditions. Rare Jurassic and more common Permian-Triassic spores and pollen occur sporadically throughout the core. These are derived from Jurassic Ferrar Group sediments, and from the Permian-Triassic Victoria Group, upper Beacon Supergroup. Higher frequencies of reworked Beacon palynomorphs and coaly organic matter below ~307 mbsf indicate greater erosion of the Beacon Supergroup for this lower part of the core. A color range from black, severely metamorphosed specimens, to light-colored, yellow (indicating low thermal alteration), reworked Permian palynomorphs, indicates local provenance in the dolerite-intruded Beacon strata of the Transantarctic Mountains, as well as areas (now sub-ice) of Beacon strata with little or no associated dolerite well inland (cratonwards) of the present Transantarctic Mountains.

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A 100-m-thick Paleocene sequence of mainly pelagic sediments at ODP Site 1121, on the eastern flanks of the Campbell Plateau, contains few to common radiolarians of relatively low diversity in the lower 40 m (Early to early Late Paleocene) and abundant, diverse radiolarian assemblages in the upper 60 m (mid-Late Paleocene). The 150 taxa recorded from the entire Paleocene interval are thought to under-represent the actual species diversity by at least one half as many morphotypes have not been differentiated below the level of genus. Assemblages in the lower 40 m are similar to those described from onland New Zealand and DSDP Site 208 (northern Lord Howe Rise); they are correlated with South Pacific radiolarian zones RP4 and RP5. Assemblages in the upper 60 m differ from other known Late Paleocene assemblages in the great abundance of plagiacanthids and cycladophorids. Similarities are noted with later Cenozoic cool-water assemblages. This upper interval is correlated with South Pacific zone RP6, as revised herein, based on comparison with faunas from Site 208 and Marlborough, New Zealand. The interval is also correlated with the upper part of North Atlantic zone RP6 (RP6b-c) based on the presence of Aspis velutochlamydosaurus, Plectodiscus circularis and Pterocodon poculum. Other species, such as Buryella tetradica and Buryella pentadica, are valuable for local correlation but exhibit considerable diachroneity between the Pacific, Indian and Atlantic Oceans. An age model for the Paleocene interval at Site 1121, based on well-constrained nannofossil and radiolarian datums, indicates that the rate of compacted sediment accumulation doubles from 15 to 30 mm/ka at the RP5/RP6 zonal boundary. In large part this is due to a sudden and pronounced increase in accumulation rates for all siliceous fossils; radiolarians and larger diatoms increase from <100 to >10 000 specimens/cm2/ka. This apparent increase in biosiliceous productivity is age-equivalent to a mid-Paleocene cooling event (57-59 Ma) identified from global stable isotope records that is associated with the heaviest delta13C values for the entire Cenozoic.

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Late Miocene to Holocene planktonic foraminifers from the subantarctic South Atlantic were investigated for their biostratigraphic and environmental significance. The calcareous planktonic microfauna are of low diversity and are very rare or absent at sites located below the CCD. The sediments recovered from Sites 703 and 704 on the Meteor Rise at about 47°S are useful for biostratigraphic and environmental studies. In the whole sequence 16 species or varieties of planktonic foraminifers were recognized. Two species occur in the uppermost Miocene. In the Pliocene the Globorotalia puncticulata population can be used to separate the early from the late Pliocene. The Pliocene/Quaternary boundary does not appear to be well distinguished in the foraminiferal assemblage. A faunal change noted at 2.5 Ma could correspond to the development of glaciation in the Northern Hemisphere and its Antarctic counterpart. At about 5.2 Ma the first increase in polar fauna near the Meteor Rise occurs. Two other cooling periods are indicated in these sequences at about 4 and 3 Ma. Moreover, the hydrologic environment became more productive at about 2.1 Ma and close to the Brunhes/Matuyama boundary.

