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Significance A substantial component of the global nitrogen cycle is the production of biologically inaccessible dinitrogen attributed to anaerobic denitrification by prokaryotes. Recent evidence identified a eukaryote, foraminifera, as new key players in this “loss” of bioavailable nitrogen. The evolution of denitrification in eukaryotes is a rare event, and the genetic mechanisms of the denitrification pathway in foraminifera are just starting to be elucidated. We present large-scale sequencing analyses of 10 denitrifying foraminiferal species, which reveals the high conservation of the foraminiferal denitrification pathway. We further find evidence for a complementation of denitrification by the foraminiferal microbiome. Together, these findings provide insights into the early evolution of a previously overlooked component in the marine nitrogen cycle. Abstract: Benthic foraminifera are unicellular eukaryotes that inhabit sediments of aquatic environments. Several foraminifera of the order Rotaliida are known to store and use nitrate for denitrification, a unique energy metabolism among eukaryotes. The rotaliid Globobulimina spp. has been shown to encode an incomplete denitrification pathway of bacterial origin. However, the prevalence of denitrification genes in foraminifera remains unknown, and the missing denitrification pathway components are elusive. Analyzing transcriptomes and metagenomes of 10 foraminiferal species from the Peruvian oxygen minimum zone, we show that denitrification genes are highly conserved in foraminifera. We infer the last common ancestor of denitrifying foraminifera, which enables us to predict the ability to denitrify for additional foraminiferal species. Additionally, an examination of the foraminiferal microbiota reveals evidence for a stable interaction with Desulfobacteraceae, which harbor genes that complement the foraminiferal denitrification pathway. Our results provide evidence that foraminiferal denitrification is complemented by the foraminifera-associated microbiome. The interaction of foraminifera with their resident bacteria is at the basis of foraminiferal adaptation to anaerobic environments that manifested in ecological success in oxygen depleted habitats.

Phosphorus is essential for all living organisms, being a component of DNA and RNA and the energy carrier ATP. Phosphogenesis is a main sink of reactive phosphorus in the oceans. The present study reports the presence of intracellular dissolved inorganic phosphate (DIP) in benthic foraminifera from the Peruvian oxygen minimum zone (OMZ). The mean intracellular DIP concentration was 28 ± 3 mM; two to three orders-of-magnitude higher than in the ambient pore waters. The biological implications of the high intracellular phosphate enrichment may be related to the synthesis of polyphosphates or phospholipids for cell-membranes. The comparative genomics analysis of multiple species of foraminifera from different environments reveals that foraminifers encode the genes required for both phospholipid and polyphosphate metabolism. Rapid phosphogenesis and phosphorite formation associated with foraminiferal tests is hypothesized due to the pre-concentration of intracellular phosphate in these organisms. The results indicate that foraminifera may play a key and previously overlooked role in the global phosphorus cycle.

Supratidal sands are vitally important for coastal defence in the German Wadden Sea. They are less affected by human activities than other areas as they are located far off the mainland shore, touristical and commercial activities are generally prohibited. Therefore, supratidal sands are of high ecological interest. Nevertheless, the faunal inventory and distribution pattern of microorganisms on these sands were studied very little. The composition of living and dead foraminiferal assemblages was therefore investigated along a transect from the supratidal sand Japsand up to Hallig Hooge. Both assemblages were dominated by calcareous foraminifera of which Ammonia batava was the most abundant species. Elphidium selseyense and Elphidium williamsoni were also common in the living assemblage, but Elphidium williamsoni was comparably rare in the dead assemblage. The high proportions of Ammonia batava and Elphidium selseyense in the living assemblage arose from the reproduction season that differed between species. While Ammonia batava and Elphidium selseyense just finished their reproductive cycles, Elphidium williamsoni was just about to start. This was also confirmed by the size distribution patterns of the different species. The dead assemblage revealed 20 species that were not found in the living assemblage of which some were reworked from older sediments (e.g., Bucella frigida) and some were transported via tidal currents from other areas in the North Sea (e.g., Jadammina macrescens). The living foraminiferal faunas depicted close linkages between the open North Sea and the mainland. Key species revealing exchange between distant populations were Haynesina germanica, Ammonia batava and different Elphidium species. All these species share an opportunistic behaviour and are able to inhabit a variety of different environments; hence, they well may cope with changing environmental conditions. The benthic foraminiferal association from Japsand revealed that transport mechanisms via tides and currents play a major ecological role and strongly influence the faunal composition at this site.

Highlights • Foraminifera responded faster to environmental perturbations than macroorganisms. • Population densities doubled and Asterigerinata mamilla bloomed under stronger flow. • Excess of threshold values and changes to new ecosystems induce acme occurrences. Abstract Benthic foraminifera are deemed sensitive indicators of environmental conditions. Triggers and magnitudes of faunal response to environmental changes are yet poorly constrained. Benthic foraminiferal faunas were monitored annually at Ria Formosa (Algarve, Portugal) coastal lagoon since 2013. Distinct environmental changes were recognised during the monitoring period. The relocation of a tidal inlet in winter 2015 effected faster flushing, higher tidal levels, and stronger currents in the Esteiro do Ancão tidal channel. The epibenthic foraminiferal species Asterigerinata mamilla increased in abundance and the population densities of the whole fauna were double as high as before inlet relocation. Enhanced sediment redeposition was recorded and extensive polychaete colonies successively replaced firmground patches with oysters. The standing stock of the foraminiferal fauna declined in the next year due to food impoverishment, while the high hydraulic energy levels and high percentages of Asterigerinata mamilla maintained. Benthic foraminifera responded much faster to environmental perturbations than macroorganisms identifying them as powerful proxies in environmental studies.

Highlights • Investigations of the benthic foraminiferal distribution in the Elbe Estuary. • Low diverse assemblages are dominated by Ammonia species. • Low salinities and high-frequency dredging confines foraminiferal proliferation. • Over 40 years, changes in hydrodynamic conditions induced assemblage modifications. Abstract For the past 200 years, estuarine environments experienced intense and rapid environmental degradations due to human interventions. In addition, Global Changes are modifying the estuarine physiography, leading to a re-structuration of marginal marine benthic communities. The aim of this study is to document, the modern assemblage composition and the species-environment relations of benthic foraminifera upstream the Elbe Estuary (southern North Sea) and to observe what has changed since the first survey in 1981. For this purpose, a surface sampling was carried out from 22 stations along the transitional area of the Elbe Estuary. Living (rose-Bengal stained) and dead foraminiferal assemblages were analysed as well as hydrological and sedimentological parameters (such as salinity, pH, grain-size, and organic matter). Living faunas are characterized by very low densities and largely dominated by Ammonia species. Dead assemblages are more diverse and dominated by Ammonia aomoriensis, Haynesina germanica, and Cribroelphidium selseyense. Salinity and grain-size seem to be the major factors influencing foraminiferal distributions in the transitional area. Under the ongoing climate changes, future strategies will be taken to foster the application of benthic foraminifera as biomonitoring tool in the Elbe Estuary, via this baseline investigation.

Heavy metal pollution from anthropogenic sources increasingly influence marine environments and biota because of their toxicity, persistence and bioaccumulation. Especially coastal environments act as natural catchments for anthropogenic pollutants because these areas are highly affected by industry, agriculture and urban sewage runoff. In nearly all natural environments that are harmed by heavy metal pollution, a combination of several pollutants occurs at the same time. In marginal seas and coastal areas, benthic foraminifera are common in meiofaunal associations, and they can be used as biomonitoring tool for changes in environmental parameters like temperature, salinity, or redox conditions. Furthermore, foraminifera take up heavy metals from the seawater and incorporate those into their carbonate shells during calcification. Moreover, foraminifera have a short life cycle and can therefore react immediately to contaminations of the environment. Here, we will present results from culturing studies with Ammonia aomoriensis, Elphidium excavatum and Ammonia batava addressing the relationship of heavy metal concentration in the seawater and in the foraminiferal tests. The partitioning factor between the ambient seawater and the calcium carbonate of the foraminifera is constrained by continuous water monitoring and laser ablation ICP – MS measurements on single chambers grown during the experiment in a manipulated culturing medium. The foraminifera were exposed to a combination of ten different heavy metals over a range of concentrations comparable to current conditions in medium to high pollutedhigh-polluted areas. A correlation between the heavy metal concentration in the culture medium and in the foraminiferal calcite was recognised for several heavy metals (e.g. Cd, Cr, Pb). Once the carbonate/seawater metal partitioning coefficients are constrained with certainty, investigations of the chemistry of benthic foraminiferal shells offer an advanced and reliable method to monitor short-term changes in the concentrations and bioavailability of toxic elements in seawater.

