851 resultados para Emergency food supply
Resumo:
Lower Miocene through upper Pleistocene benthic foraminifer assemblage records from Ocean Drilling Program Site 751 on the Southern Kerguelen Plateau (57°44'S, water depth 1634 m) were combined with benthic and planktonic foraminifer oxygen and carbon isotope records and high-resolution CaCO3 data from the same site. Implications for the Neogene productivity and paleoceanography of the southern Indian Ocean are discussed. We used distinctive features of the Miocene d18O and d13C curves for stratigraphic correlation. Coinciding with a lower middle Miocene hiatus from 14.2 to 13.4 Ma, there was a rapid increase in benthic d18O values by 1.2 per mil. This distinct increase occurs in middle Miocene benthic foraminifer oxygen isotope curves from all oceans. No major change, however, in benthic foraminifer faunal composition occurred in this period of growth of the Antarctic ice cap and cooling of deep ocean waters (14.9-14.2 Ma). A drastic change in benthic foraminifer faunas coincided with a hiatus from 8.4 to 5.9 Ma. Shortly after this hiatus, in the latest Miocene, the CaCO3 content of the sediments dropped from 75% to 0%. From that time ( 5.8 Ma) through the early Pliocene, Site 751 has been situated beneath a high biogenic siliceous productivity zone. Carbonate contents of upper Pliocene and Pleistocene sediments vary between 20% and 70%. The benthic foraminifer faunas in the uppermost Pliocene and lower Pleistocene reflect strong bottom current conditions, in contrast to those in the upper Pleistocene, which indicate calm sedimentation and high food supply. High d13C values of planktonic foraminifers compared with low values of benthic foraminifers suggest high primary productivity in the late Pleistocene. The changes in productivity were probably a result of latitudinal migration and meandering of the Polar Frontal Zone.
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In 1986 participants of the Benthos Ecology Working Group of ICES conducted a synoptic mapping of the infauna of the southern and central North Sea. Together with a mapping of the infauna of the northern North Sea by Eleftheriou and Basford (1989, doi:10.1017/S0025315400049158) this provides the database for the description of the benthic infauna of the whole North Sea in this paper. Division of the infauna into assemblages by TWINSPAN analysis separated northern assemblages from southern assemblages along the 70 m depth contour. Assemblages were further separated by the 30, 50 m and 100 m depth contour as well as by the sediment type. In addition to widely distributed species, cold water species do not occur further south than the northern edge of the Dogger Bank, which corresponds to the 50 m depth contour. Warm water species were not found north of the 100 m depth contour. Some species occur on all types of sediment but most are restricted to a special sediment and therefore these species are limited in their distribution. The factors structuring species distributions and assemblages seem to be temperature, the influence of different water masses, e.g. Atlantic water, the type of sediment and the food supply to the benthos.
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In the maritime Antarctic, brown skuas (Catharacta antarctica lonnbergi) show two foraging strategies: some pairs occupy feeding territories in penguin colonies, while others can only feed in unoccupied areas of a penguin colony without defending a feeding territory. One-third of the studied breeding skua population in the South Shetlands occupied territories of varying size (48 to >3,000 penguin nests) and monopolised 93% of all penguin nests in sub-colonies. Skuas without feeding territories foraged in only 7% of penguin sub-colonies and in part of the main colony. Females owning feeding territories were larger in body size than females without feeding territories; no differences in size were found in males. Territory holders permanently controlled their resources but defence power diminished towards the end of the reproductive season. Territory ownership guaranteed sufficient food supply and led to a 5.5 days earlier egg-laying and chick-hatching. Short distances between nest and foraging site allowed territorial pairs a higher nest-attendance rate such that their chicks survived better (71%) than chicks from skua pairs without feeding territories (45%). Due to lower hatching success in territorial pairs, no difference in breeding success of pairs with and without feeding territories was found in 3 years. We conclude that skuas owning feeding territories in penguin colonies benefit from the predictable and stable food resource by an earlier termination of the annual breeding cycle and higher offspring survivorship.
