Phylogeny and Rates of Molecular Evolution of Planktonic Foraminifera: SSU rDNA Sequences Compared to the Fossil Record

Planktonic foraminifera are marine protists, whose calcareous shells form oceanic sediments and are widely used for stratigraphic and paleoenvironmental analyses. The fossil record of planktonic foraminifera is compared here to their molecular phylogeny inferred from ribosomal DNA sequences. Eighteen partial SSU rDNA sequences from species representing all modern planktonic families (Globigerinidae, Hastigerinidae, Globorotaliidae, Candeinidae) were obtained and compared to seven sequences representing the major groups of benthic foraminifera. The phylogenetic analyses indicate a polyphyletic origin for the planktonic foraminifera. The Candeinidae, the Globorotaliidae, and the clade Globigerinidae + Hastigerinidae seem to have originated independently, at different epochs in the evolution of foraminifera. Inference of their relationships, however, is limited by substitution rates of heterogeneity. Rates of SSU rDNA evolution vary from 4.0 × 10⁻⁹ substitutions/site/year in the Globigerinidae to less than 1.0 × 10⁻⁹ substitutions/site/year in the Globorotaliidae. These variations may be related to different levels of adaptation to the planktonic mode of life. A clock-like evolution is observed among the Globigerinidae, for which molecular and paleontological data are congruent. Phylogeny of the Globorotaliidae is clearly biased by rapid rates of substitution in two species (G. truncatulinoides and G. menardii). Our study reveals differences in absolute rates of evolution at all taxonomic levels in planktonic foraminifera and demonstrates their effect on phylogenetic reconstructions. Received: 21 January 1997 / Accepted: 17 April 1997     

Paleoceanographic conditions in the western Caribbean Sea for the last 560 kyr as inferred from planktonic foraminifera

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

Geochemical investigation of gametogenic calcite addition in the planktonic foraminifera Orbulina universa

Several species of planktonic foraminifera precipitate a final layer of calcite onto the shell surface immediately prior to gamete release at the end of the foraminifera life cycle. Here, we present the results of carbon-13, oxygen-18 and thermal labeling experiments conducted under high (HL) and low light (LL) regimes that vary symbiont photosynthetic activity. Mean experimental group data show that gametogenic (gam) calcite contributes between 4–17% and 14–20% to final shell mass for high and low light experiments respectively. These data indicate that past studies of gam calcite addition have overestimated the amount of gam calcite on foraminifera shells by ~ 30–55%. Calculations indicate that the mass of gam calcite added to the O. universa shell, 4.2 ± 2.0 μg and 4.0 ± 2.4 μg, is constant in the HL and LL groups respectively. We propose that the production of gam calcite may be the result of the discharge of a relatively constant-volume cytoplasmic pool of either Ca²⁺ or alkalinity (carbon pool) that increases the calcite saturation state in the microenvironment adjacent to the foraminifera shell just prior to gamete formation and release. Results from these experiments indicate that the geochemistry of thin-walled O. universa from deep sea sediments is composed of > 80% ontogenetic calcite that was precipitated in the primary, near surface habitat of this species.     

First appearance of Globorotalia truncatulinoides: cladogenesis and immigration

The first appearance datum of the planktic foraminifer Globorotalia truncatulinoides is widely used to identify the base of the Quaternary Period. However, its appearance is not globally isochronous. We have previously shown that G. truncatulinoides evolved gradually from its ancestor Globorotalia crassaformis via the intermediate species Globorotalia tosaensis. This cladogenesis was documented in the southwest Pacific during the Late Pliocene in sympatric or parapatric populations. Based on qualitative observations, similar but younger, gradual transitions have been reported from other areas of the world's oceans. Therefore, this gradual evolutionary branching might have occurred in response to changing environments at different times in different ocean areas. To evaluate the hypothesis of a repeated, environmentally driven gradual, sympatric cladogenesis, we studied the morphological transitions of the three taxa, using image analytical techniques, in several deep-sea sections from various areas. Our study confirms that G. truncatulinoides evolved between 2.8 and 2.3 Ma sympatrically in large populations from its ancestor G. crassaformis in the southwest Pacific. Differentiated morphotypes of G. truncatulinoides subsequently immigrated into the Indian and Atlantic oceans between 2.3 and 1.9 Ma. Our morphometric data show these younger appearances outside the southwest Pacific to be punctuated.We hypothesize that the global cooling of surface waters, coinciding with the northern hemisphere glaciation, led to the formation of oceanographic barriers that could have retarded the expansion of G. truncatulinoides up to 2.3 Ma. At this time, a relative warming and subsequent transgression could have spurred the migration, possibly through the Indonesian passage. A direct link between the speciation and surface water changes linked to the northern hemisphere glaciation has not been proven so far and seems unlikely. In fact, stable isotope data in this lineage indicate that the three species' depth habitat preferences remained unchanged through the speciation and migration of G. truncatulinoides and that all three species were dominantly deep-dwellers, in agreement with their present environmental preferences.     

