Modelling the temperature dependent growth rates of planktic foraminifera

The temperature influence on foraminifera growth rate was analysed using a mechanistic formulation that take into account enzyme inactivation at extreme temperatures. Growth rates are calculated using available published and unpublished laboratory culture experiments for eight species, including Neogloboquadrina pachyderma (sinistral and dextral forms), Neogloboquadrina dutertrei, Globigerina bulloides, Globigerinoides ruber, Globigerinoides sacculifer, Globigerinella siphonifera and Orbulina universa. Modeled growth formulas readily reproduce the observed growth patterns for all species. Similar growth patterns are observed for the species that have the same symbiotic algae G. ruber, G. sacculifer, and O. universa. However, different growth patterns are observed for herbivorous species (Neogloboquadrina genus) compared to carnivorous species with or without symbionts. Our growth estimates correspond well to in situ observations from both plankton tows and sediment traps. These estimates will help to improve the quantification of the effects of environmental parameters on foraminifera species distribution and abundance.     

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).     

Late neogene biostratigraphy and paleoceanography of DSDP site 310 Central North Pacific and correlation with the Southwest Pacific

Late Neogene planktonic foraminifera have been examined at Site 310 in the Central North Pacific and their stratigraphic ranges and frequencies are presented here. Blow's (1969) zonation developed for tropical regions has been applied where applicable. Where tropical index taxa are rare or absent in this temperate region, Globorotalia crassaformis, and the evolutionary bioseries G. conoidea — G. conomiozea and G. puncticula — G. inflata have been found useful for zonal subdivisions. A correlation between stratigraphic ranges and frequency distributions of these species at Site 310 in the Central North Pacific, and Site 284 in the Southwest Pacific indicates that these species are relatively consistent biostratigraphic markers in temperate regions of both the North and South Pacific Oceans. An informal zonation for temperate latitudes of the Southwest Pacific has been established by Kennett (1973) and a similar zonal subdivision can be made at Site 310.Paleoclimatic/paleoceanographic interpretations based on coiling ratios, percent abundance, and phenotypic variations of Neogloboquadrina pachyderma indicate four major cold events during early, middle, and late Pliocene, and early Pleistocene. Faunal correlations of these events with similar events elsewhere in the Northeast and Southwest Pacific which have been paleomagnetically dated indicate the following approximate ages for these cold events: 4.7 Ma, 3.0 Ma, 2.6–1.8 Ma. and 1.2 Ma. Faunal assemblages have been divided into three groups representing cool, intermediate, and warmer water assemblages. Cool water assemblages are dominated by >60% N. pachyderma; intermediate temperature faunas are dominated by species of Globigerina and Globigerinita and contain between 20% and 30% N. pachyderma. Warmer water assemblages are dominated by species of Globorotalia and contain <10% N. pachyderma. Frequency oscillations within these groups, in addition to paleotemperature parameters evident in N. pachyderma, afford refined paleoclimatic/paleoceanographic interpretations.     

Distribution of recent planktonic foraminifera in surface sediments of the Mediterranean Sea

The Mediterranean Sea is a partillay isolated ocean where excess evaporation over precipitation results in large east to west gradients in temperature and salinity. Recent planktonic foraminiferal distributions have been examined in 66 surface sediment samples from the Mediterranean Sea. In addition to mapping the frequency distribution of 16 species, the faunal data has been subjected to cluster analysis, factor analysis and species diversity analysis. The clustering of species yields assemblages that are clearly temperature related. A warm assemblage contains both tropical and subtropical elements, while the cool assemblage can be subdivided into cool-subtropical, transitional and polar-subpolar groupings. Factor analysis is used to delineate the geographic distribution of four faunal assemblages. Factor 1 is a tropical-subtropical assemblage dominated by Globigerinoiden ruber. It has its highest values in the warmer eastern basin. Transitional species (Globorotalia inflata and Globigerina bulloides) dominate factor 2 with highest values occurring in the cooler western basin. Factor 3 reflects the distribution of Neogloboquadrina dutertrei and is considered to be salinity dependent. Subpolar species dominate factor 4 (Neoglobuquadrina pachyderma and G. bulloides), with highest values occurring in the northern part of the western basin where cold bottom water is presently being formed. The Shannon-Weiner index of species diversity shows that high diversity exists over much of the western basin and immediately east of the Strait of Sicily. This region is marked by equitable environmental conditions and relatively even distribution of individuals among the species. Conversely, in areas where temperature and salinity values are more extreme, diversity values are lower and the assemblages are dominated by one or two species.     

