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Deep-sea Cibicidoides pachyderma (forma mundulus) and related Cibicidoides spp. were cultured at in situ pressure for 1–2 d, or 6 weeks to 3 months. During that period, fluorescence analyses following BCECF-AM (2′,7′-bis(2-carboxyethyl)-5-(and-6)-carboxyfluorescein acetoxymethyl ester) or calcein (bis[N,N-bis(carboxymethyl)aminomethyl]-fluorescein) labelling revealed a persisting cytoplasmic sheet or envelope surrounding the Cibicidoides tests. Thus, the Cibicidoides shell can be considered as an internal rather than an external cell structure. A couple of days to a week after being transferred into high-pressure aquaria and adjusted to a pressure of 115 bar, the foraminifera changed from a mobile to a more or less sessile living mode. During this quasi-sessile way of life, a series of comparably thick static ectoplasmic structures developed that were not resorbed or remodelled but, except for occasional further growth, remained unchanged throughout the experiments. Three different types of these permanent structures were observed. (a) Ectoplasmic “roots” were common in adult C. pachyderma, C. lobatulus, and C. wuellerstorfi specimens. In our experiments single ectoplasmic roots grew to a maximum of 700 times the individuals' shell diameter and were presumably used to anchor the specimen in an environment with strong currents. (b) Ectoplasmic “trees” describe rigid ectoplasmic structures directed into the aquarium's water body and were used by the foraminifera to climb up and down these ectoplasmic structures. Ectoplasmic trees have so far only been observed in C. pachyderma and enabled the tree-forming foraminifera to elevate itself above ground. (c) Ectoplasmic “twigs” were used to guide and hold the more delicate pseudopodial network when distributed into prevailing currents and were, in our experiments, also only developed in C. pachyderma specimens. Relocation of a specimen usually required it to tear apart and leave behind the rigid ectoplasmic structures and eventually also the envelope surrounding the test. Apparently, these rigid structures could not be resorbed or reused.

Following early hypotheses about the possible existence of Arctic ice shelves in the past1–3, the observation of specific erosional features as deep as 1,000 metres below the current sea level confirmed the presence of a thick layer of ice on the Lomonosov Ridge in the central Arctic Ocean and elsewhere4–6. Recent modelling studies have addressed how an ice shelf may have built up in glacial periods, covering most of the Arctic Ocean7,8. So far, however, there is no irrefutable marine-sediment characterization of such an extensive ice shelf in the Arctic, raising doubt about the impact of glacial conditions on the Arctic Ocean. Here we provide evidence for at least two episodes during which the Arctic Ocean and the adjacent Nordic seas were not only covered by an extensive ice shelf, but also filled entirely with fresh water, causing a widespread absence of thorium-230 in marine sediments. We propose that these Arctic freshwater intervals occurred 70,000–62,000 years before present and approximately 150,000–131,000 years before present, corresponding to portions of marine isotope stages 4 and 6. Alternative interpretations of the first occurrence of the calcareous nannofossil Emiliania huxleyi in Arctic sedimentary records would suggest younger ages for the older interval. Our approach explains the unexpected minima in Arctic thorium-230 records9 that have led to divergent interpretations of sedimentation rates10,11 and hampered their use for dating purposes. About nine million cubic kilometres of fresh water is required to explain our isotopic interpretation, a calculation that we support with estimates of hydrological fluxes and altered boundary conditions. A freshwater mass of this size—stored in oceans, rather than land—suggests that a revision of sea-level reconstructions based on freshwater-sensitive stable oxygen isotopes may be required, and that large masses of fresh water could be delivered to the north Atlantic Ocean on very short timescales. Unexpected intervals of low 230Th concentration in marine sediment cores are explained by considering that during at least two such periods, the Arctic Ocean and Nordic seas were composed entirely of fresh water and covered by a thick ice shelf.

