Relative abundance and ranges of selected diatoms from Pliocene-Pleistocene sections of ODP Leg 177, Atlantic sector of the Southern Ocean

During Ocean Drilling Program (ODP) Leg 177, seven sites were drilled aligned on a transect across the Antarctic Circumpolar Current in the Atlantic sector of the Southern Ocean. The primary scientific objective of Leg 177 was the study of the Cenozoic paleoceanographic and paleoclimatic history of the southern high latitudes and its relationship with the Antarctic cryosphere development. Of special emphasis was the recovery of Pliocene-Pleistocene sections, allowing paleoceanographic studies at millennial or higher time resolution, and the establishment of refined biostratigraphic zonations tied to the geomagnetic polarity record and stable isotope records. At most sites, multiple holes were drilled to ensure complete recovery of the section. A description of the recovered sections and the construction of a multihole splice for the establishment of a continuous composite is presented in the Leg 177 Initial Reports volume for each of the sites (Gersonde, Hodell, Blum, et al., 1999). Here we present the relative abundance pattern and the stratigraphic ranges of diatom taxa encountered from shore-based light microscope studies completed on the Pliocene-Pleistocene sequences from six of the drilled sites (Sites 1089-1094). No shore-based diatom studies have been conducted on the Pliocene-Pleistocene sediments obtained at Site 1088, located on the northern crest of the Agulhas Ridge, because of the scattered occurrence and poor preservation of diatoms in these sections (Shipboard Scientific Party, 1999b). The data included in our report present the baseline of a diatom biostratigraphic study of Zielinski and Gersonde (2002), which (1) includes a refinement of the southern high-latitude Pliocene-Pleistocene diatom zonation, in particular for the middle and late Pleistocene, and (2) presents a biostratigraphic framework for the establishment of age models of the recovered sediment sections. Zielinski and Gersonde (2002) correlated the diatom ranges with the geomagnetic polarity record established shipboard (Sites 1090 and 1092) (Shipboard Scientific Party, 1999c, 1999d) and on shore (Sites 1089, 1091, 1093, and 1094) by Channell and Stoner (2002). The Pliocene-Pleistocene diatom zonation proposed by Zielinski and Gersonde (2002) relies on a diatom zonation from Gersonde and Bárcena (1998) for the northern belt of the Southern Ocean. Because of latitudinal differentiation of sea-surface temperature, nutrients, and salinity between Antarctic and Subantarctic/subtropical water masses, the Pliocene-Pleistocene stratigraphic marker diatoms are not uniformly distributed in the Southern Ocean (Fenner, 1991; Gersonde and Bárcena, 1998). As a consequence, Zielinski and Gersonde (2002) propose two diatom zonations for application in the Antarctic Zone south of the Polar Front (Southern Zonation, Sites 1094 and 1093) and the area encompassing the Polar Front Zone (PFZ) and the Subantarctic Zone (Northern Zonation, Sites 1089-1092). This accounts especially for the Pleistocene zonation where Hemidiscus karstenii, whose first abundant occurrence datum and last occurrence datum defines the subzonation of the northern Thalassiosira lentiginosa Zone, occurs only sporadically in the cold-water realm south of the PFZ and thus is not applicable in sections from this area. However, newly established marker species assigned to the genus Rouxia (Rouxia leventerae and Rouxia constricta) are more related to cold-water environments and allow a refinement of the Pleistocene stratigraphic zonation for the southern cold areas. A study relying on quantitative counts of both Rouxia species confirms the utility of these stratigraphic markers for the identification of sequences attributed to marine isotope Stages 6 and 8 in the southern Southern Ocean (Zielinski et al., 2002).

Planktonic and benthic stables Isotopes, age model and carbon analysis of HUD91/039 from the northernmost Baffin Bay