We describe for the first time subtropical intertidal foraminiferal assemblages from beach sands on São Vincente, Cape Verdes. Sixty-five benthic foraminiferal species were recognised, representing 47 genera, 31 families, and 8 superfamilies. Endemic species were not recognised. The new checklist largely extends an earlier record of nine benthic foraminiferal species from fossil carbonate sands on the island. Bolivina striatula, Rosalina vilardeboana and Millettiana milletti dominated the living (rose Bengal stained) fauna, while Elphidium crispum, Amphistegina gibbosa, Quinqueloculina seminulum, Ammonia tepida, Triloculina rotunda and Glabratella patelliformis dominated the dead assemblages. The living fauna lacks species typical for coarse-grained substrates. Instead, there were species that had a planktonic stage in their life cycle. The living fauna therefore received a substantial contribution of floating species and propagules that may have endured a long transport by surface ocean currents. The dead assemblages largely differed from the living fauna and contained redeposited tests deriving from a rhodolith-mollusc carbonate facies at20 m water depth. A comparison of the Recent foraminiferal inventory with other areas identified the Caribbean and Mediterranean as the most likely source regions. They have also been constrained as origin points for littoral to subtidal macroorganisms on other Cape Verdean islands. Micro-and macrofaunal evidences assigned the Cape Verde Current and North Equatorial Current as the main trajectories for faunal immigrations. The contribution from the NW African coast was rather low, a pattern that cannot be explained by the currently available information.

Highlights • Foraminiferal species were assigned to ecological groups according to AMBI procedure. • The benthic foraminifera-based Foram-AMBI was calculated using the AMBI formula. • Foram-AMBI was successfully tested on independent data sets. • Foram-AMBI correlates well with Shannon's diversity and TOC in test data sets. • Foram-AMBI reflects a stress gradient caused by increased organic carbon. The present study follows up the FOraminiferal BIo-MOnitoring (FOBIMO) working group's aim to explore methods which will improve the usefulness of benthic foraminifera in environmental monitoring. An internationally well-established marine biotic index, AMBI, commonly applied to assess ecological quality status was adapted for use on benthic foraminifera. As required by the AMBI formula, species were assigned to one of five ecological groups according to their sensitivity/tolerance to conditions along an increasing stress gradient (here increasing organic matter enrichment). For the assignments, we used 19 published data sets on fully marine NE Atlantic and Arctic fjord, continental shelf, and slope assemblages for which total organic carbon (TOC) data were available. Assignments were based on the relative abundance of the different species along associated TOC gradients. Of the 128 assigned species, the majority was assigned to Groups I–III dominating in low to moderately organic enriched environments with a high to good ecological quality status. Groups IV and V, representing strongly organically enriched environments with a moderate to poor ecological quality status, had 1 and 2 species, respectively. The resulting foraminifera-based Foram-AMBI was calculated using the AMBI formula and tested on four independent foraminiferal data sets from the same geographical region. The validation included correlations of the Foram-AMBI with Shannon's diversity (H′log2) as well as with the organic carbon content in the validation data sets. In two validation data sets from the Norwegian Skagerrak coast, a high proportion of the assemblages consisted of assigned species. The results showed a good correlation between the Foram-AMBI and both the TOC and Shannon's diversity H′log2. In two more southern validation data sets all TOC values were low and the abundance of unassigned species was too high for the Foram-AMBI to provide trustworthy results. The Foram-AMBI of the two validation data sets with high abundance of assigned species clearly reflected an increasing organic carbon-induced stress gradient. Hence, this first attempt to apply the AMBI formula on benthic foraminiferal data shows promising results. However, to improve the applicability of Foram-AMBI, there is a need to assign more species by obtaining data from studies along wide organic carbon pressure gradients, particularly from the southern North Sea and southwards.

Pleistocene and late Pliocene benthic foraminifera from the Manihiki Plateau (southwestern tropical Pacific) were studied at piston-core 34KL. A new benthic foraminiferal biozonation is proposed. It comprises the Nodogenerina sagriensis Partial Range Zone from core base to 566.5 cm and the Fissurina seminiformis Partial Range Zone from this level to core top. The boundary is defined by the last occurence of Nodogenerina sagriensis which is time equivalent to the ‘Stilostomella extinction’ in the Eastern Atlantic. High abundances of Cibicidoides wuellerstorfi indicate a strong influence of near-bottom currents. The absence of high-productivity sensitive species reveals a low flux of organic matter to the sea floor from which a considerable amount is adduced by lateral advection.

Benthic foraminifera have proven to be suitable for environmental monitoring because of their high levels of adaptation, small size and high abundance in Recent sediments and the fossil record. Foraminifera are scarcely used in monitoring studies because a standardization of methods has not been achieved to date. When particular methods were introduced and why they were applied is often hidden in the literature. This paper reviews the development of field and laboratory methods, their constraints and consequences for faunal and data analyses. Multiple and box corers and some grab samplers retrieve reliable surface sediment samples provided the bow wave is minimized as the sampler approaches the sea floor. Most disturbances are created during handling of the unit on deck and subsampling. Ethanol for preservation, rose Bengal as vital stain and a mesh size of 63 µm to wash foraminiferal samples are used extensively. Faunal analyses of a larger size fraction are occasionally necessary. The fractions >125 µm and >150 µm are often preferentially chosen even though this may artificially reduce specimen numbers and faunal diversity. Generally, a much lower level of common practice prevails in sample preparation and faunal analyses than in sampling or laboratory procedures. Increasing preference has been given to quantitative methods and the acquisition of independently revisable census data during recent decades.

Living benthic foraminiferal assemblages were monitored between January and July 2004 in the Kiel Bight, Baltic Sea. Phytoplankton blooms and the deposition of organic detritus were depicted by fluorometer hydro casts and pigment concentrations in surface sediments. Three depositional pulses of organic matter were identified by high phytoplankton concentrations above the sea floor and elevated pigment concentrations in the surface sediment. The foraminiferal assemblage composition remained rather constant but the population density of Elphidium excavatum clavatum showed a two- to six-fold increase within a few days after organic detritus deposition. The foraminiferal assemblage composition was compared to earlier studies in this area. Elphidium excavatum, Ammotium cassis and E. incertum dominated the living fauna in the 1960s and 1970s. The recent survey revealed a predominance of E. excavatum subspecies (more than 90% of the living assemblage). The average population densities were six times higher than in the 1970s. Data structure and model calculations suggested that E. excavatum clavatum is able to reproduce rapidly, with high offspring numbers at elevated food supply. The diminution of A. cassis has occurred in Kiel Bight during the last ten years and was most likely induced by a period of low deep-water salinities in the early 1990s.