Here we describe two new species of the genus Turrispirillina associated with large siliceous sponge reefs on the Karasik Seamount (Gakkel Ridge, Arctic Ocean). Careful analysis of Rose Bengal-stained samples and observations on untreated frozen sponge material revealed both species live exclusively inside siliceous sponges of the genus Geodia. More detailed information on the intra-sponge habitat was obtained from untreated frozen sponge material that showed both Turrispirillina species attach themselves to the large megascelere spicules that stabilize the choanoflagellates-harbouring subcortical crypts situated under the thick sponge cortex. Unstained specimens of both species were very abundant in the sediments surrounding the sponge. The number of Rose Bengal-stained specimens in each sample obviously depended on the penetration of a Geodia sponge, likely also the exact position of penetration with respect to the sponge. As sexual and asexual generations are observed in both taxa and sampling took place in autumn, opportunistic behavior with rapid reproduction following the spring bloom may determine the standing stock of both species. These are the first Arctic Turrispirillina species described with pseudospines.

This is the first study on the interactions between foraminifera and sponges. Although Cibicides and Hyrrokin are regarded as parasites on siliceous sponges, it is not yet clarified whether foraminifera specifically colonize sponges or are accidentally sucked in during the pelagic stage. To better elucidate these relationships, 12 sponges of different genera were examined and their foraminiferal communities analyzed. In 2018, the sponges for this study were collected with a ROV in water depths of 223 to 625 m in the Norwegian-Greenland Sea. Sponge parts were preserved in ethanol (96 %) and stained with Rose Bengal (2g l-1) to allow a differentiation between the living and dead foraminiferal fauna. Each sponge sample contained several hundred live and dead foraminiferal individuals of up to 60 different species. Even on Geodia baretti, which is able to release barettin to avoid colonalisation of other organisms, few foraminiferal individuals were observed. On all sponges, the most abundant genus was Cibicides, with Cibicides lobatulus and Cibicides refulgens as the most common taxa. Other very common species were Discorbinella bertheloti or Epistominella nipponica. Also, Hyrrokkin sarcophaga was found on different sponges and following its lifestyle, penetrating the sponge surfaces. The fact that besides adult foraminifera splendid juvenile stages were found indicate that foraminifera reproduced while inside the sponges. This reproduction might be stimulated/triggered by enhanced food availability by the pumping sponge.In summary, sponges are a special habitat for a high number of foraminiferal taxa. Their interaction ranges from parasitic lifestyle up to reproduction purposes. All these aspects highlight the importance of foraminifera-sponge interactions.

Numerous studies have addressed the possible existence of large floating ice sheets in the glacial Arctic Ocean from theoretical, modelling, or seafloor morphology perspectives. Here, we add evidence from the sediment record that support the existence of such freshwater ice caps in certain intervals, and we discuss their implications for possible non-linear and rapid behaviour of such a system in the high latitudes.We present sedimentary activities of 230Th together with 234U/238U ratios, the concentrations of manganese, sulphur and calcium in the context of lithological information and records of microfossils and their isotope composition. New analyses (PS51/038, PS72/396) and a re-analysis of existing marine sediment records (PS1533, PS1235, PS2185, PS2200, amongst others) in view of the naturally occurring radionuclide 230Thex and, where available, 10Be from the Arctic Ocean and the Nordic Seas reveal the widespread occurrence of intervals with a specific geochemical signature. The pattern of these parameters in a pan-Arctic view can best be explained when assuming the repeated presence of freshwater in frozen and liquid form across large parts of the Arctic Ocean and the Nordic Seas.Based on the sedimentary evidence and known environmental constraints at the time, we develop a glacial scenario that explains how these ice sheets, together with eustatic sea-level changes, may have affected the past oceanography of the Arctic Ocean in a fundamental way that must have led to a drastic and non-linear response to external forcing.This concept offers a possibility to explain and to some extent reconcile contrasting age models for the Late Pleistocene in the Arctic Ocean. Our view, if adopted, offers a coherent dating approach across the Arctic Ocean and the Nordic Seas, linked to events outside the Arctic.

