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.     

Benthic foraminifera off West Africa (1°N to 32°S): Do live assemblages from the topmost sediment reliably record environmental variability?

Recent benthic foraminifera (> 125 μm) were investigated from multicorer samples on a latitudinal transect of 20 stations between 1°N and 32°S along the upper slope off West Africa. Samples were selected from a narrow water depth interval, between 1200 and 1500 m, so that changes in water masses are minimized, but changes in surface productivity are important and the only significant environmental variable. Live (Rose Bengal stained) benthic foraminifera were counted from the surface sediment down to a maximum of 12 cm. Dead foraminifera were investigated in the top 5 cm of the sediment only. Five live and five dead benthic foraminiferal assemblages were identified using Q-mode principal component analysis, matching distinct primary productivity provinces, characterized by different systems of seasonal and permanent upwelling. Differences in seasonality, quantity, and quality of food supply are the main controlling parameters on species composition and distribution of the benthic foraminiferal faunas.To test the sensitivity of foraminiferal studies based on the uppermost centimeter of sediment only, a comparative Q-mode principal component analysis was conducted on live and dead foraminiferal data from the top 1 cm of sediment. It has been demonstrated that, on the upper slope off West Africa, most of the environmental signals as recorded by species composition and distribution of the “total” live and dead assemblages, i.e., including live and dead foraminifera from the surface sediment down to 12 cm and 5 cm, respectively, can be extracted from the assemblages in the top centimeter of sediment only. On the contrary, subsurface abundance maxima of live foraminifera and dissolution of empty tests strongly bias quantitative approaches based on the calculation of standing stocks and foraminiferal numbers in the topmost centimeter.     

Living foraminifera and total populations in salt marsh peat cores: Kelsey Marsh (Clinton, CT) and the Great Marshes (Barnstable, MA)

Common species of intertidal agglutinated benthic foraminifera in salt marshes in Massachusetts and Connecticut live predominantly at the marsh surface and in the topmost sediment (0–2.5 cm), but a considerable part of the fauna lives at depths of 2.5–15 cm. Few specimens are alive at depths of 15–25 cm, with rare individuals alive between 25–50 cm in the sediments. Specimens living between the sediment surface and 25 cm deep occur in all marsh settings, whereas specimens living deeper than 25 cm are restricted to cores from the lower and middle marsh, and have an irregular distribution-with-depth. Lower and middle marsh areas are bioturbated by metazoa, suggesting that living specimens reach these depths at least in part by bioturbation. High-marsh sediments in New England consist of very dense mats of Spartina patents or Distichlis spicata roots and are not bioturbated by metazoa. In this marsh region bioturbation by plant roots and vertical fluid motion may play a role in moving the foraminifera into the sediment. The depth-distribution of living specimens varies with species: living specimens of Trochammina inflata consistently occur at the deepest levels. This suggests that species have differential rates of survival in the sediment, possibly because of differential adaptation to severe dysoxia to anoxia, or because of differing food preferences. There is no simple correlation between depth-in-core and faunal diversity, absolute abundance, and species composition of the assemblages. It is therefore possible to derive a signal of faunal changes and thus the environmental changes that may have caused them from the complex faunal signal of fossil assemblages.     

Cluster analysis of recent benthic foraminifera from the northwestern Mediterranean coast of Egypt

A suite of 74 surface sediment samples, collected from two areas along the Egyptian Mediterranean coast (the Western Harbor of Alexandria and its environs, and the area comprising the Gulf of Kanayis and the Abu Hashafa Bay), have been examined for their benthic foraminiferal faunas. A total of 82 species were identified. Census data were obtained for different species in each sample and the statistically significant fractional abundances values (≥5%) were analyzed using a Q-mode cluster analysis. Samples were segregated into four clusters, each having its peculiar benthic faunal assemblage (biotope), reflecting particular environmental conditions. These clusters are: (1) the Ammonia beccarii forma tepida Biotope, found in samples located in areas with waters of low energy, characterizing semi-closed basin conditions, with muddy or sandy mud bottom sediments (Harbor Proper); (2) the Quinqueloculina spp. Biotope, found in samples collected from depths bathed by turbid inner shelf conditions with some fresh water inflow and sandy bottom sediments (Harbor’s environs); (3) the Peneroplis-Amphistegina Biotope, found in samples collected from depths characterizing marine shelf environments with calcareous algae, and medium to very coarse calcareous sands (the Gulf of Kanayis and the Abu Hashafa Bay); (4) the Triloculina trigonula-Adelosina laevigata Biotope was represented by only one sample (at the far eastern part of the Gulf of Kanayis), collected from a site exhibiting very restricted environmental conditions. The study suggests that nutrients, turbidity, light intensity, type of substrate, and salinity are the main ecological factors controllingthe distribution of benthic foraminifera.     