Planktonic foraminifera of the northern Indian Ocean: Distribution and preservation in surface sediments

In the northern Indian Ocean, planktonic foraminiferal tests accumulate in a wide variety of surface-water environments and depositional settings. This variability enables us to isolate the effects that surface-water ecology and differential dissolution have on the distribution of planktonic foraminifera from 251 geographically widespread surface sediment samples.Foraminiferal abundance varies from 0 to > 10⁴ whole foraminifera in the greater than 150 μm fraction per gram dry sediment. Values < 10 characterize the three deep basins of the equatorial Indian Ocean and the western Bay of Bengal. Foraminiferal tests are most abundant on carbonate covered Ninety-East and Carlsberg Ridges. Absolute abundance patterns are mainly controlled by non-ecological processes. Variations in dissolution resistant species (RSP) with water depth reveal that the foraminiferal lysocline (FL) varies regionally. The FL is deepest (3,800 m) in the equatorial region, rises abruptly to 3,300 m in the Arabian Sea, and varies from 2,600 m to near 2,000 m moving northward in the Bay of Bengal. Deep samples with anomalously low RSP (< 30%) suggest redeposition.Systematic geographic and depth-related variation is observed for the 17 most abundant foraminiferal species. Dissolution resistant species (G. menardii, G. tumida, G. dutertrei, P. obliquiloculata) generally exhibit a rapid and continuous increase in relative abundance at and below the FL. Susceptible species (G. ruber, G. bulloides, G. glutinata, for example) exhibit a rapid and continuous decrease in relative abundance at and below the FL. Moderately susceptible species (G. conglobatus, G. aequilateralis, G. conglomerata, for example) rapidly increase in abundance at the FL and systematically decrease with depth below the FL.Principal components analysis (PCA) of faunal data from minimally dissolved (< 30% RSP) samples reveals important ecologically related species intercorrelations. The major biogeographic gradient is the negative covarying relationship between aG. bulloides-G. glutinata species pair and a grouping ofG. sacculifer, G. conglobatus, G. aequilateralis, andG. ruber. PCA of all samples demonstrates how differential dissolution alters this and other species relationships. Species groupings that incorporateG. ruber, G. menardii, andG. dutertrei are particularly affected by dissolution.Comparison of average faunal data from minimally dissolved samples in the northern Indian Ocean with similar samples from other tropical regions suggests varying environmental factors produce distinct faunas within the tropical ocean. For example,G. bulloides, G. falconensis, andG. hexagona are significantly more abundant in northern Indian Ocean surface sediments while such species asG. ruber, G. sacculifer, G. dutertrei, andP. obliquiloculata dominate in other tropical regions.     

Standing stock,vertical distribution and flux of planktonic foraminifera in the Panama Basin

