Experimental and field evidence of Antarctic foraminiferal tolerance to anoxia and hydrogen sulfide

The common occurrence of foraminifera in strata deposited under anoxic environmental conditions throughout the geologic record raises the question of whether foraminifera actually survive anoxic or reducing conditions. To test this, benthic foraminifera from McMurdo Sound, Antarctica were exposed to anoxic or reducing conditions for 30 days. These Antarctic forms are known to be exposed to anoxic events in their natural environment. An adenosine triphosphate (ATP) assay was used to determine foraminiferal survival, and their ultrastructure was examined using transmission electron microscopy (TEM) to evaluate any possible effect that might have resulted from exposure to anoxic or reducing conditions. The experimental treatments did not have any statistically significant effects upon the foraminifers with regards to survival or average ATP content per individual including the four most common living species. Some ultrastructural evidence for encystment (fibrillar membrane-bound bodies in the cytoplasm of many specimens) was observed. Field data include the observation thatGlobocassidulina cf.G. biora resides in sediments to depths of 7 cm, as evidenced by ultrastructural investigations. A specimen ofGlobocassidulina cf.G. biora collected from sediments containing anoxic pore waters showed numerous bacteria within the confines of the organic lining. The occurrence of cytoplasmic inclusions similar in appearance to car☐ysomes within these bacteria suggest possible affinities to the chemolithotrophicThiobacillus. Bacteria of this type were not observed in specimens collected from oxygenated pore waters. These field and laboratory findings suggest that certain foraminifera may be facultative anaerobes able to withstand reducing conditions.     

Palaeoenvironmental conditions preceding the Messinian Salinity Crisis: A case study from Gavdos Island

The Messinian pre-evaporitic sedimentary succession of Gavdos Island (Metochia section) is a nearly uninterrupted succession of marine sediments, dominated by finely laminated diatomaceous marls, which are cyclically alternating with clayey diatomites and white diatomites. The qualitative and quantitative analysis of the planktonic foraminiferal fauna allowed the recognition of nine bioevents, which have been astronomically dated for the Mediterranean. The base of the diatomitic succession in Gavdos Island is dated at 6.722 Ma and the top at 6.015 Ma. The studied section contains benthic foraminiferal genera characteristic of an outer shelf to slope environment. The qualitative and quantitative analysis of this microfauna revealed three benthic foraminiferal fossil assemblages and the occurrence of allochthonous species transported into the bathyal environment by current activity. The cyclical pattern of the benthic foraminifera assemblages indicates that the studied sediments have been affected by repeated episodes of basin restriction characterized by low diversity benthic foraminifera populations, and a limited planktonic foraminifer association typified by shallow, surface-dwelling forms. This restriction was partly due to Antarctic cooling, which produced palaeo-Mediterranean sea-level oscillations during the Early Messinian, as a prelude to closure of the Atlantic connections. The relative impact of climatic versus tectonic control on sedimentation patterns within this basin is discussed.     

Paleoceanographic implications of smaller benthic and planktonic foraminifera from the Eocene Pazin Basin (Coastal Dinarides,Croatia)

Summary Smaller benthic and planktonic foraminifera from the clastic sediments of the Pazin Basin (Istria, Croatia) were studied in order to obtain more data about paleoceanographic conditions that existed in the Middle Eocene Dinaric foreland basin. The succession investigated corresponds to the Middle Eocene planktonic foraminiferal zones Globigerapsis kugleri/Morozovella aragonensis (P11), Morozovella lehneri (P12), and Globigerapsis beckmanni (P13). Benthic foraminiferal assemblages from the clastic succession are dominated by epifaunal trochospiral genera suggesting oligotrophic to mesotrophic conditions and moderately oxygenated bottom waters. Planktonic foraminiferal assemblages indicate mesotrophic to eutrophic conditions of the surface waters, with increased eutrophication in the upper part of the section. Water depth, based on the ratio between planktonic and epifaunal benthic foraminifera and on the recognized species of cosmopolitan benthic foraminifera, was estimated to have been between about 900 and 1200 m. The basin was elongated and open to marine currents on both sides allowing good circulation and ventilation of the bottom water.     

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     

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.     