Distribution of recent benthic foraminifera on the Sunda Shelf (South China Sea)

In this study we investigate the species composition and spatial distribution patterns of Rose Bengal stained and unstained benthic foraminifera from the central part of the Sunda Shelf in the south-western South China Sea in relation to environmental factors. The uppermost centimetre of the surface sediments (> 150 μm) from 45 sites from inner (60 m) to outer shelf (226 m) water depths revealed 584 species including 443 stained species.The univariate analyses of individual species abundances and community parameters and next canonical correspondence analysis were used to relate the faunal data to a set of measured environmental parameters. Four biofacies recognised on the Sunda Shelf are most strongly correlated to water depth, primary production and sediment type of the habitat. The inner shelf biofacies (CCA cluster A), defined by Ammomassilina alveoliniformis and Asterorotalia pulchella, occurs in fine grained sediments classified as modern terrigenous mud in the region with the highest primary production values. The high-energy inner shelf biofacies (CCA cluster B), defined by Heterolepa dutemplei and Textularia lythostrota, occurs in modern terrigenous sand and silt dominated sediments, northeast from the Natuna Island. The high-energy outer shelf biofacies (CCA cluster C), defined by Cibicidoides pachyderma and Textularia bocki, is sandwiched between assemblages of biofacies D. It occurs in the region characterised by neritic relict sand. In the shallow-waters on the Sunda Shelf the relationship of benthic foraminiferal faunal composition to grain size of sediments indirectly signals the prevailing bottom hydrodynamic conditions. The dominance of the epibenthic foraminifera attached to bigger particles (e.g. Cibicides lobatulus, Planulina arimiensis) and much higher abundances of empty tests suggest greater current velocities northeast of Natuna Island. The outer shelf biofacies (CCA cluster D) is defined by Facetocochlea pulchra and Bulimina marginata. It occurs in an area covered with modern terrigenous silt and mud and is characterised by lower annual primary production, but seasonally influenced by weak upwelling.     

Effects of carbon dioxide sequestration on California margin deep-sea foraminiferal assemblages

Deep-sea sequestration of CO₂ is being considered as a possible mitigation tool to decrease atmospheric CO₂ concentrations and its associated negative effects. This study investigated potential effects of liquid carbon dioxide (CO₂) injection on deep-sea foraminiferal assemblages. Foraminifera are ideal for this ecological impact investigation because of differing test composition (calcareous and non-calcareous) and thickness, and diverse epifaunal and infaunal depth preferences. The experiment was conducted on August–September 2003, at 3600 m off the coast of Monterey Bay, California, aboard the R/V Western Flyer using the ROV Tiburon. The pH of the site was monitored throughout the experiment. Sediment push-cores were collected (both from the experimental and control sites) and stained to distinguish live (stained) from dead (unstained) individuals. Effects of CO₂ injection on assemblages have been tracked both vertically (to 10 cm depth below sea floor) and horizontally (up to 10 m from CO₂ injection sites), as well as between live and dead individuals. Within corrals (containing the injected CO₂) and their underlying sediments, severe pH changes (near 4.0 units) were recorded. This compares with a record of small average reductions in ocean pH (− 0.05 units) combined with large episodic excursions (− 1.7 units) over the experimental area due to the injection of CO₂. Exposure to this gradient of low pH caused increased mortality and dissolution of calcareous forms within corrals, as far as 5 m from the injection site, and to at least 10 cm depth in the sediments.This experiment revealed several major effects of CO₂ injection on foraminiferal assemblages in surficial sediments: 1) total number of foraminifera in a sample decreases; 2) foraminiferal species richness decreases in both stained and unstained specimens; and 3) relative percentage of stained (live) forms in the remaining tests increases. Down-core trends (to 10 cm below sea floor) have revealed: 1) percent agglutinated forms decline and calcareous forms increase with depth; 2) agglutinated species richness decreases with depth; and 3) experimental core assemblages become increasingly similar with depth to those in control cores not subjected to CO₂ injection. These results imply almost complete initial mortality and dissolution in the upper 10 cm throughout the corrals following liquid CO₂ injection. Since calcareous foraminifera represent more than 50% of the total assemblages, this clearly indicates that emplacement of CO₂ will result in negative effects to diversity and survivorship of the deep-sea benthic meiofauna.     