The relationships between planktonic foraminifera in the upper 2000 m of the water column and those in sediment traps and deep-sea sediments of the Panama Basin were investigated as part of the Composition, Flux and Transfer Experiments (CFATE) and the Sediment Trap Intercomparison Experiment (STIE) July–Dec. 1979. Planktonic foraminifera larger than 333 μm, sampled during the July–August trap deployment cruise, occurred most abundantly in the euphotic zone with maximal populations associated with the chlorophyll and primary production maxima both located in the upper thermocline just below the mixed layer. Juvenile abundances exceeded those of the >333 μm adults by 3–4 orders of magnitude and their depth distribution systematics indicated that most foraminifera reproduction occurs prior to sinking of the adults from the euphotic zone. Macroscopic aggregates and fecal pellets were identified as major carriers of smaller sized shells from the euphotic zone.During the July–August cruise, the species changed from an assemblage dominated byG. theyeri to one dominated byG. ruber; the liveG. theyeri population sank passively from the euphotic zone in early August with a mean settling speed of 150 m day^?1 while theG. ruber population grew in. The abundance of empty shells of all species in the upper 200 m varied by a factor of 40 during the deployment cruise and peaked on July 30, 1979. This variability was reflected in foraminifera collection rates of sediment traps deployed at 305 m for 24- and 6-h periods. The trap and watercolumn data were combined to calculate mean sinking speeds (>333 μm fraction) of several dominant foraminiferal species (G. ruber andG. theyeri, 500 m day^?1;G. dutertrei, 2000 m day^?1). Size and weight analysis of empty shells from plankton tows yielded values of empty shell density (calculated on a spherical basis using maximum shell dimension) ranging from 0.12 and 0.15 g cm^?3 (G. theyeri andG. ruber) to 0.60 g cm^?3 (G. dutertrei). These speeds and shell densities were found to be consistent with the laboratory sinking experiments of Berger and Piper (1972) and Fok-Pun and Komar (1983). It was concluded that shell fluxes through the water column may be calculated with reasonable accuracy using a modified version of the Bishop et al. (1977) settling model and empty shell size distribution data from pump and plankton tow collections. The Fok-Pun and Komar (1983) settling model is recommended for future modelling efforts but requires a more extensive set of measurements of each foraminiferal specimen to be made.The interpretation of foraminiferal flux profiles, based on the collections of sediment traps, plankton tows, and pumps deployed on time scales shorter than several days, must take into account the very short residence times of empty shells in the water column and short term temporal variability in empty shell production rates.Populations of >333 μm foraminifera sampled in Nov.–Dec. 1979 were reduced by a factor of six relative to the July–Aug. values. The cruise-to-cruise differences were due to reduced primary production and greater mixed layer depth.Planktonic foraminifera were well preserved in sediment traps deployed for 112 days at 665, 935 and 1770 m, although aragonitic pteropod shells were partially dissolved. The fluxes of many species into the 935 m trap could not be explained based on the standing stock data from the July–August cruise alone and it was noted that species fluxes during STIE were comparable to or exceeded maximal values measured by Thunell and Reynolds (1984) in bi-monthly time series traps deployed at the same location during the whole of 1980. The moored trap data could only be explained by higher mean foraminifera shell fluxes in the interval between trap deployment and recovery, consistent with the Bishop and Marra (1984) model of primary production and carbon flux at the STIE site.Primary differences between core top samples and trap samples, were explained by the loss from the sediments of dissolution prone species,G. theyeri andG. ruber leaving an assemblage dominated by the dissolution resistant species,G. dutertrei.     

Changes in the oceanic 13C/12C ratio during the last 140 000 years: High-latitude surface water records

Detailed records of the carbon and oxygen isotopic ratios of Neogloboquadrina pachyderma are compared between nine high-latitude sediment cores, from the Northern and Southern Hemispheres, covering the last 140 000 yrs. The strong analogies between the δ¹³C records permit to define a δ¹³C stratigraphic scale, with three clear cut transitions simultaneous with the oxygen isotopic transitions 6/5 (125 kyrs.), 5/4 (65 kyrs.), and 2/1 (13 kyrs.). The δ¹³C records of N. pachyderma in the high-latitude cores, which follow the changes in δ¹³C of the surface water TCO₂ near areas of deep water formation present trends similar to the benthic foraminifera δ¹³C records in cores V19–30 and M12-392, although amplitudes of the isotopic shifts are different. This implies that a large part of the observed variations represents global changes in the carbon distribution between biosphere and ocean.The ¹³C/¹²C ratios of N. pachyderma in the North Atlantic cores display larger regional variations at 18 kyrs. B.P. than at present. To explain these differences, we have plotted the 18 kyrs. B.P. δ¹³C values of N. pachyderma from 17 cores distributed N of 40°N. Comparison with published surface water temperature distribution at 18 kyrs. B.P. indicates that a strong divergent cyclonic cell, centered approximatively 55°N and 15°W, was active during most of the last ice-age maximum This hydrology, analogous to the present Weddell Sea, explains the published evidences of bottom water formation, if located on the northern flank of the gyre, and the strong polar front on the southern flank, probable location of intermediate water formation.     