The seafloor morphology of the Arctic Ocean and the Nordic Seas show specific capped structures in water depths of up to 1000 m. Together with modelling results, they strongly point to the existence of ice shelves covering the interior of the Arctic Ocean in past glacials[1,2]. In an attempt to link these major events to the sedimentary archive, we study a selection of new and existing data from many parts of the Arctic Ocean. We present activities of 230Th together with 234U/238U ratios, the concentrations of manganese, sulphur and calcium in the context of lithological information, and we show records of microfossils and their isotope composition. New high resolution analyses (PS51/038, PS72/396) and a re-analysis of existing marine sediment records (PS1533, PS1235, PS2185, PS2200, amongst others) in view of the naturally occurring radionuclide 230Thex and, where available, 10Be from the Arctic Ocean and the Nordic Seas reveal the widespread occurrence of intervals that are best explained by the temporary absence of sea water. [3]. We present a scenario of glacial conditions in which Bering Strait is closed and the Greenland-Scotland Ridge partly blocked by ice, leading to an accumulation of freshwater in liquid and solid form in the Arctic. This would explain how floating Arctic ice sheets, together with eustatic sea-level changes, have affected the past situation of the Arctic Ocean in a fundamental way that must have led to a drastic and non-linear response to certain threshold values, in agreement with sudden events in the climate record that were missing plausible explanations so far. This concept can reconcile contrasting age models for the Late Pleistocene in the Arctic Ocean. Our view, if adopted, would allow a coherent dating approach across the Arctic Ocean and the Nordic Seas, also offering a new perspective on climate records from lower latitudes.

Deciphering the evolution of marine plankton is typically based on the study of microfossil groups. Cryptic speciation is common in these groups, and large intragenomic variations occur in ribosomal RNA genes of many morphospecies. In this study, we correlated the distribution of ribosomal amplicon sequence variants (ASVs) with paleoceanographic changes by analyzing the high-throughput sequence data assigned to Neogloboquadrina pachyderma in a 140,000-year-old sediment core from the Arctic Ocean. The sedimentary ancient DNA demonstrated the occurrence of various N. pachyderma ASVs whose occurrence and dominance varied through time. Most remarkable was the striking appearance of ASV18, which was nearly absent in older sediments but became dominant during the last glacial maximum and continues to persist today. Although the molecular ecology of planktonic foraminifera is still poorly known, the analysis of their intragenomic variations through time has the potential to provide new insight into the evolution of marine biodiversity and may lead to the development of new and important paleoceanographic proxies.

This study aims to emphasize the ecological importance of foraminifera contained within the > 1000 µm sediment fraction. Stones and gravel offer a habitat for diverse and rich epilithic foraminiferal communities. However, gravel-rich sediments > 1000 µm are usually not the subject of quantitative benthic foraminifera studies, because most foraminifera species used as proxies or bioindicators are sediment-dwelling and found within smaller sediment size fractions. Therefore, there is a current lack of knowledge about the biology and ecology of epilithic foraminifera, specifically about agglutinated species.During a cruise at the Gullmarfjord/Sweden in September 2018, we retrieved a core at a station at 7 m water depth, which contained organic-rich material and a relatively large portion of gravel and shell fragments, which were densely populated by monothalamus, agglutinated foraminifera.A faunal analysis of foraminifera in the > 1000 µm sediment fraction showed, that the most abundant species (> 85 %, 54 ind. 10 cm-3) in > 1000 µm consisted of Tholosina vesicularis, an unicellular agglutinated species that can reach up to 4 mm diameter. SEM-analysis revealed, that large quantities of partially decomposed diatom frustules were embedded within the protoplasm of T. vesicularis individuals, which were supposedly the remains of the foraminiferal diet. The sediment fraction of 125 - 1000 µm was dominated by Ammonia species (58 %, 190 ind. 10 cm-3), a genus known for its fast turnover of diatom-derived carbon and as key-players in benthic nutrient fluxes. Preliminary biovolume analysis of the two genera (T. vesicularis: n = 74, Ammonia spp.: n = 110) resulted in far higher values for T. vesicularis (853 ± 944 µm-3 10 cm-3, med. = 506 µm-3 10 cm-3), than for Ammonia spp. (117 ± 56 µm-3 10 cm-3, med. = 96 µm-3 10 cm-3), within the uppermost sediment layer of 0 - 1 cm. Therefore, T. vesicularis could be a main driver of benthic carbon turnover in gravel-dominated sediments. We hypothesize, that the epilithic fauna, when present is at least as relevant as sediment-dwelling species.Additionally, biodiversity indices (species diversity, Shannon-Index, Evenness), show a slightly higher diversity and a more even distribution within the softer substrate, compared to the stones.This is most likely due to the motility of the free-living forms of the smaller size fraction, which allows them a flexible response to spacial competition. However, niche separation between hard and soft-substrate allows the co-existence of the opportunistic species T. vesicularis and Ammonia spp..In conclusion, this study shows, that if present, the > 1000 µm fraction can contain important information for the interpretation of ecological studies on the communities of live foraminifera and their role in marine ecosystems.

