Cenozoic deep-sea ostracods from Maud Rise, Weddell Sea, Antarctica (ODP Site 689): a palaeoceanographical perspective

Cenozoic palaeoceanography of the Maude Rise, Weddell Sea, Antarctica, has been investigated using Palaeocene to Quaternary deep-sea ostracod faunas from 23 samples of ODP Site 689. The abundance of ostracods is high enough only during the Palaeogene (Palaeocene-Oligocene) to allow palaeoceanographical inferences based on changes in diversity, dominance, endemism and faunal turnover (first and last occurrences). The abundance is particularly high throughout the Palaeocene and Eocene, but declines irreversibly near the Eocene/Oligocene boundary. The diversity increases more or less continuously from the Early Palaeocene to the Middle Eocene, and then it generally decreases throughout the remaining part of the Palaeogene (Middle Eocene-Oligocene); an exception is a positive peak in the Shannon-Weaver index in a single sample in the Late Oligocene. No positive peaks in diversity and taxa originations (first occurrences) at c. 40-38 Ma, occurs at Site 689; so the site provides no evidence for the establishment of the psychrosphere at this time. This corroborates similar regional results from an earlier study of benthonic foraminifera. Explanations for this may be related to Late Eocene-Early Oligocene changes in sedimentology and clay-mineralogy (associated with the progressive cooling of the Antarctica) which could have negatively affected abundance and diversity locally at Site 689. Alternatively, by this time, the ostracod fauna could also have been subjected to selective removal (with possible local extinction) of taxa (due to increased ventilation) or to thanatocoenosis dissolution (due to a decrease in temperature and availability of CaCO₃). A further possibility may be related to the fact that Site 689 was at intermediate water depths and may have remained within older water masses near the Eocene/Oligocene boundary. Failing these explanations, the results could indicate that the Late Eocene-Early Oligocene palaeoenvironmental changes in the world oceans were more gradual and occurred over a longer time interval than the global ostracod data show, at least at southern high latitudes.     

Biochronology and paleoclimatic implications of Middle Eocene to Oligocene planktic foraminiferal faunas

Planktic foraminiferal assemblages have been analyzed quantitatively in six DSDP sites in the Atlantic (Site 363), Pacific (Sites 292, 77B, 277), and Indian Ocean (Sites 219, 253) in order to determine the nature of the faunal turnover during Middle Eocene to Oligocene time. Biostratigraphic ranges of taxa and abundance distributions of dominant species are presented and illustrate striking similarities in faunal assemblages of low latitude regions in the Atlantic, Pacific and Indian oceans. A high resolution biochronology, based on dominant faunal characteristics and 55 datum events, permits correlation between all three oceans with a high degree of precision. Population studies provide a view of the global impact of the paleoclimatic and paleoceanographic changes occurring during Middle Eocene to Oligocene time.Planktic foraminiferal assemblage changes indicate a general cooling trend between Middle Eocene to Oligocene time, consistent with previously published oxygen isotope data. Major faunal changes, indicating cooling episodes, occur, however, at discrete intervals: in the Middle Eocene 44-43 Ma (P13), the Middle/Late Eocene boundary 41-40 Ma ( ), the Late Eocene 39-38 Ma ( ), the Eocene/Oligocene boundary 37-36 Ma (P18), and the Late Oligocene 31-29 Ma ( ). With the exception of the boundary, faunal changes occur abruptly during short stratigraphic intervals, and are characterized by major species extinctions and first appearances. The Eocene/Oligocene boundary cooling is marked primarily by increasing abundances of cool water species. This suggests that the boundary cooling, which marks a major event in the oxygen isotope record affected planktic faunas less than during other cooling episodes. Planktic foraminiferal faunas indicate that the boundary event is part of a continued cooling trend which began during the Middle Eocene.Two hiatus intervals are recognized in low and high latitude sections at the Middle/Late Eocene boundary and in the Late Eocene ( ). These hiatuses suggest that vigorous bottom water circulation began developing in the Middle Eocene, consistent with the onset of the faunal cooling trend, and well before the development of the psychrosphere at the boundary.     

