Live benthic foraminiferal faunas along a bathymetrical transect (140-4800 m) in the Bay of Biscay (NE Atlantic)

In a 10-stations bathymetrical transect in the Bay of Biscay, we observed important changes in the density, composition and microhabitats of live foraminiferal faunas from the outer continental shelf to the abyssal plain. Four zones are recognised: (1) at the upper continental shelf (140 m water depth), foraminiferal densities are very high and the superficial sediment is occupied by Bolivina subaenariensis and Valvulineria bradyana. Globobulimina spp., Chilostomella oolina and Nonion fabum dominate the infaunal niches, which are positioned close to the sediment-water interface due to a strong compaction of the vertical succession of redox zones. (2) At the upper continental slope stations (300-1000 m), foraminiferal densities are high and the superficial sediments are dominated by Uvigerina mediterranea/peregrina. Deeper in the sediment, intermediate infaunal niches are occupied by Melonis barleeanus. Due to a deeper oxygen penetration, the deep infaunal taxa Globobulimina spp. and C. oolina live at a considerable depth in the sediment. (3) At the mid and lower slope stations (1000-2000 m) in the superficial sediment Cibicidoides kullenbergi and Hoeglundina elegans progressively replace U. mediterranea. U. peregrina is still a dominant taxon, reflecting its preference for a somewhat intermediate organic flux level. Deep infaunal taxa become increasingly rare. (4) At the lower slope and abyssal plane stations (deeper than 2000 m), faunal densities are very low and the fauna is composed exclusively by shallow infaunal species, such as Nuttallides umboniferus and Melonis pompilioides. The foraminiferal data together with the pore water data in the sediment give evidence of the presence of a trophic gradient from very eutrophic settings at the upper continental shelf towards oligotrophic settings at the abyssal area.     

Stable oxygen and carbon isotopes of live benthic foraminifera from the Bay of Biscay: Microhabitat impact and seasonal variability

We determined the stable oxygen and carbon isotopic composition of live (Rose Bengal stained) benthic foraminifera (> 150 μm size fraction) of seven taxa sampled along a downslope transect between 140 to 2000 m water depth in the Bay of Biscay. At the five stations, Hoeglundina elegans, Cibicidoides pachydermus, Uvigerina peregrina, Uvigerina mediterranea preferentially occupy shallow infaunal niches, whereas Melonis barleeanus and Uvigerina elongatastriata occupy an intermediate infaunal microhabitat, and Globobulimina spp. live in a deep infaunal niche close to the zero oxygen boundary.When compared with δ¹⁸O values of calcite formed in equilibrium with bottom waters, U. peregrina forms its test in close equilibrium with bottom water δ¹⁸O. All other foraminiferal taxa calcify with a constant offset to calculated equilibrium calcite. There is no systematic relationship between the foraminiferal microhabitat depth and the Δδ¹⁸O between foraminiferal and equilibrium calcite. We calculated correcting factors for the various taxa, which are needed for constructing multispecies-based oxygen isotope records in paleoceanographic studies of the study area.The δ¹³C values of foraminiferal taxa investigated in this study do neither record bottom water δ¹³C_DIC in a 1 : 1 relationship nor with a constant offset, but appear to be mainly controlled by microhabitat effects. The increase of δ¹³C values of shallow infaunal taxa with increasing water depth reflects the decrease of the exported flux of organic carbon along the bathymetric transect and early diagenetic processes in the surface sediment. This is particularly the case for the shallow infaunal U. peregrina. The δ¹³C values of deep infaunal Globobulimina spp. are much less dependent on the exported organic matter flux. We suggest that the Δδ¹³C between U. peregrina and Globobulimina spp. can shed light on the various pathways of past degradation of organic detritus in the benthic environments.At a station in 550 m water depth, where periodic eutrophication of sediment surface niches was demonstrated previously, we performed a two-year seasonal survey of the isotopic composition of foraminiferal faunas. No marked seasonal changes of the stable carbon isotopic composition of shallow, intermediate and deep infaunal foraminiferal taxa were observed. Thus, the δ¹³C values of foraminiferal individuals belonging to the > 150 μm fraction may result from rather long-term calcification processes lasting for several weeks or months, which limit the impact of ephemeral ¹²C enrichment of shallow infaunal niches on the isotope chemistry of adult individuals during eutrophic periods. Only highly opportunistic taxa reproducing or calcifying during phytoplankton bloom periods and the subsequent deposits of phytoplankton remains in the benthic environment may exhibit a particularly low δ¹³C, indicative of such short productive periods.     