Pleistocene agglutinated foraminifera from the Lomonosov Ridge and Amundsen Basin, Arctic Basin. Initial report on piston cores 2177-5 (KAL) and 2176-3 (KAL)

Noncalcareous Pleistocene sediments of the Central Arctic Ocean contain sparse benthic foraminiferal assemblages consisting entirely of agglutinated taxa. Deep water agglutinated foraminifera are studied from two piston cores collected from the Lomonosov Ridge and Amundsen Basin [Cores PS 2177-5 (KAL) and 2176-3 (KAL)]. Core PS 2177-5 (KAL) contains an assemblage of 10 species, dominated by Cyclammina pusilla Brady, and is interpreted to reflect a bathyal environment with variable organic flux and nutrition levels. Core PS 2176-3 (KAL) in the Amundsen Basin yielded a very depauperate benthic foraminiferal assemblage. It is assumed that the environment was inhospitable for agglutinated foraminifera.     

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.     

Foraminiferal turnover across the Eocene–Oligocene transition at Fuente Caldera, southern Spain: No cause–effect relationship between meteorite impacts and extinctions

We studied planktic and small benthic foraminifera from the Fuente Caldera section, southern Spain, across the Eocene–Oligocene transition. Benthic foraminifera indicate lower bathyal depths for the late Eocene and earliest Oligocene. Detailed high-resolution sampling and biostratigraphical data allowed us to date precisely layers with evidence for meteorite impact (Ni-rich spinel), which occur in the lower part of the planktic foraminiferal Globigerapsis index Biozone and in the middle part of the small benthic foraminiferal Cibicidoides truncanus (BB4) Biozone (middle Priabonian, late Eocene). Major turnovers of foraminifera occur at the Eocene/Oligocene boundary, only. The impact did not occur at a time of planktic or benthic foraminiferal extinction events, and the late Eocene meteorite impacts did thus not cause extinction of foraminifera. The most plausible cause of the Eocene/Oligocene boundary extinctions is the significant cooling, which generated glaciation in Antarctica and eliminated most of the warm and surface-dwelling foraminifera.     

A Decline in Benthic Foraminifera following the Deepwater Horizon Event in the Northeastern Gulf of Mexico

Sediment cores were collected from three sites (1000–1200 m water depth) in the northeastern Gulf of Mexico from December 2010 to June 2011 to assess changes in benthic foraminiferal density related to the Deepwater Horizon (DWH) event (April-July 2010, 1500 m water depth). Short-lived radioisotope geochronologies (²¹⁰Pb, ²³⁴Th), organic geochemical assessments, and redox metal concentrations were determined to relate changes in sediment accumulation rate, contamination, and redox conditions with benthic foraminiferal density. Cores collected in December 2010 indicated a decline in density (80–93%). This decline was characterized by a decrease in benthic foraminiferal density and benthic foraminiferal accumulation rate (BFAR) in the surface 10 mm relative to the down-core mean in all benthic foraminifera, including the dominant genera (Bulimina spp., Uvigerina spp., and Cibicidoides spp.). Cores collected in February 2011 documented a site-specific response. There was evidence of a recovery in the benthic foraminiferal density and BFAR at the site closest to the wellhead (45 NM, NE). However, the site farther afield (60 NM, NE) recorded a continued decline in benthic foraminiferal density and BFAR down to near-zero values. This decline in benthic foraminiferal density occurred simultaneously with abrupt increases in sedimentary accumulation rates, polycyclic aromatic hydrocarbon (PAH) concentrations, and changes in redox conditions. Persistent reducing conditions (as many as 10 months after the event) in the surface of these core records were a possible cause of the decline. Another possible cause was the increase (2–3 times background) in PAH’s, which are known to cause benthic foraminifera mortality and inhibit reproduction. Records of benthic foraminiferal density coupled with short-lived radionuclide geochronology and organic geochemistry were effective in quantifying the benthic response and will continue to be a valuable tool in determining the long-term effects of the DWH event on a larger spatial scale.     