Paleoceanography of the Mauritanian margin during the last two climatic cycles: From planktonic foraminifera to African climate dynamics

The last 220 ka of the MD03-2705 (18° 05.81′ N–21° 09.19′ W) sedimentary sequence, retrieved off the Cape Verde islands, was investigated using a multiproxy approach. Planktonic foraminifera assemblage analyses, coupled with isotopic measurements (δ¹⁸O) from benthic (Planulina wuellerstorfi) and planktonic (Globigerinoides ruber) foraminifera monospecific samples were conducted along the topmost 11 m of the sequence. High resolution X-ray fluorescence measurements (0.5 cm resolution), giving access to major element ratio, have completed the geochemical analyses along the core. Seasonal and annual past sea surface temperatures (SST) were quantitatively reconstructed. Local sea-surface salinity (SSS) changes were then estimated by coupling SST with planktonic δ¹⁸O data. Our data provide a set of both oceanic and continental markers of environmental changes along the north-western African margin. The major changes detected in our record are discussed in the light of the regional paleoceanographic and paleoclimatic history of the last 220 ka. Coupled oceanographic and atmospheric processes portray the climatic evolution of the area, and show strong links among the regional oceanography (water mass advection), the upwelling dynamics and the Intertropical Convergence Zone (ITCZ) migration. An increased upwelling activity (or influence of upwelling filaments) is noted at the end of the two last glacial periods, probably in response to a more southerly position of the ITCZ. Higher SSS are recorded over the area during arid intervals and were tentatively interpreted as signing a southward shift of the Cape Verde Frontal Zone. A detailed coupling between dust advection and SSS values over the site of study was noted during MIS6.5.     

Late Quaternary variations in water mass in the Shatsky Rise area, northwest Pacific Ocean

Oxygen isotope analysis of planktonic and benthic foraminifera in piston core S-2 collected from the Shatsky Rise (33°21.75N, 159°07.70E; water depth 3107 m) provides a paleoceanographic record for the last 540 000 years in the northwestern Pacific Ocean. Although peaks in the abundance of sinistral Neogloboquadrina pachyderma occur during Marine Isotope Stage 2, and particularly 6 and 12, the southward shifting of the Subarctic front did not reach the core site during these glacial periods. However, mass accumulation rates of total organic carbon, biogenic opal, and terrigenous matter contents indicate that surface productivity increased during cold periods. In addition, the C/N ratio analyzed in organic matter reached values of up to 10 during glacial periods. These results imply that delivery of eolian dust to this site was enhanced by strengthened westerly winds during glacial periods. Down-core fluctuations in δ¹³C values of Globigerinoides ruber and Globorotalia inflata nearly overlap, particularly during the period from 540 to 260 ka. This latter trend suggest that the subtropical surface water mass prevailed at the core site throughout that period, based upon the very small vertical δ¹³C gradient through water column in modern Kuroshio Current water.     

