Reconstruction of Indian monsoon variability over the past 230,000 years: Planktic foraminiferal evidence from the NW Arabian Sea open-ocean upwelling area

We determined the faunal composition and total number of tests (#/g) of planktic foraminifera (> 125 μm) in core KH00-05 GOA 6 near Oman in order to decipher monsoon-induced variability of oceanographic productivity in the open-ocean upwelling area in the northwest Arabian Sea. The core contains a continuous record of sedimentation over the last 230 kyr, with the age model based on oxygen isotope and accelerator mass spectrometry ¹⁴C dates. We focused on species (Globigerina bulloides and Globigerinita glutinata) typical for SW monsoonal upwelling and species typical for NE monsoon conditions (Neogloboquadrina incompta, Neogloboquadrina dutertrei, Globigerinoides ruber, and Globigerinoides sacculifer). The changes in relative abundance of these monsoonal indicators suggest that the open-ocean upwelling area was dominated by the SW monsoon during interglacial periods, but by the NE monsoon during glacial periods.Increases in total test abundance during glacial periods confirmed that the NE monsoon rather than SW monsoon contributes largely to planktic foraminiferal productivity in this area. We argue that three types of circumstances resulted in high productivity, with nine high productivity events occurring at a 23-kyr frequency. The first type caused high productivity events at 102 and 199 ka (interglacial periods), characterized by the dominance of upwelling species, indicating high productivity during strong SW monsoons, correlated with high July insolation at 45° N. An exceptional high productivity event occurred at 37 ka during interglacial marine isotope stage (MIS) 3, with contributions from both SW and NE monsoons. The second type of high productivity event occurred at 61, 147, and 175 ka, during glacial periods, characterized by dominance of NE monsoon species, and correlated with low January insolation at 45° N. In addition, a high productivity event at 85 ka (interglacial period) also was induced by enhanced NE monsoons. The last two high productivity events occurred during transitional periods from glacial to interglacial (MIS 6/5.5 and 2/1), were characterized by the replacement of NE monsoon species with upwelling species, and corresponded to abrupt climate warming, suggesting that they are related to both accelerated SW monsoon systems and reduced NE monsoon systems.     

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.     

Speciation chronology of rockhopper penguins inferred from molecular, geological and palaeoceanographic data

Aim The Southern Ocean is split into several biogeographical provinces between convergence zones that separate watermasses of different temperatures. Recent molecular phylogenies have uncovered a strong phylogeographic structure among rockhopper penguin populations, Eudyptes chrysocome sensu lato, from different biogeographical provinces. These studies suggested a reclassification as three species in two major clades, corresponding, respectively, to warm, subtropical and cold sub‐Antarctic watermasses rather than to geographic proximity. Such a phylogeographic pattern, also observed in plants, invertebrates and fishes of the Southern Ocean, suggests that past changes in the positions of watermasses may have affected the evolutionary history of penguins. We calculated divergence times among various rockhopper penguin clades and calibrated these data with palaeomagmatic and palaeoceanographic events to generate a speciation chronology in rockhopper penguins. Location Southern Ocean. Methods Divergence times between populations were calculated using five distinct mitochondrial DNA loci, and assuming a molecular clock model as implemented in mdiv . The molecular evolution rate of rockhopper penguins was calibrated using the radiochronological age of St Paul Island and Amsterdam Island in the southern Indian Ocean. Separations within other clades were correlated with palaeoceanographic data using this calibrated rate. Results The split between the Atlantic and Indian populations of rockhopper penguins was dated as 0.25 Ma, using the date of emergence of St Paul and Amsterdam islands, and the divergence between sub‐Antarctic and subtropical rockhopper penguins was dated as c. 0.9 Ma (i.e. during the mid‐Pleistocene transition, a major change in the Earth’s climate cycles). Main conclusions The mid‐Pleistocene transition is known to have caused a major southward shift in watermasses in the Southern Ocean, thus changing the environment around the northernmost rockhopper penguin breeding sites. This ecological isolation of northernmost populations may have caused vicariant speciation, splitting the species into two major clades. After the emergence of St Paul and Amsterdam islands in the subtropical Indian Ocean 0.25 Ma, these islands were colonized by penguins from the subtropical Atlantic, 6000 km away, rather than by penguins from the sub‐Antarctic Indian Ocean, 5000 km closer.     