A multiproxy study of palaeoceanographic and climatic changes in northernmost Baffin Bay shows that major environmental changes have occurred since the deglaciation of the area at about 12 500 cal. yr BP. The interpretation is based on sedimentology, benthic and planktonic foraminifera and their isotopic composition, as well as diatom assemblages in the sedimentary records at two core sites, one located in the deeper central part of northernmost Baffin Bay and one in a separate trough closer to the Greenland coast. A revised chronology for the two records is established on the basis of 15 previously published AMS 14C age determinations. A basal diamicton is overlain by laminated, fossil-free sediments. Our data from the early part of the fossiliferous record (12 300 - 11 300 cal. yr BP), which is also initially laminated, indicate extensive seasonal sea-ice cover and brine release. There is indication of a cooling event between 11 300 and 10 900 cal. yr BP, and maximum Atlantic Water influence occurred between 10 900 and 8200 cal. yr BP (no sediment recovery between 8200 and 7300 cal. yr BP). A gradual, but fluctuating, increase in sea-ice cover is seen after 7300 cal. yr BP. Sea-ice diatoms were particularly abundant in the central part of northernmost Baffin Bay, presumably due to the inflow of Polar waters from the Arctic Ocean, and less sea ice occurred at the near-coastal site, which was under continuous influence of the West Greenland Current. Our data from the deep, central part show a fluctuating degree of upwelling after c. 7300 cal. yr BP, culminating between 4000 and 3050 cal. yr BP. There was a gradual increase in the influence of cold bottom waters from the Arctic Ocean after about 3050 cal. yr BP, when agglutinated foraminifera became abundant. A superimposed short-term change in the sea-surface proxies is correlated with the Little Ice Age cooling.

Diatom biostratigraphy and datum levels of sediments from ODP Leg 167 sites

Ocean Drilling Program Leg 167 represents the first time since 1978 that the North American Pacific margin was drilled to study ocean history. More than 7500 m of Quaternary to middle Miocene (14 Ma) sediments were recovered from 13 sites, representing the most complete stratigraphic sequence on the California margin. Diatoms are found in most samples in variable abundance and in a moderately well-preserved state throughout the sequence, and they are often dominated by robust, dissolution-resistant species. The Neogene North Pacific diatom zonation of Yanagisawa and Akiba (1998, doi:10.5575/geosoc.104.395) best divides the Miocene to Quaternary sequences, and updated ages of diatom biohorizons estimated based on the geomagnetic polarity time scale of Cande and Kent (1995, doi:10.1029/94JB03098) are slightly revised to adjust the differences between the other zonations. Most of the early middle Miocene through Pleistocene diatom datum levels that have been proven to be of stratigraphic utility in the North Pacific appear to be nearly isochronous within the level of resolution constrained by sample spacing. The assemblages are characterized by species typical of middle-to-high latitudes and regions of high surface-water productivity, predominantly by Coscinodiscus marginatus, Stephanopyxis species, Proboscia barboi, and Thalassiothrix longissima. Latest Miocene through Pliocene assemblages in the region of the California Current, however, are intermediate between those of subarctic and subtropical areas. As a result, neither the existing tropical nor the subarctic (high latitude) zonal schemes were applicable for this region. An interval of pronounced diatom dissolution detected throughout the Pliocene sequence apparently correspond to a relatively warmer paleoceanographic condition resulting in a slackening of the southward flow of the California Current.

Diatom assamblage in samples from sediment traps across the Pacific Sector of the Southern Ocean

An array of four sediment trap moorings recorded the particulate flux across the Antarctic Circumpolar Current (ACC) at 170 °W, between November 1996 and January 1998, as part of the US JGOFS-Antarctic Environment and Southern Ocean Process Study (AESOPS) program. The trap locations represent sampling within the Polar Frontal Zone, the Antarctic Polar Front, the Antarctic Zone and the Southern Antarctic Zone. Here we report observations from 1000 m below the sea-surface compared to seafloor and surface water distributions. Sub-sample splits from each trap were obtained and total diatom flux and species composition were determined. The diatom fluxes were quantified using both a dilution and a 'spike' method to allow for the rapid repeatability of measurements. Diatom flux was found to be highly seasonal across the ACC particularly at higher latitudes. Marine snow aggregates of intact diatom cells and chains were the major components of the biogenic flux. Siliceous particle size was noted to decrease with increasing latitude, which could be aligned with a shift of the diatom assemblage to small-size species/sea-ice affiliated species. A 'double-structured' diatom flux was recorded at the location of the Antarctic Polar Front trap, with a shift in the diatom assemblage from larger to smaller diatoms in the second flux episode. The sediment trap assemblage shows deviations from the surface water assemblage, while surface sediment samples indicate that significant dissolution occurs after 1000 m and at the sediment-water interface. Estimation of diatom biovolumes across the ACC shows that large diatoms have the potential to greatly impact biogenic fluxes to the ocean interior despite their low fluxes. Small species of the genus Fragilariopsis could potentially export as much Corg as Fragilariopsis kerguelensis near the retreating ice edge. However, their low abundance in the surface sediments also suggests that these diatoms are a shallow export species.