Planktonic and benthic foraminifera are the most significant providers of information on the state of surface and deep oceans in the past. Many foraminiferal proxies rely on the knowledge of ecological preferences of individual species and the assumption that these remained similar through time. Consequently, the applicability of such proxies is limited in time by the extent of the modern fauna. By analysing extensive datasets of species occurrences, we show that the modern oceanic foraminifer fauna originated during the Neogene. This occurred during two distinct diversification pulses: one in the Middle Miocene (17–14 Ma) and the second at the Miocene/Pliocene transition (7–4 Ma). The first diversification coincides with the time of a major change in the frequency of the dominant climate cycles during the Miocene Climatic Optimum. The environmental driver of the second diversification could be related to an increased provincialism induced by the closure of the Panama Seaway, but the exact link is not clear, particularly for the plankton. Surprisingly, major changes of ocean circulation due to the growth of Antarctic ice-sheet and closure of low-latitude seaways appear to have caused mainly extinctions. Given the age of the latest diversification and extinction pulses that shaped the modern foraminiferal fauna, we conclude that calibrated proxies based on assemblage properties should not be interpreted quantitatively in sediments older than the late Pliocene. Much of what we know about the Cenozoic oceans comes from the study of fossil foraminifera. These microscopic protozoans occur in nearly all marine environments – from the bottom of the deepest trenches (Todo et al. 2005) to brine channels in Antarctic sea-ice (Dieckmann et al. 1991) – and their ornate shells are readily preserved in sediments. Foraminiferal species are sensitive to a range of environmental parameters and the chemical and isotopic composition of their calcite skeleton records the properties of ambient seawater and the nature of metabolic and kinetic processes that take place during calcification. This remarkable capacity to reflect the state of their habitat, coupled with the abundance of their fossils in marine cores, made foraminifera the main tool for reconstructions of past oceans and climate. Benthic foraminifera convey information about conditions prevailing on the ocean floor. Early research on benthic foraminifera was qualitative, aimed at assessing the faunal inventory, and obtaining a general understanding of past environmental conditions. Recent studies focused on the development of quantitative tools, in particular for the reconstruction of organic matter flux to the sea floor (Herguera & Berger 1991; de Rijk et al. 2000), oxygen content of near-bottom waters (Kaiho 1994), and intensity of bottom currents (Schonfeld 2002). Planktonic foraminiferal assemblages have long been used to estimate sea-surface temperature (Imbrie & Kipp 1971; Kucera et al. 2005), upwelling intensity (Thiede 1975; Conan et al. 2002) and productivity (Ivanova et al. 2003), and the modification of their assemblages on the sea floor has been used to reconstruct past bottom-water carbonate ion concentration (Conan et al. 2002). The elemental and isotopic chemistry of calcareous tests from both benthic and planktonic foraminifera forms the basis of an ever-increasing battery of proxies used to reveal the age, origin, chemical and physical properties of ocean waters (Fischer & Wefer 1999; Henderson 2002). The application of foraminifera as palaeoceanographic proxies is based on the recognition of ecological relationships in the modern ocean and their translation into the fossil realm. This method relies on a number of assumptions (e.g. Birks 1995), the most pertinent one being the stationarity through time of the individual ecological relationships. Some foraminiferal proxies are based on determination of broad functional and ecological types (benthic vs. planktonic) or preservational state (fragmentation) and their temporal applicability is therefore less of an issue. However, most quantitative proxies rely on species-specific calibrations and the interpretation of almost all chemical From: WILLIAMS, M., HAYWOOD, A. M., GREGORY, F. J. & SCHMIDT, D. N. (eds) Deep-Time Perspectives on Climate Change: Marrying the Signal from Computer Models and Biological Proxies. The Micropalaeontological Society, Special Publications. The Geological Society, London, 409–425. 1747-602X/07/$15.00 # The Micropalaeontological Society 2007. signals in foraminifera requires knowledge of the habitat and phenology of the analysed species. Because of circular evidence, the stability of the ecological preferences of taxa through time is often difficult to assess. Therefore, morphological similarity has been used as a first approximation of ecology and the fossil range of a given species has been taken to represent the duration of its specific, present-day ecological behaviour. This approximation is inevitably crude, as it is known that species’ ecology can change with little or no shift in morphology (Norris et al. 1996; Kucera & Kennett 2002), but it provides a consistent and objective means to estimate the maximum range of modern environmental calibrations. The purpose of this study is to review data on the age of extant species of planktonic and benthic foraminifera, determine the diversification history of the groups and assess its links with oceanographic events in the Cenozoic. A further aim is to produce objective guidelines for the use of foraminiferal proxies based on modern environmental calibrations in ‘deep time’. This review will only deal with oceanic foraminifera; shelf seas, brackish facies, marshes and other marginal environments will not be considered. Origin of the modern foraminiferal fauna The number of living benthic foraminiferal species is estimated at approximately 4000 (Thies 1991), and about 45 living planktonic species are commonly recognized (Hemleben et al. 1989). The number of all benthic and planktonic species since the Cambrian is estimated at about 10,000 (Vickermann 1992). This estimate would suggest that foraminiferal faunas are dominated by conservative, long-ranging taxa and that the modern fauna should have its roots in deep time. A direct assessment of the antiquity of extant foraminiferal species would require a thorough analysis of species occurrences on a global scale. However, information on the ranges of living benthic foraminifera is rather scattered. Biostratigraphic compendia typically cover a limited number of stages or a subsystem, have a regional bias and concentrate on stratigraphically important taxa. Only a few studies in the framework of the Deep Sea Drilling Project and Ocean Drilling Program go beyond that scope (see Berggren & Miller 1989; Thomas 1992; and Hayward 2001 for further discussion). Jones (1994) gives a list of stratigraphic ranges for 305 recent, mostly deep-sea benthic foraminifera and 26 planktonic species figured by Brady (1884). Less than 10% of the presumed living benthic species and only a half of the living planktonic foraminifera are covered by this compilation. Nonetheless, the dataset is methodologically and taxonomically consistent and provides the first approximation of the distribution of ages of modern oceanic foraminifera (Fig. 1). As many as 13 (4.2%) of the living benthic species listed by Jones (1994) have originated in the Cretaceous; the oldest was reported as being of Valanginian age. The number of survivors from the Palaeogene is with 58 (19.0%) still low. Clearly, the Miocene was the crucial time for the diversification of the modern benthic fauna. More than 50% of the recent benthic species included in Jones’ (1994) compilation have originated during this stage. Remarkably, almost a quarter of the modern species appear to have originated during the last 5 Ma in the Pliocene or Pleistocene. 0 20 40 60 80 100 120 140 160 180 Pleistocene Pliocene Miocene Oligocene Eocene Palaeocene Cretaceous O rig in s of r ec en t f or am in ife ra Planktonic species

An important tool for deep-sea temperature reconstruction is Mg/Ca paleothermometry applied to benthic foraminifera. Foraminifera of the genus Melonis appear to be promising candidates for temperature reconstructions due to their wide geographical and bathymetric distribution, and their infaunal habitat, which was suggested to reduce secondary effects from carbonate ion saturation (Δ[CO3 2−]). Here, we make substantial advances to previous calibration efforts and present new multi-lab Mg/Ca data for Melonis barleeanum and Melonis pompilioides from more than one hundred core top samples spanning in situ bottom temperatures from −1 to 16 °C, coupled with morphometric analyses of the foraminifer tests. Both species and their morphotypes seem to have a similar response of Mg/Ca to growth temperature. Compilation of new and previously published data reveals a linear dependence of temperature on Mg/Ca, with a best fit of Mg/Ca (mmol/mol) = 0.113 ± 0.005 ∗ BWT (°C) + 0.792 ± 0.036 (r2 = 0.81; n = 120; 1σ SD). Salinity, bottom water Δ[CO3 2−], and varying morphotypes have no apparent effect on the Mg/Ca-temperature relationship, but pore water Δ[CO3 2−] might have had an influence on some of the samples from the tropical Atlantic.

The faunal composition and diversity pattern of Recent benthic foraminifera from the shelf and slope of the South Western Approaches (Celtic Sea) were assessed. The sampling stations cover a depth range from 100 to 500 m. A total number of 294 species was recorded, of which 89 were found exclusively in the living fauna and 118 only in the dead assemblage, whereas 87 species were found in both assemblages. The faunal composition revealed a distinct bisection of the living fauna on the shelf. While certain distribution patterns of living dominant species were recognized along a NE-SW trending transect towards the shelf edge, the living fauna changed within small depth intervals and geographic position on the slope. Causes for these structured slope assemblages were probably along-slope currents of varying strengths, as well as variations in topography and bottom sediments. Analyses of population densities and diversity patterns determined high densities along the shelf edge and at one slope station, as well as an increasing diversity with water depth. A comparison with literature data from the same area yields distinct differences in faunal composition on the shelf and slope. The diversity was similar on the shelf, but higher at corresponding stations on the slope. We conclude that the faunal composition was highly influenced by the time and methods of sampling. In order to obtain consistent results in regional studies, all samples should be taken in a short time interval and by using the same sampling device. In addition, we depicted the influences of environmental parameters on dead faunal assemblages and their consequences for paleoenvironmental interpretations of fossil foraminiferal assemblages.

The present study investigated the combined effects of ocean acidification, temperature, and salinity on growth and test degradation of Ammonia aomoriensis. This species is one of the dominant benthic foraminifera in near-coastal habitats of the southwestern Baltic Sea that can be particularly sensitive to changes in seawater carbonate chemistry. To assess potential responses to ocean acidification and climate change, we performed a fully crossed experiment involving three temperatures (8, 13, and 18uC), three salinities (15, 20, and 25) and four pCO2 levels (566, 1195, 2108, and 3843 matm) for six weeks. Our results highlight a sensitive response of A. aomoriensis to undersaturated seawater with respect to calcite. The specimens continued to grow and increase their test diameter in treatments with pCO2 ,1200 matm, when Vcalc .1. Growth rates declined when pCO2 exceeded 1200 matm (Vcalc ,1). A significant reduction in test diameter and number of tests due to dissolution was observed below a critical Vcalc of 0.5. Elevated temperature (18uC) led to increased Vcalc, larger test diameter, and lower test degradation. Maximal growth was observed at 18uC. No significant relationship was observed between salinity and test growth. Lowered and undersaturated Vcalc, which results from increasing pCO2 in bottom waters, may cause a significant future decline of the population density of A. aomoriensis in its natural environment. At the same time, this effect might be partially compensated by temperature rise due to global warming.

Past river run-off is an important measure for the continental hydrological cycle and the assessment of freshwater input into the ocean. However, paleosalinity reconstructions applying different proxies in parallel often show offsets between the respective methods. Here, we compare the established foraminiferal Ba/Ca and View the MathML source salinity proxies for their capability to record the highly seasonal Orinoco freshwater plume in the eastern Caribbean. For this purpose we obtained a data set comprising Ba/Ca and View the MathML source determined on multiple species of planktonic foraminifera from core tops distributed around the Orinoco River mouth. Our findings indicate that interpretations based on either proxy could lead to different conclusions. In particular, Ba/Ca and View the MathML source diverge in their spatial distribution due to different governing factors. Apparently, the Orinoco freshwater plume is best tracked by Ba/Ca ratios of G. ruber (pink and sensu lato morphotypes), while View the MathML source based on the same species is more related to the local precipitation-evaporation balance overprinting the riverine freshwater contribution. Other shallow dwelling species (G. sacculifer, O. universa) show a muted response to the freshwater discharge, most likely due to their ecological and habitat preferences. Extremely high Ba/Ca ratios recorded by G. ruber are attributed to Ba2+-desorption from suspended matter derived from the Orinoco. Samples taken most proximal to the freshwater source do not show pronounced Ba/Ca or View the MathML source anomalies. Here, the suspension loaded freshwater lid developing during maximum discharge suppresses foraminiferal populations. Both proxies are therefore biased towards dry season conditions at these sites, when surface salinity is only minimally reduced.