Paleoenvironmental changes across the Cenomanian/Turonian Boundary Event (Oceanic Anoxic Event 2) as indicated by benthic foraminifera from the Demerara Rise (ODP Leg 207)

This study is based on Cenomanian to lower Turonian sediments of Ocean Drilling Program (ODP) Sites 1258, 1259, 1260, and 1261 from the Demerara Rise (Leg 207, western tropical Atlantic, off Suriname) that are oriented along a paleodepth transect. Studied sediments include the Cenomanian/Turonian Boundary Event (CTBE) or Oceanic Anoxic Event 2 (OAE 2) and consist of laminated black shales with TOC values between 5% and 10% below and above OAE 2 and up to 29% within the OAE 2 interval. Benthic foraminiferal assemblages in this eutrophic environment are generally characterized by low diversities and strong fluctuations of abundances, indicating oxygen depletion and high organic matter fluxes. Dominant taxa at all sites are Bolivina anambra, Gavelinella dakotensis, Tappanina sp., Praebulimina prolixa, and Neobulimina albertensis. Based on the positive stable carbon isotope excursion characteristics of OAE 2 we subdivided the studied successions into three intervals: (1) the interval below OAE 2; (2) the carbon isotope excursion reflecting OAE 2; and (3) the interval above OAE 2. In the bathymetrically shallower Sites 1260 and 1261 benthic foraminiferal assemblages indicate anoxic to sometimes slightly dysoxic bottom-water conditions below the OAE 2 interval. The bathymetrically deepest Site 1258, in contrast, reflects more oxygenated bottom waters with an almost continuous occurrence of benthic foraminifera. It is therefore suggested that the shallower sites were located within the oxygen minimum zone (OMZ), whereas Site 1258 was below the OMZ. During OAE 2 anoxic conditions prevailed at the shallower sites. At Site 1258 benthic foraminifera indicate severe dysoxic but not anoxic conditions. This pattern is proposed to reflect a strengthening of the OMZ possibly related to increased primary production during OAE 2. A short-term repopulation event of benthic foraminifera in the lower third of the OAE 2 interval was observed at all sites, reflecting a brief bottom-water oxygenation event. This short-lived event parallels a surface-water cooling and is probably equivalent to the “Plenus Cool Event” in Europe and the “benthonic zone” in the U.S. Western Interior. The benthic foraminifera of a ~0.5 Ma interval following OAE 2 still indicate oxygen depletion of bottom waters. Subsequently, however, a strong increase in benthic foraminiferal abundance and diversity reflects better oxygenation of the bottom-water masses, probably related to a weakening of the OMZ due to decreasing organic matter flux.     

Ontogenetic effects on stable carbon and oxygen isotopes in tests of live (Rose Bengal stained) benthic foraminifera from the Pakistan continental margin