Diatom biostratigraphy and sea ice concentration reconstruction from core DA06-139G, Vaigat Strait in Disko Bugt, West Greenland

A diatom-based sea-ice concentration (SIC) transfer function is developed using 72 surface samples from west of Greenland and around Iceland, and through comparison with the associated modern SIC. Canonical correspondence analysis on surface sediment diatoms and monthly average of SIC reveals that April SIC is the most important environmental factor controlling the distribution of diatoms in the area, and permits the development of a diatom-based SIC transfer function. The consistency between reconstructed SIC based on diatoms from West Greenland and the instrumental and documentary data during the last ~75 years demonstrates that the diatom-based SIC reconstruction is reliable for studying the palaeoceanography off West Greenland. Relatively warm conditions with strong influence of the Irminger Current (IC) are indicated for the early part of the record (~5000-3860 cal. yr BP), corresponding in time to the latest part of the Holocene Thermal Maximum. The April SIC oscillated around the mean value between 3860 and 1510 cal. yr BP and was above mean afterwards, particularly during the time interval 1510-1120 cal. yr BP and after 650 cal. yr BP, indicating more extensive sea-ice cover in Disko Bugt. A high degree of consistency between the reconstructed April SIC and changes in the diatom species suggests that the sea-ice condition in Disko Bugt is strongly influenced by variations in the relative strength of two components of the West Greenland Current, i.e. the cold East Greenland Current and the relatively warm IC.

Radiocarbon ages and diatom abundance in cores GA306-BC4 and GA306-GC4 obtained during Galathea 3 cruise, West Greenland

Diatom assemblages from Holsteinsborg Dyb on the West Greenland shelf were analysed with high temporal resolution for the last 1200 years. A high degree of consistency between changes in frequency of selected diatom species and instrumental data from the same area during the last 70 years confirms the reliability of diatoms (particularly sea-ice species and warm-water species) for the study of palaeoceanographic changes in this area. A general cooling trend with some fluctuations is marked by an increase in sea-ice species throughout the last 1200 years. A relatively warm period with increased influence of Atlantic water masses of the Irminger Current (IC) is found at AD 750-1330, although with some oceanographic variability after AD 1000. A pronounced oceanographic shift occurred at AD 1330, corresponding in time to the transition from the so-called 'Medieval Warm Period' (MWP) to the 'Little Ice Age' (LIA). The LIA cold episode is characterized by three intervals with particularly cold sea-surface conditions at AD 1330-1350, AD 1400-1575 and AD 1660-1710 as a result of variable influence of Polar waters in the area. During the last 70 years, two relatively warm periods and one cold period (the early 1960s to mid-1990s) are indicated by changes in the diatom components. Our study demonstrates that sedimentary records on the West Greenland shelf provide valuable palaeoenvironment data that confirm a linkage between local and large-scale North Atlantic oceanographic and atmospheric oscillations.

Diatom biostratigraphy of sediments from the Kerguelen Plateau, Southern Ocean

The biostratigraphic distribution and abundance of lower Oligocene to Pleistocene diatoms is documented from Holes 747A, 747B, 748B, 749B, and 751A drilled during Ocean Drilling Program Leg 120 on the Kerguelen Plateau in the southeast Indian Ocean. The occurrence of middle and upper Eocene diatoms is also documented, but these are rare and occur in discrete intervals. The recovery of several Oligocene to Pleistocene sections with minimal coring gaps, relatively good magnetostratigraphic signatures, and mixed assemblages of both calcareous and siliceous microfossils makes the above four Leg 120 sites important biostratigraphic reference sections for the Southern Ocean and Antarctic continent. A high-resolution diatom zonation divides the last 36 m.y. into 45 zones and subzones. This zonation is built upon an existing biostratigraphic framework developed over the past 20 yr of Southern Ocean/Antarctic deep-sea coring and drilling. After the recent advances from diatom biostratigraphic studies on sediments from Legs 113, 114, 119, and 120, a zonal framework for the Southern Ocean is beginning to stabilize. The potential age resolution afforded by the high-diversity diatom assemblages in this region ranks among the highest of all fossil groups. In addition to the 46 datum levels that define the diatom zones and subzones, the approximate stratigraphic level, age, and magnetic anomaly correlative of more than 150 other diatom datums are determined or estimated. These total 73 datum levels for the Pliocene-Pleistocene, 67 for the Miocene, and 45 for the Oligocene. Greater stratigraphic resolution is possible as the less common and poorly documented species become better known. This high-resolution diatom stratigraphy, combined with good to moderately good magnetostratigraphic control, led to the recognition of more than 10 intervals where hiatuses dissect the Oligocene-Pleistocene section on the Kerguelen Plateau. We propose 12 new diatom taxa and 6 new combination