Spatial distribution of live benthic foraminifera in the Rhône prodelta: Faunal response to a continental–marine organic matter gradient

Benthic foraminifera were collected in the Rhône prodelta (Gulf of Lions, Mediterranean Sea), an enriched zone with high organic matter content. In June 2005, sediment cores were sampled at depths ranging from 20 to 100 m. Four distinct foraminiferal assemblages were determined in the study area, reflecting the geographical distribution of the impact of river supply. The living foraminiferal faunas present a typical picture, with strongly impoverished faunas composed exclusively of stress-tolerant taxa (Fursenkoina fusiformis, Bulimina aculeata, Leptohyalis scottii, and Adelosina longirostra) in the immediate vicinity of the river mouth. This assemblage is well adapted to a high input of continental organic matter and a minimum oxygen penetration depth into the sediment. To the southwest, under the main corridor followed by the river plume, high organic input with a dominantly terrestrial signature (more refractory) may be stressful for many taxa which need organic matter of a more labile quality. In this area, Nonion scaphum, Nonionella turgida and Rectuvigerina phlegeri are present in low densities. On the edge of this area, these taxa show much higher densities. A greater proportion of marine organic carbon could explain their increasing abundances in this area. Towards the east and towards the deepest stations, in the outer part of the enriched zone, biodiversity increases. Faunas at these stations have intermediate densities and contain a number of taxa (Cassidulina carinata, Epistominella vitrea, Valvulineria bradyana, Nonionella iridea/bradyi) at the deepest stations; Bolivina dilatata/spathulata and Textularia porrecta at the eastern stations) that seem to benefit from more marine organic matter. The comparison of geochemical measurements and foraminiferal data strongly suggests that the spatial distribution of foraminifera in the Rhône prodelta is mainly governed by the quality and the quantity of organic matter reaching the sediment–water interface. Since bottom waters are well oxygenated (215–260 µmol/L), and oxygen penetration into the sediment is less than 1 cm at all stations, benthic ecosystem oxygenation appears to have only a minor impact on regional differences in faunal distribution.     

Response of deep-sea benthic foraminifera to Miocene paleoclimatic events, DSDP Site 289

Changes in the Miocene deep-sea benthic foraminifera at DSDP Site 289 closely correlate to the climatically induced variations in deep and bottom waters in the Pacific Ocean. In early Miocene time, oxygen and carbon isotopes indicate that bottom waters were relatively warm and poorly oxygenated. Benthic foraminiferal assemblages are characterized by various species inherited from the Oligocene. Expansion of the Antarctic icecap in the early middle Miocene, 14–16 m.y. ago, increased oxygen isotope values, produced cold, more oxygenated bottom waters and lead to a turnover in the benthic foraminifera. An Oligocene—early Miocene assemblage was replaced by a cibicidoid-dominated assemblage. Some species became extinct and benthic faunas became more bathymetrically restricted with the increased stratification of deep waters in the ocean. In mid-Miocene time, Epistominella exigua and E. umbonifera, indicative of young, oxygenated bottom waters, are relatively common at DSDP Site 289. Further glacial expansion 5–9 m.y. ago lowered sealevel, increased oceanic upwelling and associated biological productivity and intensified the oxygen minima. Abundant hispid and costate uvigerines become a dominant faunal element at shallow depths above 2500 m as E. umbonifera becomes common to abundant below 2500 m. By late Miocene time, benthic faunas similar in species composition and proportion to modern faunas on the Ontong-Java plateau, had become established.     