We determined the stable oxygen and carbon isotopic composition of live (Rose Bengal stained) tests belonging to different size classes of two benthic foraminiferal species from the Pakistan continental margin. Samples were taken at two sites, with water depths of about 135 and 275 m, corresponding to the upper boundary and upper part of the core region of the oxygen minimum zone (OMZ). For Uvigerina ex gr. Uvigerina semiornata and Bolivina aff. Bolivina dilatata, δ¹³C and δ¹⁸O values increased significantly with increasing test size. In the case of Uvigerina ex gr. U. semiornata, δ¹³C increased linearly by about 0.105‰ for each 100-μm increment in test size, whereas δ¹⁸O increased by 0.02 to 0.06‰ per 100 μm increment. For Bolivina aff. B. dilatata the relationship between test size and stable isotopic composition is better described by logarithmic equations. A strong positive linear correlation is observed between δ¹⁸O and δ¹³C values of both taxa, with a constant ratio of δ¹⁸O and δ¹³C values close to 2:1. This suggests that the strong ontogenetic effect is mainly caused by kinetic isotope fractionation during CO₂ uptake. Our data underline the necessity to base longer δ¹⁸O and δ¹³C isotope records derived from benthic foraminifera on size windows of 100 μm or less. This is already common practice in down-core isotopic studies of planktonic foraminifera.     

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.     

Ammonite-benthic Foraminifera turnovers across the Lower-Middle Jurassic transition in the Lusitanian Basin (Portugal)

This paper describes and characterises the co-occurrence of ammonite and benthic foraminiferal assemblages across the São Gião outcrop (Central Portugal), a reference section for the Lower-Middle Jurassic boundary in the Lusitanian Basin. The upper Toarcian-lower Aalenian marls and marly-limestones in this section provide a precise and detailed ammonite-based biostratigraphic zonation, with a mixed assemblage of northwest European and Mediterranean faunal elements, associated with benthic foraminifera assemblages with northern hemisphere affinities, both correlatable with the Aalenian GSSP at the Fuentelsaz section (Iberian Cordillera, Spain). A total of 447 well-preserved ammonite specimens and 13.116 foraminifera have been studied; no evidence was detected of any taphonomic processes that could have changed the original assemblages. From a biostratigraphic point of view, the ammonite record has enabled four biostratigraphic units to be recognised (the Mactra and Aalensis subzones of the Aalensis Biozone in the upper Toarcian, and the Opalinum and Comptum subzones of the Opalinum Biozone in the lower Aalenian). With regard to the benthic foraminifera, the taxa identified have enabled the Astacolus dorbignyi Zone and 11 bioevents to be identified, most of which representing local biostratigraphic proxies. However, the increase in the relative abundance of Lenticulina exgaleata Dieni from the upper part of the Opalinum Subzone to the lower part of the Comptum Subzone has a regional value. The constant and continuous ammonite record of northwest European taxa, together with typical Mediterranean taxa – namely Grammoceratinae – throughout the section, the high relative abundance of Miliolina representatives – generally interpreted as foraminifers typical of shallow waters – and the absence of foraminiferal forms typical of cool waters, do not support the inference of cool seawater temperatures attributed to the Early Aalenian, or the global character of the “Comptum cooling event”, at least with reference to the Lusitanian Basin.     

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.     

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.     

Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight

Cold-water coral ecosystems are characterised by a high diversity and population density. Living and dead foraminiferal assemblages from 20 surface sediment samples from Galway and Propeller Mounds were analysed to describe the distribution patterns of benthic foraminifera on coral mounds in relation to different sedimentary facies. Hard substrates were examined to assess the foraminiferal microhabitats and diversities in the coral framework. We recognised 131 different species, of which 27 prefer an attached lifestyle. Epibenthic species are the main constituents of the living and dead foraminiferal assemblages. The frequent species Discanomalina coronata was associated with coral rubble, Cibicides refulgens showed preference to the off-mound sand veneer, and Uvigerina mediterranea displayed abundance maxima in the main depositional area on the southern flank of Galway Mound, and in the muds around Propeller Mound. The distribution of these species is rather governed by their specific ecological demands and microhabitat availability than by the sedimentary facies. Benthic foraminiferal assemblages from coral mounds fit well into basin-wide-scale distribution patterns of species along the western European continental margin. The diversity of the foraminiferal faunas is not higher on the carbonate mounds as in their vicinity. The living assemblages show a broad mid-slope diversity maximum between 500 and 1,300 m water depth, which is the depth interval of coral mound formation at the Celtic and Amorican Margin. The foraminiferal diversity maximum is about 700 m shallower than comparable maxima of nematodes and bivalves. This suggests that different processes are driving the foraminiferal and metazoan diversity patterns.     