The oxygen isotope composition of planktonic foraminifera from the Cariaco Basin,Venezuela: Seasonal and interannual variations

Material collected during a three-year sediment trapping experiment in the Cariaco Basin, Venezuela (January 1997 to December 1999) is used to examine both temporal and inter-species variability in the oxygen isotope composition of planktonic foraminifera. Specifically, this study compares the oxygen isotope composition of six species of planktonic foraminifera (Globigerinoides ruber (pink), Globigerina bulloides, Neogloboquadrina dutertrei, Orbulina universa, Globorotalia menardii and Globorotalia crassaformis) with the climatology and hydrography of the region, and evaluates the application of each species for use in paleoceanographic reconstructions. The isotope results are consistent with known depth habitats for all six species. The lowest δ¹⁸O values (− 1 to − 2‰) were measured on G. ruber (pink) and G. bulloides, two species that live in the surface mixed layer. Values for deeper-dwelling species such as N. dutertrei, G. menardii and G. crassaformis are higher, predominantly ranging from 0 to − 0.5‰. Temperature estimates derived using species-specific paleotemperature equations indicate that G. ruber (pink) accurately estimates sea surface temperatures (SSTs) throughout the year, while G. bulloides temperature estimates are similar to measured surface temperatures only during the upwelling season (January–April). For the remainder of the year, the δ¹⁸O-derived temperatures for G. bulloides typically are lower than the measured SST. Although the maximum flux of all species occurs during upwelling, the flux-weighted annual mean isotopic composition of the six species indicates that only G. bulloides is biased towards this season. Therefore, we conclude that the sediment δ¹⁸O record of G. ruber (pink) is most suitable for estimating past values of mean annual SST, while G. bulloides provides information on conditions during spring upwelling. The depth of calcification of N. dutertrei varies seasonally in response to changes in the depths of the thermocline and chlorophyll maximum. As a result, the δ¹⁸O difference between G. ruber (pink) and N. dutertrei provides an estimate of the annual surface to thermocline temperature gradient in the basin.     

Comparison of seasonal flux variations of planktonic foraminifera in sediment traps on both sides of the Ryukyu Islands,Japan

Variations in the fluxes of planktonic foraminifera were analyzed based on sediment trap data collected over a period of 10 months, from October 1994 to August 1995, at stations JAST01 and JAST02, located on both sides of the Ryukyu Islands, Japan. Station JAST01 (latitude 27°23′N, longitude 126°44′E) was deployed at a depth of 1000 m in the Okinawa Trough, East China Sea, along the flow axis of the Kuroshio Current, west of the islands, whereas station JAST02 (latitude 25°4′N, longitude 127°34′E) was deployed at a depth of 3000 m in the Ryukyu Trench, along the western margin of the northwest Pacific, east of the Islands. The total planktonic foraminiferal fluxes (TFFs) in the eastern station were high in winter, when the surface-water column was vertically well mixed. In contrast, the TFF peaks did not show significant seasonality in the western station, but values fluctuated with the northwest–southeast oscillation of the Kuroshio axis. Among the 36 planktonic foraminiferal species identified in the trap samples, Globigerina bulloides, Globigerinita glutinata, Neogloboquadrina dutertrei, Pulleniatina obliquiloculata, Globigerinoides ruber, Globigerinoides sacculifer and Globigerina falconensis exhibited a greater shell flux. On both sides of the Ryukyu Island Arc, the fluxes of G. ruber and G. sacculifer increased synchronously with the seasonal warming of surface waters, whereas that of G. falconensis increased during winter, when the water column was vertically well mixed as a result of the NW monsoon. In contrast to these species, which exhibited similar seasonal flux variation patterns at both stations, the species G. bulloides, G. glutinata, N. dutertrei, and P. obliquiloculata displayed different flux variations on both sides of the islands. These fluxes seem to be regulated by phytoplankton productivity, which is controlled by the vertical structure of the water column on the eastern Ryukyu Trench side and by the oscillation of the Kuroshio axis on the western Okinawa Trough side. Lateral transport of suspended planktonic foraminiferal shells to the Okinawa Trough might exist, but is not prominent enough to wipe out the original features of the planktonic foraminiferal flux. The species G. falconensis is an indicator of winter mixing on both sides of the Ryukyu Islands, and may possibly be used as a proxy to trace the intensity of the paleo-winter monsoon.     