PLEISTOCENE EXTINCTIONS OF DEEP-SEA BENTHIC FORAMINIFERA: THE SOUTH ATLANTIC RECORD

Abstract:  Sixty-two species and 19 genera of elongate, cylindrical benthic foraminifera disappeared from the deep-sea in the south-east Atlantic (ODP Sites 1082 and 1083) and the Atlantic sector of the Southern Ocean (ODP Site 1088) during the Early and Middle Pleistocene as part of the global extinction of the families Pleurostomellidae, Stilostomellidae and portions of the Nodosariidae. During the mid-Pleistocene Climate Transition (1·2–0·6 Ma) in the Southern Ocean, these extinct taxa exhibited three pulses of glacial decline in abundance and diversity separated by partial interglacial recoveries. Beneath the high-productivity Benguela Current upwelling region (Sites 1082, 1083), glacial declines in the extinct taxa were suppressed by favourable high organic-carbon flux and consequent low-oxygen bottom conditions. Here two major pulses of diversity loss occurred at c.  1·3–1·2 Ma and 1·0–0·7 Ma. At all three locations, the most dramatic decline in abundance and diversity occurred c.  0·85–0·80 Ma (marine isotope stage 20), and the final disappearance of Extinction Group taxa was completed by 0·67 Ma beneath the Benguela Current and 0·60 Ma in the Southern Ocean. We speculate that this period of enhanced global extinctions was linked to a pulsed decline in glacial temperatures and/or increase in ventilation of deep and intermediate water masses, associated with polar ice cap growth since the late Pliocene.     

Benthic ostracods and deep water-masses in the Atlantic Ocean

Ostracod faunas at six locations are compared, and related to distributions in an Atlantic Ocean-wide data base. Five, widely developed, vertical faunal sequences are recognised at particular levels within deep water-masses: Henryhowella Fauna (lower part of Antarctic Intermediate Water); Krithe Fauna (Upper North Atlantic Deep Water); Poseidonamicus-Bosquetina Fauna (upper part of Lower NADW); Dutoitella Fauna (lower part of Lower NADW); Legitimocythere Fauna (Antarctic Bottom Water). These faunas are correlated with previously established deep water benthic foraminiferal assemblages, and their possible palaeo-oceanographic use is discussed.     

Pliocene benthic foraminifera from the Blake Plateau: Faunal assemblages and paleocirculation

Relative abundance of benthic foraminifera have been analyzed from core V26-145 from the Blake Plateau. The investigated sequence represents the time interval between 1.8 and 4.6 Ma. In order to determine how different sieve sizes influence the relative abundance patterns, three sediment size fractions were studied separately. It becomes difficult to maintain consistent taxonomic concepts in the fraction 63–125 μm, partly because this fraction contains high abundances of juvenile forms. However, the 63–125 μm fraction holds high abundances of the important small speciesEpistominella exigua. Due to these reasons only the two larger fractions (125–250 μm and >250 μm) were considered meaningful to analyze for relative abundance patterns. An analysis of the two larger fractions (>125 μm; >250 μm) shows no consistency in relative abundance patterns.The relative abundance patterns for the 34 most common species in the size fraction >125 μm were analyzed by means of correspondence analysis. Three benthic foraminiferal assemblages (I, II, and III) were recognized and these can be associated with water masses. Assemblage I is associated with the Florida Current and consists of shallow water species (Amphistegina gibbosa, Compressigerina sp. A,Discorbinella biconcavus, Islandiella teretis, Reussella atlantica, andSiphonina pulchra). Assemblage II contains key species for North Atlantic Deep Water (NADW) (Cibicidoides kullenbergi, Epistominella exigua, Globocassidulina subglobosa, Lenticulina peregrina, Oridorsalis umbonatus, andPlanulina wuellerstorfi). The third assemblage (III) contains species associated with the Antilles Current (Bolivina rhomboidalis, Cassidulina obtusa, Cassidulina vortex, andNuttallides umbonifera). The correspondence analysis reveals an alternation in dominance between Assemblage I and Assemblage II prior to 3.3 Ma, suggesting lateral oscillations between the Florida Current and NADW. At about 3.3 Ma Assemblage I disappears and Assemblage III increases in importance, suggesting an increasing influence of the Antilles Current in the upper part of the record.     