Environmental and climatic changes in the central Mediterranean Sea (Siculo–Tunisian Strait) during the last 30 ka based on dinoflagellate cyst and planktonic foraminifera assemblages

A high resolution micropalaeontological study of the core MD 04-2797 CQ recovered in the Sicilian–Tunisian Strait provides insights into the paleoclimatic history of the Mediterranean Sea at the transition between the western and eastern basin over the last 30 ka. Using the analysis of dinoflagellate cyst and planktonic foraminiferal assemblages, we reconstruct the paleoenvironmental changes that took place in this region. High abundances of cold temperate dinocyst species (Nematosphaeropsis labyrinthus, Spiniferites elongatus, Bitectatodinium tepikiense) and the polar planktonic foraminifera Neogloboquadrina pachyderma (left coiling) reveal three major cooling events synchronous with North Atlantic Henrich events 1 and 2 (H1 and H2) and the European and North Atlantic Younger Dryas event. During the Holocene, the presence of warm dinocyst species (Spiniferites mirabilis and Impagidinium aculeatum) and planktonic foraminifera (Globorotalia inflata and Globigerinoides ruber), reflects a significant increase of sea surface temperatures in the western Mediterranean basin, but a full warming was not recorded until 1500 years after the onset of the Holocene. Moreover, our results show that the Holocene was interrupted by at least four brief cooling events at ~ 9.2 ka, ~ 8 ka, ~ 7 ka and ~ 2.2 ka cal. BP, which may be correlated to climatic events recorded in Greenland ice cores and in the Atlantic Ocean.     

Foraminifera and coccolithophorid assemblage changes in the Panama Basin during the last deglaciation: Response to sea-surface productivity induced by a transient climate change

The responses of community assemblages of planktonic and benthonic foraminifera and coccolithophorids to transient climate change are explored for the uppermost 2 m of cores ODP677B (1.2°N; 83.74°W, 3461 m) and TR163-38 (1.34°S; 81.58°W, 2200 m), for the last ∼ 40 ka. Results suggest that the deglaciation interval was a time of increased productivity and a major reorganization of planktonic trophic webs. The succession in dominance between the planktonic foraminifera species Globorotalia inflata, Globigerina bulloides, and Neogloboquadrina pachyderma denote four periods of oceanographic change: (1) advection (24-20 ka), (2) strong upwelling (20-15 ka), (3) weak upwelling (14-8 ka) and (4) oligotrophy (8 ka to present). Strong upwelling for the deglaciation interval is supported by the low Florisphaera profunda/other coccolithophorids ratio and the high percentage abundance of Gephyrocapsa oceanica. Benthonic foraminifera assemblage changes are different in both cores and suggest significant regional variations in surface productivity and/or oxygen content at the seafloor, and a decoupling between surface productivity and export production to the seafloor. This decoupling is evidenced by the inverse relationship between the percentage abundance of infaunal benthonic foraminifera and the percentage abundance of N. pachyderma. The terrigenous input of the Colombian Pacific rivers, particularly the San Juan River, is suggested as a possible mechanism. Finally, the Globorotalia cultrata/Neogloboquadrina dutertrei ratio is used to reconstruct the past influence of the Costa Rica Dome-Panama Bight and cold tongue upwelling systems in the Panama Basin. A northern influence is suggested for the late Holocene (after 5 ka) and the last glacial (before 20 ka), whereas a southern influence is suggested for the 20-5 ka interval. There is a correspondence between our reconstructed northern and southern influences and previously proposed positions of the Intertropical Convergence Zone (ITCZ) after the Last Glacial Maximum (LGM).     

Coiling dimorphism within a genetic type of the planktonic foraminifer Globorotalia truncatulinoides

The coiling direction—sinistral (left-coiled) or dextral (right-coiled)—of planktonic foraminiferal shells is a classical proxy used to assess past environmental changes and to understand their evolutionary patterns. Globorotalia truncatulinoides is composed of five different genetic types (I to V), each with a specific biogeographic distribution. So far, type II is the only type within which both coiling types have been frequently found. Here we examine the coiling direction as a dimorphic variation in a single genetic type, and show that the two forms with different coiling directions differ in ecology. The studied 140 left- and 137 right-coiled specimens from eight depth layers at four stations in the Sargasso Sea all belonged to Type II, based on the phylogenetic analyses of the ITS (internal transcribed spacer) region of ribosomal DNA. The distributions of left- and right-coiled Type II (Type II-L and II-R) differed strongly: Type II-L dominated between 400 and 200 m depth in the central water mass, whereas Type II-R was common in the top 200 m of the subtropical gyre. The left- and right-coiled forms are thus associated with different water masses. However, the vertical distribution along a temperature gradient indicates that their coiling direction is not determined by temperature, and might have some genetic basis. Our molecular phylogenetic analyses showed that the two forms probably share the same gene pool through their dispersals between different water masses. Moreover, the lineage of G. truncatulinoides is young (< 3 Ma), which might not be long enough to evolve into two lineages with opposite coiling direction. We discuss the practical and theoretical usefulness of a simple genetic model for coil dimorphism, in order to build testable hypotheses regarding the evolutionary history of planktonic foraminifera at the population level.     