Kleptoplastidic benthic foraminifera from aphotic habitats: insights into assimilation of inorganic C,N and S studied with sub-cellular resolution

The assimilation of inorganic compounds in foraminiferal metabolism compared to predation or organic matter assimilation is unknown. Here, we investigate possible inorganic-compound assimilation in Nonionellina labradorica, a common kleptoplastidic benthic foraminifer from Arctic and North Atlantic sublittoral regions. The objectives were to identify the source of the foraminiferal kleptoplasts, assess their photosynthetic functionality in light and darkness and investigate inorganic nitrogen and sulfate assimilation. We used DNA barcoding of a ~ 830 bp fragment from the SSU rDNA to identify the kleptoplasts and correlated transmission electron microscopy and nanometre-scale secondary ion mass spectrometry (TEM-NanoSIMS) isotopic imaging to study ¹³C-bicarbonate, ¹⁵N-ammonium and ³⁴S-sulfate uptake. In addition, respiration rate measurements were determined to assess the response of N. labradorica to light. The DNA sequences established that over 80% of the kleptoplasts belonged to Thalassiosira (with 96%–99% identity), a cosmopolitan planktonic diatom. TEM-NanoSIMS imaging revealed degraded cytoplasm and an absence of ¹³C assimilation in foraminifera exposed to light. Oxygen measurements showed higher respiration rates under light than dark conditions, and no O₂ production was detected. These results indicate that the photosynthetic pathways in N. labradorica are not functional. Furthermore, N. labradorica assimilated both ¹⁵N-ammonium and ³⁴S-sulfate into its cytoplasm, which suggests that foraminifera might have several ammonium or sulfate assimilation pathways, involving either the kleptoplasts or bona fide foraminiferal pathway(s) not yet identified.     

Stable isotopic study of Oligocene-Miocene sediments from DSDP Site 354, Equatorial Atlantic

The oxygen- and carbon-isotope compositions of planktic and benthic foraminifera and calcareous nannofossils from Middle Oligocene-Early Miocene Equatorial Atlantic sediments (DSDP Site 354) indicate two important paleoceanographic changes, in the Late Oligocene (foraminiferal Zone P.21) and in the Early Miocene (foraminiferal Zone N.5). The first change, reflected by a δ¹⁸O increase of 1.45‰ inGlobigerina venezuelana, affected only intermediate pelagic and not surface, deep or bottom waters. The second change affected surface and intermediate waters, whereas deep and bottom waters showed only minor fluctuations. In the case of the former the isotope effect of the moderate ice accumulation on the Antarctic continent is amplified in the Equatorial Atlantic by changes in the circulation pattern. The latter paleoceanographic change, reflected by a significant increase in¹⁸O in both planktic and benthic forms (about 1.0‰ and 0.5‰, respectively), may have been caused by ice volume increase and temperature decrease. Both oxygen- and carbon-isotope compositions indicate a marked depth-habitat stratification for planktic foraminifera and calcareous nannofossils. Three different dwelling groups are recognized: shallowGlobigerinoides, Globoquadrina dehiscens, Globorotalia mayeri and nannofossils; intermediateGlobigerina venezuelana; and deepCatapsydrax dissimilis. The comparison of foraminifera and calcareous nannofossils suggests that the isotopic compositions of nannofossils are generally controlled by the same parameters which control the isotopic composition of shallow-dwelling foraminifera, but the former are more enriched in¹⁸O.     

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.     

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.     

Stepwise postglacial migration of benthic foraminifera into the abyssal northeastern Norwegian Sea

Accumulation rates of individual species (SpecAR) and relative abundances (percentages) of benthic foraminifera of an AMS ¹⁴C-dated high resolution sediment core from the Norwegian Seas (water depth: 2707 m) provide a record of the faunal fluctuations from the last glacial maximum across the Weichselian deglaciation to the Holocene. During glacial times, the total foraminifera accumulation remains at a very low level (< 100 specimens cm⁻² kyr⁻¹) and is dominated by two endofaunal species: Oridorsalis umbonatus (Reuss) and Siphotextularia rolshauseni (Phleger and Parker) and reworked specimens of the genus Elphidium. The following deglaciation period exhibits an increase of the AR of the total fauna at 14 kyr B.P. The species distribution is marked by the last appearance of S. rolshauseni and the first postglacial appearance of suspension feeding Cibicidoides wuellerstorfi (Schwager) 13 kyr B.P. The absolute maximum of benthic foraminiferal AR (2750 spec cm⁻² kyr⁻¹ occurred near 9 kyr B.P. at the end of the deglaciation. This maximum also marks the re-appearance of the agglutinating species Cribrostomoides subglobosus (Sars). The post-glacial interval is characterized by a twofold reduction of the total accumulation of benthic foraminifera. The species distribution shows two new species: Ammobaculites agglutinans (d'Orbigny, at 6 kyr B.P.) and Epistominella exigua (Brady, at 3.5 kyr B.P.). The total AR indicates benthic activity during glacial times was at a low level. It was significantly higher during the Holocene with an abrupt increase of benthic foraminiferal abundance from 10 to 9 kyr B.P. The Stepwise re-invasion into the postglacial deep-sea environment maybe related to specific habitat preference.     