High resolution paleoceanography of the Guaymas Basin, Gulf of California, during the past 15000 yeras

Deep Sea Drilling Project Site 480 (27°54.10'N, 111°39.34'W; 655 m water depth) contains a high resolution record of paleoceanographic change of the past 15000 years for the Guaymas Basin, a region of very high diatom productivity within the central Gulf of California. Analyses of diatoms and silicoflagellates were completed on samples spaced every 40-50 yr, whereas ICP-AES geochemical analyses were completed on alternate samples (sample spacing 80-100 yr). The Bolling-Allerod interval (14.6-12.9 ka) (note, ka refers to 1000 calendar years BP throughout this report) is characterized by an increase in biogenic silica and a decline in calcium carbonate relative to surrounding intervals, suggesting conditions somewhat similar to those of today. The Younger Dryas event (12.9-11.6 ka) is marked by a major drop in biogenic silica and an increase in calcium carbonate. Increasing relative percentage contributions of Azpeitia nodulifera and Dictyocha perlaevis (a tropical diatom and silicoflagellate, respectively) and reduced numbers of the silicoflagellate Octactis pulchra are supportive of reduced upwelling of nutrient-rich waters. Between 10.6 and 10.0 ka, calcium carbonate and A. nodulifera abruptly decline at DSDP 480, while Roperia tesselata, a diatom indicative of winter upwelling in the modern-day Gulf, increases sharply in numbers. A nearly coincident increase in the silicoflagellate Dictyocha stapedia suggests that waters above DSDP 480 were more similar to the cooler and slightly more saline waters of the northern Gulf during much of the early and middle parts of the Holocene (~10 to 3.2 ka). At about 6.2 ka a stepwise increase in biogenic silica and the reappearance of the tropical diatom A. nodulifera marks a major change in oceanographic conditions in the Gulf. A winter shift to more northwesterly winds may have occurred at this time along with the onset of periodic northward excursions (El Nino-driven?) of the North Equatorial Countercurrent during the summer. Beginning between 2.8 and 2.4 ka, the amplitude of biogenic silica and wt% Fe, Al, and Ti (proxies of terrigenous input) increase, possibly reflecting intensification of ENSO cycles and the establishment of modern oceanographic conditions in the Gulf. Increased numbers of O. pulchra after 2.8 ka suggest enhanced spring upwelling.

Diatom abundance in ODP Leg 178 holes

This paper documents the biostratigraphic distribution and abundance of diatoms from sites drilled during Ocean Drilling Program Leg 178, off the Pacific margin of the Antarctic Peninsula. Drift sediments cored on the continental rise at Sites 1095, 1096, and 1101 have good recovery and a well-defined paleomagnetic record. Well-preserved diatoms are present throughout the upper Miocene to middle Pliocene and in the upper Quaternary section of these sites. The stratigraphic occurrence of diatom species through these intervals defines numerous datum levels. Diatom events are given absolute age estimates through direct correlation to the established paleomagnetic stratigraphy of Sites 1095, 1096, and 1101. Leg 178 diatom biostratigraphic results enable the development of a regional stratigraphic framework for the Pacific sector of the Southern Ocean and record the interaction of open-ocean and shelf-margin diatom floras.