Living benthic foraminifera of the Okhotsk Sea: Faunal composition,standing stocks and microhabitats

Live (Rose Bengal stained) benthic foraminifera were investigated in surface sediment samples from the Okhotsk Sea to reveal the relationship between faunal characteristics and environmental parameters. Live benthic foraminifera were quantified in the size fraction > 125 µm in the upper 8 cm of replicate sediment cores, recovered with a multicorer at five stations along the Sakhalin margin, and at three stations on the southwestern Kamchatka slope. The stations are from water depths between 625 to 1752 m, located close or within the present Okhotsk Sea oxygen minimum zone, with oxygen levels between 0.3 and 1.5 ml l^- 1. At the high-productivity and ice-free Kamchatka stations, live benthic foraminifera are characterized by maximal standing stocks (about 1700-3700 individuals per 50 cm²), strong dominance of calcareous species (up to 87-91% of total live faunas), and maximal habitat depths (down to 5.2-6.7 cm depth). Vertical distributions of total faunal abundances exhibit a clear subsurface maximum in sediments. At the Sakhalin stations, which are seasonally ice-covered and less productive, live benthic foraminifera show lower standing stocks (about 200-1100 individuals per 50 cm²), lower abundance of calcareous species (10-64% of total live faunas), and shallower habitat depths (down to 2.5-5.4 cm depth). Faunal vertical distributions are characterized by maximum in the uppermost surface sediments. It is suggested that 1) lower and strongly seasonal organic matter flux, caused by the seasonal sea ice cover and seasonal upwelling, 2) lower bottom water oxygenation (0.3-1.1 ml l^- 1), and 3) more pronounced influence of carbonate undersaturated bottom water along the Sakhalin margin are the main factors responsible for the observed faunal differences. According to species downcore distributions and average living depths, common calcareous species were identified as preferentially shallow, intermediate and deep infaunal. Foraminiferal microhabitat occupation correlates with the organic matter flux and the depth of the oxygenated layer in sediments.     

Foraminiferal oxygen and carbon isotopes during the last 34 kyr off northern Japan,northwestern Pacific

Oxygen and carbon isotopes of foraminifera were analyzed in core PC4, water depth 1366 m, off northern Japan, near the east side of the Tsugaru Strait (130 m depth) between the open northwestern Pacific Ocean and the Japan Sea. At present, the site is at the confluence of the Tsugaru Warm Current which flows eastwards out of the Sea of Japan through the Tsugaru Strait, the subarctic Oyashio Current and the subtropic Kuroshio Current. During the Last Glacial Maximum (LGM), the Oyashio Current penetrated further to the South and outflow from the Japan Sea was restricted by glacio-eustatic sea level lowering.The isotopic values of the planktic foraminifer Neogloboquadrina pachyderma (sinistral) and the benthic foraminifer Uvigerina akitaensis reflect rapid millennial-scale paleoceanographic changes between 34 and 6 ka. Hydrographic changes during deglaciation were related to events at high northern latitudes, but Holocene hydrographic changes were dominated by local effects, such as the development of the outflow of the Tsugaru Warm Current. High values of planktic δ¹⁸O during the LGM reflect the southward advance of the Oyashio Current. These values decreased by 0.3‰ from 19.4 to 18.9 ka, then increased by 0.5‰ at 18 ka, with highest values between 17.5 and 15 ka. The δ¹⁸O oscillations between 19.4 and 15 ka may reflect millennial-scale warm–cold oscillations during Heinrich event 1. Planktic microfossil data indicate that cold Oyashio waters flowed from the northwestern Pacific into the Japan Sea via the Tsugaru Strait between 17 and 16 ka, consistent with the occurrence of the highest planktic δ¹⁸O values in core PC4. Planktic δ¹⁸O values rapidly decreased by 0.9‰ at 15 ka, possibly reflecting the effects of both a rapid increase in fresh water flux and rising temperatures in the subarctic North Pacific. During the Younger Dryas, cold event planktic δ¹⁸O values increased by 0.5‰, followed by a gradual decrease by 1‰ from the early to middle Holocene, reflecting a gradual increase in eastward outflow via the Tsugaru Strait with sea level rise. Both planktic and benthic foraminiferal δ¹³C values oscillated between 34 and 10 ka, at relatively large amplitudes (about 0.5‰), then remained relatively stable during the last 10 kyr. Several negative planktic and benthic (∼ − 0.7‰) δ¹³C excursions were present in sediment dated between the precipitation of secondary carbonates during episodic methane release possibly associated with methane release from continental margin sediments.     