Thanetian–Ilerdian coralline algae and benthic foraminifera from northeast India: microfacies analysis and new insights into the Tethyan perspective

Thanetian–Ilerdian carbonate deposits from the Lakadong Limestone in Assam Shelf, Meghalaya, northeast India, are studied with respect to microfacies distributions and controlling ecological factors on dominant biogenic components. Palaeoenvironmental implications are inferred following the detailed analysis of microfacies characterized by rich assemblages of coralline red algae and benthic foraminifera. The carbonate sediments have been interpreted as lagoonal to outer shelf facies. It is envisaged that the analysed benthic communities thrived in a meso‐oligotrophic regime above the fair‐weather wave base. The Lakadong Limestone constitutes a well‐preserved record of Late Palaeocene–earliest Eocene shallow marine carbonate ecosystem and has high potential to decipher its response to an interval of distinct changes in climate and tectonic settings. The abundance of oligotrophic larger benthic foraminifera in the Lakadong Limestone is comparable to the foraminiferal assemblages of west Tethys. The phylogenetic changes (‘Larger Foraminiferal Turnover’, LFT) and subsequent rapid radiation of typical Eocene larger benthic foraminifera (Alveolina, Nummulites) usually observed in the west Tethys have also been observed in the upper part of the Lakadong Limestone. The eastward migration of Eocene foraminifera from the west coincided with the India‐Asia collision and global warming events at the Palaeocene–Eocene boundary that may have produced a wide array of modifications in biogeography, seasonal run‐offs and ocean circulation pathways. The data indicate that rapid rate of migration from west before the onset of geographic barriers and/or timely restoration of pan‐Tethyan environmental conditions ensured the incidence of these forms in the earliest Eocene sediments.     

Heterosteginapapyracea Seguenza du Miocène du Corridor Sud-Rifain (Maroc) : biométrie et particularités paléoécologiques

Biometric study of the inner features of Heterostegina specimens preserved in tortonian sediments of the oued Yhoudi member allows to confirm the presence of species Heterostegina papyracea Seguenza, 1880. Analysis of both large and small components of the foraminiferal assemblage in all examined samples establishes the dominant character of species Heterostegina papyracea Seguenza (more than 60% of total benthic foraminifera). The assemblage of small benthic foraminifera associated with the Heterostegina is different from that described from Recent sediments, as well as from Miocene sediments of Calabria and Spain. In order to explain this result, two hypotheses can be put forward: (1) the Tortonian Heterostegina from Morocco proliferated abundantly in very shallow environments in association with small foraminifera (i.e. abundant Ammonia beccarii). (2) Because of powerful tidal currents in the South Rifian Corridor, Heterostegina tests were probably transferred after death. This transfer could possibly be due to the narrowness of the South Rifian Corridor in the Early Late Tortonian and its position between the Atlantic and the Mediterranean. Similar cases foraminiferal displacements are also known from modern basins. This study illustrates the difficulties in reconstructing the paleogeography of the studied area and the importance of considering all available components of the assemblage.     

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.     

Deep-sea benthic foraminiferal recolonization of the 1991 Mt. Pinatubo ash layer in the South China Sea

A census count of Rose Bengal stained benthic foraminifera from the surface area on top of a 2 to 6 cm thick ashfall layer at three deep water stations along the western margin of the Philippines exhibits a unique assemblage composition of benthic foraminifera. The total number of benthic foraminifera is low and the ratio of living individuals to empty tests is high. Specific diversity is low, with a significant dominance of infaunal morphotypes including species of the genus Reophax (R. scorpiurus, R. bilocularis and R. dentaliniformis), which are regarded as successful recolonizers. Assemblages below the ash layers are diverse and contain many epifaunal suspension-feeding agglutinated and calcareous foraminifera. The 1991 Mt. Pinatubo eruption caused mass mortality of benthic foraminifera in a vast area of the eastern South China Sea followed by step-wise recolonization of the ash substrate. Three years after the eruption, the benthic foraminiferal community structure is still far from recovery to background levels.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.     

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.