Linkages between surface and deep circulations in the southern Japan Sea during the last 27,000 years: Evidence from planktic foraminiferal assemblages and stable isotope records

We analyzed planktic foraminiferal assemblages, oxygen and carbon isotope records, and the presence or absence of laminations to reconstruct the paleoenvironments of the southern Japan Sea since the last glacial period. Data were collected from two well-dated cores. One core (water depth 999 m) included thinly laminated mud layers, the other (water depth 283 m) contained nonlaminated sediments. Tephrochronology and accelerator mass spectrometry ¹⁴C dating of 14 horizons revealed that the two cores contained continuous records of the last 27 cal kyr. A total of 13 planktic foraminiferal species belonging to six genera were identified in down-core samples. The typical indicators of the Tsushima Current water, Globigerinoides ruber, Neogloboquadrina dutertrei, Pulleniatina obliquiloculata, Globigerinoides tenellus, and Globigerinita glutinata occurred since 9.3 cal kyr BP. Neogloboquadrina incompta, which was the dominant species in the Tsushima Current region of the modern Japan Sea, first occurred at 8.2 cal kyr BP and dominated the assemblage since 7.3 cal kyr BP. These results clearly indicate that the warm Tsushima Current started to inflow into the Japan Sea at 9.3 cal kyr BP, and the modern surface conditions in the southern Japan Sea were essentially established at 7.3 cal kyr BP. Our data and comparison of the presence or absence of laminated sediments in three locations from the southern Japan Sea suggest that deep circulation during the deglacial period was weaker than that at present. In addition, deep circulation in the modern Japan Sea, which supplies oxygen-rich water to the entire basin, started probably in association with the first inflow of the Tsushima Current beginning at 9.3 cal kyr BP.     

Late pleistocene paleoclimatology: Planktonic foraminiferal analyses of sediment cores from the central North Atlantic

Planktonic foraminiferal analyses of six deep-sea sediment cores from the central North Atlantic east of the Azores Islands between 37°N and 40°N show distinct oscillations in planktonic foramineferal assemblages during the last 300,000 years. A paleoclimatic curve has been constructed using “Total Fauna Analysis” that reveals three glacial and four interglacial episodes. Relatively minor climatic oscillations are superimposed upon the major glacial-interglacial episodes. The paleoclimatic curve is similar to previous paleoclimatic curves from the Atlantic and adjacent areas. Minor paleoclimatic fluctuations are more distinct in paleoclimatic curves from high latitudes of the Atlantic.The faunal assemblages are transitional between subarctic and subtropical assemblages. During portions of the interglacial episodes, the assemblage is dominated by Globorotalia inflata. Neogloboquadrina pachyderma (dextral-coiling) or Globigerina bulloides dominate during the remainder of the interglacial episodes and during the glacial episodes. Glacial episodes are also marked by particularly high frequencies of Globigerina quinqueloba and Globorotalia scitula. Interglacial episodes are also marked by increases in Globorotalia truncatulinoides, Globigerinoides ruber, and Globigerinella aequilateralis.The planktonic foraminiferal faunal oscillations in the cores are complex and cannot be entirely explained by temperature variation. Other parameters such as salinity, nutrients and biological competitin must influence the faunal oscillations.The faunas suggest no major planktonic foraminiferal faunal boundaries migrated across the area between 37°N and 40°N in the central North Atlantic during the last 300,000 years.     