Paleoenvironmental evolution of the eastern Mediterranean during the Messinian: Constraints from integrated microfossil data of the Pissouri Basin (Cyprus)

Integrated data of calcareous nannofossils, as well as planktonic and benthic foraminifera from the Pissouri Motorway section on Cyprus allow the reconstruction of surface- and bottom-water paleoenvironments of the eastern Mediterranean during the interval preceding the Messinian salinity crisis (MSC). Contrary to deeper-water locations, where benthic foraminifera faunas are suppressed or absent just after the Tortonian–Messinian boundary, sediments deposited at intermediate water depths do contain benthic assemblages. From the earliest Messinian onwards, a development towards increasingly unfavourable paleoenvironments is reflected in the planktonic and benthic microfossil records of the Pissouri section and proceeds with rather discrete time steps that can be correlated to sequences throughout the Mediterranean. Shortly after the Tortonian–Messinian boundary a transition is recorded in the sedimentology and the open marine, deeper-water taxa disappear from the benthic foraminifera assemblages; subsequently, the diversity of all fauna groups diminishes. The changes recorded at species level in both surface-water and sea-floor dwelling taxa suggest decreasing circulation of the bottom waters, associated with changes in the surface waters, most likely due to increasing stratification. From ∼6.73 Ma onwards, our data indicate a prominent change to more restricted conditions and increasing salinity at the sea floor together with intermittently rising surface water salinity. The dominance of oligotypic and monospecific assemblages and the frequent shifts in assemblage compositions of all microfossil groups indicate severely stressed environments after ∼6.4 Ma, probably related to increased salinity. The major changes in paleoenvironmental conditions, including oxygen deprivation due to stagnation and hypersalinity, can be explained by hydrographical changes in the Mediterranean basin, which are probably caused by tectonic movements in the Rif Corridor acting in concert with astronomical cyclicity. Evaluation of the paleodepth proxies indicates that the depth of the Pissouri Basin remained rather constant at ∼300–500 m, with a minimum of 200 m, until deposition of the “barre jaune”, the transitional interval towards the evaporites and that early shallowing to neritic depths, as was proposed before, is highly unlikely.     

The spatial and vertical distribution of living (stained) benthic foraminifera from a tropical,intertidal environment,north Queensland,Australia

We investigated the distribution of living (stained) benthic foraminifera across a tropical, intertidal shoreline adjacent to Cocoa Creek, Queensland, Australia for the purpose of better understanding the nature of test production and ultimately fossil assemblage development within such environments. Short cores (up to 1 m) were collected during the wet and dry season, along an elevational gradient comprising non-vegetated intertidal mudflat and higher-intertidal mangrove forest environments. The distribution of stained specimens can be broadly delineated into assemblages characterising ‘upper mangrove’ (2.64–2.91 m above Lowest Astronomical Tide (LAT)) and ‘low mangrove-mudflat’ (1.62–2.18 m above LAT) environments. Agglutinated species were generally limited to upper mangrove stations. Calcareous species occurred within all of the intertidal environments examined but differ in their composition between upper and lower intertidal settings. Upper mangrove faunas were characterised by the agglutinated species Arenoparrella mexicana, Haplophragmoides wilberti, Miliammina fusca, Miliammina obliqua and Trochammina inflata and the calcareous species Helenina anderseni. Live (stained) assemblages at lower intertidal elevations were dominated by the calcareous species Ammonia aoteana, as well as Rosalina spp., Elphidium oceanicum, Triloculina oblonga, Ammonia pustulosa and Shackoinella globosa.     

Benthic foraminiferal population fluctuations related to anoxia: Santa Barbara Basin

The pore-water geochemistry and benthic foraminiferal assemblages of sediments from two slope sites and within the central portion of the Santa Barbara Basin were characterized between February 1988 and July 1989. The highest foraminiferal numerical densities (1197 cm^–3 as determined by an ATP assay) occurred at a slope site in June 1988 (550 m) in partially laminated sediments. In continuously laminated sediments from the central basin, foraminifera were found living (as determined by ATP assay) in October 1988 to depths of 4 cm, and specimens prepared for transmission electron microscopy were found with intact organelles to 3 cm, indicating their inhabitation of anoxic pore waters. Ultrastructural data from Nonionella stella is consistent with the hypothesis that this species can survive by anaerobic respiration. However, the benthic foraminifera appear unable to survive prolonged anoxia. The benthic foraminiferal population was completely dead in July 1989 when bottom water O₂ was undetectable.     