Benthic foraminiferal turnover across the Cretaceous/Paleogene boundary at Agost (southeastern Spain): paleoenvironmental inferences

We studied Upper Cretaceous and Lower Paleogene benthic foraminifera from the Agost section (southeastern Spain) to infer paleobathymetrical changes and paleoenvironmental turnover across the Cretaceous/Paleogene (K/P) transition. Benthic foraminifera indicate uppermost bathyal depths at Agost during the Abathomphalus mayaroensis Biochron (from about 400 kyr before the K/P boundary) through the early Plummerita hantkeninoides Biochron (about 120–150 kyr before that boundary). The depth increased to middle bathyal for the remainder of the Cretaceous, and remained so for the Danian part of the studied section (Parasubbotina pseudobulloides Biochron, at least 200 kyr after the K/P boundary). There were no perceivable bathymetrical changes at the K/P boundary, where 5% of the species became extinct, and the species composition of the benthic foraminiferal fauna changed considerably. Below the boundary, infaunal morphogroups constitute up to 65–73% of the faunas. Directly above the boundary, in the black clays of the lower Guembelitria cretacea Biozone, benthic foraminifera are rare. Several opportunistic taxa (e.g. the agglutinant Haplophragmoides sp.) have short peaks in relative abundance, possibly reflecting low-oxygen conditions as well as environmental instability, with benthos receiving food from short-lived, local blooms of primary producers. Above the clays through the end of the studied interval, epifaunal morphogroups dominate (up to 70% of the assemblages) or there is an even mixture or epifaunal and infaunal morphogroups. Infaunal groups do not recover to pre-extinction relative abundances, indicating that the food supply to the benthos did not recover fully over the studied interval (about 200 kyr after the K/P boundary). The benthic foraminiferal faunal changes are compatible with the direct and indirect effects of an asteroid impact, which severely destabilized primary producers and the oceanic food web that was dependent upon them.     

Environmental perturbation in the southern Tethys across the Paleocene/Eocene boundary (Dababiya,Egypt): Foraminiferal and clay mineral records

Foraminiferal and clay mineral records were studied in the upper Paleocene to lower Eocene Dababiya section (Egypt). This section hosts the GSSP for the Paleocene/Eocene boundary and as such provides an expanded and relatively continuous record across the Paleocene/Eocene Thermal Maximum (PETM). Deposition of illite–smectite clay minerals is interpreted as a result of warm and arid conditions in the southern Tethys during the latest Paleocene. Benthic foraminiferal assemblages are indicative of seasonal variation of oxygen and food levels at the seafloor. A sea-level fall occurred in the latest Paleocene, followed by a rise in the earliest Eocene. Foraminiferal diversity and densities decreased strongly at the P/E boundary, coinciding with the level of global extinction of benthic foraminifera (BEE) and start of the Carbon Isotope Excursion (CIE) and PETM. In the lower CIE, the seafloor of the stratified basin remained (nearly) permanently anoxic and azoic. A sudden increase in mixed clay minerals (kaolinite and others) suggests that warm and perennial humid conditions prevailed on the continent. High levels of TOC and phosphathic concretions in the middle CIE are evidence for increased organic fluxes to the sea floor, related to upwelling and to augmented continental runoff. Low densities of opportunistic taxa appeared, indicating occasional ephemeral oxygenation and repopulation of the benthic environment. The planktic community diversified, although conditions remained poor for deep-dwelling taxa. An increase in illite–smectite dominated clay association is considered to mark the return of a seasonal signature on climatic conditions. During the late CIE environmental conditions changed to seasonally fluctuating mesotrophic conditions and diverse and rich benthic and planktic foraminiferal communities developed. Post-CIE planktic faunas consisted of both deep and shallow-dwelling taxa and buliminid-dominated benthic assemblages reflect fluctuating mesotrophic conditions.The frequent environmental perturbations during the CIE/PETM at Dababiya provided a rather specialized group of foraminiferal taxa (i.e., Anomalinoides aegyptiacus) the opportunity to repopulate, survive and subsequently dominate by a hypothesized capacity to switch to an alternative life strategy (population dynamics, habitat shift) or different metabolic pathway. The faunal record of Dababiya provides insight into the cause and development of the BEE: various severe global changes during the PETM (e.g., ocean circulation, CaCO₃-dissolution, productivity and temperature changes) disturbed a wide range of environments on a geologically brief timescale, explaining together the geographically and temporally variable character of the BEE. This allowed a number of specific but different foraminiferal assemblages composed of stress-tolerant and opportunistic taxa to be successful during and after the periods of environmental perturbations associated with the PETM.     