The oxygen isotope composition of planktonic foraminifera from the Guaymas Basin,Gulf of California: Seasonal,annual, and interspecies variability

Sediment trap samples collected over a seven-year period (February 1991–October 1997) from Guaymas Basin in the Gulf of California were used to study the oxygen isotope composition of five species of planktonic foraminifera, Globigerinoides ruber (white), Globigerina bulloides, Neogloboquadrina dutertrei, Pulleniatina obliquiloculata, and Globorotalia menardii. The δ¹⁸O data were analyzed for temporal and interspecies variability and were compared to local hydrography to evaluate the use of each species in reconstructing past oceanographic applications. The two surface dwelling species, G. ruber and G. bulloides displayed the lowest δ¹⁸O values (～ 0.0 to ? 5.0‰), while δ¹⁸O values for the thermocline dwelling N. dutertrei, P. obliquiloculata, and G. menardii were higher (～ 0.0 to ? 2.0‰). The δ¹⁸O of G. ruber most accurately records measured sea surface temperatures (SSTs) throughout the year. G. bulloides δ¹⁸O tracks SSTs during the winter–spring upwelling period but for the remainder of the year records slightly colder, subsurface temperatures. The difference between the δ¹⁸O of the surface dwelling species, G. ruber and G. bulloides, and that of the thermocline dwelling species, N. dutertrei, P. obliquiloculata, and G. menardii, was used to estimate the surface to thermocline temperature gradient. During the winter these δ¹⁸O differences are small (～ 0.50‰) reflecting a well-mixed water column. These interspecies δ¹⁸O differences increase during the summer (～ 1.90‰) in response to the strong thermal stratification that exists at this time of year.     

Regional and interannual productivity of biogenic components and planktonic foraminiferal fluxes in the northwestern Pacific Basin

Time-series sediment trap experiments at subtropical (WCT-1) and subarctic (WCT-2) stations in the northwestern Pacific indicate seasonal, latitudinal and depth variations in total particulate, biogenic and foraminiferal fluxes. At the subtropical station, the average total mass flux was 19.4 mg m⁻² day⁻¹ in the shallow trap (1060 m) and 21.5–26.1 mg m⁻² day⁻¹ in the deep trap (3930 m) during the sampling period. At subarctic station, these values were 91.5–176.9 mg m⁻² day⁻¹ in the shallow and 68.6–112.3 mg m⁻² day⁻¹ in the deep trap. We recognized 12 and 15 planktonic foraminiferal species at Station WCT-1 and Station WCT-2, respectively. The planktonic foraminiferal flux and species turnover are related to seasonal and interannual changes in source water and water column conditions at both stations. At Station WCT-1, the highest flux was recorded during the summer, with a peak in mid to late June associated with similar flux patterns of the dominant species, Globigerinoides ruber and Globigerinita glutinata. The total flux of foraminiferal tests at the shallow and deep traps is similar in numbers and magnitude. At Station WCT-2, the peaks of total flux of foraminiferal tests at the two trap depths differ in number, and their magnitude in the deep trap is almost half of that in the shallow trap. A distinctive seasonal pattern occurred in the shallow and the deep trap, with a peak in total foraminiferal flux in mid June to mid July. Globigerina quinqueloba, Neogloboquadrina pachyderma and Neogloboquadrina dutertrei dominate the planktonic population throughout the year.Subtropical Station WCT-1 was characterized by low total foraminiferal fluxes and low total mass flux, which is dominated by calcium carbonate and depleted in opal, whereas high foraminiferal fluxes and a high total mass flux dominated by high biogenic opal, and less calcium carbonate and organic matter characterize subarctic Station WCT-2. The foraminiferal carbonate that reaches the seafloor accounts for an average 20–27% and 22–23% of the total calcium carbonate at Station WCT-1 and Station WCT-2, respectively. The primary reason for the difference in flux at both stations thus lies in the different contributions of siliceous and calcareous planktonic assemblages. The seasonal variation in biogenic particulate flux at both stations implies that temporal changes in biological productivity are governed by large-scale seasonal climatic variability and local hydrography.     