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Although it is well known that the Paleocene/Eocene thermal maximum (PETM) coincided with a major benthic foraminiferal extinction event, the detailed pattern of the faunal turnover has not yet been clarified. Our high-resolution benthic foraminiferal and carbon isotope analyses at the low latitude Pacific Ocean Shatsky Rise have revealed the following record of major faunal transitions: (1) An initial turnover which involved the benthic foraminiferal extinction event (BFE). The BFE, marked by a sharp transition from Pre-extinction fauna to Disaster fauna represented by small-sized Bolivina gracilis, expresses the onset of the PETM and the abrupt extinction of about 30% of taxa. This faunal transition lasted about 45-74 kyr after the initiation of the PETM and was followed by: (2) the appearance of Opportunistic fauna represented by Quadrimorphina profunda, which existed for about 74-91 kyr after the initiation of the PETM. These two faunas, which appeared after the extinction event, are characterized by low diversity and dwarfism, possibly due to lowered oxygen condition and decreased surface productivity. The second pronounced turnover involved the gradual recovery from Opportunistic Fauna to the establishment of Recovery fauna, which coincided with the recovery about 83-91 kyr after its initiation.

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During Leg 123, abundant and well-preserved Neocomian radiolarians were recovered at Site 765 (Argo Abyssal Plain) and Site 766 (lower Exmouth Plateau). The assemblages are characterized by a scarcity or absence of Tethyan taxa. The Berriasian-early Aptian radiolarian record recovered at Site 765 is unique in its density of well-preserved samples and in its faunal contents. Remarkable contrasts exist between radiolarian assemblages extracted from claystones of Site 765 and reexamined DSDP Site 261, and faunas recovered from radiolarian sand layers of Site 765. Clay faunas are unusual in their low diversity of apparently ecologically tolerant species, whereas sand faunas are dominated by non-Tethyan species that have never been reported before. Comparisons with Sites 766 and 261, as well as sedimentological observations, lead to the conclusion that this faunal contrast results from a difference in provenance, rather than from hydraulic sorting. Biostratigraphic dating proved difficult principally because of the paucity or even absence of (Tethyan) species used in published zonations. In addition, published zonations are contradictory and do not reflect total ranges of species. Radiolarian assemblages recovered from claystones at Sites 765 and 261 in the Argo Basin reflect restricted oceanic conditions for the latest Jurassic to Barremian time period. Neither the sedimentary facies nor the faunal associations bear any resemblance to sediment and radiolarian facies observed in typical Tethyan sequences. I conclude that the Argo Basin was paleoceanographically separated from Tethys during the Late Jurassic and part of the Early Cretaceous by its position at a higher paleolatitude and by enclosing landmasses, i.e., northeastern India and the Shillong Block, which were adjacent to the northwestern Australian margin before the opening. Assemblages recovered from radiolarian sand layers are dominated by non-Tethyan species that are interpreted as circumantarctic. Their sudden appearance in the late Berriasian/early Valanginian pre-dates the oceanization of the Indo-Australian break-up (Ml 1, late Valanginian) by about 5 m.y., but coincides with a sharp increase in margin-derived pelagic turbidites. The Indo-Australian rift zone and its adjacent margins probably were submerged deeply enough to allow an intermittent "spillover" of circumantarctic cold water into the Argo Basin, creating increased bottom current activity. Circumantarctic cold-water radiolarians transported into the Argo Basin upwelled along the margin and died en masse. Concomitant winnowing by bottom currents led to their accumulation in distinct radiolarite layers. High rates of faunal change and the sharp increase of bottom current activity are thought to be synchronous with the two pronounced late Berriasian-early Valanginian lowstands in sea level. Hypothetically, both phenomena might have been caused by a glaciation on the Antarctic-Australian continent, which was for the first time isolated from the rest of Gondwana by oceanic seaways as a result of Jurassic and Early Cretaceous seafloor spreading. The absence of typical Tethyan radiolarian species during the late Valanginian to late Hauterivian period is interpreted as reflecting a time of strong influx of circumantarctic cold water following oceanization (Mil) and rapid spreading between southeast India and western Australia. The reappearance and gradual increase in abundance and diversity of Tethyan forms along with the still dominant circumantarctic species are thought to result from overall more equitable climatic conditions during the Barremian and early Aptian and may have resulted from the establishment of an oceanic connection with the Tethys Ocean during the early Aptian.