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A multidisciplinary study was undertaken at the Qijurittuq Site (IbGk-3) on Drayton Island in Low-Arctic Quebec (Canada) to document the relationships between climatic, environmental, and cultural changes and the choice of Thule/Inuit dwelling style in the eastern Arctic. Several marine terraces were 14C-dated with shells in order to reconstruct the area's uplift (glacioisostatic rebound) curve. Plant macrofossil analysis of peat was conducted to reconstruct past vegetation and, indirectly, past climate. Archaeological surveys and excavations characterized the structure of subterranean sod houses at the Qijurittuq Site and were supplemented with open interviews with Inuit elders for a better understanding of site location and the use of household space. The sites selected for habitation were well-drained sandy marine terraces in a valley sheltered from prevailing winds. Sod houses were in turn made possible by the abundance of driftwood on the island and the presence of nearby peatland. Thule/Inuit people used semi-subterranean houses rather than igloos at the Qijurittuq Site during the dry, cold conditions toward the end of the Little Ice Age. Stable environmental conditions and food supply during winter possibly explain the use of those semipermanent houses on Drayton Island. However, it does not exclude the use of igloos during short expeditions on ice.
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Bacterial abundance, biomass and cell size were studied in the oligotrophic sediments of the Cretan Sea (Eastern Mediterranean), in order to investigate their response to the seasonal varying organic matter (OM) inputs. Sediment samples were collected on a seasonal basis along a transect of seven stations (ranging from 40 to 1570 m depth) using a multiple-corer. Bacterial parameters were related to changes in chloroplastic pigment equivalents (CPE), the biochemical composition (proteins, lipids, carbohydrates) of the sedimentary organic matter and the OM flux measured at a fixed station over the deep basin (1570 m depth). The sediments of the Cretan Sea represent a nutrient depleted ecosystem characterised by a poor quality organic matter. All sedimentary organic compounds were found to vary seasonally, and changes were more evident on the continental shelf than in deeper sediments. Bacterial abundance and biomass in the sediments of the Cretan Sea (ranging from 1.02 to 4.59 * 10**8 cells/g equivalent to 8.7 and 38.7 µgC/g) were quite high and their distribution appeared to be closely related to the input of fresh organic material. Bacterial abundance and biomass were sensitive to changes in nutrient availability, which also controls the average cell size and the frequency of dividing cells. Bacterial abundance increased up to 3-fold between August '94 and February '95 in response to the increased amount of sedimentary proteins and CPE, indicating that benthic bacteria were constrained more by changes in quality rather than the quantity of the sedimentary organic material. Bacterial responses to the food inputs were clearly detectable down to 10 cm depth. The distribution of labile organic compounds in the sediments appeared to influence the vertical patterns of bacterial abundance and biomass. Cell size decreased significantly with water depth. Bacterial abundance and biomass were characterised by clear seasonal changes in response to seasonal OM pulses. The strong coupling between protein flux and bacterial biomass together with the strong bacterial dominance over the total biomass suggest that the major part of the carbon flow was channelled through the bacteria and the benthic microbial loop.