Diatom studies in surface and downcore sediments off southern Sumatra and Java

A wealth of sedimentary records aimed at reconstructing late Quaternary changes in productivity and temperature have been devoted to understanding linkages between the Indo-Pacific Warm Pool (IPWP) and other distant oceanic areas. Most of these reconstructions are based, however, on biogeochemical and sedimentological proxies, with comparatively less attention devoted to microfossils. A high-resolution (<1 ka) study of diatom concentrations and the community at site GeoB10038-4, recovered off southern Sumatra (ca. 6°S, 103°E), closely tracks the variations of diatom concentrations in the westernmost IPWP during the last glacial-interglacial cycle. The diatom record provides evidence that diatom paleoproductivity was highest during interglacials, primarily due to the input of lithogenics and nutrients following the rise in sea level after full glacials. In addition, the co-variation of total diatom concentration and Northern Hemisphere forcing for Marine Isotope Stage 5 suggests a direct response of diatom productivity and upwelling intensity to boreal summer insolation. Temporal shifts of the diverse diatom community at site GeoB10038-4 correspond well with the present-day seasonal monsoon pattern and the strengthening and weakening phases of upwelling along the southern coast of Sumatra. Resting spores of Chaetoceros, typical of nutrient-rich waters, were dominant during periods of highest diatom paleoproductivity and responded to the strengthening of the SE monsoon, while diatoms of oligotrophic to mesotrophic waters characterized intermonsoon periods. The close correspondence between the dominance of upwelling diatoms and the boreal summer insolation resembles the present-day dynamics of diatom production. The observed interglacial highs and glacial lows of diatom productivity at site GeoB10038-4 is a unique pattern in the late Quaternary tropics.

Neogene diatoms of the Japan Sea

A series of excellent upper Miocene through Quaternary diatomaceous sequences recovered at four sites during Leg 127 was examined for diatoms. The diagenetic transition from opal-A to opal-CT is a diachronic horizon from the uppermost part of the Denticulopsis katayamae Zone (8.5 Ma) at Hole 797B to the uppermost part of the Neodenticula kamtschatica Zone (5.73 Ma) at Hole 795A. The diatom zonation of Koizumi (1985) best divides the upper Miocene to Quaternary sequences above the opal-A/opal-CT boundary and also is useful to date carbonate concretions including diatoms below the boundary. Forty diatom datum levels were evaluated biostratigraphically based on the sediment accumulation rate curve, and several isochronous datum levels are newly proposed for the Japan Sea area. A warm-water current did not penetrated into the Japan Sea through the Tsushima strait during the late Miocene and Pliocene time, because subtropical warm-water diatoms are essentially not present in such sediment samples. The occurrences of diatom are cyclic throughout the Quaternary sediments and are affected by eustatic sea level changes.

Miocene to Holocene diatom biostratigraphy of ODP Leg 105 sites

Diatoms occur sporadically in lower Miocene to Holocene sediments recovered at ODP Site 645 and in upper Pliocene to Holocene sediments at ODP Site 646. The diatom assemblage at Site 645 contains rare stratigraphic indicators. Fragmentation of frustules and the occurrence of species characteristic of high-latitude shelf, upper-slope environments suggest current transportation from the shelf. The diatom abundance and preservation at Site 646 probably reflect climatic changes and are also affected by dissolution, but it is not possible to detect the dominant factor. Therefore, the stratigraphic ranges of the primary and secondary biostratigraphic indicators are often unreliable.

High-resolution sedimentological and paleontological record of ODP Site 167-1019

Holocene and latest Pleistocene oceanographic conditions and the coastal climate of northern California have varied greatly, based upon high-resolution studies (ca. every 100 years) of diatoms, alkenones, pollen, CaCO3%, and total organic carbon at Ocean Drilling Program (ODP) Site 1019 (41.682°N, 124.930°W, 980 m water depth). Marine climate proxies (alkenone sea surface temperatures [SSTs] and CaCO3%) behaved remarkably like the Greenland Ice Sheet Project (GISP)-2 oxygen isotope record during the Bølling-Allerod, Younger Dryas (YD), and early part of the Holocene. During the YD, alkenone SSTs decreased by >3°C below mean Bølling-Allerod and Holocene SSTs. The early Holocene (ca. 11.6 to 8.2 ka) was a time of generally warm conditions and moderate CaCO3 content (generally >4%). The middle part of the Holocene (ca. 8.2 to 3.2 ka) was marked by alkenone SSTs that were consistently 1-2°C cooler than either the earlier or later parts of the Holocene, by greatly reduced numbers of the gyre-diatom Pseudoeunotia doliolus (<10%), and by a permanent drop in CaCO3% to <3%. Starting at ca. 5.2 ka, coastal redwood and alder began a steady rise, arguing for increasing effective moisture and the development of the north coast temperate rain forest. At ca. 3.2 ka, a permanent ca. 1°C increase in alkenone SST and a threefold increase in P. doliolus signaled a warming of fall and winter SSTs. Intensified (higher amplitude and more frequent) cycles of pine pollen alternating with increased alder and redwood pollen are evidence that rapid changes in effective moisture and seasonal temperature (enhanced El Niño-Southern Oscillation [ENSO] cycles) have characterized the Site 1019 record since about 3.5 ka.