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The Middle Eocene diatom and silicoflagellate record of ODP Site 1260A (Demerara Rise) is studied quantitatively in order to throw light on the changes that siliceous phytoplankton communities experienced during a Middle Eocene warming event that occurred between 44.0 and 42.0 Ma. Both Pianka's overlap index, calculated per couple of successive samples, and cluster analysis, point to a number of significant turnover events highlighted by changes in the structure of floristic communities. The pre-warming flora, dominated by cosmopolitan species of the diatom genus Triceratium, is replaced during the warming interval by a new and more diverse assemblage, dominated by Paralia sulcata (an indicator of high productivity) and two endemic tropical species of the genus Hemiaulus. The critical warming interval is characterized by a steady increase in biogenic silica and a comparable increase in excess Ba, both reflecting an increase in productivity. In general, it appears that high productivity not only increased the flux of biogenic silica, but also sustained a higher diversity in the siliceous phytoplankton communities. The microflora preserved above the critical interval is once again of low diversity and dominated by various species of the diatom genus Hemiaulus. All assemblages in the studied material are characterized by the total absence of continental and benthic diatoms and the relative abundance of neritic forms, suggesting a transitional depositional environment between the neritic and the oceanic realms.

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Faunal analyses of planktonic foraminifera and upper-water temperature reconstructions with the modern analog technique are studied and compared to themagnetic susceptibility and gamma ray logs of ODP Core 999A (western Caribbean) for the past 560 kyr in order to explore changes in paleoceanographic conditions in the western Caribbean Sea. Long-term trends in the percentage abundance of planktonic foraminifera inODP Core 999Asuggest two hydrographic scenarios: before and after 480 ka.High percentage abundances of Neogloboquadrina pachyderma and Globorotalia inflata, low abundances of Globorotalia menardii and Globorotalia truncatulinoides, low diversity, and sea-surface temperatures (SST) under 24 °C are typical characteristics occurring from 480 to 560 ka. These characteristics suggest a "shallow" well-oxygenated upper thermocline and the influx of nutrients by either seasonal upwelling plumes and/or eddy-mediated entrainment. The second scenario occurred after 480 ka, and it is characterized by high and fluctuating percentage abundances of Neogloboquadrina dutertrei, G. truncatulinoides, G. menardii, Globigerinita glutinata, Globigerinella siphonifera, and Globigerinoides ruber; a declining trend in diversity; and large SSTs. These characteristics suggest a steady change from conditions characterized by a "shallow" thermocline and chlorophyll maximum to conditions characterized by a "deep" thermocline (mainly during glacial stages) and by more oligotrophic conditions. The influence of the subtropical North Atlantic on the upper thermocline was apparently larger during glacial stages, thus favoring a deepening of the thermocline, an increase in sea-surface salinity, and a dramatic reduction of nutrients in the Guajira upwelling system. During interglacial stages, the influx of nutrients from the Magdalena River is stronger, thus resulting in a deep chlorophyll maximumand a fresher upper ocean. The eddy entrainment of nutrients is the probable mechanism responsible of transport from the Guajira upwelling and Magdalena River plumes into ODP 999A site.