Benthic foraminiferal extinctions linked to late Pliocene–Pleistocene deep-sea circulation changes in the northern Indian Ocean (ODP Sites 722 and 758)

During the late Pliocene–middle Pleistocene, 63 species of elongate, bathyal–upper abyssal benthic foraminifera (Extinction Group = Stilostomellidae, Pleurostomellidae, some Nodosariidae) declined in abundance and finally disappeared in the northern Indian Ocean (ODP Sites 722, 758), as part of the global extinction of at least 88 related species at this time. The detailed record of withdrawal of these species differs by depth and geography in the Indian Ocean. In northwest Indian Ocean Site 722 (2045 m), the Extinction Group of 54 species comprised 2–15% of the benthic foraminiferal fauna in the earliest Pleistocene, but declined dramatically during the onset of the mid-Pleistocene Transition (MPT) at 1.2–1.1 Ma, with all but three species disappearing by the end of the MPT (∼0.6 Ma). In northeast Indian Ocean Site 758 (2925 m), the Extinction Group of 44 species comprised 1–5% of the benthic foraminiferal fauna at ∼3.3–2.6 Ma, but declined in abundance and diversity in three steps, at ∼2.5, 1.7, and 1.2 Ma, with all but one species disappearing by the end of the MPT. At both sites there are strong positive correlations between the accumulation rate of the Extinction Group and proxies indicating low-oxygen conditions with a high organic carbon input. In both sites, there was a pulsed decline in Extinction Group abundance and species richness, especially in glacial periods, with some partial recoveries in interglacials. We infer that the glacial declines at the deeper Site 758 were a result of increased production of colder, well-ventilated Antarctic Bottom Water (AABW), particularly in the late Pliocene and during the MPT. The Extinction Group at shallower water depths (Site 722) were not impacted by the deeper water mass changes until the onset of the MPT, when cold, well-ventilated Glacial North Atlantic Intermediate Water (GNAIW) production increased and may have spread into the Indian Ocean. Increased chemical ventilation at various water depths since late Pliocene, particularly in glacial periods, possibly in association with decreased or more fluctuating organic carbon flux, might be responsible for the pulsed global decline and extinction of this rather specialised group of benthic foraminifera.     

Diversity and extinction in the Cenozoic history of Caribbean reefs

Occurrences of reef corals are examined at Caribbean fossil localities to determine how biodiversity has changed within the region over the past 50 million years. Analyses of 294 species (66 genera) at 58 fossil localities show that Caribbean generic diversity rose to 44 between 50–22 Ma, ranged from 32–39 between 22–2 Ma, and dropped to 25 afterwards. Regional species diversity was high at 40–36 Ma, 28–22 Ma, and 5–2 Ma. Origination rates were elevated throughout each high diversity interval, but extinction was concentrated near the end of each interval. Regional highs of origination and extinction, therefore, differed in timing and duration, causing the observed regional diversity increases during the three remarkably long intervals of turnover. Highs of generic origination decreased in magnitude as immigration from the Mediterranean ceased, but speciation highs increased in association with emergence of the Central American isthmus. Peaks of extinction coincided with regional changes in climate and oceanic circulation. Maximum species diversities within assemblages increased to 40–60 between 50–36 Ma, and have remained relatively constant ever since. Assemblage compositions differed among localities having similar ages and environments, suggesting that the timing and pattern of turnover varied across the region. Stable diversities but variable compositions within assemblages suggest that dispersal and recruitment influenced the pattern of faunal change during turnover. Accepted: 22 August 1999     

Impact of the Middle Miocene climate transition on elongate,cylindrical foraminifera in the subtropical Pacific