Sensitivity of south‐east Atlantic planktonic foraminifera to mid‐Pleistocene climate change

Abstract: The last one million years are important in terms of climate development during the so‐called Mid‐Pleistocene Transition when amplification of the glacial–interglacial cycles occurred. This study describes abundance changes in fossil planktonic foraminifera in sediments from Core T89‐40, retrieved from the Walvis Ridge in the south‐east Atlantic, across this time period. Cycles between upwelling and subtropical planktonic foraminiferal assemblages are shown to mirror changes between glacial and interglacial periods, respectively. During interglacial marine isotopic stages (MIS) 9, 11 and 31, however, anomalously high abundances of the polar left‐coiled Neogloboquadrina pachyderma occur, presumably linked to unusual seasonal upwelling waters. The planktonic foraminiferal abundance record shows 41‐ky cyclic variations in the regional oceanography linked to cycles in insolation influenced by changes in the Earth’s axial tilt (obliquity). These orbitally induced oscillations in oceanographic change occurred throughout the entire record. The most conspicuous feature of the planktonic foraminiferal record is the near absence of left‐coiled Globorotalia truncatulinoides between 960 and 610 ka (MIS 26‐15). The abrupt disappearance of this species is synchronous with the onset of the Mid‐Pleistocene Transition in MIS 26.     

Distribution of living planktonic foraminifera in the Ross Sea and the Pacific sector of the Southern Ocean (Antarctica)

In order to determine the factors controlling the distribution of planktonic foraminifera as a proxy for reconstruction of paleoenvironments, we present data on live assemblages collected in the Southern Ocean. Plankton tows and hydrographic measurements were taken in the upper 400 m of the water column at different sites in the Ross Sea (site B) and at the Polar Front of the Pacific Ocean (site O) during austral summers from 1998 to 2003.Based on qualitative micropaleontological observations we discriminated between Neogloboquadrina pachyderma dextral (dex) and N. pachyderma sinistral (sin). In addition for N. pachyderma (sin) we distinguished four morphs: the first one (1) has a thickened test and depressed sutures; the second morph (2) is represented by specimens characterized by a subspheric and heavily encrusted test; the third morph (3) has a thin and lobate walled test; the fourth one (4) represent the juvenile stage of N. pachyderma (sin) and is characterized by a smaller average size.The microfauna collected in the Ross Sea (site B) is characterized by the dominance of N. pachyderma (sin) (morphs 1 and 2), whereas low occurrences of Turborotalia quinqueloba, N. pachyderma (dex) and Neogloboquadrina dutertrei were noted in the first 50 m of the water column. The water column at this station is characterized by a marked and shallow stratification and a marked thermocline during the sampling season.At the ocean station (site O), the assemblage shows increasing diversification: T. quinqueloba, G. bulloides, N. pachyderma (dex) and few specimens of Globigerinita uvula characterize the planktonic microfauna. There is a predominance of non-encrusted morphs and juvenile specimens (3 and 4). At this station the mixed surface layer is deeper than in the Ross Sea (60–70 m), the pycnocline and the thermocline less marked.The depth and the intensity of the Deep Chlorophyll Maximum (DCM) influence foraminiferal distribution: N. pachyderma (sin) shows abundance peaks at or just below the DCM while G. bulloides peaks above the DCM. Coiling direction of N. pachyderma seems to be not controlled exclusively by Sea Surface Temperature (SST): probably the two coiling types are genetically different.Results document that diversity of planktonic foraminifera, number of specimens and variations in test morphology are related to regional differences in water properties (temperature, salinity, and DCM depth).     