Distribution of Pliocene diatom faunas in the Northwest Pacific

High-resolution quantitative diatom data are tabulated for the early part of the late Pliocene ( 3.25 to 2.08 Ma ) at DSDP Site 580 in the northwestern Pacific. Sample spacing averages 11 k.y. between 3.1 and 2.8 Ma, but increases to 14 to 19 k.y. prior to 3.1 Ma and after 2.8 Ma. Q-mode factor analysis of the middle Pliocene assemblage reveals four factors which explain 92.4% of the total variance of the 47 samples studied between 3.25 and 2.55 Ma. Three of the factors are closely related to modern subarctic, transitional, and subtropical elements, while the fourth factor, which is dominated by Coscinodiscus marginatus and the extinct Pliocene species Neodenticula kamtschatica, appears to correspond to a middle Pliocene precursor of the subarctic water mass. Knowledge of the modern and generalized Pliocene paleoclimatic relationships of various diatom taxa is used to generate a paleoclimate curve ("Twt") based on the ratio of warm-water (subtropical) to cold-water diatoms with warm-water transitional taxa (Thalassionema nitzschioides, Thalassiosira oestrupii, and Coscinodiscus radiatus) factored into the equation at an intermediate (0.5) value. The "Twt" ratios at more southerly DSDP Sites 579 and 578 are consistently higher (warmer) than those at Site 580 throughout the Pliocene, suggesting the validity of the ratio as a paleoclimatic index. Diatom paleoclimatic data reveal a middle Pliocene (3.1 to 3.0 Ma) warm interval at Site 580 during which paleotemperatures may have exceeded maximum Holocene values by 3 °- 5.5 °C at least three times. This middle Pliocene warm interval is also recognized by planktic foraminifers in the North Atlantic, and it appears to correspond with generalized depleted oxygen isotope values suggesting polar warming. The diatom "Twt" curve for Site 580 compares fairly well with radiolarian and silicoflagellate paleoclimatic curves for Site 580, planktic foraminiferal sea-surface temperature estimates for the North Atlantic, and benthic oxygen isotope curves for late Pliocene, although higher resolution studies on paired samples are required to test the correspondence of these various paleoclimatic indices.

Diatom abundance in middle-to-late Pleistocene sediments of IODP Site 303-U1304

We examined diatom assemblages in a series of remarkable laminated diatomaceous ooze (LDO) horizons in the marine sediments from Integrated Ocean Drilling Program (IODP) Site U1304 to reconstruct the middle-to-late Pleistocene paleoceanographic evolution of the northern North Atlantic Ocean. Four confirmed diatom biohorizons combined with calcareous nannofossil and paleomagnetic stratigraphies established the chronological framework for the material. The planktonic, araphid, needle-like species Thalassiothrix longissima was the greatest contributor to the LDO facies. From the results of a principal component analysis using the percent abundances of 65 significant (p = 5%) diatom taxa, except for Tx. longissima, which was extremely dominant in almost all horizons observed, we identified two principal component (PC) axes. Taxa probably associated with the stratigraphic distribution of the major zonal marker Neodenticula seminae (ranging from 1.26 to 0.84 Ma in this ocean) loaded on PC1 with a high value. PC2 was related to the ocean surface temperature. The stratigraphic variability of the PC2 score indicated that switching between warm- and cold-water assemblages occurred concurrently with LDO deposition (or extreme Tx. longissima dominance) episodes in several horizons (particularly after 0.84 Ma), suggesting that the Subarctic Convergence (SAC) oceanic front passed over Site U1304 during Pleistocene glacial/interglacial cycles. Our floral evidence supports the model of nearly monospecific LDO formation caused by the enhanced physical accumulation of particular diatoms such as Tx. longissima. On the other hand, Nd. seminae, which probably contributes to spring phytoplankton blooms in the modern ocean, was present only between 1.26 and 0.84 Ma in this area. Thus, we infer that the main contributor of export flux in the regional annual primary production cycle would have shifted drastically from one of a spring phytoplankton bloom leader (Nd. seminae) to minor but mass dump assemblages (Tx. longissima etc.) in the mid-Pleistocene.