Fifty-eight species of elongate, cylindrical benthic foraminifera (here referred to as the Extinction Group) belonging to genera that became extinct during the mid-Pleistocene Climate Transition (MPT), were documented (~ 50 kyr resolution) through the early middle Miocene (15–13 Ma) in two sites on opposite sides of the subtropical Pacific Ocean (ODP Sites 1146, South China Sea; ODP Site 1237, southeast Pacific). The study was undertaken to investigate the response of the Extinction Group (Ext. Gp) to the major cooling during the middle Miocene Climate Transition (MCT) to look for clues that might explain the causes of the extinction during the glacials of the mid-Pleistocene Climate Transition. Ext. Gp faunal differences between the two sites (attributed to regional and bathymetric differences in food supply to the seafloor) are greater than those that occurred through the 2 myr time span at either site. The middle Miocene Climate Transition was not an interval of enhanced species turnover or a decline in Ext. Gp abundance, in contrast to the major extinctions that occurred during the mid-Pleistocene Climate Transition. Distinct changes in the composition of the Ext. Gp faunas did occur through this time (more pronounced in Site 1237). At both sites the pre-middle Miocene Climate Transition faunas were transformed into their post-middle Miocene Climate Transition composition during the period of major cooling (14.0–13.7 Ma). During this transition interval the faunal composition swung back and forth between the two end member faunas. These faunal changes are attributed to changes in productivity (decrease in South China Sea, increase in southeast Pacific), brought about by major changes in global climate and continental aridity.     

Benthic foraminiferal response to the emergence of the Isthmus of Panama and coincident paleoceanographic changes

Late Cenozoic benthic foraminiferal faunas from the Caribbean Deep Sea Drilling Project (DSDP) Site 502 (3052 m) and East Pacific DSDP Site 503 (3572 m) were analyzed to interpret bottom-water masses and paleoceanographic changes occurring as the Isthmus of Panama emerged. Major changes during the past 7 Myr occur at 6.7–6.2, 3.4, 2.0, and 1.1 Ma in the Caribbean and 6.7–6.4, 4.0–3.2, 2.1, 1.4, and 0.7 Ma in the Pacific. Prior to 6.7 Ma, benthic foraminiferal faunas at both sites indicate the presence of Antarctic Bottom Water (AABW). After 6.7 Ma benthic foraminiferal faunas indicate a shift to warmer water masses: North Atlantic Deep Water (NADW) in the Caribbean and Pacific Deep Water (PDW) in the Pacific. Flow of NADW may have continued across the rising sill between the Caribbean and Pacific until 5.6 Ma when the Pacific benthic foraminiferal faunas suggest a decrease in bottom-water temperatures. After 5.6 Ma deep-water to intermediate-water flow across the sill appears to have stopped as the bottom-water masses on either side of the sill diverge.The second change recorded by benthic foraminiferal faunas occurs at 3.4 Ma in the Caribbean and 4.0-3.2 Ma in the Pacific. At this time the Caribbean is flooded with cold AABW, which is either gradually warmed or is replaced by Glacial Bottom Water (GBW) at 2.0 Ma and by NADW at 1.1 Ma. These changes are related to global climatic events and to the depth of the sill between the Caribbean and Atlantic rather than the rising Isthmus of Panama. Benthic foraminiferal faunas at East Pacific Site 503 indicate a gradual change from cold PDW to warmer PDW between 4.0 and 3.2 Ma. The PDW is replaced by the warmer, poorly oxygenated PIW at 2.1 Ma. Although the PDW affects the faunas during colder intervals between 1.4 and 0.7 Ma, the PIW remains the principal bottom-water mass in the Guatemala Basin of the East Pacific.     

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.     

Deep-sea benthonic foraminiferal faunal turnover near the Eocene/Oligocene boundary