Foraminiferal oxygen and carbon isotopes during the last 34 kyr off northern Japan, northwestern Pacific

Oxygen and carbon isotopes of foraminifera were analyzed in core PC4, water depth 1366 m, off northern Japan, near the east side of the Tsugaru Strait (130 m depth) between the open northwestern Pacific Ocean and the Japan Sea. At present, the site is at the confluence of the Tsugaru Warm Current which flows eastwards out of the Sea of Japan through the Tsugaru Strait, the subarctic Oyashio Current and the subtropic Kuroshio Current. During the Last Glacial Maximum (LGM), the Oyashio Current penetrated further to the South and outflow from the Japan Sea was restricted by glacio-eustatic sea level lowering.The isotopic values of the planktic foraminifer Neogloboquadrina pachyderma (sinistral) and the benthic foraminifer Uvigerina akitaensis reflect rapid millennial-scale paleoceanographic changes between 34 and 6 ka. Hydrographic changes during deglaciation were related to events at high northern latitudes, but Holocene hydrographic changes were dominated by local effects, such as the development of the outflow of the Tsugaru Warm Current. High values of planktic δ¹⁸O during the LGM reflect the southward advance of the Oyashio Current. These values decreased by 0.3‰ from 19.4 to 18.9 ka, then increased by 0.5‰ at 18 ka, with highest values between 17.5 and 15 ka. The δ¹⁸O oscillations between 19.4 and 15 ka may reflect millennial-scale warm–cold oscillations during Heinrich event 1. Planktic microfossil data indicate that cold Oyashio waters flowed from the northwestern Pacific into the Japan Sea via the Tsugaru Strait between 17 and 16 ka, consistent with the occurrence of the highest planktic δ¹⁸O values in core PC4. Planktic δ¹⁸O values rapidly decreased by 0.9‰ at 15 ka, possibly reflecting the effects of both a rapid increase in fresh water flux and rising temperatures in the subarctic North Pacific. During the Younger Dryas, cold event planktic δ¹⁸O values increased by 0.5‰, followed by a gradual decrease by 1‰ from the early to middle Holocene, reflecting a gradual increase in eastward outflow via the Tsugaru Strait with sea level rise. Both planktic and benthic foraminiferal δ¹³C values oscillated between 34 and 10 ka, at relatively large amplitudes (about 0.5‰), then remained relatively stable during the last 10 kyr. Several negative planktic and benthic ( − 0.7‰) δ¹³C excursions were present in sediment dated between the precipitation of secondary carbonates during episodic methane release possibly associated with methane release from continental margin sediments.     

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.     

Surface-water cooling and salinity decrease during the Middle Miocene climate transition at Southern Ocean ODP Site 747 (Kerguelen Plateau)

Paleoceanographic variability at southern high latitude Ocean Drilling Program (ODP) Site 747 was investigated in this study through the interval which spans the Middle Miocene Climate Transition (MMCT). Between 15.0 and 12.2 million years ago (Ma), foraminiferal δ¹⁸O records derived from both benthic (Cibicidoides spp.) and planktonic taxa (Globorotalia praescitula and Globigerina bulloides) reveal a history of changes in water column thermal and salinity structure and a strong imprint of seasonality. Prior to the MMCT, in the interval between 14.35 and 13.9 Ma, G. bulloides displays relatively high δ¹⁸O values similar to those of G. praescitula, interpreted to indicate weakening of the thermocline and/or increased seasonality with cooler early-spring and/or late-fall temperatures. Following this interval, G. bulloides δ¹⁸O values diverge significantly from benthic and G. praescitula values, with G. bulloides values remaining relatively low for at least 600 kyr following the benthic foraminiferal δ¹⁸O shift during the MMCT at ~ 13.9 Ma. This divergence in δ¹⁸O records occurs in direct association with the Mi3 cooling and glaciation event and may suggest: (1) a strengthening of the vertical temperature gradient, with greater cooling of deep waters than surface waters, (2) changes in the depth habitat of G. bulloides, (3) changes in the dominant season of G. bulloides calcification, (4) modification of surface-water δ¹⁸O values in association with enhanced sea-ice formation, (5) increased surface-water carbonate ion concentration, and/or (6) a significant decrease in surface-water salinity across the MMCT. The first of these possible scenarios is not likely, particularly in light of recent Mg/Ca evidence for significant surface-water cooling in the Southern Ocean associated with the MMCT. Of the remaining possibilities, we favor a change in surface salinity to explain the observed trends in δ¹⁸O values and hypothesize that surface salinity may have decreased by up to 2 salinity units at ~ 13.9 Ma. In this scenario, the development of a lower-salinity Antarctic surface layer coincided with regional cooling of both surface and deep waters of the Southern Ocean during the Mi3 glaciation of East Antarctica, and contributed into the dominance of Neogloboquadrina spp. between 13.8 and 13.2 Ma. Additionally, the distinct patterns observed in planktonic foraminiferal δ¹⁸O records spanning the MMCT correspond with changes in the vertical δ¹³C gradient between planktonic and benthic foraminiferal records and major changes in planktonic foraminiferal assemblages at Site 747, providing further evidence of the environmental significance of this climatic transition.     