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Sannai-Maruyama is one of the most famous and best-researched mid-Holocene (mid-Jomon) archaeological sites in Japan, because of a large community of people for a long period. Archaeological studies have shown that the Jomon people inhabited the Sannai-Maruyama site from 5.9-4.2 +/- 0.1 cal. kyr B.P. However, a continuous record of the terrestrial and marine environments around the site has not been available. Core KT05-7 PC-02, was recovered from Mutsu Bay, only 20 km from the site, for the reconstruction of high-resolution time series of environmental records, including sea surface temperature (SST). C37 alkenone SSTs showed clear fluctuations, with four periods of high (8.4-7.9, 7.0-5.9, 5.1-4.1, and 2.3-1.4 cal. kyr B.P.) and four of low (-8.4, 7.9-7.0, 5.9-5.1, and 4.1-2.3 cal. kyr B.P.) SST. Thus, each SST cycle lasted 1.0-2.0 kyr, and the amplitude of fluctuation was about 1.5-2.0 °C. Total organic carbon (TOC) and C37 alkenone contents, and the TOC/total nitrogen ratio indicate that marine biogenic production was low before 7.0 cal. kyr B.P., but was clearly increased between 5.9 and 4.0 cal. kyr B.P., because of stronger vertical mixing. During the period when the community at the site prospered (between 5.9 and 4.2 +/- 0.1 cal. kyr B.P.), the terrestrial climate was relatively warm. The high relative abundance of pollen of both Castanea and Quercus subgen. Cyclobalanopsis supports the interpretation that the local climate was optimal for human habitation. Between 5.9 and 5.1 cal. kyr B.P., in spite of warm terrestrial climates, the C37 alkenone SST was low; this apparent discrepancy may be attributed to the water column structure in the Tsugaru Strait, which differed from the modern condition. The evidence suggests that at about 5.9 cal. kyr B.P, high productivity of marine resources such as fish and shellfish and a warm terrestrial climate led to the establishment of a human community at the Sannai-Maruyama site. Then, at about 4.1 +/- 0.1 cal. kyr B.P., abrupt marine and terrestrial cooling, indicated by a decrease of about 2 °C in the C37 alkenone SST and an increase in pollen of taxa of cooler climates, led to a reduced terrestrial food supply, causing the people to abandon the site. The timing of the abandonment is consistent with the timing (around 4.0-4.3 cal. kyr B.P.) of the decline of civilizations in north Mesopotamia and along the Yangtze River. These findings suggest that a temperature rise of ~2 °C in this century as a result of global warming could have a great impact on the human community and especially on agriculture, despite the advances of contemporary society.
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The impact of an asteroid at the Cretaceous/Paleogene (K/Pg) boundary triggered dramatic biotic, biogeochemical and sedimentological changes in the oceans that have been intensively studied. Paleo-biogeographical differences in the biotic response to the impact and its environmental consequences, however, have been less well documented. We present a high-resolution analysis of benthic foraminiferal assemblages at Southern Ocean ODP Site 690 (Maud Rise, Weddell Sea, Antarctica). At this high latitude site, late Maastrichtian environmental variability was high, but benthic foraminiferal assemblages were not less diverse than at lower latitudes, in contrast to those of planktic calcifiers. Also in contrast to planktic calcifiers, benthic foraminifera did not suffer significant extinction at the K/Pg boundary, but show transient assemblage changes and decreased diversity. At Site 690, the extinction rate was even lower (~3%) than at other sites. The benthic foraminiferal accumulation rate varied little across the K/Pg boundary, indicating that food supply to the sea floor was affected to a lesser extent than at lower latitude sites. Compared to Maastrichtian assemblages, Danian assemblages have a lower diversity and greater relative abundance of heavily calcified taxa such as Stensioeina beccariiformis and Paralabamina lunata. This change in benthic foraminiferal assemblages could reflect post-extinction proliferation of different photosynthesizers (thus food for the benthos) than those dominant during the Late Cretaceous, therefore changes in the nature rather than in the amount of the organic matter supplied to the seafloor. However, severe extinction of pelagic calcifiers caused carbonate supersaturation in the oceans, thus might have given competitive advantage to species with large, heavily calcified tests. This indirect effect of the K/Pg impact thus may have influenced the deep-sea dwellers, documenting the complexity of the effects of major environmental disturbance.