Eocene-Oligocene deep-sea benthonic foraminifera in D.S.D.P. Site 277 in the southwest Pacific have been analyzed to determine the benthonic foraminiferal response to the development of the psychrosphere near the Eocene/Oligocene boundary. Biostratigraphic ranges of 41 taxa show that 23 taxa are found throughout the Late Eocene to Early Oligocene sequence, while 18 taxa exhibit first or last occurrences. Comparison of the faunal changes in Site 277 with a benthonic foraminiferal oxygen isotope record shows that the development of the psychrosphere did not have a profound effect upon the benthonic foraminifera, and the overall faunal change preceding and subsequent to the bottom-water circulation event occurred gradually. The inferred water-mass event affected the relative abundance of one species, Epistominella umbonifera. The lack of major faunal changes at the Eocene/Oligocene boundary in Site 277 probably reflects either wide environmental tolerances of the benthonic foraminifera, or a bottom-water temperature change less than 3°C.Examination of previously published benthonic foraminiferal biostratigraphic data from D.S.D.P. Sites 167, 171, 357, 360, 363, and 400A, and deep-sea ostracode data from D.S.D.P. Leg 3 show faunal changes occurred during discrete intervals in the Middle Eocene-Early Oligocene. The faunal patterns from these data and from Site 277 show that the Eocene/Oligocene cooling event did not cause rapid, catastrophic changes of the benthonic faunas of the open ocean, although significant faunal changes are associated with the water mass event in Sites 167, 171 and 400A.The benthonic faunal changes in Middle Eocene-Early Oligocene time are consistent with the gradual decrease of inferred bottom-water temperatures, based on previously published oxygen isotopic data. The δ ¹⁸O Eocene/Oligocene enrichment of 0.76‰ is a major event in the Southern Ocean oxygen isotopic record, but is considerably less in magnitude than the 1.75-2.00‰ change that occurred gradually from mid-Early Eocene to the Eocene/Oligocene boundary. The benthonic foraminiferal and isotopic data indicate that bottom-water circulation may have developed during the Middle Eocene to Early Oligocene interval, with the 3°C bottom-water cooling near the Eocene/Oligocene boundary representing part of this development.     

Late Cretaceous and Cainozoic bathyal Ostracoda from the Central Pacific (DSDP Site 463)

Cainozoic deep-sea ostracod assemblages from the summits of Mid-Pacific guyots point to high levels of endemism possibly as a result of their bathymetric separation from the surrounding sea floor. However, the interpretation of these fossil assemblages is hampered by the paucity of comparative material from surrounding non-guyot sites. Fifteen ostracod assemblages from DSDP Site 463 (Late Cretaceous-Pleistocene) were studied to compare with those from nearby guyots. Three distinct faunal assemblages are recognised at Site 463: Assemblage A (Maastrichtian-Eocene), Assemblage B (Oligocene-Upper Miocene) and Assemblage C (Upper Miocene-Pleistocene) although the palaeoenvironmental significance of these units is unclear. Sixty-two ostracod species are identified, the thirteen most abundant are discussed in the taxonomic section, five of which are described as new. Between 30 and 100% of the species encountered in each sample are considered as endemic to Site 463, while some of the remaining species were previously thought to be endemic to individual guyots. Similarly high levels of endemism on nearby guyots probably reflect an incomplete knowledge of deep-sea ostracod faunas rather than the establishment of geographically or bathymetrically restricted populations. The presence of globally pandemic and geographically widespread taxa on sites such as the Mid-Pacific Mountains, surrounded by abyssal depths which lie below the CCD, indicates that some faunal exchange or migration of ostracods does take place. This must be achieved within the intermediate waters and probably occurs passively.     

Paleocene-Pleistocene benthic foraminiferal evidence of major paleoceanographic events in the eastern South Atlantic (DSDP Site 525, Walvis Ridge)

Benthic foraminifers in the size-fraction greater than 0.073 mm were studied in 88 Paleocene to Pleistocene samples from Deep Sea Drilling Project Site 525 (Hole 525A, Walvis Ridge, eastern south Atlantic). Clustering of the samples on the basis of the 86 most abundant foraminifers (in total, 331 taxa were identified) allowed separating two major assemblage zones: the Paleocene to Eocene interval, and the Oligocene to Pleistocene interval. Each of these, in turn, were subdivided into three minor subzones as follows: lower upper Paleocene (approx. 62.4 to 57.8 Ma); upper upper Paleocene (56.6 to 56.2 Ma); lower and middle Eocene (55.3 to 46.8 Ma); upper Oligocene to middle Miocene (25.3 to 16 Ma); middle Miocene to Pliocene (15.7 to 4.2 Ma); and lower Pleistocene (0.4 to 0.02 Ma), with only minor differences with the previous zone. Some very abundant taxa span most of the column studies (Bolivina huneri, Cassidulina subglobosa, Eponides bradyi, E. weddellensis, Gavelinella micra, Oridorsalis umbonatus, etc.). Several of the faunal breaks recorded coincide with conspicuous minima in the specific diversity curve, thus suggesting that the corresponding turnovers signal the final stages of periods of faunal impoverishment. At least one major bottom-water temperature drop (as derived from δ¹⁸O data) is synchronous with a decrease in the foraminiferal specific diversity. On the other hand, a specific diversity maximum in the middle Miocene might be associated with a δ¹³C increase at approx. 16 to 12 Ma. Highest foraminiferal abundances (up to 600–800 individuals per gram of dry sediment) occurred in the late Paleocene and in the early Pleistocene, in coincidence with the lowest diversity figures calculated. The magnitude of the most important faunal turnover recorded, between the middle Eocene and the late Oligocene, is magnified in our data set by the large hiatus which separates the middle Eocene from the upper Oligocene sediments. Considerably smaller overturns occurred within the late Paleocene (in coincidence with changes in the specific diversity, absolute abundance of foraminiferal tests, and δ¹³C), and in the middle Miocene (in coincidence with a specific diversity maximum and a δ¹³C excursion). New information on the morphology and the stratigraphic ranges of several species is furnished. For all the taxa recorded the number of occurrences, total number of individuals identified and first and last appearances are listed.     