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.     

Distribution of recent foraminiferal assemblages near the Ombrone River mouth (Northern Tyrrhenian Sea, Italy)

Benthic foraminiferal assemblages in 127 samples, collected at water depth, ranging between 15 and 184 m on the Southern Tuscany continental shelf off the Ombrone River were analysed. Statistical analyses (Cluster and Principal Component Analysis) performed on the 48 most abundant species (>5%) and sedimentological data led to the identification of three clusters related to the size of grain sediment (sand, silt or clay). Q-mode cluster analysis singled out six groups, corresponding to six distinct foraminiferal assemblages: a typical infralittoral assemblage (15-39 m) on sandy silt, clayey silt or silty clay, dominated by Ammonia and Elphidium species, with Eggerelloides scabrus, Rectuvigerina phlegeri and Valvulineria bradyana; a second assemblage (24-78.5 m) associated with vegetated environments or sandy bottoms (Elphidium crispum, Rosalina bradyi, Asterigerinata mamilla, Neoconorbina terquemi, and Tretomphalus concinnus); a third assemblage recognised between 30 and 90 m water depth and characterised by the dominance of the opportunistic species V. bradyana (related to silty bottoms), with Bulimina marginata, R. phlegeri, Ammonia inflata and Ammonia beccarii as other common taxa; an upper circalittoral assemblage (70-100 m) on silty clays, containing B. marginata, Cassidulina carinata and V. bradyana; a lower circalittoral assemblage (95-177 m) on clayey bottoms, with B. marginata, Textularia bocki and Uvigerina mediterranea; and finally, a second lower circalittoral assemblage (104-184 m) on clayey sediments, dominated by two Uvigerina species (U. mediterranea and Uvigerina peregrina), with Sphaeroidina bulloides and B. marginata. The typical V. bradyana assemblage, characterised by relatively low diversity and high dominance, marks the most eutrophicated area running parallel to the coast. The spatial distribution of assemblages is closely associated with sea-bottom sedimentary environments and bathymetry but it is also probably influenced by the outflow of the Ombrone River. The composition, structure and distribution of V. bradyana assemblage suggest an environmental model, useful for paleogeographic reconstruction in areas characterised by a river mouth and a closed morphological setting typical of a nutrient-trap.     

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.     

Faunal turnovers in central Pacific benthic foraminifera during the Paleocene-Eocene thermal maximum

Although it is well known that the Paleocene/Eocene thermal maximum (PETM) coincided with a major benthic foraminiferal extinction event, the detailed pattern of the faunal turnover has not yet been clarified. Our high-resolution benthic foraminiferal and carbon isotope analyses at the low latitude Pacific Ocean Shatsky Rise have revealed the following record of major faunal transitions: (1) An initial turnover which involved the benthic foraminiferal extinction event (BFE). The BFE, marked by a sharp transition from Pre-extinction fauna to Disaster fauna represented by small-sized Bolivina gracilis, expresses the onset of the PETM and the abrupt extinction of about 30% of taxa. This faunal transition lasted about 45-74 kyr after the initiation of the PETM and was followed by: (2) the appearance of Opportunistic fauna represented by Quadrimorphina profunda, which existed for about 74-91 kyr after the initiation of the PETM. These two faunas, which appeared after the extinction event, are characterized by low diversity and dwarfism, possibly due to lowered oxygen condition and decreased surface productivity. The second pronounced turnover involved the gradual recovery from Opportunistic Fauna to the establishment of Recovery fauna, which coincided with the recovery about 83-91 kyr after its initiation.     