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The Bounty Trough, east of New Zealand, lies along the southeastern edge of the present-day Subtropical Front (STF), and is a major conduit via the Bounty Channel, for terrigenous sediment supply from the uplifted Southern Alps to the abyssal Bounty Fan. Census data on 65 benthic foraminiferal faunas (>63 µm) from upper bathyal (ODP 1119), lower bathyal (DSDP 594) and abyssal (ODP 1122) sequences, test and refine existing models for the paleoceanographic and sedimentary history of the trough through the last 150 ka (marine isotope stages, MIS 6-1). Cluster analysis allows recognition of six species groups, whose distribution patterns coincide with bathymetry, the climate cycles and displaced turbidite beds. Detrended canonical correspondence analysis and comparisons with modern faunal patterns suggest that the groups are most strongly influenced by food supply (organic carbon flux), and to a lesser extent by bottom water oxygen and factors relating to sediment type. Major faunal changes at upper bathyal depths (1119) probably resulted from cycles of counter-intuitive seaward-landward migrations of the Southland Front (SF) (north-south sector of the STF). Benthic foraminiferal changes suggest that lower nutrient, cool Subantarctic Surface Water (SAW) was overhead in warm intervals, and higher nutrient-bearing, warm neritic Subtropical Surface Water (STW) was overhead in cold intervals. At lower bathyal depths (594), foraminiferal changes indicate increased glacial productivity and lowered bottom oxygen, attributed to increased upwelling and inflow of cold, nutrient-rich, Antarctic Intermediate Water (AAIW) and shallowing of the oxygen-minimum zone (upper Circum Polar Deep Water, CPDW). The observed cyclical benthic foraminiferal changes are not a result of associations migrating up and down the slope, as glacial faunas (dominated by Globocassidulina canalisuturata and Eilohedra levicula at upper and lower bathyal depths, respectively) are markedly different from those currently living in the Bounty Trough. On the abyssal Bounty Fan (1122), faunal changes correlate most strongly with grain size, and are attributed to varying amounts of mixing of displaced and in-situ faunas. Most of the displaced foraminifera in turbiditic sand beds are sourced from mid-outer shelf depths at the head of the Bounty Channel. Turbidity currents were more prevalent during, but not restricted to, glacial intervals.
Resumo:
Deep-sea benthic foraminifera show important but transient assemblage changes at the Cretaceous/Paleogene (K/Pg) boundary, when many biota suffered severe extinction. We quantitatively analyzed benthic foraminiferal assemblages from lower bathyal-upper abyssal (1500-2000 m) northwest Pacific ODP Site 1210 (Shatsky Rise) and compared the results with published data on assemblages at lower bathyal (~ 1500 m) Pacific DSDP Site 465 (Hess Rise) to gain insight in paleoecological and paleoenvironmental changes at that time. At both sites, diversity and heterogeneity rapidly decreased across the K/Pg boundary, then recovered. Species assemblages at both sites show a similar pattern of turnover from the uppermost Maastrichtian into the lowermost Danian: 1) The relative abundance of buliminids (indicative of a generally high food supply) increases towards the uppermost Cretaceous, and peaks rapidly just above the K/Pg boundary, coeval with a peak in benthic foraminiferal accumulation rate (BFAR), a proxy for food supply. 2) A peak in relative abundance of Stensioeina beccariiformis, a cosmopolitan form generally more common at the middle than at the lower bathyal sites, occurs just above the buliminid peak. 3) The relative abundance of Nuttallides truempyi, a more oligotrophic form, decreases at the boundary, then increases above the peak in Stensioeina beccariiformis. The food supply to the deep sea in the Pacific Ocean thus apparently increased rather than decreased in the earliest Danian. The low benthic diversity during a time of high food supply indicates a stressed environment. This stress might have been caused by reorganization of the planktic ecosystem: primary producer niches vacated by the mass extinction of calcifying nannoplankton may have been rapidly (<10 kyr) filled by other, possibly opportunistic, primary producers, leading to delivery of another type of food, and/or irregular food delivery through a succession of opportunistic blooms. The deep-sea benthic foraminiferal data thus are in strong disagreement with the widely accepted hypothesis that the global deep-sea floor became severely food-depleted following the K/Pg extinction due to the mass extinction of primary producers ("Strangelove Ocean Model") or to the collapse of the biotic pump ("Living Ocean Model").