The Middle Miocene climatic transition in the Southern Ocean: Evidence of paleoclimatic and hydrographic changes at Kerguelen plateau from planktonic foraminifers and stable isotopes

Middle Miocene (14.8–11.9 Ma) deep-sea sediments from ODP Hole 747A (Kerguelen Plateau, southern Indian Ocean) contain abundant, well-preserved and diverse planktonic foraminiferal assemblages. A detailed study of the climatic and hydrographic changes that occurred in this region during the Middle Miocene Climatic Transition led to the identification of an intense cooling phase (the Middle Miocene Shift). Abundance fluctuations of planktonic foraminiferal species with different paleoclimatic affinities, and oxygen and carbon stable isotopes have been integrated in a multi-proxy approach. Reconstruction of changes in foraminiferal faunal composition and diversity through time were the basis for identification of three foraminiferal biofacies. The most prominent faunal change took place at 13.8 Ma, when a fauna with warm-water affinity (marked by high abundance of Globorotalia miozea group and Globoturborotalita woodi plexus) was replaced by an oligotypic, opportunistic fauna with typical polar characters and dominated by neogloboquadrinids. This faunal change is interpreted as the result of foraminiferal migration from adjacent bioprovinces, caused by modifications in climate and hydrography. A positive 2.0‰ shift in δ¹⁸O (interpreted as the Mi3 event) and a related positive 1.0‰ shift in δ¹³C (corresponding to the CM6 event) accompanied this faunal turnover. These are interpreted to reflect substantial reorganization of Southern Ocean waters, the northward migration of the Polar Front and a strong increase in primary productivity. The second faunal change took place at 12.9 Ma and was characterized by the gradual decrease in abundance of the neogloboquadrinids and the recovery of Globorotalia praescitula/scitula group and Globigerinita glutinata. A positive 1.5‰ shift in δ¹⁸O (interpreted as the Mi4 event) and a concurrent gradual negative shift in δ¹³C accompanied this faunal change, witnessing further modifications of the climate/ocean system. Variations in sea surface temperature, considered as the main factor causing changes of surface hydrography at the Kerguelen Plateau, seem to have been driven by obliquity and long-term eccentricity, thus suggesting a key role played by the astronomical forcing on the evolution of Southern Ocean dynamics during the Middle Miocene. Also an evident 1.2 Myr modulation of the δ¹³C record suggests a main control of the long-term obliquity cycles on the carbon cycle dynamics. Particularly, the Mi3/CM6 events exactly fit with a node of the 1.2 Myr modulation cycles. This confirms the key role played by orbital parameters on high-latitude temperatures and Antarctic ice volume, and indirectly on global carbon burial and/or productivity. This climatic transition was marked also by changes in surface hydrography. From 14.8 to 13.8 Ma an intermediate-strength thermocline controlled by seasonality developed just below the photic zone. Weaker seasonality characterized the interval from 13.8 to 12.9 Ma, when the thermocline became shallower and sharper and favored intermediate-water foraminifers. From 12.9 Ma, seasonality increased again and an intermediate-strength thermocline re-developed.     