The aim of the present study was to compare preservation, staining and preparation techniques to assess the influence of different sample treatments and analyses on the accuracy of benthic foraminiferal assemblage data from NE Atlantic shelf seas. Replicate surface samples from the SE North Sea were preserved with ethanol–rose Bengal or formalin, some were stained after processing, or foraminifera were concentrated by flotation. Coloration of living specimens was different between samples treated with an ethanol–rose Bengal solution and those stained after washing. In the latter case, only the last two or three chambers were stained. The aliquot sample preserved with formalin showed dissolution features in agglutinated and porcellaneous species. Population density varied between different preservation, picking modes and investigators. The accuracy of picking was in the range of ±2 % (1σ), while external reproducibility ranged from −34 to +16 %. There was no significant difference between wet and dry picking. Samples that were concentrated by flotation generally yielded a lower number of specimens. Agglutinated species were under-represented in samples that were stained after washing and in the flotation concentrate. Size fractions showed a reduction of population density and Fisher alpha diversity index with increasing mesh size. Only half of the specimens and less than two-thirds of the species are captured if the >125 µm rather than >63 µm fraction is analysed. In oxygen minimum zones, where small-sized species dominate the assemblage, the recovery in larger size fractions could be lower.

It is expected that the calcification of foraminifera will be negatively affected by the ongoing acidification of the oceans. Compared to the open oceans, these organisms are subjected to much more adverse carbonate system conditions in coastal and estuarine environments such as the southwestern Baltic Sea, where benthic foraminifera are abundant. This study documents the seasonal changes of carbonate chemistry and the ensuing response of the foraminiferal community with bi-monthly resolution in Flensburg Fjord. In comparison to the surface pCO2, which is close to equilibrium with the atmosphere, we observed large seasonal fluctuations of pCO2 in the bottom and sediment pore waters. The sediment pore water pCO2 was constantly high during the entire year ranging from 1244 to 3324 μatm. Nevertheless, in contrast to the bottom water, sediment pore water was slightly supersaturated with respect to calcite as a consequence of higher alkalinity (AT) for most of the year. Foraminiferal assemblages were dominated by two calcareous species, Ammonia aomoriensis and Elphidium incertum, and the agglutinated Ammotium cassis. The one-year cycle was characterised by seasonal community shifts. Our results revealed that there is no dynamic response of foraminiferal population density and diversity to elevated sediment pore water pCO2. Surprisingly, the fluctuations of sediment pore water undersaturation (Ωcalc) co-vary with the population densities of living Ammonia aomoriensis. Further, we observed that most of the tests of living calcifying foraminifera were intact. Only Ammonia aomorienis showed dissolution and recalcification structures on the tests, especially at undersaturated conditions. Therefore, the benthic community is subjected to high pCO2 and tolerates elevated levels as long as sediment pore water remains supersaturated. Model calculations inferred that increasing atmospheric CO2 concentrations will finally lead to a perennial undersaturation in sediment pore waters. Whereas benthic foraminifera indeed may cope with a high sediment pore water pCO2, the steady undersaturation of sediment pore waters would likely cause a significant higher mortality of the dominating Ammonia aomoriensis. This shift may eventually lead to changes in the benthic foraminiferal communities in Flensburg Fjord, as well as in other regions experiencing naturally undersaturated Ωcalc levels.

A high-resolution record, covering 9.3–0.2 ka BP, from the sub-arctic Stjernsund (70°N) was studied for benthic foraminiferal faunas and stable isotopes, revealing three informally named main phases during the Holocene. The Early- to Mid-Holocene (9.3–5.0 ka BP) was characterized by the strong influence of the North Atlantic Current (NAC), which prevented the reflection of the Holocene Climatic Optimum (HCO) in the bottom-water temperature. During the Mid-Holocene Transition (5.0–2.5 ka BP), a turnover of benthic foraminiferal faunas occurred, Atlantic Water species decreased while Arctic-Polar species increased, and the oxygen isotope record showed larger fluctuations. Those variations correspond to a period of global climate change, to spatially more heterogeneous benthic foraminiferal faunas in the Nordic Seas region, and to regionally diverging terrestrial temperatures. The Cool Late Holocene (2.5–0.2 ka BP) was characterized by increased abundances of Arctic-Polar species and a steady cooling trend reflected in the oxygen isotopes. In this period, our record differs considerably from those on the SW Barents Sea shelf and locations farther south. Therefore, we argue that regional atmospheric cooling triggered the late Holocene cooling trend. Several cold episodes centred at ∼8.3, ∼7.8, ∼6.5, ∼4.9, ∼3.9 and ∼3.3 ka BP were identified from the benthic foraminiferal faunas and the δ18O record, which correlated with marine and atmospherically driven proxy records. This suggests that short-term cold events may result from reduced heat advection via the NAC or from colder air temperatures.

Palaeoceanography relies on the assumption that parameters measureable in sediment cores correlate reliably to hydrographic parameters. Areas of dynamic watermass mixing, which generally have steep hydrographic gradients, therefore provide both a tempting target (large spatial and temporal differences in temperature and salinity) and a significant challenge (high flow velocities and high “noise”) for palaeoceanographers. Here, we investigate the ability of parameters measured on core tops to be used as the basis for simulating regional hydrography within one globally important area of watermass mixing, the Gulf of Cadiz. Using grainsize, sediment composition, benthic foraminiferal assemblage and benthic foraminiferal stable isotope analysis it is possible to build an effective framework for qualitative constraint of the position of the modern Mediterranean Outflow Water (MW) Plume. We propose that the Gulf of Cadiz slope can be sub-divided into 8 hydrographic zones reflecting these parameters; Proximal MW, Core MW1 (steep slopes), Core MW1 (gentle slopes), Distal MW1, Distal MW2, Lower Limit of MW plume, North Atlantic Deep Water and Ambient Atlantic Water. We anticipate that compilation of time-slice data will reveal hydrographic sub-divisions of the slope for the past in a similar manner, improving our understanding of past changes in the size and position of the MW plume, but more work needs to be done before a secure protocol for quantitative reconstruction can be created. Stable isotope analysis alone is insufficient for the task, with δ13C behaving in a non-conservative manner and δ18Ocalcite acting in an ambiguous way due to the competing influences of δ18Owater and temperature. It is likely that if a single parameter that unambiguously determines the difference between Mediterranean Water and Atlantic Water can be identified, combination with δ18Ocalcite will allow extension of our qualitative analysis into a quantitative means of palaeohydrographic reconstruction for this region. Similar complexity to that found in this study would be expected in any mixing region, and in particular on the majority of sediment drifts. We recommend that core top surveys similar to this study are performed on these sediment drift systems before any form of quantitative palaeohydrology is attempted.

Benthic foraminifera are unicellular eukaryotes inhabiting sediments of aquatic environments. Several species were shown to store and use nitrate for complete denitrification, a unique energy metabolism among eukaryotes. The population of benthic foraminifera reaches high densities in oxygen-depleted marine habitats, where they play a key role in the marine nitrogen cycle. However, the mechanisms of denitrification in foraminifera are still unknown, and the possibility of a contribution of associated bacteria is debated. Here, we present evidence for a novel eukaryotic denitrification pathway that is encoded in foraminiferal genomes. Large-scale genome and transcriptomes analyses reveal the presence of a denitrification pathway in foraminifera species of the genus Globobulimina. This includes the enzymes nitrite reductase (NirK) and nitric oxide reductase (Nor) as well as a wide range of nitrate transporters (Nrt). A phylogenetic reconstruction of the enzymes' evolutionary history uncovers evidence for an ancient acquisition of the foraminiferal denitrification pathway from prokaryotes. We propose a model for denitrification in foraminifera, where a common electron transport chain is used for anaerobic and aerobic respiration. The evolution of hybrid respiration in foraminifera likely contributed to their ecological success, which is well documented in palaeontological records since the Cambrian period.