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The iterative evolutionary radiation of planktic foraminifers is a well-documented macroevolutionary process. Here we document the accompanying size changes in entire planktic foraminiferal assemblages for the past 70 My and their relationship to paleoenvironmental changes. After the size decrease at the Cretaceous/Paleogene (K/P) boundary, high latitude assemblages remained consistently small. Size evolution in low latitudes can be divided into three major phases: the first is characterized by dwarfs (65-42 Ma), the second shows moderate size fluctuations (42-14 Ma), and in the third phase, planktic foraminifers have grown to the unprecedented sizes observed today. Our analyses of size variability with paleoproxy records indicate that periods of size increase coincided with phases of global cooling (Eocene and Neogene). These periods were characterized by enhanced latitudinal and vertical temperature gradients in the oceans and high diversity (polytaxy). In the Paleocene and during the Oligocene, the observed (minor) size changes of the largely low-diversity (oligotaxic) assemblages seem to correlate with productivity changes. However, polytaxy per se was not responsible for larger test sizes.

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We report the first microbiological characterization of a terrestrial methane seep in a cryo-environment in the form of an Arctic hypersaline (~24% salinity), subzero (-5 C), perennial spring, arising through thick permafrost in an area with an average annual air temperature of -15 C. Bacterial and archaeal 16S rRNA gene clone libraries indicated a relatively low diversity of phylotypes within the spring sediment (Shannon index values of 1.65 and 1.39, respectively). Bacterial phylotypes were related to microorganisms such as Loktanella, Gillisia, Halomonas and Marinobacter spp. previously recovered from cold, saline habitats. A proportion of the bacterial phylotypes were cultured, including Marinobacter and Halomonas, with all isolates capable of growth at the in situ temperature (-5 C). Archaeal phylotypes were related to signatures from hypersaline deep-sea methane-seep sediments and were dominated by the anaerobic methane group 1a (ANME-1a) clade of anaerobic methane oxidizing archaea. CARD-FISH analyses indicated that cells within the spring sediment consisted of ~84.0% bacterial and 3.8% archaeal cells with ANME-1 cells accounting for most of the archaeal cells. The major gas discharging from the spring was methane (~50%) with the low CH4/C2 + ratio and hydrogen and carbon isotope signatures consistent with a thermogenic origin of the methane. Overall, this hypersaline, subzero environment supports a viable microbial community capable of activity at in situ temperature and where methane may behave as an energy and carbon source for sustaining anaerobic oxidation of methane-based microbial metabolism. This site also provides a model of how a methane seep can form in a cryo-environment as well as a mechanism for the hypothesized Martian methane plumes.

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Recent drilling on the Kerguelen Plateau (Ocean Drilling Program Leg 183) has provided a unique and exciting high latitude record of palaeoceanographic change during the Cenomanian-Turonian in the Southern Ocean. The benthic foraminiferal succession at Site 1138 records the evolution of the Kerguelen Plateau from a subaerially exposed platform in the Cenomanian to a bathyal, pelagic environment in the early Turonian, following a major transgressive pulse and increased thermal subsidence of the Kerguelen Plateau, which led to a sea-level rise of possibly several hundred metres. Diversified benthic foraminiferal assemblages indicate an upper bathyal, mesotrophic setting after the peak of the transgression. The assemblages exhibit strong similarities to temperate, shelf and slope assemblages in the Northern Hemisphere. This bimodal distribution reflects the existence of open oceanic gateways and a dynamic trans-hemispheric global circulation. Equatorial assemblages are characterized by a low-diversity, high carbon flux biofacies. Assemblages from Alaska demonstrate high organic productivity and low oxygen conditions and the prevalence of elevated temperatures on the flooded shelf of the North Slope. Our results show that the distribution of upper bathyal benthic foraminifera was strongly modulated by carbon flux and oxygenation fluctuations, and not by physical migration barriers.