Living planktonic foraminifera in the Fram Strait (Arctic): absence of diel vertical migration during the midnight sun

The timing of vertical migration in planktonic foraminifera (ex. ontogenetic, diel) is still an open debate. This work aims to investigate the diel vertical migration (DVM) of Neogloboquadrina pachyderma (N. pachyderma) and Turborotalita quinqueloba (T. quinqueloba) in the Arctic during the midnight sun. N. pachyderma and T. quinqueloba dominate the total assemblage in the cold Polar Water and warmer North Atlantic Water masses, respectively. Foraminifera were collected at several depths along the Fram Strait. Afterwards sampling was performed at the same station for 24 h at continuous and discrete time intervals. Results show no evidence of planktonic foraminifera DVM since there was no significant variability in the abundance and size distribution during the 24-h collection period. This finding provides information to improve the interpretation of foraminifera in paleoclimatic works. This is especially relevant in the Fram Strait as paleoclimatic studies in this region are fundamental to investigating the history of the Atlantic water inflow into the Arctic Ocean.     

Influence of setting,sieve size,and sediment depth on multivariate and univariate assemblage attributes of coral reef-associated mollusc death assemblages from the Gulf of Aqaba

Measures of diversity and ecology of marine invertebrate assemblages depend on a variety of factors including environmental conditions and methodological decisions. In this study, the influence of such factors on multi- and univariate assemblage parameters of molluscan death assemblages from the Gulf of Aqaba (Red Sea, Jordan) was evaluated. Sediment samples were collected at two coral reef types, a patch reef at 13 m of water depth characterized by fine-grained sediments and a Millepora-fringing reef with coarse-grained sediments at 5 m of water depth. The upper and lower 10 cm of the sediment column were separately removed and sieved with mesh sizes of 1 and 2 mm. A large dataset of 6400 bivalve and gastropod shells was compiled to evaluate how setting, sediment depth, and sieve size influenced taxonomic composition and species richness, species-abundance patterns and the Shannon–Wiener index, the number of drilled shells per species and drilling frequency (DF) of the assemblage. Setting had the strongest impact on all aspects, followed by sieve size, but sediment depth was insignificant, probably due to complete homogenization of the sediments by reworking and bioturbation. Multivariate assemblage parameters distinguished much better between categories (setting, sieve size) than univariate measures. Sieve size-related disagreements recognized between the two higher taxa are mostly due to the underlying difference in body-size distribution of bivalve and gastropod assemblages. We conclude that species richness and other ecological characteristics of molluscan death assemblages in coral reef-associated sediments will most strongly reflect habitat complexity of the sites chosen, are significantly influenced by methodological decisions (i.e., sieve size), will only poorly preserve temporal patterns, and the results will differ between bivalves and gastropods.     