Differential extinction and the contrasting structure of polar marine faunas

Background

The low taxonomic diversity of polar marine faunas today reflects both the failure of clades to colonize or diversify in high latitudes and regional extinctions of once-present clades. However, simple models of polar evolution are made difficult by the strikingly different faunal compositions and community structures of the two poles.

Methodology/Principal Findings

A comparison of early Cenozoic Arctic and Antarctic bivalve faunas with modern ones, within the framework of a molecular phylogeny, shows that while Arctic losses were randomly distributed across the tree, Antarctic losses were significantly concentrated in more derived families, resulting in communities dominated by basal lineages. Potential mechanisms for the phylogenetic structure to Antarctic extinctions include continental isolation, changes in primary productivity leading to turnover of both predators and prey, and the effect of glaciation on shelf habitats.

Conclusions/Significance

These results show that phylogenetic consequences of past extinctions can vary substantially among regions and thus shape regional faunal structures, even when due to similar drivers, here global cooling, and provide the first phylogenetic support for the “retrograde” hypothesis of Antarctic faunal evolution.     

Middle to late Miocene fluctuations in the incipient Benguela Upwelling System revealed by calcareous nannofossil assemblages (ODP Site 1085A)

Middle to late Miocene calcareous nannofossil data of ODP Site 1085 from the eastern South Atlantic off Namibia were analysed to document spatial and temporal changes in surface-ocean circulation, upwelling initiation, and associated productivity.

Our data show that calcareous nannofossils constitute a significant part of the carbonate fraction throughout the investigated interval from 12.5 to 7.7 million years (Ma). Highest numbers of calcareous nannofossils (up to 38,000 × 10⁶ nannofossils g^− 1 sediment) were observed during the intervals 9.9 to 9.7 and 8.7 to 8.0 Ma. These elevated numbers of calcareous nannofossils may generally be linked to the initiation of upwelling at about 10 Ma in the studied region. In contrast, diminished numbers of calcareous nannoplankton, as in the interval 9.6 to 9.0 Ma, probably characterise time intervals of weaker productivity resulting in a decrease of nannofossil carbonate contents in the sediments of Site 1085. This decrease in nannofossil production could be one possible explanation for the major CaCO₃ depression in between 9.6 and 9.0 Ma. Coccoliths of the genus Reticulofenestra are the most abundant taxa. Their occurrences are characterised by changes in the investigated time interval. In addition, Coccolithus pelagicus, Calcidiscus leptoporus and Umbilicosphaera spp. contribute a common part of the assemblage. Calcareous nannofossils account for more than half of the carbonate, with peak contribution up to 80% at 8.8 Ma.     

Bacteria beneath the West Antarctic Ice Sheet

Subglacial environments, particularly those that lie beneath polar ice sheets, are beginning to be recognized as an important part of Earth's biosphere. However, except for indirect indications of microbial assemblages in subglacial Lake Vostok, Antarctica, no sub-ice sheet environments have been shown to support microbial ecosystems. Here we report 16S rRNA gene and isolate diversity in sediments collected from beneath the Kamb Ice Stream, West Antarctic Ice Sheet and stored for 15 months at 4°C. This is the first report of microbes in samples from the sediment environment beneath the Antarctic Ice Sheet. The cells were abundant (∼10⁷ cells g⁻¹) but displayed low diversity (only five phylotypes), likely as a result of enrichment during storage. Isolates were cold tolerant and the 16S rRNA gene diversity was a simplified version of that found in subglacial alpine and Arctic sediments and water. Although in situ cell abundance and the extent of wet sediments beneath the Antarctic ice sheet can only be roughly extrapolated on the basis of this sample, it is clear that the subglacial ecosystem contains a significant and previously unrecognized pool of microbial cells and associated organic carbon that could potentially have significant implications for global geochemical processes.     