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The Indo-Pakistan Continental Margin represents an extreme habitat for benthic foraminifera since (1) high fluxes of organic matter offer a high food supply, (2) an intensified oxygen minimum Zone (OMZ) develops from the base of the euphotic Zone to water depths over 1000 m and (3) the monsoon causes seasonal oscillations within the biogeochemical cycle. At three stations from the uppermost (233 m), the central (658 m) and the deeper part (902 m) of the OMZ, living benthic foraminiferal assemblages were analyzed within the uppermost 10 cm of the sediment column. The ecologic structure of foraminiferal faunas is characterized by high abundances at the sediment surface and a rapid decrease within the uppermost 2 cm of the sediment column. Despite dysoxic to suboxic bottom-water conditions, stained benthic foraminifera occurred in all cores down to the base of the sampled interval. High surface abundances, a high dominance by few endobenthic calcareous taxa and a low diversity, which may result from specific physiological adaptations to almost anoxic conditions and the absence of predators, are recognized in the central part of the OMZ. The upper and lower margins of the OMZ are characterized by higher diversities and lower abundances. The shallowest part of the OMZ is dominated by calcareous foraminifera, whereas agglutinated species are the most common taxa in the deeper part. Comparisons with previous studies show that benthic foraminiferal assemblages, that are influenced by seasonal oscillations controlling food supply and/or the availability of oxygen, show variations in faunal density and species composition. Since there is strong evidence that oxygen is not a limiting factor for some taxa, it seems more likely that the distribution pattern of benthic foraminifera is preferentially controlled by trophic conditions.

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In the late Paleocene to early Eocene, deep sea benthic foraminifera suffered their only global extinction of the last 75 million years and diversity decreased worldwide by 30-50% in a few thousand years. At Maud Rise (Weddell Sea, Antarctica; Sites 689 and 690, palaeodepths 1100 m and 1900 m) and Walvis Ridge (Southeastern Atlantic, Sites 525 and 527, palaeodepths 1600 m and 3400 m) post-extinction faunas were low-diversity and high-dominance, but the dominant species differed by geographical location. At Maud Rise, post-extinction faunas were dominated by small, biserial and triserial species, while the large, thick-walled, long-lived deep sea species Nuttallides truempyi was absent. At Walvis Ridge, by contrast, they were dominated by long-lived species such as N. truempyi, with common to abundant small abyssaminid species. The faunal dominance patterns at the two locations thus suggest different post-extinction seafloor environments: increased flux of organic matter and possibly decreased oxygen levels at Maud Rise, decreased flux at Walvis Ridge. The species-richness remained very low for about 50 000 years, then gradually increased. The extinction was synchronous with a large, negative, short-term excursion of carbon and oxygen isotopes in planktonic and benthic foraminifera and bulk carbonate. The isotope excursions reached peak negative values in a few thousand years and values returned to pre-excursion levels in about 50 000 years. The carbon isotope excursion was about -2 per mil for benthic foraminifera at Walvis Ridge and Maud Rise, and about -4 per mil for planktonic foraminifera at Maud Rise. At the latter sites vertical gradients thus decreased, possibly at least partially as a result of upwelling. The oxygen isotope excursion was about -1.5 per mil for benthic foraminifera at Walvis Ridge and Maud Rise, -1 per mil for planktonic foraminifera at Maud Rise. The rapid oxygen isotope excursion at a time when polar ice-sheets were absent or insignificant can be explained by an increase in temperature by 4-6°C of high latitude surface waters and deep waters world wide. The deep ocean temperature increase could have been caused by warming of surface waters at high latitudes and continued formation of the deep waters at these locations, or by a switch from dominant formation of deep waters at high latitudes to formation at lower latitudes. Benthic foraminiferal post-extinction biogeographical patterns favour the latter explanation. The short-term carbon isotope excursion occurred in deep and surface waters, and in soil concretions and mammal teeth in the continental record. It is associated with increased CaC03-dissolution over a wide depth range in the oceans, suggesting that a rapid transfer of isotopically light carbon from lithosphere or biosphere into the ocean-atmosphere system may have been involved. The rapidity of the initiation of the excursion (a few thousand years) and its short duration (50 000 years) suggest that such a transfer was probably not caused by changes in the ratio of organic carbon to carbonate deposition or erosion. Transfer of carbon from the terrestrial biosphere was probably not the cause, because it would require a much larger biosphere destruction than at the end of the Cretaceous, in conflict with the fossil record. It is difficult to explain the large shift by rapid emission into the atmosphere of volcanogenic CO2, although huge subaerial plateau basalt eruptions occurred at the time in the northern Atlantic. Probably a complex combination of processes and feedback was involved, including volcanogenic emission of CO2, changing circulation patterns, changing productivity in the oceans and possibly on land, and changes in the relative size of the oceanic and atmospheric carbon reservoirs.