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Tropical climate is variable on astronomical time scale, driving changes in surface and deep-sea fauna during the Pliocene-Pleistocene. To understand these changes in the tropical Indian Ocean over the past 2.36 Myr, we quantitatively analyzed deep-sea benthic foraminifera and selected planktic foraminifera from >125 µm size fraction from Deep Sea Drilling Project Site 219. The data from Site 219 was combined with published foraminiferal and isotope data from Site 214, eastern Indian Ocean to determine the nature of changes. Factor and cluster analyses of the 28 highest-ranked species distinguished four biofacies, characterizing distinct deep-sea environmental settings. These biofacies have been named after their most dominant species such as Stilostomella lepidula-Pleurostomella alternans (Sl-Pa), Nuttallides umbonifer-Globocassidulina subglobosa (Nu-Gs), Oridorsalis umbonatus-Gavelinopsis lobatulus (Ou-Gl) and Epistominella exigua-Uvigerina hispido-costata (Ee-Uh) biofacies. Biofacies Sl-Pa ranges from ~2.36 to 0.55 Myr, biofacies Nu-Gs ranges from ~1.9 to 0.65 Myr, biofacies Ou-Gl ranges from ~1 to 0.35 Myr and biofacies Ee-Uh ranges from 1.1 to 0.25 Myr. The proxy record indicates fluctuating tropical environmental conditions such as oxygenation, surface productivity and organic food supply. These changes appear to have been driven by changes in monsoonal wind intensity related to glacial-interglacial cycles. A shift at ~1.2-0.9 Myr is observed in both the faunal and isotope records at Site 219, indicating a major increase in monsoon-induced productivity. This coincides with increased amplitude of glacial cycles, which appear to have influenced low latitude monsoonal climate as well as deep-sea conditions in the tropical Indian Ocean.
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In order to evaluate bioturbation in abyssal Arabian-Sea sediments of the Indus fan profiles of 210Pb (half-life: 22.3 yr) and 234Th (half-life: 24.1 d) were measured in cores collected during September and October 1995 and April 1997, respectively. The density and composition of epibenthic megafauna and lebensspuren were determined in vertical seafloor photographs during April 1997. Mean eddy-diffusive mixing coefficients according to the distribution of excess 210Pb ( 210Pb-DB) were 0.072±0.028, 0.068±0.055, 0.373±0.119, 0.037±0.009 and 0.079±0.119 cm**2 yr**-1 in the northern, western, central, eastern and southern abyssal Arabian sea, respectively. Mean eddy-diffusive mixing coefficients according to the distribution of excess 234Th (234Th-DB) were 0.53, 1.64 and 0.47 cm**2 yr**-1 in the northern, western and central abyssal Arabian Sea, respectively. Mobile epibenthic megafauna at the western, northern, central and southern study sites were dominated by ophiuroids, holothurians, ophiuroids and natant decapods (the respective densities were 100, 82, 29 and 6 individuals 1000 m**-2). The northern study site was characterized by a high abundance of spoke traces and fecal casts. The central site showed spoke traces and many tracks. The southern site displayed the highest abundance of spoke traces, whereas at the western site hardly any lebensspuren were observed. There is evidence for at least two functional endmember communities in the Arabian Sea. In the northwestern Arabian Sea (WAST) vertical particle displacement seems to be dominated by macrofauna and primarily eddy-diffusive. In the southern Arabian Sea (SAST) non-local and 'incidental' mixing due to spoke-trace producers might become more important and superimpose reduced eddy-diffusive mixing. With respect to biological data CAST is an intermediate location. Given the biological data, average 210Pb-DB is higher and decimeter-scale variability of 210Pb-DB smaller at CAST than expected. These findings indicate that in a mixture of both endmember communities the organisms may interact in way that increases values of biodiffusivity, as reflected by 210Pb-DB, and reduces decimeter-scale 210Pb-DB heterogeneity in comparison to the simple sum of the isolated effects of the endmembers. For time scales <100 years there was no evidence for a relationship between food supply (POC flux) and bioturbation intensity, as reflected by 210Pb-DB and 234Th-DB. Bioturbation intensity should be controlled primarily by the composition of the benthic fauna, its specific adaptation to the environmental setting, and the abundance of each species of the benthic community. Food supply can have only an indirect influence on bioturbation intensity. In certain parts of the ocean the a priori overall positive relationship between POC flux and biodiffusivity might include restricted intervals displaying no or even negative relations.
Resumo:
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 order to examine the long-term development of offshore macrozoobenthic soft-bottom communities of the German Bight, four representative permanent stations (MZB-SSd, -FSd, -Slt, -WB) have been sampled continuously since 1969. Inter-annual variability and possible long-term trends were analysed based on spring-time samples from 1969 until 2000. This is part of the ecological long-term series of the AWI and is supplemented by periodic large-scale mapping of the benthos. The main factors influencing the development of the benthic communities are biological interactions, climate, food supply (eutrophication) and the disturbance regime. The most frequent disturbances are sediment relocations during strong storms or by bottom trawling, while occasional oxygen deficiencies and extremely cold winters are important disturbance events working on a much larger scale. Benthic communities at the sampling stations show a large inter-annual variability combined with a variation on a roughly decadal scale. In accordance with large-scale system shifts reported for the North Sea, benthic community transitions occurred between roughly the 1970ies, 80ies and 90ies. The transitions between periods are not distinctly marked by strong changes but rather reflected in gradual changes of the species composition and dominance structure.