Middle Miocene paleoceanography of the western equatorial Pacific (DSDP site 289) and the evolution ofGloborotalia (Fohsella)

Evolution of the planktic foraminiferal lineageGloborotalia (Fohsella) occurred during the Miocene between 23.7 and 11.8 Ma and forms the basis for stratigraphic subdivision of the early middle Miocene (Zones N10 through N12). Important morphologic changes within theG. (Fohsella) lineage included a marked increase in test size, a transition from a rounded to an acute periphery, and the development of a keel in later forms. We found that the most rapid changes in morphology ofG. (Fohsella) occurred between 13 and 12.7 Ma and coincided with an abrupt increase in the δ¹⁸O ratios of shell calcite. Comparison of isotopic results ofG. (Fohsella) with other planktic foraminifers indicate that δ¹⁸O values of the lineage diverge from surface-dwelling species and approach deep-dwelling species after 13.0 Ma, indicating a change in depth habitat from the surface mixed layer to intermediate depth near the thermocline. Isotopic and faunal evidence suggests that this change in depth stratification was associated with an expansion of the thermocline in the western equatorial Pacific. After adapting to a deeper water habitat at 13.0 Ma, theG. (Fohsella) lineage became extinct abruptly at 11.8 Ma during a period when isotopic and faunal evidence suggest a shoaling of the thermocline. Following the extinction ofG. (Fohsella), the ecologic niche of the lineage was filled by theGloborotalia (Menardella) group, which began as a deep-water form and later evolved to an intermediate-water habitat. We suggest that the evolution ofG. (Fohsella) andG. (Menardella) were tightly linked to changes in the structure of the thermocline in the western equatorial Pacific.     

Living benthic foraminifera of the Okhotsk Sea: Faunal composition, standing stocks and microhabitats

Live (Rose Bengal stained) benthic foraminifera were investigated in surface sediment samples from the Okhotsk Sea to reveal the relationship between faunal characteristics and environmental parameters. Live benthic foraminifera were quantified in the size fraction > 125 µm in the upper 8 cm of replicate sediment cores, recovered with a multicorer at five stations along the Sakhalin margin, and at three stations on the southwestern Kamchatka slope. The stations are from water depths between 625 to 1752 m, located close or within the present Okhotsk Sea oxygen minimum zone, with oxygen levels between 0.3 and 1.5 ml l^- 1. At the high-productivity and ice-free Kamchatka stations, live benthic foraminifera are characterized by maximal standing stocks (about 1700-3700 individuals per 50 cm²), strong dominance of calcareous species (up to 87-91% of total live faunas), and maximal habitat depths (down to 5.2-6.7 cm depth). Vertical distributions of total faunal abundances exhibit a clear subsurface maximum in sediments. At the Sakhalin stations, which are seasonally ice-covered and less productive, live benthic foraminifera show lower standing stocks (about 200-1100 individuals per 50 cm²), lower abundance of calcareous species (10-64% of total live faunas), and shallower habitat depths (down to 2.5-5.4 cm depth). Faunal vertical distributions are characterized by maximum in the uppermost surface sediments. It is suggested that 1) lower and strongly seasonal organic matter flux, caused by the seasonal sea ice cover and seasonal upwelling, 2) lower bottom water oxygenation (0.3-1.1 ml l^- 1), and 3) more pronounced influence of carbonate undersaturated bottom water along the Sakhalin margin are the main factors responsible for the observed faunal differences. According to species downcore distributions and average living depths, common calcareous species were identified as preferentially shallow, intermediate and deep infaunal. Foraminiferal microhabitat occupation correlates with the organic matter flux and the depth of the oxygenated layer in sediments.     