To date, observations on a single location indicate that cryogenic gypsum (Ca[SO4]⚫2H2O) may constitute an efficient but hitherto overlooked ballasting mineral enhancing the efficiency of the biological carbon pump in the Arctic Ocean. In June–July 2017 we sampled cryogenic gypsum under pack ice in the Nansen Basin north of Svalbard using a plankton net mounted on a remotely operated vehicle (ROVnet). Cryogenic gypsum crystals were present at all sampled stations, which suggested a persisting cryogenic gypsum release from melting sea ice throughout the investigated area. This was supported by a sea ice backtracking model, indicating that gypsum release was not related to a specific region of sea ice formation. The observed cryogenic gypsum crystals exhibited a large variability in morphology and size, with the largest crystals exceeding a length of 1 cm. Preservation, temperature and pressure laboratory studies revealed that gypsum dissolution rates accelerated with increasing temperature and pressure, ranging from 6 % d−1 by mass in polar surface water (−0.5 ∘C) to 81 % d−1 by mass in Atlantic Water (2.5 ∘C at 65 bar). When testing the preservation of gypsum in formaldehyde-fixed samples, we observed immediate dissolution. Dissolution at warmer temperatures and through inappropriate preservation media may thus explain why cryogenic gypsum was not observed in scientific samples previously. Direct measurements of gypsum crystal sinking velocities ranged between 200 and 7000 m d−1, suggesting that gypsum-loaded marine aggregates could rapidly sink from the surface to abyssal depths, supporting the hypothesized potential of gypsum as a ballasting mineral in the Arctic Ocean.

Amino acid racemization (AAR) geochronology is a powerful tool for dating Quaternary marine sediments across the globe, yet its application to Arctic Ocean sediments has been limited. Anomalous rates of AAR in foraminifera from the central Arctic were reported in previously published studies, indicating that either the rate of racemization is higher in this area, or inaccurate age models were used to constrain the sediment ages. This study investigates racemization rates in foraminifera from three well-dated sediment cores taken from the Yermak Plateau during the 2015 TRANSSIZ (TRansitions in the Arctic Seasonal Sea Ice Zone) expedition on RV Polarstern. D and L isomers of the amino acids aspartic acid (Asp) and glutamic acid (Glu) were separated in samples of the planktic foraminifer Neogloboquadrina pachyderma and the benthic species Cassidulina neoteretis to quantify the extent of racemization. In total, 241 subsamples were analysed, extending back to marine oxygen isotope stage (MIS) 7. Two previously published power functions, which relate the extent of racemization of Asp and Glu in foraminifera to sample age are revisited, and a comparison is made between the ages predicted by these calibrated age equations and independent geochronological constraints available for the cores. Our analyses reveal an excellent match between ages predicted by a global compilation of racemization rates for N. pachyderma and confirm that a proposed Arctic-specific calibration curve is not applicable at the Yermak Plateau. These results generally support the rates of AAR determined for other cold bottom water sites and further highlight the anomalous nature of the purportedly high rate of racemization indicated by previous analyses of central Arctic sediments.