Abyssal NE Atlantic benthic foraminifera during the last 15 kyr: Relation to variations in seasonality of productivity

Benthic foraminiferal faunas (> 63 μm) and stable isotopes from the last 15 kyr were studied in BENGAL programme (high-resolution temporal and spatial study of the BENthic biology and Geochemistry of a north-eastern Atlantic abyssal Locality) kasten core 13078#16 from the Porcupine Abyssal Plain, NE Atlantic (48°49.91 N, 16°29.94 W, water depth 4844 m). Changes occurred in the accumulation rates, species composition, diversity, and stable isotopes during the last 15 kyr. Today, the area is strongly influenced by seasonal inputs of phytodetritus following the spring blooms in surface water primary productivity. Variations in the relative abundance of the two most abundant species, Epistominella exigua and Alabaminella weddellensis, which today show significant increases in abundance with the presence of phytodetritus on the sea-floor, are interpreted as resulting from changes in the seasonality of productivity. Seasonal productivity was higher during the Holocene than during the last deglaciation and Younger Dryas, probably coinciding with the retreat of the polar front to higher latitudes. This hypothesis is consistent with simultaneous decreases in the percentage of the polar planktic foraminifera Neogloboquadrina pachyderma (s), and increases in the percentage of Globigerina bulloides, a warmer water planktic foraminifera indicative of phytoplankton blooms and enhanced productivity. The relative abundance of the ‘phytodetritus species’ (E. exigua and A. weddellensis) covary between 14.7 and 8.1 kyr, but not between 7.8 and 1.2 kyr. Major decreases in the numbers per gram and accumulation rates of planktic and benthic foraminifera occurred at ∼ 12–8.5 kyr and at ∼ 4 kyr which correspond to decreases in the % sediment coarse fraction and published data on inorganic carbon contents suggesting that dissolution may have increased at these times. Relationships between benthic foraminiferal faunas and benthic stable isotope records suggest no simple relationship between faunal abundances and test isotope chemistry. For example, the abundances of phytodetritus species do not show strong correlations with either the δ¹³C values of E. exigua or the Δδ¹³C _{E. exigua − P. wuellerstorfi} record, which have previously been suggested as indicative of seasonality of productivity.     

Distribution of the Eelpout Lycodes soldatovi(Pisces: Zoarcidae) over the Continental Slope of the Sea of Okhotsk

The eelpout Lycodes soldatovi was found at depths of 153 to 1005 m in the Sea of Okhotsk; however, it usually inhabits the depth range of 400 to 800. The mean and maximum abundance and the biomass of this species were 198.5 and 1037 individuals per km² and 173.7 and 1275 kg/km², respectively. In view of the data on the distribution of young fish (up to 30 cm in total body length) and the reports on the absence of Lycodes larvae, a hypothesis was developed that this species spawns predominantly in the waters off western Kamchatka and eastern Sakhalin at the depth range of 700 to 900 m. Large individuals (>50 cm) exhibit higher migratory activity and are more tolerant of environmental conditions. That is why they are encountered throughout the distribution area of this species, even in subzero temperature areas. Lycodes soldatovi seems to be a typical inhabitant of the Sea of Okhotsk, except for the waters off the South and Central Kuril Islands and the Pacific waters off the North Kuril Islands.     