The biostratigraphy and paleobiology of Oligocene planktonic foraminifera from the equatorial Pacific Ocean (ODP Site 1218)

Planktonic foraminifera from a continuous Oligocene succession with clear magnetochronology and sediment cycles at Ocean Drilling Program Site 1218 (equatorial Pacific Ocean) were studied in the interval from 27 to 30 Ma. Paragloborotalia taxa are common and we examined their size, relative abundance, and stable isotopes. Multispecies stable isotope data indicate the depth habitats of Oligocene planktonic foraminifera and suggest that “Globoquadrina” venezuelana and Dentoglobigerina globularis were probably mixed-layer dwellers, with paragloborotaliids recording heavier δ¹⁸O signatures consistent with a thermocline habitat. Cyclic variations in the abundance of Paragloborotalia match eccentricity (100 kyr) variations in percent carbonate and δ¹³C, suggesting orbitally forced upwelling in the equatorial Pacific Ocean and that Paragloborotalia were responding directly to changes in surface water productivity. The high-resolution biostratigraphy calibrated to the magnetochronology constrains the extinction of Paragloborotalia opima which marks the top of Planktonic Foraminifera Biozone O5 (P21b) at 27.456 Ma. The highest occurrence of P. opima is associated with a 50% size decrease in Paragloborotalia pseudocontinuosa taxa within Chron 9n. In addition, we find the extinction of Chiloguembelina cubensis is consistent with other deep-sea sections within Chron 10n at 28.426 Ma marking the O4/O5 (P21a/P21b) boundary.     

Habitat modelling of <Emphasis Type="Italic">Electrona antarctica</Emphasis> (Myctophidae,Pisces) in Kerguelen by generalized additive models and geographic information systems

Electrona antarctica is one of the most abundant mesopelagic fishes in the oceanic zone surrounding the Kerguelen Archipelago in the Indian sector of the Southern Ocean. Generalized additive models (GAM) combined with geographical information systems (GIS) were used to predict and map the abundance of this species according to three environmental variables: sea surface temperature, bathymetry and surface chlorophyll a. The model was applied on the Antarctic Polar Front in the eastern part of Kerguelen Archipelago. E. antarctica seems to be linked to areas presenting low chlorophyll a concentrations, depths greater than 500 m and temperatures lower than 5°C. The model was then applied to the Kerguelen’s plateau for three different years: 1998, 1999 and 2000. The position of Antarctic Polar Front and the intensity of an upwelling play an important role in the abundance variability of E. antarctica. Furthermore, the model allows the understanding of the habitat of E. antarctica and its trophic place in the pelagic ecosystem.     

THE PELAGIC DISTRIBUTION OF SEA-BIRDS IN THE WESTERN INDIAN OCEAN

The present paper summarises observations made during the International Indian Ocean Expedition on board the R.R.S.' Discovery, from August to November 1963 and from February to September 1964 in the Indian Ocean north of 20° S and west of 70° E. In 1963 work was carried out in the Somali Basin. In 1964 a series of transects were made over the ocean, the main purpose of which was to investigate the system of equatorial currents and the changes in the sea associated with the onset of the Southwest Monsoon in May. The Tropical Indian Ocean is briefly described. Except for local concentrations, the surface layers are poor in nutrient salts and plankton. There is a seasonal reversal of winds and surface currents in the Arabian Sea, but seasonal changes become less marked further south. Previous ornithological observations in the western Indian Ocean are mostly confined to the Arabian Sea or to the island groups. Thus, existing information on the pelagic range of pantropical species is incomplete. Observations made on each transect across the Indian Ocean in 1964 are summarised and compared with oceanographic data collected at the same time. General conclusions are not possible on the basis of so little information, though there appeared to be some relationship between the distribution of certain species and wind or current belts. The only marked discontinuity recorded was a concentration of Puffinus pacificus and Gygis alba at the northern edge of the Equatorial Counter-current at 58° E in March. Observations made on transects that were repeated before and after the onset of the Southwest Monsoon suggested that Sterna fuscata concentrates in the equatorial region as the monsoon develops. In general, both plankton and sea-birds were more abundant at 58° E than at 67¹/₄° E. An analysis of the presence or absence of sea-birds during each observation period, which lasted an average of one hour, established the difference between pelagic species and those largely restricted to within 50 miles of their breeding stations. There was no evidence of any correlation with zooplankton abundance, though birds of all species taken together were commonest where flying-fish were most abundant and S. fuscata appeared to be commonest in cool-water areas with strong winds, i.e. the Trade-wind belts. It was not possible to sample the food organisms of sea-birds quantitatively. Sea-bird observations in the Indian Ocean more than 200 miles from the continental coasts are summarized and compared with previous observations. The little information collected in the Indian Ocean agrees with previous work in the Pacific Ocean, where sea-birds are commonest in areas of convergence, and not where local upwelling and an associated concentration of plankton occur, such as on the equator. This may be due to the fact that populations of organisms on which sea-birds feed develop or concentrate a considerable time after upwelling of nutrient-rich water occurs. Since few sea-birds were seen feeding, it seems likely that available food is scarce and that much time is required to locate areas where it is abundant. Finally, evidence is presented to indicate how some sea-bird species may avoid or reduce competition by feeding at different distances offshore, or in different geographical areas.     