Ammonia and Elphidium collected in the Kiel Fjord for the present study were first identified on morphological bases as Ammonia beccarii (Linne´, 1758) and Elphidium excavatum (Terquem, 1876). Phylogenetic analyses based on partial SSU rDNA and LSU rDNA sequences show that Ammonia specimens sampled in the Kiel Fjord belong to the phylotype T6, which has a disjunct distribution (Wadden and Baltic Seas/China and Japan) and has been identified as Ammonia aomoriensis (Asano, 1951). Partial SSU rDNA sequence analyses indicate that Elphidium specimens from the Kiel Fjord belong to the clade E. excavatum, confirming the morphological identification. This clade can be further divided in three subclades. Kiel Fjord Elphidium belong to two of these subclades and were identified morphologically as the subspecies E. excavatum excavatum (Terquem, 1876) and E. e. clavatum Cushman, 1930.

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.

Living benthic foraminifera of Flensburg Fjord were surveyed in June 2006. The muddy and organic-rich sediments of the inner fjord were dominated by Elphidium incertum. E. incertum and E. excavatum were frequent in muds and sandy muds of the fjord loop around Holnis Peninsula and in the outer part. Gelting Bay yielded a different biofacies, indicating a brackish and sandy habitat, poor in food supply and with microfauna dominated by Ammonia beccarii and E. albiumbilicatum. The central fjord and nearshore zones of the loop were characterized by sandy muds, relatively poor in food and occupied by A. beccarii, E. incertum and E. excavatum subspecies. High abundances of E. excavatum were encountered in the innermost fjord, with fine-grained and organic-rich muddy sediments.A comparison with previous studies revealed the profound changes in species composition in the outer Flensburg Fjord since the 1970s. A decline in numbers of Ammotium cassis and flourishing of Ammonia beccarii in Gelting Bay were recognized. These changes are most likely associated with decreased intensity and frequency of salt-water inflows into the Baltic Sea since the 1960s. It is inferred that the decline of A. cassis is similar to that of Eggerelloides scaber, which currently is found only in depressions of Kiel Bight with higher salinity.

The diatom spring bloom and response of the benthic foraminifera was monitored between January and July 2004 in the Kiel Bight, western Baltic Sea. Phytoplankton production, the deposition of organic detritus, and feeding of Elphidium excavatum clavatum and Reophax dentaliniformis were monitored by pigment concentrations and pigment composition in water samples, surface sediments, and foraminiferal protoplasm. The population dynamics of the dominant species, Elphidium e. clavatum, was assessed by using their size distributions. The main spring bloom was triggered by a sudden rise in the three-day average insolation to above 170 W m(-2) d(-1) and took place during late February. Chlorophyll a concentrations in the protoplasm of Elphidium e. clavatum followed those in the surface water. The chlorophyll a / fucoxanthin ratio was the same as in bacillariophycean diatoms suggesting a "bloom-feeding" behaviour of Elphidium e. clavatum. Alternatively, the pigment composition in Reophax dentaliniformis only occasionally mirrored that of diatoms, indicating prochlorophycean and cyanophycean algae as this species' preferred food source. The size distributions and the abundances of living Elphidium e. clavatum revealed two reproduction events, which were 17 days apart. Both events coincided with an increase in sedimentary chlorophyll a concentrations. The reproduction was preceded by growth and feeding of adult specimens and it was followed by distinct growth phases of young specimens, which lasted a few days only. One reproductive cycle covered the spring bloom and produced many more offspring than the subsequent cycle. The light carbon isotopic signal from diatoms was only transiently recorded in foraminiferal calcite and with a low offset during the spring bloom. The oxygen isotopic composition of foraminiferal calcite mirrors seasonal delta(18)O-seawater-salinity relationships rather than temperature.

The Adra River deltaic system, southeast Iberian Peninsula, shows a steep topography and is subjected to strong climatic seasonality. This system has been affected by alternating wet and dry periods, and it has also undergone numerous anthropogenic activities such as deforestation, mining activities, river channel deviations and dam construction, particularly during the last two centuries. Two sediment cores were retrieved off the Adra River, from the western (MS_V9) and eastern (MS_V4) lobes of the subaqueous deltaic deposit. A multi-proxy study was carried out, including grain size, benthic foraminiferal assemblages, magnetic susceptibility and geochemical element analyses, in order to understand the sedimentary expression of recent climatic cycles and anthropogenic interventions in the river basin. Periods of increased deposition of coarse-grained sediments, low absolute abundance of benthic foraminifera and high elemental ratios indicative of terrigenous contributions, were interpreted as periods of increased sediment supply to the shelf. Four flooding events were recorded in core MS_V9 and three events on core MS_V4, of which two were observed in both cores. They were related to periods with major floods that were documented on the southern Iberian Peninsula around 1770e1810 and 1860 e1870 AD. On the other hand, sediment core intervals exhibiting increasing proportion of fine-grained sediments and higher abundances of foraminiferal species assigned as successful colonizers (Textularia earlandi) and opportunistic species that feed on bacteria or terrestrial organic matter (Bolivina ordinaria, Bulimina elongata, Eggerelloides scaber and Ammonia beccarii or tepida), indicate the establishment of new environments with new ecological constraints. They were related to significant decreases of terrigenous sediment input during low rainfall or dry periods. The increase of opportunistic species feeding on fresh phytodetritus (Nonionella iridea, Nonionella stella, Nonionella sp., Brizalina dilatata, Epistominella vitrea and Bolivinellina pseudopuntata), under more stable environmental conditions, also point to a stronger marine influence on the prodeltaic environments during these periods. The human interventions on the river basin after 1872 AD, with the deviation of the main river channel to the east, led to a drastic reduction of the sediment exported to the western delta lobe. This study showed that the sedimentation on the Adra subaqueous deltaic deposit was mainly controlled by rainfall variability from 1663 to 1872 AD, and afterwards by anthropogenic interventions., SI

Phases with massive abundances of Uvigerina species recorded in North Atlantic deep-sea cores were previously attributed to water mass properties or to extended high productivity during the last Glacial. We investigated the Uvigerina distribution in 53 sediment cores and 555 surface samples from the north-eastern Atlantic ranging from 308 to 708N and 1 to 4410 m water depth. Our results showed that Uvigerina pigmea dominated in late Pleistocene assemblages whereas Uvigerina peregrina and Uvigerina peregrina parva were common during the Holocene. Uvigerina pigmea lives today in the Gulf of Guinea and South Atlantic only (98N to 178S), but was present up to 548N during the Last Glacial Maximum. The species retreated north- and southwards during Termination I and disappeared in the north-eastern Atlantic off northern Portugal 6100 years ago. Uvigerina peregrina and U. peregrina parva spread out from isolated occurrences along the continental slope off western Iberia at the onset of Termination I. Uvigerina peregrina exhibited Recent abundance maxima at Goban Spur, Plateau des Landes, and the Iberian Margin. These areas are subjected to strong lateral advection by downslope transport and sediment focusing. The actual flux rate of particulate organic matter is locally raised by a factor of 4 to 5. Comparable flux rates were otherwise recorded at U. peregrina maxima under the NW-African coastal upwelling regime. The Recent distribution of U. pigmea and evidence from sediment cores show an organic carbon flux of 3.5 g C m-2 yr-1 as minimum for a sustainable population of this species. Alkenone, pigment and planktonic foraminiferal records from cores MD952039 and MD952040 revealed that a spring bloom dominated productivity dynamics and a balanced food particle composition of haptophyteae algae and diatoms were limiting environmental factors for U. pigmea during the Pleistocene. A widespread change from glacial to modern productivity characteristics during the last Deglaciation may have triggered the early Holocene disappearance of U. pigmea from the north-eastern Atlantic.

Living and dead benthic foraminiferal assemblages from two size fractions, 63–250 μm and >250 μm, were studied at 128 stations from the shelves down to abyssal plains of the Gulf of Guinea in the eastern equatorial Atlantic. The sample size ranged between 300 and 400 cm2 of the sediment/water interface, with a penetration depth of 1 cm. Environmental parameters were quantified for sediment composition, food webs, bottom-water oxygen concentrations, and benthic oxygen respiration. Seven major station groupings (named G1–G7) are subdivided by factor analysis. The three most abundant species therein are discussed with respect to their ecological requirements, for both size classes and for living and dead counts. For 49 living species, the range of environmental gradients is established, and for five species live observations are reported. Five station groupings can be attributed toward stepwise ranges scaled by bathymetry and flux rates of organic carbon (G4, 27–75 m; G5, 68–269 m; G6, 250–740 m; G2, 674–2007 m; G1, 1475–4970 m). Two geographically restricted station groupings locally interfinger with other groupings in the upper bathyal and abyssal water depths (G7, 82–451 m; G3, 1002–4658 m). G7 shows lowered oxygen concentrations and benthic oxygen respirations, whereas hemipelagic sediments are typical of G3. The interfingering of G7 and G3, however, is not purely a result of the increasing number of species most perfectly adapted to these environments but rather that these groupings are structured by the decrease of species with lower tolerance towards environmental perturbation. In fact, the more common species have an environmental range much broader than the environmental frame of the groupings they dominate. This observation holds for all size classes and for live and dead counts. We conclude that species-specific preferences and thresholds provide a more valuable and more consistent tool in environmental research than the environmental ranges observed for assemblages.