Glacial Holocene environment of the southeastern Okhotsk Sea: evidence from geochemical and palaeontological data

Environmental conditions and productivity changes in the southeastern Okhotsk Sea have been reconstructed for the last 20 ka using planktonic and benthic foraminiferal oxygen isotope records and calcium carbonate, organic carbon and opal content data from two sediment cores. Species variability in benthic foraminiferal and diatom assemblages provides additional palaeoceanographic evidence. AMS radiocarbon dating of the sediments and oxygen isotope stratigraphy serve as the basis for the age models of the cores for the last 20 ¹⁴C kyr and for correlation between environmental variations in the Okhotsk Sea, and regional and global climate changes. Benthic foraminiferal assemblages in the two cores (depth 1590 and 1175 m) varied with time, so that we could recognise seven zones with different species composition. Changes in the benthic foraminiferal assemblages parallel major environmental and productivity variations. During the last glaciation, fluxes of organic matter to the sea floor showed strong seasonal variations, indicated by the presence of abundant A. weddellensis and infaunal Uvigerina spp. Benthic foraminiferal assemblages changed with warming at 12.5–11 and 10–8 ¹⁴C kyr BP, when productivity blooms and high organic fluxes were coeval with global meltwater pulses 1A and 1B. Younger Dryas cooling caused a decline in productivity (11–10 kyr BP) affecting the benthic faunal community. Subsequent warming triggered intensive diatom production, opal accumulation and a strong oxygen deficiency, causing significant changes in benthic fauna assemblages from 5.26–4.4 kyr BP to present time.     

Comparison of radiolarian and sedimentologic paleoproductivity proxies in the latest Miocene–Recent Benguela Upwelling System

Estimating past ocean productivity from ocean sediments often gives different results depending on the measurement used. We have examined a suite of paleoproductivity proxies in latest Miocene–Recent sediments from DSDP Site 532 and ODP Site 1084, two deep-sea sections underlying the Benguela Upwelling System off the Atlantic coast of southern Africa. The productivity history of this system has been previously established via organic carbon concentration, diatom floras and alkenone based estimates of surface water temperature, and shows a change from low productivity in the early Pliocene to sustain high productivity in the late Pliocene–Recent. Each of our samples was split and simultaneously analysed for several proxies of ocean productivity, including organic carbon (TOC%), carbonate, abundance of opaline radiolarians, accumulation rate of benthic foraminifera (BFAR); the radiolarian faunal composition indices Upwelling Radiolarian Index (URI) and the Water Depth Ecology index (WADE); other proxies for opal and carbonate dissolution, plus stable isotopes of benthic foraminifera. Comparisons between proxies in the same measured samples, between sites in downcore plots and to the published productivity record for this region suggest that TOC and radiolarian faunal composition, particularly the WADE index, are good indicators of past productivity, albeit with different sensitivities (log–linear correlation WADE–TOC% r = 0.78, n = 65, p < 0.01). In contrast, carbonate, and carbonate-based proxies such as BFAR primarily reflect changes in dissolution. Radiolarian faunal composition indices do not appear to be affected by bulk opal accumulation or changes in opal preservation. WADE analysis of radiolarian faunas and TOC% measurements appear to be useful proxies for productivity in late Neogene sediments, particularly for sections where opal or carbonate dissolution is significant.     

Influence of the mode of macrofauna-mediated bioturbation on the vertical distribution of living benthic foraminifera: First insight from axial tomodensitometry