Amino acid racemization (AAR) geochronology is a powerful tool for dating Quaternary marine sediments across the globe, yet its application to Arctic Ocean sediments has been limited. Anomalous rates of AAR in foraminifera from the central Arctic were reported in previously published studies, indicating that either the rate of racemization is higher in this area, or inaccurate age models were used to constrain the sediment ages. This study investigates racemization rates in foraminifera from three well-dated sediment cores taken from the Yermak Plateau during the 2015 TRANSSIZ Expedition on RV Polarstern. D and L isomers of the amino acids, aspartic acid (Asp) and glutamic acid (Glu), were separated in samples of the planktic foraminifera, Neogloboquadrina pachyderm and the benthic species, Cassidulina neoteretis to quantify the extent of racemization. In total, 241 subsamples were analysed, extending back to Marine Isotope Stage (MIS) 7. Two previously published power functions, which relate the extent of racemization of Asp and Glu in foraminifera to sample age are revisited, and a comparison is made between the ages predicted by these calibrated age equations and independent geochronological constraints available for the cores. Our analyses reveal an excellent match between ages predicted by a global compilation of racemization rates for N. pachyderma, and confirm that a proposed Arctic-specific calibration curve is not applicable at the Yermak Plateau. These results generally support the rates of AAR determined for other cold bottom water sites, and further highlight the anomalous nature of the purportedly high rate of racemization indicated by previous analyses of central Arctic sediments.

Mineral ballasting enhances carbon export from the surface to the deep ocean; however, little is known about the role of this process in the ice-covered Arctic Ocean. Here, we propose gypsum ballasting as a new mechanism that likely facilitated enhanced vertical carbon export from an under-ice phytoplankton bloom dominated by the haptophyte Phaeocystis. In the spring 2015 abundant gypsum crystals embedded in Phaeocystis aggregates were collected throughout the water column and on the sea floor at a depth below 2 km. Model predictions supported by isotopic signatures indicate that 2.7 g m−2 gypsum crystals were formed in sea ice at temperatures below −6.5 °C and released into the water column during sea ice melting. Our finding indicates that sea ice derived (cryogenic) gypsum is stable enough to survive export to the deep ocean and serves as an effective ballast mineral. Our findings also suggest a potentially important and previously unknown role of Phaeocystis in deep carbon export due to cryogenic gypsum ballasting. The rapidly changing Arctic sea ice regime might favour this gypsum gravity chute with potential consequences for carbon export and food partitioning between pelagic and benthic ecosystems.

Most palaeo-deep-water reconstructions are based on geochemical information stored in the calcareous shells of Cibicidoides species but hardly anything is known about their life cycle, population dynamics or ecology. The number of specimens of a single Cibicidoides species can locally be very limited and species may be lacking completely during certain intervals in the geological past. As a consequence, geochemical analyses are often carried out on lumped Cibicidoides spp. assuming that they share the same epizoic to epifaunal habitat and precipitated their shell in comparable offsets to surrounding bottom water mass properties. However, there is a growing body of evidence that particularly Cibicidoides pachyderma and its morphotypes C. mundulus and C. kullenbergi, may not be reliable bottom water recorders. We have recently developed aquaria that allowed, for the first time, observations of Cibicidoides pachyderma var. C. mundulus under in situ pressure and temperature. Experiments were carried out with and without artificial sediments to simulate soft sediments and rocks, respectively. Seawater was set to pH 8 and pH 7.4 to simulate more or less particulate carbon export or more or less ventilation of bottom water. Our experiments demonstrate that C. mundulus may opt for an epifaunal or an infaunal habitat depending on elapsed time following physical disturbance, pH, current activity, the availability of sediments and growth. The specimen's initial response following transfer from atmospheric pressure into the high-pressure aquaria was to immerse into the sediment or to cover more or less parts of the test with aggregated sediments or algae. However, within 24 h a strong rheotaxis became apparent and most specimens moved to sites of increased current activity under normal pH conditions (pH 8). Only few specimens remained in algae cysts or in the sediment in the pH-8 experiment. On the contrary, all specimens under pH 7.4 agglutinated a firm sediment cyst around their test and remained infaunal throughout the experimental period of three months. Independent of pH, growth was only observed in specimens that lived within an agglutinated cyst or infaunal. A solid thick cyst covered the specimens of the pH 7.4 experiment throughout the experiment and possibly restricted water exchange between the in-cyst water and the surrounding artificial bottom water mass. We suggest that a more fragile and possibly more porous sedimentary envelope may, at least temporally, have covered the infaunal specimens under pH 8 but no evidence for this was found upon termination of the experiment.