Recent benthic foraminiferal assemblages were analyzed in the Gulf of Cadiz, northeastern Atlantic, to study the impact of the Mediterranean Outflow Water (MOW) undercurrent on the benthic environment. Foraminiferal counts and the analysis of specimens attached to hard substrates from 26 surface samples reveal a relationship of epibenthic assemblages with sedimentary and hydrodynamic environment. Epibenthic species make up as much as 60% of the living assemblage at proximal sites with high current velocities and 3–18% in distal areas or near the margins of the MOW flow paths at low velocities. These foraminifers inhabit elevated substrates only within the MOW, which evidently provides an ecological niche for opportunistic suspension feeders. They adapt their settling elevation dynamically and occur at greater heights above the ambient sediment surface under stronger currents. Mobility, fixation strength, suspension feeding, and reproduction efficiency emerge as individual capabilities promoting the occupation of elevated substrates by certain epibenthic species. The active microhabitat selection is pursued as basic strategy of these foraminifers to optimize their food acquisition. A better access to food sources stimulates reproduction and leads to a greater contribution of foraminiferal tests to the surface sediments. Elevated epibenthos percentages from the dead assemblage and current velocities prevailing at the sampling sites are closely correlated. A compilation including other data from southern Portugal, Florida Straits, and the English Channel infers an exponential relationship between epibenthic abundances and flow strength implying that endobenthic species prevail even under high current velocities. A linear model provides a significantly better fit for the Gulf of Cadiz data however. This relation is used for a case study in order to estimate near-bottom current strengths for the late Holocene Peak III contourite in core M39008-3. Trends and absolute values of current velocities, inferred from the benthic foraminiferal proxy, are the same scale as estimates obtained from sediment grain-size distribution and hydrodynamic models. Epibenthic foraminifera thus bear a high potential as proxy for palaeocurrent studies that even may overcome objections by predetermined grain-size distributions in deep-sea cores.

The benthic environment in the Gulf of Cadiz, north-eastern Atlantic, is strongly affected by the Mediterranean outflow water undercurrent (MOW) which flows northwards along the western Iberian Margin at 500–1500 m water depth. Foraminiferal census counts of living and dead assemblages from 27 surface samples ranging from 103 to 1917 m water depth, and the examination of hard substrates reveal a close correlation of the fauna with the local hydrography and sediment facies. Four different faunal groups are separated by factor analysis of the living fauna. Assemblage 1 contains typical lower slope species and dominates samples from the lower MOW core layer and in the North Atlantic deep water below. Shelf edge foraminifera are common in assemblage 2a which shows the highest proportions in samples from 103 to 272 m. Assemblage 2b is dominated by upper slope species and suspension-feeders that are frequent in the upper MOW core layer and in distal settings between 396 and 901 m. Species from assemblage 3 prefer epibenthic habitats and are recorded with high proportions exclusively in the immediate flow paths of the upper MOW between 496 and 881 m. Colonisation structures and species composition of epibenthic assemblages from the proximal facies largely differ from those in distal settings. In general, epibenthic foraminifers only use elevated substrates under the influence of near-bottom flow. Under high current velocities, epibenthic foraminifers prefer large and heavy objects. They colonise high attachment levels where a maximum yield of advected food particles can be achieved. In distal settings at lower flow velocities, the elevation height does not exceed 20 mm above the surrounding sediment surface. This level is related to a hydrologic transition layer with high concentrations of suspended particles. The comparison of microhabitat preferences and faunal structure under high and low current velocities reveal that substrate stability may be a confining environmental variable for endobenthic and shallow epibenthic foraminifers. The observations also indicate that the preferential settling height of epibenthic foraminifera is related to the highest lateral flux rates of food particles within reach from the sea floor. A dynamic selection of elevated microhabitats is only used by 7.8% of all species recognised in the Gulf of Cadiz area.

Calcifying foraminifera are expected to be endangered by ocean acidification; however, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 6 months of incubation, foraminiferal assemblages were kept and treated in natural sediment with pCO2-enriched seawater of 430, 907, 1865 and 3247 μatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months of incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Ωcalc was much higher in the sediment than in the water column in nearly all pCO2 treatments and remained close to saturation. As a result, the life cycle (population density, growth and reproduction) of living assemblages varied markedly during the experimental period, but was largely unaffected by the pCO2 treatments applied. According to the size–frequency distribution, we conclude that foraminifera start reproduction at a diameter of 250 μm. Mortality of living Ammonia aomoriensis was unaffected, whereas size of large and dead tests decreased with elevated pCO2 from 285 μm (pCO2 from 430 to 1865 μatm) to 258 μm (pCO2 3247 μatm). The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of CaCO3 weight. Living individuals had a calcium carbonate production rate of 0.47 g m−2 a−1, whereas dead empty tests accumulated a rate of 0.27 g m−2 a−1. Although Ωcalc was close to 1, approximately 30% of the empty tests of Ammonia aomoriensis showed dissolution features at high pCO2 of 3247 μatm during the last 2 months of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. Our results emphasize that the sensitivity to ocean acidification of the endobenthic foraminifera Ammonia aomoriensis in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

The depth distribution of living (Rose Bengal-stained) benthic foraminifera in near-surface sediments was analyzed along 4 short cores from the western Iberian Margin and Gulf of Cadiz at water depths from 800 to 1920 m. Pore-water oxygen, Chloroplastic Pigment Equivalent (CPE), and total organic carbon (C org ) concentrations were measured in the same or adjacent cores. These values were used to constrain the limiting oxygen and nutrient levels that defined the ranges in sediment depth and abundance maxima of foraminiferal species. The population density showed a good correlation with pore-water oxygenation indicating that oxygen is a limiting factor for endobenthic foraminiferal assemblages at mid-depths between 800 and 1200 m. At the deep-water site, however, the fauna appeared nutrient limited. More than half of the benthic species occupied extended sediment depth habitats and a wide range of oxygen levels among the sites studied. Thirty-nine of 103 species occurred in two or more cores and showed distinct abundance maxima, 36 in the uppermost 2.0 cm where the CPE concentrations were high. Living foraminifers were not found at depths where the CPE-values were lower than 0.1 μg/g. This level indicates trophic conditions critical for the survival of endobenthic species. The frequency distribution of the lower oxygen range limits of all species showed maxima at the high oxic/low oxic (3 ml/l), low oxic/suboxic (1.5 ml/l), and suboxic/dysoxic (0.3 ml/l) boundaries and thus depicted the ecological significance of these previously recognized ecolimits. Only 30 % of the species display consistent patterns in relation to pore water oxygen. Dysoxic indicators were Globobulimina affinis, Chilostomella ovoidea and Bathysiphon capillare. Many species previously designated as suboxic indicators were ubiquitous or preferred microhabitats at oxic levels. Boundary conditions for the dysoxic indicator G. affinis were pelagic flux rates higher than 3.5 g C m −2 yr −1 , a habitat depth below the homogeneously bioturbated layer in areas outside of high productivity zones, and CPE concentrations of 0.1 to 1.0, preferentially 0.7 to 0.8 μg/g. The distribution pattern suggested that G. affinis thrives mainly on dysoxic bacteria and actively seeks this nutrient resource.

Records of benthic foraminiferal assemblage variations and benthic δ13C along 12 sediment cores from the western Iberian Margin, between 36° and 42°N at water depths from 820 to 3580 m, are used to monitor fluctuations of the Mediterranean Outflow Water (MOW) during the past 30 ka. The chronostratigraphy of the cores is based on planktonic δ18O records, 14C AMS-dating, and the recognition of Heinrich Events H1 through H4. Increased abundances of suspension feeding benthic foraminifers, denoted as ’Epibenthos Group‘, closely match areas where the recent MOW core layers impinge on the continental slope at 800 and 1300 m water depth, and near-bottom current velocities are enhanced. Elevated ‘Epibenthos Group’ abundances, increased benthic δ13C, and sedimentological evidence for winnowing and erosion are found in glacial sections up to the earliest Termination I in cores at water depths between 1600 and 2200 m off southern Portugal. The combined evidence reveals enhanced current activity at these depths due to a deep glacial MOW. The MOW advection at the Portuguese margin during the last Glacial was about 700 m deeper than today, conceivably forced by increased MOW density due to higher salinity and colder temperatures of Mediterranean waters. The deep MOW current gradually decreased in strength and shoaled to 1300 m water depth during the Termination and early Holocene. A shallow MOW core layer became active with the onset of Termination I at depths between 600 and 1000 m. Both the shallow and deep MOW current culminated during the Younger Dryas period. The present flow pattern with two MOW core layers centred at 800 and 1300 m water depth was established between 7.5 and 5.5 ka.