We investigated the influence of bioturbation by macrofauna on the vertical distribution of living (stained) benthic foraminifera in marine intertidal sediments. We investigated the links between macrofaunal bioturbation and foraminiferal distribution, by sampling from stations situated on a gradient of perturbation by oyster-farming, which has a major effect on benthic faunal assemblages. Sediment cores were collected on the French Atlantic coast, from three intertidal stations: an oyster farm, an area without oysters but affected by oyster biodeposits, and a control station. Axial tomodensitometry (CT-scan) was used for three-dimensional visualization and two-dimensional analysis of the cores. Biogenic structure volumes were quantified and compared between cores. We collected the macrofauna, living foraminifera, shells and gravel from the cores after scanning, to validate image analysis. We did not investigate differences in the biogenic structure volume between cores. However, biogenic structure volume is not necessarily proportional to the extent of bioturbation in a core, given that many biodiffusive activities cannot be detected on CT-scans. Biodiffusors and larger gallery-diffusors were abundant in macrofaunal assemblage at the control station. By contrast, macrofaunal assemblages consisted principally of downward-conveyors at the two stations affected by oyster farming. At the control station, the vertical distribution of biogenic structures mainly built by the biodiffusor Scorbicularia plana and the large gallery-diffusor Hediste diversicolor was significantly correlated with the vertical profiles of living foraminifera in the sediment, whereas vertical distributions of foraminifera and downward-conveyors were not correlated at the station affected by oyster farming. This relationship was probably responsible for the collection of foraminifera in deep sediment layers (> 6 cm below the sediment surface) at the control station. As previously suggested for other species, oxygen diffusion may occur via the burrows built by S. plana and H. diversicolor, potentially increasing oxygen penetration and providing a favorable microhabitat for foraminifera in terms of oxygen levels. By contrast, the absence of living foraminifera below 6 cm at the stations affected by oyster farming was probably associated with a lack of biodiffusor and large gallery-diffusor bioturbation. Our findings suggest that the effect of macrofaunal bioturbation on the vertical distribution of foraminiferal assemblages in sediments depends on the effects of the macrofauna on bioirrigation and sediment oxidation, as deduced by Eh values, rather than on the biogenic structure volume produced by macrofauna. The loss of bioturbator functional diversity due to oyster farming may thus indirectly affect infaunal communities by suppressing favorable microhabitats produced by bioturbation.     

Perturbation at the sea floor during the Paleocene–Eocene Thermal Maximum: Evidence from benthic foraminifera at Contessa Road, Italy

Detailed analyses of the benthic foraminiferal assemblages extracted with the cold acetolyse method together with high resolution geochemical and mineralogical investigations across the Paleocene/Eocene (P/E) boundary of the classical succession at Contessa Road (western Tethys), allowed to recognize and document the Paleocene–Eocene Thermal Maximum (PETM) interval, the position of the Benthic Extinction Event (BEE) and the early recovery of benthic faunas in the aftermath of benthic foraminiferal extinction. The stratigraphical interval spanning the P/E boundary consists of dominantly pelagic limestones and two prominent marly beds. Benthic foraminifera indicate that these sediments were deposited at lower bathyal depth, not deeper than 1000–1500 m. The Carbon Isotope Excursion (CIE) interval is characterized by high barite abundance with a peak at the base of the same stratigraphic interval, indicating a complete, although condensed record of the early CIE. A succession of events and changes in the taxonomic structure of benthic foraminifera has been recognized that may be of use for supra-regional stratigraphic correlation across the P/E boundary interval. The composition of the benthic foraminiferal assemblages, dominated by infaunal taxa, indicates mesotrophic and changing conditions on the sea floor during the last  45 kyr of the Paleocene. The BEE occurs at the base of the CIE within the lower marly bed and it is recorded by the extinction of several deep-water cosmopolitan taxa. Then, the lysocline/CCD rose and severe carbonate dissolution occurred. Preservation deteriorated, the faunal density and simple diversity dropped to minimum values and a peak of Glomospira spp. has been observed. Stress-tolerant and opportunistic groups, represented mainly by bi-and triserial taxa, dominate the low-diversity post-extinction assemblages, indicating a benthic foraminiferal recovery under environmental unstable conditions, probably within a context of sustained food transfer to the bottom. A three-phase pattern of faunal recovery is recognizable. At first the lysocline/CCD started to descend and then recovered. Small-sized “Bulimina”, Oridorsalis umbonatus, and Tappanina selmensis rapidly repopulated the severely stressed environment. Later on, Siphogenerinoides brevispinosa massively returns, dominating the assemblage together with other buliminids, Nuttallides truempyi, and Anomalinoides sp.1. Finally, a marked drop in abundance of S. brevispinosa is followed by a bloom of the opportunistic and recolonizer agglutinated Pseudobolivina that, for the first time, is recorded within the main CIE. A second interval of dissolution, but less severe than the previous one, has been recognized within the upper marly bed (uppermost part of the main CIE interval) and it is interpreted as a renewed, less pronounced shoaling of the lysocline/CCD that interrupted the recovery of benthic faunas. This further rise likely represents a response to persistent instability of ocean geochemistry in this sector of the Tethys before the end of the CIE. In the CIE recovery and post CIE intervals, the composition of the benthic foraminiferal assemblages suggests mesotrophic and unstable conditions at the sea floor. According to the geochemical proxy for redox conditions, the deposition of the PETM sediments at Contessa Road occurred in well-oxygenated waters, leading out a widespread oxygen depletion as major cause of the BEE. Changing oceanic productivity, carbonate corrosivity and global warming appear to have played a much more important role in the major benthic foraminiferal extinction at the P/E boundary.     