In field studies of active hydrocarbon seeps the carbon isotopic composition of Rose Bengal stained benthic foraminiferal tests (δ13Ctest) and bottom water DIC (δ13CDIC) deviates from their normal marine ratios. This circumstance led to ongoing discussions on whether aerobic foraminifers like Cibicides wuellerstorfi are capable of living at seepage sites and, more importantly, if their tests reflect the low δ13C values of emanating methane. To evaluate the discrepancy between δ13CDIC and δ13Ctest, we conducted methane seepage-emulating culture experiments on undepressurized sediments from the Håkon Mosby Mud Volcano, a modern methane seepage structure that hosts living C. wuellerstorfi with distinct negative δ13C values. The collected sediments were cultured at a site-alike pressure and mean bottom water methane concentration using newly developed high-pressure aquaria. Over an experimental period of 5 months our novel technology enabled a successful reproduction of all calcareous deep-sea benthic foraminiferal species living at that site, notably the first C. wuellerstorfi cultured in the laboratory. To show the influence of methane on δ13Ctest, we ran parallel experiments with > 99% 12C- and 99% 13C-methane in the experimental “bottom water”. During the experimental running time methanotrophs in the water column obviously converted the experimentally added methane source to δ13C-enriched and -depleted DIC, respectively. Since whole sediment cores were cultured, it was impossible to keep δ13CDIC constant over the 5-month duration, which is reflected in a variability of δ13Ctest in foraminiferal shells. Irrespective of that, the methane source is reflected in δ13Ctest of foraminiferal shells, and for the natural seep-conditions simulating 12C-experiment the mean δ13CDIC and δ13Ctest in C. wuellerstorfi were equal. Although for future culturing experiments improvements of the experimental conditions are advisable, our first results are evidence that persistent methane emanation impacts the carbon isotopic composition of deep-sea benthic foraminifera.

Scientific Reports 6: Article number: 38529; published online: 06 December 2016; updated: 17 May 2017 In Figure 1, the latitudes ‘80.5 N’ and ‘81.0 N’ were incorrectly given as ‘81.0 N’ and ‘81.5 N’ respectively. In addition, the scale between 0 km and 40 km was incorrectly given as between 0 km and80 km.

Published version. Source at http://doi.org/10.1038/srep38529. License CC BY 4.0. Slope failure like in the Hinlopen/Yermak Megaslide is one of the major geohazards in a changing Arctic environment. We analysed hydroacoustic and 2D high-resolution seismic data from the apparently intact continental slope immediately north of the Hinlopen/Yermak Megaslide for signs of past and future instabilities. Our new bathymetry and seismic data show clear evidence for incipient slope instability. Minor slide deposits and an internally-deformed sedimentary layer near the base of the gas hydrate stability zone imply an incomplete failure event, most probably about 30000 years ago, contemporaneous to or shortly after the Hinlopen/Yermak Megaslide. An active gas reservoir at the base of the gas hydrate stability zone demonstrate that over-pressured fluids might have played a key role in the initiation of slope failure at the studied slope, but more importantly also for the giant HYM slope failure. To date, it is not clear, if the studied slope is fully preconditioned to fail completely in future or if it might be slowly deforming and creeping at present. We detected widespread methane seepage on the adjacent shallow shelf areas not sealed by gas hydrates.