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Radiocarbon dating series, bulk sediment, and organic carbon flux from various Atlantic deep-sea regions reveal that the thickness of the bioturbated zone increases by 2 cm if food supply increases by 1 gC/m**2/yr (r = 0.8). Bulk sediment accumulation rates do not influence the depth of bioturbational mixing under normal pelagic sedimentary conditions. We believe that this relationship between nutrient supply and benthic mixing can be used for a quantitative and time-variable unmixing procedure to improve high-resolution stratigraphic correlations and paleoclimatic interpretations of deep-sea records.
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Canonical correspondence analysis indicates that the distribution of Neogene benthic foraminiferal faunas (>63 µm) in seven DSDP and ODP sites (500-4500 m water depth) east of New Zealand (38-51°S, 170°E-170°W) is most strongly influenced by depth (water mass stratification), and secondly by age (palaeoceanographic changes influencing faunal composition and biotic evolution). Stratigraphic faunal changes are interpretted in terms of the pulsed sequential development of southern, and later northern, polar glaciation and consequent cooling of bottom waters, increased vertical and lateral stratification of ocean water masses, and increased overall and seasonal surface water productivity. Oligocene initiation of the Antarctic Circumpolar Current and Deep Western Boundary Current (DWBC), flowing northwards past New Zealand, resulted in extensive hiatuses throughout the Southwest Pacific, some extending through into the Miocene. Planktic foraminiferal fragmentation index values indicate that carbonate dissolution was significant at abyssal depths throughout most of the Neogene, peaking at upper abyssal depths in the late Miocene (11-7 Ma), with the lysocline progressively deepened thereafter. Miocene abyssal faunas are dominated by Globocassidulina subglobosa and Oridorsalis umbonatus, with increasing Epistominella exigua after 16 Ma at upper abyssal depths. Peak abundances of Epistominella umbonifera indicate increased input of cold Southern Component Water to the DWBC at 7-6 Ma. Faunal association changes imply establishment of the modern Oxygen Minimum Zone (upper Circumpolar Deep Water) in the latest Miocene. Significant latitudinal differences between the benthic foraminiferal faunas at lower bathyal depths indicate the existence of an oceanic front along the Chatham Rise (location of present Subtropical Front), since the early late Miocene at least, with more pulsed productivity (higher E. exigua) along the south side. Modern Antarctic Intermediate Water faunal associations were established north of the Chatham Rise at 10-9 Ma, and south of it at 3-1.5 Ma. Middle-upper bathyal faunas on the Campbell Plateau are dominated by reticulate bolivinids during the early and middle Miocene, indicative of sustained productivity above relatively sluggish, suboxic bottom waters. Faunal changes and hiatuses indicate increased current vigour over the Campbell Plateau from the latest Miocene on. Surface water productivity (food supply) appears to have increased in three steps (at times of enhanced global cooling) marked by substantially increased relative abundance of: (1) Abditodentrix pseudothalmanni, Alabaminella weddellensis, Cassidulina norvangi (16-15 Ma, increased pulsed productivity); (2) Bulimina marginata f. aculeata, Nonionella auris, Trifarina angulosa, Uvigerina peregrina (3-1.5 Ma, increased overall productivity); and (3) Cassidulina carinata (1-0.5 Ma, increased overall productivity). Three intervals of deep-sea benthic foraminiferal taxonomic turnover are recognised (16-15, 11.5-10, 2-0.5 Ma) corresponding to intervals of enhanced global cooling and possible productivity changes. The late Pliocene-middle Pleistocene extinction, associated with increasing Northern Hemisphere glaciation, culminating in the middle Pleistocene climatic transition, was more significant in the study area than the earlier Neogene turnovers.