Late Pliocene benthic foraminifera and mollusks from the Atsipades Section, Central Crete; Palaeoecological distribution and use in palaeoenvironmental assessment

Changes in benthic foraminiferal and mollusk assemblages from the lower part of the Upper Pliocene of the Atsipades Section (Iraklion Basin, central Crete) were studied. The Atsipades Section represents a shallowing-upward sequence from outer shelf blue-gray clays at the bottom of the sequence, deposited below the storm wave base, to shallow inner shelf deposits affected by storm waves at the top. The foraminiferal assemblage at the bottom of the sequence is dominated by Bolivina spathulata, Bolivina dilatata and Uvigerinidae, a microfossil assemblage corresponding to the deepest deposits formed under dysoxic sea-floor conditions. Foraminiferal assemblages of the middle part of the section are highly diversified, predominantly Haynesina depressula, Cassidulina carinata and Reusella spinulosa. The top of the section is mainly characterised by Asterigerinata planorbis, Bolivina pseudoplicata, Cibicides lobatulus and Elphidium sp., a typically epiphytic foraminiferal assemblage which can be correlated with the presence of an algal covered sea-bottom. Within this general environmental trend, a minor shallowing cycle can be differentiated. The boundaries of this cycle can be inferred, based on a substantial microfossil assemblage change and on the coincidence of species diversity maximum and a planktonic/benthic (P/P + B) ratio peak. Nonetheless, upwelling currents and/or over-abundance of nutrients due to continental outflow could also contribute to increased diversity and P/P + B ratio. The character of the mollusk assemblages is in accordance with these trends. Moreover, the increase in diversity and in sculpture constitutes a clear indication of an increase in hydrodynamic energy related to a shallowing-upward trend.     

Morphology and microhabitat preferences of benthic foraminifera from the northwest Atlantic Ocean

The distribution of Rose Bengal stained calcareous benthic foraminifera was determined in six ☐ cores raised from water depths between 200 and 3000 m on the Nova Scotian continental margin and Gulf of Maine. The taxa can be separated into four microhabitats within the surficial sediments. Epifaunal taxa are generally found in the top cm, intermediate infaunal taxa are found from about 1 to 4 cm and deep infaunal taxa are found at > 4 cm sediment depth in at least one ☐ core. A fourth group, shallow infaunal taxa, is found in the top 2 cm and is inferred to be infaunal based on wall porosity characteristics and test shapes similar to infaunal taxa. The epifaunal, shallow infaunal and intermediate infaunal taxa maintain their positions within the sediments from core to core, whereas the deep infaunal taxa are found at progressively shallower sediment depths in cores within increasing organic carbon contents from shallower water depths.Each microhabitat category has distinct morphological characteristics. Epifaunal taxa have plano-covex or biconvex cross sections, trochospiral coiling and large pores absent or found on only one side. Shallow infaunal taxa have uniserial, triserial, or planispiral coiling, with surface ornamentation present on a number of taxa. The intermediate infaunal taxa have rounded peripheries, pores over the entire test and planispiral coiling, with the exception ofCibicidoides bradyi which has trochospiral coiling. The deep infaunal taxa have, in general, planispiral or triserial coiling with cylindrical or ovate shaped tests.     