Neoplasia detection in Macoma balthica from the Gulf of Gdansk: comparison of flow cytometry, histology and chromosome analysis

A flow cytometry protocol was applied for the detection of neoplasia in Macoma balthica L. from the Gulf of Gdansk (Baltic Sea, Poland). A simple method, based on an osmotic shock, was used to permeabilise gill cells. The cytometric pattern of normal clams consisted of 2 peaks, a major peak B and a smaller peak C. The cytometric pattern of affected clams consisted of 2 peaks named B' and C'. Two parameters were used to define the stages of abnormalities in M. balthica clams based on the percentage of cells in peaks B, C, B' and C' and on the ratio between the fluorescence value of peaks B, C, B' and C' in all individuals. Three stages of neoplasia were clearly distinguished by flow cytometry considering peak C'. Stage 1 was characterised by a major population of cells in peak B' and more than 10% of cells in the C' peak. Stage 2 consisted of a lower percentage of cells in peak B' and more than 25% of cells in peak C'. Stage 3 of the neoplasia was characterised by a further reduction in peak B' and more than 40% of cells in peak C'. Flow cytometry allowed for objective detection of neoplasia and provided a rapid method for measuring the DNA content of thousands of cells per individual. The accuracy of flow cytometry was assessed by comparing with standard histological techniques, used here as a reference technique for the detection of neoplasia, and with chromosome analysis. All individuals were analysed in parallel using the 3 techniques. The proportion of normal and affected individuals diagnosed using flow cytometry was comparable to the proportion determined by histology and chromosome analysis.     

Uvigerina proboscidea abundances and paleoceanography of the northern Indian Ocean DSDP Site 214 during the Late Neogene

This study attempts to understand the significance of Uvigerina proboscidea in paleoceanographic reconstructions at the northern (tropical) Indian Ocean DSDP Site 214 from the Late Miocene through the Pleistocene. In this interval at this site, U. proboscidea is the most abundant species of the benthic assemblage and shows abrupt frequency changes (about 1–74%). Based on relative percentages of U. proboscidea calibrated with oxygen and carbon isotope record and the sediment accumulation rates, the modern distribution of the species in the Indian Ocean, and other evidence, the peaks of abundance of U. proboscidea are inferred to represent times of high-surface productivity. This productivity is related to intensified trade winds during strong southwest (SW) Indian monsoons, causing widespread upwelling along equatorial divergence in the Indian Ocean. The sudden increase of U. proboscidea abundance at approximately 8.5–7.5 Ma reflects significant upwelling at the equatorial divergence. This event corresponds to the permanent build-up of West Antarctic ice sheets, and a major increase in SW Indian monsoons related upwelling in the northwestern Indian Ocean. The Chron-6 carbon shift at approximately 6.2 Ma is marked by another peak of abundance, reflecting widespread ocean fertility. The highest abundances of U. proboscidea and highest sediment accumulation rates occur between 5.8 and 5.1 Ma, which coincides with the greatest development of Antarctic ice sheets and strong southwest monsoons. The higher percentages at 3.2–3.1 Ma, approximately 2.4 Ma, and 1.6 Ma all represent phases of high productivity at the equatorial divergence.     

Late Miocene—early Pliocene planktonic foraminiferal biostratigraphy and paleoceanography of low-latitude marine sequences

Plate reorganization and development of polar glaciation are closely associated with the changing climatic conditions of the Cenozoic. The planktonic foraminiferal fauna in three low-latitude DSDP sites (224, Ninety East Ridge; 317B, Manihiki Plateau; 366A, Sierra Leone Rise) has been examined to determine how these regions responded to the late Miocene climatic cooling that has been previously observed in high-latitude regions. It has been proposed that an expansion of Antarctic glaciation and a resultant eustatic regression are associated with this cooling, with the latter being at least partially responsible for the Messinian “salinity crisis” in the Mediterranean.Only one of the sites, 366A, shows any significant faunal change during the late Miocene and early Pliocene (approximately 8-3 Ma). During the late Miocene, there is a decrease in the abundance of species considered to be tropical-subtropical, suggesting the incursion of a cool water mass into the Sierra Leone Rise region during this time. The lack of any major faunal changes at Sites 214 and 317B indicates that the water masses of the low-latitude regions of the Indo-Pacific were relatively unaffected by this cooling in the late Miocene.The three sites also show no evidence for a change in the level of the CCD between the late Miocene and early Pliocene; however, this is most likely due to their equatorial position and shallow water depth. Higher-latitude sites from both the Atlantic and the Pacific reveal a definite shoaling of the CCD during the late Miocene.