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Analysis of 66 samples from DSDP Site 263 (Cores 263-4R-4 to 263-29R-4) reveals a unique faunal composition with a predominance of agglutinated taxa, many of them previously unrecorded from any other DSDP and ODP Indian Ocean sites. A total of 66 agglutinated and 31 calcareous taxa are documented and five new species are described: Hippocrepina gracilis n.sp., "Textuluriopsis" elegans n.sp., Aaptotoichus challengeri n.sp., "Gaudryinopsis" pseudobettenstaedti n.sp. and "Gaudryina" cuvierensis n.sp. Three assemblages are recognized based on changes in the composition of dominant taxa and occurrences of stratigraphically important species: (1) a high-diversity Valanginian to Barremian Bulbobaculites-Recurvoides Assemblage (Cores 263-29R to - 18R), comprised of numerous elongate agglutinated forms, rare nodosariids, and variable numbers of tubes and ammodiscids; (2) a moderately diverse Aptian to Albian Rhizammina-Ammodiscus-Glomospira Assemblage (Cores 263-18R to -7R) with highly fluctuating numbers of the nominate taxa and Haplophragmoides, Trochammina, Verneuilinoides spp., and Verneuilina howchini; (3) a very low diversity Albian or younger Assemblage (Cores 263-6R to -4R) containing sparse agglutinated foraminifera, rare nodosariids and rotaliids. We interpret the assemblages as shelf to lower slope and consider them to reflect a deepening palaeobathymetry as the Cuvier margin subsided after the initial breakup of East Gondwana during the Valanginian. Our interpretation is in sharp contrast with initial palaeodepth estimates of less than 100 m, as well as with original chronostratigraphic interpretations based on foraminifera and nannofossils which correlated the base of the recovered interval with the Aptian. The absence of many cosmoplitan forms, despite high diversity suggests strong faunal differentiation in the Austral realm or endemisn within the Cuvier Basin during the Early Cretaceous.

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Late Neogene biostratigraphy of planktonic foraminifers has been investigated from 13 sites cored during Ocean Drilling Program Leg 167 off the coast of California. The planktonic foraminiferal biostratigraphy of six of these sites is presented here at higher stratigraphic resolution for the interval that encompasses the late early Pliocene through the Quaternary (~3.5 Ma to present day). The sites form a transect along the California margin from 31°N to 41°N within the California Current system. A new planktonic foraminiferal zonation has been established largely on evolutionary changes within the Neogloboquadrina plexus, supported by other taxa. A total of eight zones are recognized, most of which are broadly applicable throughout the region, thus providing a biostratigraphic zonation of the sequence at ~0.5-m.y. intervals. The new zonation appears to be unique to the California Current system. The diversity of planktonic foraminiferal assemblages during the late Neogene appears to have remained relatively constant despite large-scale paleoclimatic change. The assemblages are consistently dominated by few taxa that almost always include the neogloboquadrinids and Globigerina bulloides. Low diversity and high dominance of the assemblages favored these and other taxa well adapted to upwelling systems exhibiting high seasonal surface ocean variability. Apparently the oceanographic conditions that favor such assemblages have persisted at least for the duration of the late Neogene (~3.5 Ma to present day). The biostratigraphically important forms have been illustrated with scanning electron micrographs.