A conceptual model explaining benthic foraminiferal microhabitats

We present a conceptual model which explains benthic foraminiferal microhabitat preferences in terms of differences in the downward organic flux. We argue that under oligotrophic conditions the microhabitat depth is controlled by the availability of metabolizable food particles in the sediment. Under more eutrophic conditions, the ecosystem is no longer food-controlled, but instead, a critical oxygen level determines down to what depth we find a living fauna. Under food-limited conditions, anaerobic degradation of organic matter may provide an additional food source around the redox front, which could explain deep infaunal maxima reported in the literature. In a sample transect through the Adriatic Sea, both microhabitat controls (food-limited and oxygen-limited) are present. On the shelf and the upper part of the slope, the rather shallow depth of the microhabitat is controlled by a critical oxygen level. In the 1250 m deep southern basin and on the lower part of the slope, on the contrary, the availability of metabolizable organic matter, and not a critical oxygen level, determines down to what depth living foraminifera are found.     

Carbon and oxygen isotope geochemistry of live (stained) benthic foraminifera from the Aleutian Margin and the Southern Australian Margin

Comparisons of ambient bottom-water geochemistry and stable isotopic values of the tests of living (stained) calcareous benthic foraminifera from the North Pacific (on the Aleutian Margin, water depth 1988 m) and Murray Canyons group in the Southern Indian Ocean (Australian Margin, water depths 2476 m and 1634 m) provide modern environmental analogs to calibrate paleoenvironmental assessments. Consistent with the hypothesis that microhabitat preferences influence foraminiferal isotopic values, benthic foraminifera from both margins were depleted in ¹³C with respect to bottom-water dissolved inorganic carbon (DIC). The carbon isotope values of deep infaunal foraminifera (Chilostomella oolina, Globobulimina pacifica) showed greater differences from estimates of those of DIC than shallow benthic foraminifera (Bulimina mexicana, Bolivinita quadrilatera, Pullenia bulloides). This study provides new isotopic and ecological information for B. quadrilatera. The mean Δδ¹³C value, defined as foraminiferal δ¹³C values minus estimated ambient δ¹³C values from the Aleutian Margin, is 0.97‰ higher for G. pacifica than the mean from the Murray Canyon. This difference may result either from genetic or biological differences between the populations or from differences in environmental isotopic influences (such as pore water differences) that were not accounted for in the equilibrium calculations. These analyses provide calibration information for the evaluation of bottom water conditions and circulation patterns of ancient oceans based on fossil foraminiferal geochemistry.     

Benthic foraminiferal paleoenvironmental reconstruction from the Upper Coniacian-Lower Campanian succession in north-western Tunisia

Quantitative analysis of benthic foraminifera is used to characterize the paleoenvironments of the Upper Coniacian-Lower Campanian succession in the Jbil section of north-western Tunisia. Foraminiferal parameters and benthic foraminiferal assemblages show that the studied section includes four distinct paleoenvironmental phases. From oldest to youngest, these are as follows: (1) an interval with a Praebulimina reussi assemblage with infaunal ratios as high as 96.1%. High abundances of P. reussi, reflecting an increase in organic matter flux to the seafloor (meso-to eutrophic) under oxygenated bottom-water conditions. (2) An interval characterized by a Gavelinella costulata assemblage with mixed infaunal/epifaunal foraminifera with higher Fisher's alpha values (ranging from 4 to 15.2), reflecting mesotrophic conditions in an outer shelf environment. (3) An interval with a Gaudryina laevigata assemblage indicative of a middle to outer shelf environment; there is a considerable increase in infaunal agglutinated foraminifera, as well as a relatively abundant and moderately diversified oxic/suboxic foraminifera. (4) The final interval occurs in the lower Campanian (the Globotruncana ventricosa Zone) and includes a Bolivinoides decoratus assemblage reflecting an outer shelf to upper bathyal environment. It contains a higher planktonic percentage and biodiversity with a slight increase in dysoxic species; the mixed infaunal/epifaunal content (57.6 to 73.3%) reflects mesotrophic conditions. Four well-recognized major sea-level falls are matched by the dual signatures of eustatic sea-level changes. These are coincident with the results of this study, which represent the first documentation of these events in Tunisian faunal and paleoenvironmental changes, at the following boundaries: Coniacian/Santonian, intra-Santonian, Santonian/Campanian, and intra-early Campanian.