Aim The Arctic Ocean is one of the last near-pristine regions on Earth, and,although human activities are expected to impact on Arctic ecosystems, we knowvery little about baseline patterns of Arctic Ocean biodiversity. This paper aims todescribe Arctic Ocean-wide patterns of benthic biodiversity and to explore factorsrelated to the large-scale species diversity patterns.Location Arctic Ocean.Methods We used large ostracode and foraminiferal datasets to describe thebiodiversity patterns and applied comprehensive ecological modelling to test thedegree to which these patterns are potentially governed by environmental factors,such as temperature, productivity, seasonality, ice cover and others. To testenvironmental control of the observed diversity patterns, subsets of samples forwhich all environmental parameters were available were analysed with multipleregression and model averaging.Results Well-known negative latitudinal species diversity gradients (LSDGs)were found in metazoan Ostracoda, but the LSDGs were unimodal with anintermediate maximum with respect to latitude in protozoan foraminifera. Depthspecies diversity gradients were unimodal, with peaks in diversity shallower thanthose in other oceans. Our modelling results showed that several factors aresignificant predictors of diversity, but the significant predictors were differentamong shallow marine ostracodes, deep-sea ostracodes and deep-seaforaminifera.Main conclusions On the basis of these Arctic Ocean-wide comprehensivedatasets, we document large-scale diversity patterns with respect to latitude anddepth. Our modelling results suggest that the underlying mechanisms causingthese species diversity patterns are unexpectedly complex. The environmentalparameters of temperature, surface productivity, seasonality of productivity,salinity and ice cover can all play a role in shaping large-scale diversity patterns,but their relative importance may depend on the ecological preferences of taxaand the oceanographic context of regions. These results suggest that amultiplicity of variables appear to be related to community structure in thissystem.KeywordsArctic Ocean, biodiversity, deep sea, depth diversity gradients, ecosystem,latitudinal diversity gradients, macroecology, meiobenthos, shallow marine.

To investigate a possible influence of submarine methane seepage on benthic foraminiferal communities, Rose Bengal stained (“live”) and empty tests of benthic foraminifera were studied from the sediment surface down to 15 cm sub-bottom depth of 12 sites at the Hakon Mosby mud volcano (HMMV). In addition, one reference site well away from the seep sites, but from similar water depths and the same general hydrographic setting was occupied for comparison. The HMMV is located at 1265 m water depth on the SW Barents Sea continental slope. Distinct living foraminiferal associations at the HMMV are linked to specific sedimentary, microbial, and macrofaunal habitats. In the center of the crater, and in crater areas completely covered by bacterial mats, Cassidulina reniforme is the only, albeit rare, living species. Below the top few millimeters, sediments are anoxic and devoid of living specimens. At the rim of the mud volcano, at sites densely populated by pogonophoran tube worms, three benthic foraminiferal associations are found; (i) a Fontbotia wuellerstorfi–Lobatula lobatula association living attached to the upper parts of pogonophoran tubes, which protrude into oxic water, (ii) a diverse Cassidulina neoteretis association populating dysoxic sediments of the surface centimeter, and (iii) a species-poor Bolivina pseudopunctata association colonizing the subsurface sediments down to four centimeters. Generally, we did not find endemic or seep indicative species or associations at the HMMV. However, the HMMV live faunas dominated by B. pseudopunctata are not found at the reference site nor are they described from comparable water depths and environments without gas seepages from the Norwegian-Greenland Seas. In the center and outer rim of the mud volcano, a C. neoteretis–Reophax guttifer dead association, similar to the one at the reference site, characterizes an assemblage of strongly corroded and partly displaced tests. At bacterial mat sites, a C. reniforme dead association corresponds to the living one. Thus both the living and the dead associations are indicative of a specific bacterial mat environment at the HMMV.

Investigations on the benthic foraminiferal fauna of sediment cores from the Eurasian Basin (Arctic Ocean) continental margin, reveal well correlated biodiversity maxima and minima during the last 24 kyrs. The temporal variable biodiversities observed at the core sites, on a large scale, reflect the palaeoclimatic evolution of the high northern latitudes. Herein sediments deposited during interglacials and interstadials reveal highest species richness, whereas, glacials and stadials are documented by lower species numbers. In high-resolution core PS2837 different periodicities in species richness in sediments from the Late Weichselian and Holocene can be detected. Biodiversity periodicities of 1.57 kyrs and 0.76 kyrs characterize sediments of the Late Weichselian, whereas, with the retreat of glacial ice sheets and shorter seasonal ice coverage, Holocene sediments reveal shorter periodicities of 1.16 kyrs and 0.54 kyrs. With the establishment of modern hydrographic conditions at about 4000 BP, 4 kyrs, a significant increase in the amplitudes of species richness can be observed. Although sediments deposited during warmings reveal highest species richness, we doubt that water temperature determines the abundance and distribution of most benthic foraminifera in the Arctic Ocean. We rather suggest that the temporal variabilities in species richness reflect changes in the availability of food, which, in the Arctic Ocean, mainly depends on extend and duration of seasonal sea-ice retreat. In the study area the latter one, besides the seasonal varying insolation, is essentially determined by the advection-rates and temperature of Atlantic water, entering the Arctic Ocean via the West Spitsbergen Current. Because both, flowrate and temperature of the West Spitsbergen Current were increased during interstadials and interglacials, a predominantly indirect influence of palaeotemperature on the biodiversity of benthic foraminifera is suggested. Yet, rare species, like ‘Atlantic species' may indeed have a temperature sensitive metabolism. © 2007 Elsevier B.V. All rights reserved.