Size-fractionated biomass and productivity of phytoplankton and particulate organic carbon in the southern ocean

During the austral summer of 1989/1990, surface samples were obtained of size-fractionated biomass, and the productivity of phytoplankton, its cell abundance, the composition of the dominant species, the concentration of particulate organic carbon (POC) and the related environmental surface parameters were measured in a large-scale survey primarily of the Atlantic and Indian Sectors. The results showed that the southern atlantic sector is the most fertile; chlorophylla (Chla) concentration averaged over 2 μg l⁻¹, average cell abundance was about 41.0 × 10³ cell l⁻¹, and average POC concentration was also the highest (>100 μg l⁻¹), but was lower in the Drake Passage and the southern Indian sector. The results for size-fractionated Chla showed that netplankton (>20 μm) in the South Atlantic Ocean, having abundant nutrients, accounted for the highest proportion (average 65%) of biomass. In the infertile South Indian Ocean, picoplankton (<2 μm) accounted for the highest proportion, averaging 47%. The results for size-fractionated productivity showed that the contribution of picoplankton to total productivity was the largest in the South Atlantic Ocean and Drake Passage, those of nanoplankton (2–20 μm) and netplankton being about equal. The relatively high photosynthesis assimilation number of picoplankton demonstrates their importance in the marine ecosystems of Antarctic water. In comparison with the Antarctic water, the subantarctic and subtropical waters are infertile.     

Radiation and speciation of pelagic organisms during periods of global warming: the case of the common minke whale, Balaenoptera acutorostrata

How do populations of highly mobile species inhabiting open environments become reproductively isolated and evolve into new species? We test the hypothesis that elevated ocean‐surface temperatures can facilitate allopatry among pelagic populations and thus promote speciation. Oceanographic modelling has shown that increasing surface temperatures cause localization and reduction of upwelling, leading to fragmentation of feeding areas critical to pelagic species. We test our hypothesis by genetic analyses of populations of two closely related baleen whales, the Antarctic minke whale (Balaenoptera bonaerensis) and common minke whale (Balaenoptera acutorostrata) whose current distributions and migration patterns extent are largely determined by areas of consistent upwelling with high primary production. Phylogeographic and population genetic analyses of mitochondrial DNA control‐region nucleotide sequences collected from 467 whales sampled in four different ocean basins were employed to infer the evolutionary relationship among populations of B. acutorostrata by rooting an intraspecific phylogeny with a population of B. bonaerensis. Our findings suggest that the two species diverged in the Southern Hemisphere less than 5 million years ago (Ma). This estimate places the speciation event during a period of extended global warming in the Pliocene. We propose that elevated ocean temperatures in the period facilitated allopatric speciation by disrupting the continuous belt of upwelling maintained by the Antarctic Circumpolar Current. Our analyses revealed that the current populations of B. acutorostrata likely diverged after the Pliocene some 1.5 Ma when global temperatures had decreased and presumably coinciding with the re‐establishment of the polar–equatorial temperature gradient that ultimately drives upwelling. In most population samples, we detected genetic signatures of exponential population expansions, consistent with the notion of increasing carrying capacity after the Pliocene. Our hypothesis that prolonged periods of global warming facilitate speciation in pelagic marine species that depend on upwelling should be tested by comparative analyses in other pelagic species.     