Living Neogloboquadrina pachyderma sin and its distribution in the sediments from Oman and Somalia upwelling areas

Microfaunal analysis of plankton nets collected offshore Oman/Yemen and sediment traps offshore Somalia shows that Neogloboquadrina pachyderma sin is abundant during the SW monsoon upwelling. The sediment traps recorded the highest flux (136 specimens m⁻² day⁻¹) and the highest relative frequency (3.55% of the total foraminifera assemblage) of the species during this period. During the intermonsoon it became less abundant and decreased in size, and only very few N. pachyderma sin were found in the water column during the NE monsoon. Sediment trap and Recent sediment data collected along a downslope transect off Somalia show that the species frequency decreases offshore. The highest concentrations of N. pachyderma sin in plankton nets off Oman/Yemen were found at a depth of 300–500 m. However, the oxygen isotope compositions of N. pachyderma sin at the depth of 300–500 m and from the surface 8 m are identical, and also similar to that of Globigerina bulloides which reflects properties of the upper 25 m of the water column. The carbon isotope values are relatively consistent within the studied samples from both areas except for the specimens from the surface water samples offshore Oman/Yemen. The data show that the δ¹⁸O ratio of N. pachyderma sin is in equilibrium with the ambient water while the δ¹³C values are at least 0.8‰ lower. We infer that N. pachyderma sin in the Arabian Sea reproduces, grows and calcifies in the upper 25 m of the water column, but possibly descends into deeper waters later during its life cycle.     

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.     

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).     

Fluxes and isotopic composition of planktonic foraminifera off Hainan Island,northern South China Sea: Implications for paleoceanographic studies

Planktonic foraminifera collected from a sediment trap deployed off Hainan in the northwestern South China Sea (SCS-NW) between July 2012 and April 2013 were studied to evaluate their seasonal variability and ecology as well as to infer the factors controlling their shell fluxes. The total planktonic foraminifera flux, as well as the fluxes of the dominant species (Globigerinoides ruber, Globigerinoides sacculifer and Neogloboquadrina dutertrei), showed three distinct maxima during SW-monsoon in August 2012, the SW-NE intermonsoon in October 2012 and the NE-monsoon in December 2012–February 2013. These periods were characterized by upwelling, aerosol fallout, and intense wind mixing, respectively, from which the foraminiferal assemblages benefitted, as indicated by the close correlation between wind speed, sea surface temperature (SST), chlorophyll a concentration (Chl-a), δ¹⁸O of G. ruber and the shell fluxes. The correlation also suggests that temperature and food availability might have been the primary drivers of the observed changes in foraminiferal abundance. The offset between the SST deduced from flux-weighted of G. ruber δ¹⁸O and annual mean SST is only ∼0.3 °C, much lower than ∼5.2 °C between the summer and winter temperature, indicating a balanced seasonality bias in the shell flux. The linear regression between the satellite-derived sea surface temperature and G. ruber δ¹⁸O reveals the strong potential of this species, at least in the studied region, as an ecological indicator for past oceanic environments.     

THE PHLOEM OF ETAPTERIS LECLERCQII AND BOTRYOPTERIS TRIDENTATA

The phloem of Etapteris leclercqii and Botryopteris tridentata petioles is described from Lower Pennsylvanian coal balls. Petioles of B. tridentata are characterized in transverse section by an omega-shaped xylem trace, a phloem zone which extends from 2-10 cells in width, and 2-parted cortex. Etapteris leclercqii petioles exhibit a 4–9 cell-wide phloem zone surrounding the central clepsydroid xylem mass, and a 3-parted cortex. In both taxa a 1–2 cell layer parenchyma sheath separates the xylem from the extra-xylary tissues. The phloem of both species consists of sieve elements that average about 20 μm in diam by 200 μm in length in Botryopteris, and 100 μm in length in Etapteris, with horizontal-slightly oblique end walls. In transmitted light, the radial walls of the sieve elements form an irregular reticulate pattern enclosing elliptical lighter areas. With the scanning electron microscope, these areas appear as horizontal-slightly oblique furrows on the cell wall, with many small indentations lining the furrows. These indentations, because of their regular occurrence and size (from a few fractions of a micron up to 1.0 μm in diam), are interpreted as sieve pores, and the elliptical areas that enclose them as sieve areas. The phloem of E. leclercqii and B. tridentata is compared with that described for other fossil genera and with that of extant ferns.