Breaching biogeographic barriers: the invasion of <Emphasis Type="Italic">Haynesina germanica</Emphasis> (Foraminifera,Protista) in the Bahía Blanca estuary,Argentina

Large populations of the living benthic foraminifera Haynesina germanica (Ehrenberg 1840) are reported for the first time from marsh and intertidal mud flat sediments of the Bahía Blanca estuary (Argentina). Maximum abundance of living specimens was recorded in shallow intertidal environments. The species was previously recorded from many European and North American shallow-water coastal settings, but has not been documented from Argentina. Comparative faunal assemblage analysis from dated core sample material from within the Bahía Blanca estuary shows that the species has not been present for at least the last 8,200 years. This supports the hypothesis that the species has been accidentally introduced outside its natural range as a probable result of ballast water and/or shipping activities. This study is the first report of a successful invasion of non-indigenous benthic foraminifera to the South Atlantic coast of Argentina. The introduction of nonnative foraminiferal species provides new evidence that human-mediated breaching of biogeographic barriers, will ultimately result in the biotic homogenization of foraminiferal intertidal faunal assemblages.     

Importance of foraminifera for the formation and maintenance of a coral sand cay: Green Island, Australia

 CaCO₃ production by reef-building organisms on Green Island Reef in the Great Barrier Reef of Australia is estimated and compared with the contribution of benthic foraminifera to the sediment mass of the vegetated sand cay. Major constituents of the cay are benthic foraminifera (mainly Amphistegina lessonii, Baculogypsina sphaerulata, and Calcarina hispida), calcareous algae (Halimeda and coralline algae), hermatypic corals, and molluscs. Among these reef-building organisms, benthic foraminifera are the single most important contributor to the sediment mass of the island (ca. 30% of total sediments), although their production of CaCO₃ is smaller than other reef-building organisms. Water current measurements and sediment traps indicate that the velocity of the current around Green Island is suitable for transportation and deposition of foraminiferal tests. Abundant foraminifera presently live in association with algal turf on the shallow exposed reef flat, whose tests were accumulated by waves resulting in the formation and maintenance of the coral sand cay. Accepted: 30 June 1999     

Reef habitats and associated sessile-benthic and fish assemblages across a euphotic–mesophotic depth gradient in Isla Desecheo, Puerto Rico

Quantitative surveys of sessile benthos and fish populations associated with reef habitats across a 15–50 m depth gradient were performed by direct diver observations using rebreathers at Isla Desecheo, Puerto Rico. Statistically significant differences between depths were found for total live coral, total coral species, total benthic algae, total sponges and abiotic cover. Live coral cover was higher at the mid-shelf (20 m) and shelf-edge (25 m) stations, whereas benthic algae and sponges were the dominant sessile-benthic assemblage at mesophotic stations below 25 m. Marked shifts in the community structure of corals and benthic algae were observed across the depth gradient. A total of 119 diurnal, non-cryptic fish species were observed across the depth gradient, including 80 species distributed among 7,841 individuals counted within belt-transects. Fish species richness was positively correlated with live coral cover. However, the relationship between total fish abundance and live coral was weak. Abundance of several numerically dominant fish species varied independently from live coral cover and appeared to be more influenced by depth and/or habitat type. Statistically significant differences in the rank order of abundance of fish species at euphotic vs mesophotic stations were detected. A small assemblage of reef fishes that included the cherubfish, Centropyge argi, sunshine chromis, Chromis insolata, greenblotch parrotfish, Sparisoma atomarium, yellowcheek wrasse, Halichoeres cyanocephalus, sargassum triggerfish, Xanthichthys ringens, and the longsnout butterflyfish, Chaetodon aculeatus was most abundant or only present from stations deeper than 30 m, and thus appear to be indicator species of mesophotic habitats.