Latest Eocene–Early Oligocene climate change and Southern Ocean fertility: inferences from sediment accumulation and stable isotope data

The earliest Oligocene (∼33.5 Ma) is marked by a major step in the long-term transition from an ice-free to glaciated world. The transition, characterized by both cooling and ice-sheet growth, triggered a transient but extreme glacial period designated Oi-1. High-resolution isotope records suggest that Oi-1 lasted for roughly 400,000 yr (the duration of magnetochron 13N) before partially abating, and that it was accompanied by an ocean-wide carbon isotope anomaly of ∼0.75‰. One hypothesis relates the carbon isotope anomaly to enhanced export production brought about by climate-induced intensification of wind stress and upwelling, particularly in the Southern Ocean. To understand how this climatic event affected export production in the Southern Ocean, biogenic silica (opal) and carbonate accumulation rates were computed for the sub-polar Indian Ocean using deep-sea cores from ODP Site 744, Kerguelen Plateau. Our findings suggest that net productivity in this region increased by several fold in response to the Oi-1 glaciation. In addition, calcareous primary producers dominant in the Late Eocene were partially replaced by opaline organisms suggesting a trend toward seasonally greater surface divergence and upwelling in this sector of the Southern Ocean. We attribute these changes to intensification of atmospheric/oceanic circulation brought about by high-latitude cooling and the appearance of a full-scale continental ice-sheet on East Antarctica. Higher terrigenous sediment accumulation rates support the idea that wind-induced changes in regional productivity were augmented by an increased supply of glacial dust and debris that provided limiting micro-nutrients (e.g., iron-rich dust particles). We speculate that the rapid changes in biogenic sediment accumulation in the Southern Ocean and other upwelling-dominated regions contributed to the ocean-wide positive carbon isotope anomaly by temporarily increasing the burial rate of organic carbon relative to carbonate carbon. The changes in burial rates, in turn, may have produced a positive feedback on climate by briefly drawing down atmospheric p_CO2.     

Past and present distribution, densities and movements of blue whales Balaenoptera musculus in the Southern Hemisphere and northern Indian Ocean

• 1 Blue whale locations in the Southern Hemisphere and northern Indian Ocean were obtained from catches (303 239), sightings (4383 records of ≥8058 whales), strandings (103), Discovery marks (2191) and recoveries (95), and acoustic recordings.
• 2 Sighting surveys included 7 480 450 km of effort plus 14 676 days with unmeasured effort. Groups usually consisted of solitary whales (65.2%) or pairs (24.6%); larger feeding aggregations of unassociated individuals were only rarely observed. Sighting rates (groups per 1000 km from many platform types) varied by four orders of magnitude and were lowest in the waters of Brazil, South Africa, the eastern tropical Pacific, Antarctica and South Georgia; higher in the Subantarctic and Peru; and highest around Indonesia, Sri Lanka, Chile, southern Australia and south of Madagascar.
• 3 Blue whales avoid the oligotrophic central gyres of the Indian, Pacific and Atlantic Oceans, but are more common where phytoplankton densities are high, and where there are dynamic oceanographic processes like upwelling and frontal meandering.
• 4 Compared with historical catches, the Antarctic (‘true’) subspecies is exceedingly rare and usually concentrated closer to the summer pack ice. In summer they are found throughout the Antarctic; in winter they migrate to southern Africa (although recent sightings there are rare) and to other northerly locations (based on acoustics), although some overwinter in the Antarctic.
• 5 Pygmy blue whales are found around the Indian Ocean and from southern Australia to New Zealand. At least four groupings are evident: northern Indian Ocean, from Madagascar to the Subantarctic, Indonesia to western and southern Australia, and from New Zealand northwards to the equator. Sighting rates are typically much higher than for Antarctic blue whales.
• 6 South‐east Pacific blue whales have a discrete distribution and high sighting rates compared with the Antarctic. Further work is needed to clarify their subspecific status given their distinctive genetics, acoustics and length frequencies.
• 7 Antarctic blue whales numbered 1700 (95% Bayesian interval 860–2900) in 1996 (less than 1% of original levels), but are increasing at 7.3% per annum (95% Bayesian interval 1.4–11.6%). The status of other populations in the Southern Hemisphere and northern Indian Ocean is unknown because few abundance estimates are available, but higher recent sighting rates suggest that they are less depleted than Antarctic blue whales.

     

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.     