The distribution of trace fossils and benthic foraminiferal assemblages in two sediment cores from 2160 m water depth at the northern and 1100 m at the southern Portuguese continental margin was used to reconstruct the bottom-water oxygenation during the last 40 ka. Emphasis was given to the Heinrich events, during which enhanced meltwater flux associated with the iceberg surges reduced or even halted deep water formation in the North Atlantic. Condensed trace-fossil tiering structures and high proportions of low-oxygen-tolerant benthic foraminifera suggest a major drawdown of bottom-water oxygenation during Heinrich events. Using the benthic foraminiferal oxygenation index (BFOI) of Kaiho (1991, 1994) that links benthic foraminiferal communities to ambient bottom-water oxygenation, we extracted apparent oxygen concentrations from benthic assemblages. BFOI values suggest low-oxic conditions during Heinrich events 1 and 4 with mean oxygen levels of 2.6 to 2.8 ml l −1 and high-oxic conditions with 4.1 ml l −1 during Heinrich event 2 at the shallower site. High proportions of low-oxygen-tolerant foraminiferal species and BFOI minima are recorded a few centimetres below the maximum concentrations of ice-rafted debris, indicating that the original microhabitat depth of dysoxic species is preserved beneath the Heinrich layers even though bioturbation continued during the events. Trace-fossil tiering calibrated with the redox boundary in low-oxic environments and dysoxic ichnotaxa commonly associated with black shales indicate oxygen levels lower than those implied by the BFOI. Oxygenation estimates derived from trace-fossil tiering denote low-oxic conditions during H1 (13.3 ka) with oxygen concentrations slightly above 1 ml l −1 , and dysoxic conditions during H2 (20.6 ka), H3 (27 ka) and H4 (33.5 ka) with oxygen concentrations between 0.1 and 1 ml l −1 . During the Younger Dryas, bottom waters likely were low oxic. Negative excursions of benthic carbon isotope values provide independent control on the oxygen drawdown during Heinrich events. Considering a bottom-water oxygenation of 6.1 to 6.5 ml l −1 during the last glacial, the benthic δ 13 C minima reveal an oxygen depletion to values between 3.4 and 5.0 ml l −1 during H1 through H4 at the shallower core site. Differences between estimates from benthic δ 13 C, BFOI and trace-fossil tiering suggest that benthic foraminiferal communities are rather robust to oxygen depletion if the level does not fall below 3 ml l −1 , whereas burrowing organisms react sensitively to depth variations of the redox boundary in near-surface sediments due to even small changes of bottom-water oxygenation.

Benthic foraminifera and surface sediments were analyzed from 23 box-core and core-top samples from 250 to 3600 m depth at the southern Portuguese continental margin between 37 and 38 °N. This area is affected by the MOW which flows northwards along the slope between 600 and 1500 m waterdepth. Sediment structures, grain-size distribution and clay mineral assemblages of surface sediments indicate active winnowing in the upper part of the MOW, and deposition of the fine fraction near its lower boundary. R-mode cluster analyses of foraminiferal census data from the > 250 μm fraction revealed four associations. The ‘Shelf Edge Association’ occurs on glauconitic shelf and upper slope sands down to 268 m. The ‘Upper Slope Association’ is recognized between 498 and 1300 m where sand-silt-clays to clayey silts are encountered. The ‘Lower Slope Association’ is found between 1405 and 2985 m and differs markedly from assemblages below 3000 m (‘Deep Water Association’) where the near-surface sediments are soft hemipelagic clays. The ‘Boundary Layer Association’ shows no significant depth limits but abundance maxima both in the density interface above the MOW and in the interval of decreasing turbulence below it. The boundaries of the ‘Shelf Edge’ and ‘Upper Slope Association’ correspond to these hydrographic boundary layers whereas the boundary between ‘Lower Slope’ and ‘Deep Water Assemblage’ is related to threshold values of nutrient flux to the benthic community. Examination of hard-substrates reveals 27 different species of epibenthic foraminifera. Six of these, the ‘Epibenthos Group’, were found attached to elevated substrates only within the MOW. Above the MOW, elevated substrates were inhabited by different species, whilst below the MOW they were not used by any epibenthic foraminifers. Evidently, the hydrodynamic environment of the MOW current provides an ecological niche which is efficiently used by opportunistic suspension feeders. The ‘Epibenthos Group’ is more abundant in the middle part of the study area between 37 °20′ and 37 °40′N where the sand content of surface sediments is also higher. Both indicate an enhanced sediment-water interaction in this area which results from an obstruction of the MOW flow by the ‘Principes d'Avis’ basement spur.

The discovery that foraminifera are able to use nitrate instead of oxygen as an electron acceptor for respiration has challenged our understanding of nitrogen cycling in the ocean. It was thought before that only prokaryotes and some fungi are able to denitrify. Rate estimates of foraminiferal denitrification have been very sparse and limited to specific regions in the oceans, not comparing stations along a transect of a certain region. Here, we present estimates of benthic foraminiferal denitrification rates from six stations at intermediate water depths in and below the Peruvian oxygen minimum zone (OMZ). Foraminiferal denitrification rates were calculated from abundance and assemblage composition of the total living fauna in both surface and subsurface sediments, as well as from individual species specific denitrification rates. A comparison with total benthic denitrification rates as inferred by biogeochemical models revealed that benthic foraminifera probably account for the total denitrification in shelf sediments between 80 and 250 m water depth. The estimations also imply that foraminifera are still important denitrifiers in the centre of the OMZ around 320 m (29–50% of the benthic denitrification), but play only a minor role at the lower OMZ boundary and below the OMZ between 465 and 700 m (2–6% of total benthic denitrification). Furthermore, foraminiferal denitrification has been compared to the total benthic nitrate loss measured during benthic chamber experiments. The estimated foraminiferal denitrification rates contribute 2 to 46% to the total nitrate loss across a depth transect from 80 to 700 m, respectively. Flux rate estimates range from 0.01 to 1.3 mmol m−2 d−1. Furthermore we show that the amount of nitrate stored in living benthic foraminifera (3 to 3955 μmol L−1) can be higher by three orders of magnitude as compared to the ambient pore waters in near-surface sediments sustaining an important nitrate reservoir in Peruvian OMZ sediments. The substantial contribution of foraminiferal nitrate respiration to total benthic nitrate loss at the Peruvian margin, which is one of the main nitrate sink regions in the world ocean, underpins the importance of the previously underestimated role of benthic foraminifera in global biogeochemical cycles.

The objective of this work was to review the distribution of benthic foraminiferal species at the western European continental margin from 43–58uN, determine their diversity, and generate a standardized taxonomy based on 44 publications (1913–2010) and unpublished information. Qualitative and quantitative data based upon foraminiferal occurrences and species abundances were included together with supplementary sedimentological and hydrographical data. From the species inventory, as well as from differences in morphological, physical, and hydrographic conditions in the study area, we defined six regions. The investigation of 2902 stations revealed 1486 species, of which 26% are synonymous. Most of the species have a hyaline test and live free, on or in the sediment. We recorded 608 species whose distributions were confined only to one of the six regions. Quantitative faunal data showed a general diversity increase from shelf to slope and two mid-slope diversity maxima, one located on the Basque continental margin at 550–850-m water depth and the other west of Ireland at 700–1100 m. In addition, the number of living species on the shelf generally increased from N–S. The latitudinal vs. depth distribution of six dominant species showed an irregular, lobate distribution pattern for the shelf regions. These species displayed similar distribution patterns on the continental slope, despite different modes of life, and different food and substrate preferences. This suggests that they have the same ecohabitat throughout their depth range. The faunal distribution pattern revealed close relationships between the different regions despite their varying hydrologic regimes.A delineation of one or two regions based on faunal criteria has not been attempted to date. There were no major distinctions in diversity among the six regions of the NE Atlantic, but the whole area exhibits an interregional diversity (c-diversity of 16), similar in magnitude to that of the Gulf of Mexico. In some parts of the study area, gaps in data coverage and differences in foraminiferal taxonomy and hydrographic conditions prevented the calculation of diversity indices.

Same-area comparison between patterns of Recent (living) and Holocene benthic foraminiferal assemblage composition is essential to validate their utility as proxies for paleoenvironmental reconstructions. Such reconstructions have scarcely been attempted in shelf environments. In this study, we compared living (stained), dead and Holocene benthic foraminiferal assemblages from the Gulf of Cadiz continental shelf off the Guadiana River. On average, 99% of the living benthic foraminiferal species were preserved in the dead assemblage and 95% in the fossil record. Several common species were assessed as indicators for certain environmental factors, including river discharge, supply of terrestrial organic matter, heavy metal concentrations, oxygen levels, substrate properties, hydrodynamic energy levels, sea-level rise and human impact. The paleoenvironmental reconstruction depicted five stages of Holocene environmental evolution of the Guadiana shelf over the last 11,500 cal yr BP. The characteristics of the stages, and the transitions between them, were in particular driven by changes in sea-level, influx of terrestrial organic matter and limits of productivity.