High-resolution sea surface temperature and salinity dynamics in the northern Okinawa Trough over the last 24 kyr

Based on the high-resolution Mg/Ca and oxygen isotope ratio (δ¹⁸O) of planktonic foraminifera Globigerinoides ruber, sea surface temperature (SST) and salinity (SSS) were reconstructed in Core PC-1 from the northern Okinawa Trough during the last 24 kyr. From the last glacial maximum to Holocene, SST varied from 20.2°C to 24.6°C. Millennial-scale climatic events of the SST during the last glacial, such as the Heinrich events, Bølling-Allerød warming, and Younger Dryas, have been identified, which are synchronous with the climate changes in the North Atlantic, suggesting a teleconnection between the northwestern Pacific and the North Atlantic. During the last 24 kyr, the SSS variation can be divided into three parts: (1) during 21–15.5 ka, there was a significantly low SSS; (2) during 15.5–11.7 ka, the SSS increased obviously; (3) since 11.7 ka, the SSS is relatively stable. In order to discuss the main factors influencing the SSS in the northern Okinawa Trough, the relative abundance of Pulleniatina obliquiloculata and the oxygen isotope difference between the northern and middle Okinawa Trough (Δδ¹⁸O_sw) have been used to indicate the intensity of the Kuroshio Current and Changjiang freshwater discharge, respectively. The Δδ¹⁸O_sw result shows that there is a large amount of Changjiang freshwater emptied into the northern Okinawa Trough during 18–15.5 ka, which is caused by the subtropical monsoon rain band lingering in the lower reaches of the Changjiang River drainage, supporting the Jet Transition Hypothesis. As the Kuroshio Current strengthened since 15.5 ka as indicated by the increased relative abundance of P. obliquiloculata, the variation of Kuroshio Current became the more important contribution to the SSS in the study region. The results of our study indicate that the key factor influencing the SSS in the northern Okinawa Trough is variable during the last 24 kyr.     

The Holocene Pulleniatina Minimum Event revisited: Geochemical and faunal evidence from the Okinawa Trough and upper reaches of the Kuroshio current

The Holocene Pulleniatina Minimum Event (PME) is characterized by a very low abundance of the planktonic foraminifer Pulleniatina obliquiloculata between  4.5 and 3 ka. The PME occurs widely in the Okinawa Trough and the South China Sea, and can be correlated throughout this area; it has been related to variability in the Kuroshio current. To further explore the nature of the PME, we studied cores obtained from the southern Okinawa Trough and the upper reaches of the Kuroshio current. Faunal census data indicate that all cores record the PME between  4.5 and  3 ka. The relative abundance of Neogloboquadrina dutertrei is negatively correlated to that of P. obliquiloculata in the southern Okinawa Trough, but not in the sites at the upper reaches. Mg/Ca and δ¹⁸O measurements on Globigerinoides ruber shells from the southern Okinawa Trough indicate that there was no change in sea surface temperature or sea surface salinity during the PME. The vertical structure of the water column as reconstructed by multispecies δ¹⁸O and δ¹³C profiles shows no consistent anomalies in the southern Okinawa Trough and western Philippine Sea during the PME. These observations suggest that: (1) the PME was not restricted to marginal seas, but widespread in the western North Pacific. (2) The high abundance of N. dutertrei during the PME in the Okinawa Trough may be a result of higher food-availability in the absence of P. obliquiloculata. (3) No distinctive, consistent anomalies in the paleoceanographic proxies are associated with the PME, implying there were no changes in hydrography and productivity. The absence of a linkage between faunal variation and paleoceanographic proxies indicates that we do not yet understand what causes changes in planktonic foraminiferal assemblages. This lack of understanding implies that we cannot always trust fauna-based paleothermometry at millennial timescales.