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

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

Diatoms in surface sediments of the Indonesian Archipelago and their relation to hydrography

Diatoms from 53 surface sediments (water depth 350–7200 m) of the Indonesian Archipelago were studied to determine the recent distribution of important assemblages. Three significant assemblages were distinguished, each related to hydrographic parameter(s) of the overlying water mass. The first assemblage is related to the warm saline surface waters of Pacific and Indian Ocean origin; the second assemblage reflects the low-salinity lobe in Makassar Strait; and the third corresponds to major seasonal upwelling areas in the Arafura Sea and south of Java.The normalized ratio of typically Pacific Ocean and Indian Ocean species reflects the fluctuations of the Pacific versus Indian Ocean inflow during the Late Quaternary.Absolute diatom abundances (ADA; in valves g^–1 carbonate-free dry wt) and diatom accumulation rates (DAR; in valves CM^–2 ka^–1) of autochthonous species did not correspond to daily rates of primary production in the photic zone and consequently cannot be used for paleoproductivity estimates.Besides vertical transport some lateral transport of diatoms occurs, as was demonstrated by the presence of three groups of allochthonous species. They are indicators of productivity in the littoral environment, bottom currents and river outflow.     

Rotifer distributions in the coastal waters of the northeast Pacific Ocean

Absolute abundance of rotifers was assessed from 5 to 80 km across the continental shelf off of the southern Oregon coast (U.S.A.) in the northeast Pacific Ocean. A total of 97 vertically stratified water samples were collected at 49 stations from two depths, 3 and 30 m. Coastal upwelling conditions were indicated, with decreased temperature, increased salinity and higher chlorophyll-a concentrations closer to shore. Two rotifer genera, Synchaeta and Trichocercaoccurred within 16 km of shore with densities increasing closer to shore. Synchaeta reached densities of 64 inds l^–1 while Trichocerca was sparse (<1 inds. l^–1). Rotifers were most abundant at 3 m and the densest aggregations appear to be associated with estuary outlets, suggesting that estuaries may be important in exporting rotifers to nearshore coastal waters.     

Long-term upwelling evolution in tropical and equatorial Pacific during the last 800 kyr as revealed by coccolithophore assemblages

The coccolithophore assemblages in two ODP Sites (1237 and 1238) are studied in order to reconstruct the paleoenvironmental conditions in the tropical and equatorial Pacific during the last 800 kyr. Both ODP Sites are located in the two most significant upwelling zones of the tropical and equatorial Pacific: Peru and Equatorial upwelling, respectively. The two sites are considered to have had similar evolutions. The coccolith relative abundance, the nannofossil accumulation rate (NAR) and the N ratio (namely, the proportion of < 3 μm placoliths in relation to Florisphaera profunda) allow us to identify three different intervals. Interval I (0.86-0.45 Ma) and interval III (0.22-0 Ma) are related to weak upwelling and weak Trade Winds, as suggested by coccolithophore assemblages with low N ratios. Interval II (0.45-0.22 Ma), characterized by dominant Gephyrocapsa caribbeanica and very abundant “small” Gephyrocapsa and Gephyrocapsa oceanica, is conversely related to intense upwelling and enhanced Trade Winds.     

History of West Mediterranean newts,Pleurodeles (Amphibia: Salamandridae), inferred from old and recent DNA sequences

MtDNA sequences (396 bp cytochrome b and 369 bp 12S rRNA) from recent material and old museum specimens indicate Pleurodeles poireti and P. waltl form independent clades with 7.76% genetic divergence. Within P. poireti, populations from Djebel Edough, NE Algeria are very distinct with 6.12% genetic divergence from the remainder and may deserve separate species status. Away from Djebel Edough, P. poireti consists of three distinct clades (coastal NW Tunisia; central N Algeria; Constantine plus inland NW Tunisia) with a maximum genetic divergence of only 1%. P. waltl contains two clades with 2.96% genetic divergence, one in SE and E Spain plus north Morocco, the other in Portugal and SW and central Spain. Pleurodeles probably invaded NW Africa from SW Europe during the Messinian Salinity Crisis, when land contact was first established at 5.6 Ma, and then interrupted at 5.3 Ma. Molecular clocks, calibrated in the assumption that the latter event separated P. waltl and P. poireti, suggest that Pleurodeles diverged from its sister taxon, Tylototriton, at about 8.6–10 Ma. Djebel Edough P. poireti separated at about 4.2 Ma, perhaps through isolation on a temporary, now ‘fossil’, island initiated by the Messinian crisis. Differentiation in remaining P. poireti may have been caused by Pleistocene climatic fluctuations, while bifurcation in P. waltl appears to have taken place in the Pliocene approximately between 3.2 and 2 Ma. This species reached Morocco very recently, perhaps as a result of human introduction. Use in Pleurodeles of the slower divergence rates estimated in some other salamandrids results in a less parsimonious historical hypothesis that does not fit known geophysical events.     

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.     