Colonization structures of the large parasitic foraminifer Hyrrokkin sarcophaga Cedhagen, 1994 on Lophelia pertusa, Acesta excavata and Delectopecten vitreus are described from a deep-water coral reef mound on the mid-Norwegian shelf at 240 to 300 m water depth. Hyrrokkin sarcophaga is the only epibiont which is capable of attaching itself on the soft tissue-protected coral skeleton where it tends to form clusters of 3 to 8 specimens close to the tentacles of Lophelia. The foraminifer excavates a pit up to 1.5 mm deep and etches a straight channel through the skeleton of the host which ends within the soft tissue. In contrast to Lophelia, infested bivalves show a strong wound repair reaction and seal the etched channels by intense calcification. The etching is only performed by adult specimens. Substrate pitting is considered to improve the attachment strength while boring enables the parasite to secure a persistent nutrient source.

Living (stained) benthic foraminiferal faunas were analyzed in 51 surficial sediment samples taken from the northern Gulf of Cadiz continental shelf between the mouths of the Guadiana and Guadalquivir rivers. The distribution and abundance of 26 species with relative abundance .5% were related to water depth, sediment type, river discharge, water temperature, salinity, turbidity, and primary productivity. Hierarchical classification using R- and Q-mode cluster analyses, and individual distributions, allowed these taxa to be categorized into four general groups. Group 1 is represented by Bolivina ordinaria and Hopkinsina atlantica. These species are linked to river discharge, and are associated with muddy sediments. They are considered the most opportunistic species in the assemblage, and are associated with the limits of productivity. Group 2, contains Ammonia beccarii, Eggerelloides scaber, Elphidium gerthi, Quinqueloculina laevigata, and Q. stelligera, and is characteristic of shallow water depths. Its species are associated with different sediment types and their distribution is influenced by river discharge. Group 3 is dominated by Bulimina aculeata, B. elongata, Elphidium excavatum, E. cuvillieri, Epistominella vitrea, and Rectuvigerina phlegeri. These species display the highest abundances between 30–100 m water depths. They prevail in muds associated with weak hydrodynamics, low oxygenation, and high organic matter. Group 4 includes species that are more abundant in deep stations and is divided into two subgroups. Subgroup 4A includes Bolivina catanensis, B. italica, B. striatula, Cassidulina laevigata, Stainforthia sp., Nouria polymorphinoides, and Nouria sp., with higher abundances in one or two samples. Subgroup 4B, with Brizalina dilatata, B. spathulata, Bulimina marginata, Nonionella iridea, N. stella, and N. turgida, has patchy distribution. Higher abundances of Nonionella species off the Guadiana River are related to low temperature and salinity. info:eu-repo/semantics/publishedVersion

This chapter will give a brief review about the present understanding of pores in tests of benthic foraminifera. The interpretation of the pore function changed through time, and a couple of theories were proposed. The research about the functionality of pores recently became of new interest because it seems likely that they are involved in the respiration pathways of some benthic foraminifera. The fact that several benthic species are able to survive anoxia points out the importance for a better understanding of these respiration pathways and which adaptions differentiate these species from species which cannot survive in oxygen-depleted habitats. Nitrate respiration seems to be widespread among foraminifera from oxygen-depleted habitats, and thus knowledge if and in how far the pores are involved in the process of denitrification would help to understand the process of denitrification in eukaryotic foraminiferal cells.

Recent benthic foraminifera and their distribution in surface sediments were studied on a transect through the Peruvian oxygen minimum zone (OMZ) between 10 and 12°S. The OMZ with its steep gradients of oxygen concentrations allows determinations of the oxygen-dependent changes of species compositions in a relatively small area. Our results from sediments of 13 multicorer stations from 79 to 823 m water depth demonstrate that calcareous species, especially bolivinids, dominate the assemblages throughout the OMZ. The depth distribution of several species matches distinct ranges of bottom water oxygen levels. The distribution pattern inferred a proxy which allows estimating dissolved oxygen concentrations for reconstructing oxygen levels in the geological past.

This study investigates the land-ocean interactions along the northern margin of the Gulf of Cadiz in the vicinity of the Guadiana River. Benthic foraminifera and sedimentological characteristics were analysed in a sedimentary sequence spanning ca. 5000 years (core 8, 22 m water depth) retrieved from the inner shelf prodeltaic wedge. The analyses were integrated in a temporal framework based on accelerated mass spectrometry radiocarbon dating. Paleoenvironmental changes and sediment transfer mechanisms from the continent to the shelf were investigated and related to climatic oscillations and anthropogenic impact in the region. The results allowed the identification of two main periods of deposition. The first period, from ca. 5150 cal. BP (core base) to 1500-1200 cal. BP, is characterised by a mix of fine and coarse sediments, relatively constant percentages of terrigenous and bioclasts, and benthic foraminifera species characteristic of coastal environments. These features indicate an environment strongly influenced by discharge from the Guadiana River. The second period, from ca. 1500-1200 cal. BP to ca. 200 cal. BP (core top), is characterised by silt-clay fraction dominated sediments, an increase in terrigenous sediment towards the top, and benthic foraminifera species characteristic of environments with low levels of energy. An increase in the level of human occupation associated with changes in climate led to widespread erosion and soil loss to the continental shelf during this period, with possible silting up of the Guadiana eastern distributary, allowing the transport of high amounts of sediment to the shelf by the western distributary that led to the enhancement of fine sedimentation and the formation of the prodeltaic wedge in the area of the studied core. info:eu-repo/semantics/publishedVersion

[1] This paper documents the migration of the Polar Front (PF) over the Iberian margin during some of the cold climatic extremes of the last 45 ka. It is based on a compilation of robust and coherent paleohydrological proxies obtained from eleven cores distributed between 36 and 42°N. Planktonic δ18O (Globigerina bulloides), ice-rafted detritus concentrations, and the relative abundance of the polar foraminifera Neogloboquadrina pachyderma sinistral were used to track the PF position. These three data sets, compared from core to core, show a consistent evolution of the sea surface paleohydrology along the Iberian margin over the last 45 ka. We focused on five time slices representative of cold periods under distinct paleoenvironmental forcings: the 8.2 ka event and the Younger Dryas (two recent cold events occurring within high values of summer insolation), Heinrich events 1 and 4 (reflecting major episodes of massive iceberg discharges into the North Atlantic), and the Last Glacial Maximum (typifying the highest ice volume accumulated in the Northern Hemisphere). For each event, we generated schematic maps mirroring past sea surface hydrological conditions. The maps revealed that the Polar Front presence along the Iberian margin was restricted to Heinrich events. The sea surface conditions during the Last Glacial Maximum were close to those at present day, except for the northern sites which briefly experienced subarctic conditions.

The living benthic foraminiferal assemblages in Kiel Fjord (SW Baltic Sea) were investigated in the years 2005 and 2006. The faunal studies were accomplished by geochemical analyses of surface sediments. In general, sediment pollution by copper, zinc, tin and lead is assessed as moderate in comparison with levels reported from other areas of the Baltic Sea. However, the inner Kiel Fjord is still exposed to a high load of metals and organic matter due to enhanced accumulation of fine-grained sediments in conjunction with potential pollution sources as shipyards, harbours and intensive traffic. The results of our survey show that the dominant environmental forcing of benthic foraminifera is nutrients availability coupled with human impact. A comparison with faunal data from the 1960s reveals apparent changes in species composition and population densities. The stress-tolerant species Ammonia beccarii invaded Kiel Fjord. Ammotium cassis had disappeared that reflects apparently the changes in salinity over the last 10 years. These changes in foraminiferal community and a significant increase of test abnormalities indicate an intensified environmental stress since the 1960s.

Early diagenetic features are noticed in the vicinity of carbonate platforms. Planktonic foraminifera of two tropical Atlantic deep-sea sediment cores show the strict relation between micro-scale euhydral crystallites of inorganic precipitates, higher oxygen isotope values and Mg/Ca ratios, and lower Sr/Ca ratios than expected for their pelagic environment in the time interval of ~100 000–550 000 calendar years before present. Laser ablation Mg/Ca (Sr/Ca) of crystallite-bearing foraminiferal chamber walls revealed 4–6 times elevated (2–3 times depleted) ratios, when ablating the diagenetic overgrowth. Crystalline overgrowth in proportions of 10–20% are estimated to cause the observed geochemical alteration. The extent of foraminiferal Mg/Ca alteration, moreover, seems to be controlled by the composition of the bulk sediment, especially the content of high-magnesium calcite. Anomalous ratios of >6 mmol/mol only occur, when high-magnesium calcite has dissolved within the sediment. The older parts (back to ~800 kyrs) of the records are characterized by similar trends of Mg/Ca and Sr/Ca. We discuss possible scenarios to accommodate the obtained geochemical information.