Late Neogene planktonic foraminifera of the Cibao Valley (northern Dominican Republic): Biostratigraphy and paleoceanography

An assemblage of planktonic foraminifera is described from 125 samples taken from the Cercado, Gurabo, and Mao Formations in the Cibao Valley, northern Dominican Republic. The primary objectives of this study are to establish a biochronologic model for the late Neogene of the Dominican Republic and to examine sea surface conditions within the Cibao Basin during this interval. The Cercado Formation is loosely confined to Zones N17 and N18 (∼ 7.0–5.9 Ma). The Gurabo Formation spans Zones N18 and N19 (∼ 5.9–4.5 Ma). The Mao Formation is placed in Zone N19 (∼ 4.5–3.6 Ma). Changes in the relative abundances of indicator species are used to reconstruct sea surface conditions within the basin. Increasing relative abundances of Globigerinoides sacculifer and Globigerinoides ruber, in conjunction with a decreasing relative abundance of Globigerina bulloides, suggests the onset of increasing sea surface temperature and salinity in conjunction with diminishing primary productivity at ∼ 6.0 Ma. Abrupt increases in the relative abundances of G. sacculifer and G. ruber at ∼ 4.8 Ma suggest a major increase in sea surface temperature and salinity in the early Pliocene. The most likely mechanism for these changes is isolation of the Caribbean Ocean through progressive restriction of Pacific–Caribbean transfer via the Central American Seaway. Periods of high productivity associated with upwelling events are recorded in the upper Cercado Formation (∼ 6.1 Ma) and in the middle Mao Formation (∼ 4.2 Ma) by spikes in G. bulloides and Neogloboquadrina spp. respectively. The timing of major increases in sea surface salinity and temperature as well as decreasing productivity (∼ 4.8 Ma) and periods of upwelling (∼ 6.1and 4.2 Ma) in the Cibao Basin generally corroborate previously suggested Caribbean oceanographic changes related to the uplift of Panama. Changes in sea surface conditions depicted by paleobiogeographic distributions in the Cibao Basin suggest that shoaling along the Isthmus of Panama had implications in a shallow Caribbean basin as early as 6.0 Ma. Major paleobiologic changes between ∼ 4.8 and 4.2 Ma likely represent the period of final closure of the CAS and a nearly complete disconnection between Pacific and Caribbean water masses. This study illustrates the use of planktonic foraminifera in establishing some paleoceanographic conditions (salinity, temperature, productivity, and upwelling) within a shallow water basin, outlining the connection between regional and localized oceanographic changes.     

Paleoceanography of the mid-Pleistocene

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

Spatial-temporal variability of ultraplankton vertical distribution in the Antarctic frontal zones within 60–66°E, 43–46°S

The vertical distribution of heterotrophic bacteria and four ultraphytoplanktonic (<10 µm) groups (Prochlorococcus, Synechococcus, pico- and nanoeukaryotes) was investigated by flow cytometry at three process stations located in three different sub-systems belonging to the Antarctic Circumpolar Current frontal zone and to the Southern Indian Ocean (60–66°E, 43–46°S; ANTARES 4 cruise, January-February 1999): the Subtropical Zone (STZ), the Convergence Zone and the Polar Frontal Zone (PFZ). In each sub-system, short-term variability of cell abundance and flow cytometric parameters (right-angle light scatter and chlorophyll autofluorescence) was assessed through a times series of up to 24 h with a 2 h sampling frequency. The ultraphytoplankton vertical distribution exhibited a high spatial variability, with dominance of Prochlorococcus in the STZ (mean: 762.85×10¹⁰ cells m^–2), whereas picoeukaryotes (<3 µm) were dominant in the PFZ (55.46×10¹⁰ cells m^–2), a typically high-nutrient low-chlorophyll zone. Heterotrophic bacteria abundance was maximum (9.84×10¹³ cells m^–2) in the frontal zone, between the Agulhas Front and the Subtropical Front. Nanoeukaryotes showed the largest (up to 80%) variations between two consecutive sampling periods (2 h). Abundance variations could not be assigned to the same water mass during the time series due to the highly variable hydrodynamics of the study area. Trends of short-term abundance variations were opposite between the PFZ (lowest at night) and north of the Subtropical Front (highest at night). The observed spatial and short-term variations illustrate the complexity of the water masses in the Indian sector of the Southern Ocean, and highlight the challenge of extrapolating discrete measurements over space and time for use in evaluating carbon budgets in such dynamic areas.