Analyses of benthic foraminifera in sediment cores taken at about 1000 m water depth at the Yermak Plateau and the Barents Sea slope, adjacent to the position of the ice-sheet edge during the Last Glacial Maximum, show that paleoproductivity was reduced to about a third of its present level during peak glacial stadials. These reduced values are still higher than values for modern, permanently ice-covered regions, suggesting that the core locations were at least partially ice-free even during stadials. Paleoproductivity at the core locations was higher than or equal to that of today during initial deglaciations and warm substages. Peak paleoproductivity occurred in samples with low-salinity surface waters as indicated by oxygen isotope values of planktonic foraminifera, and slightly after increased abundance of ‘Atlantic species’, suggesting that enhanced advection of warmer waters from the Atlantic supported the initial ice-sheet retreat. During the Holocene Climatic Optimum productivity was much less than at present on the Yermak Plateau, high at the Barents Sea site, perhaps because of increased advection of Atlantic water to the latter site and heavier ice coverage at the former. After this optimum, paleoproductivity at both sites was similar, with slightly lower values during cold periods such as the Little Ice Age.

The benthic foraminiferal record of two sediment cores at 900 and 2500 m water depth in the marginal Arctic Ocean reflects fluctuations in paleoproductivity, ice sheet extent, and Atlantic water inflow. Highest paleoproductivity is observed in interglacial periods, and at the termination of interstadials to stadials within oxygen isotope substages (OIS) 6.3, 5.3, 5.1, 3.1, and Termination Ia. Stable and high paleoproductivity at the ice-edge during terminations is indicated by the Melonis zaandami benthic foraminiferal association. Periods of glacial ice sheet advance or retreat from the core site are dominated by benthic foraminiferal associations related to seasonal organic carbon flux. Temperate saline Atlantic water entered the Arctic Ocean within OIS 6.3, 5.5 and the Holocene. Coincident paleoproductivity maxima are related to extended seasonal ice retreat. During glacial periods of stagnating Atlantic water advection unusual low paleoproductivity values are observed in the Arctic Ocean, indicating increasing ice coverage. Deep water was poorly ventilated during these periods and the deeper site became exposed to corrosive bottom waters.

We examine the quantitative composition of benthic foraminiferal assemblages of Rose Bengal-stained surface samples from 37 stations in the Laptev Sea, and combine this data set with an existing data set along a transect from Spitsbergen to the central Arctic Ocean. Foraminiferal test accumulation rates, diversity, faunal composition and statistically defined foraminiferal associations are analysed for living (Rose Bengal-stained) and dead foraminifers. We compare the results of several benthic foraminiferal diversity indices and statistically defined foraminiferal associations, including Fisher's alpha and Shannon–Wiener diversity indices, Q-mode principal component analysis and correspondence analysis. Diversity and faunal density (standing stock) of living benthic foraminifers are positively correlated to trophic resources. In contrast, the accumulation rate of dead foraminifers (BFAR) shows fluctuating values depending on test disintegration processes. Foraminiferal associations defined by Q-mode principal component analysis and correspondence analysis are comparable. The factor values of the correspondence analysis allow a quantitative correlation between the foraminiferal fauna and the local carbon flux, which may be used as a tool to estimate changes in primary productivity.

The vertical distribution of living (Rose Bengal stained) benthic foraminifers was determined in the upper 15 cm of sediment cores taken along transects extending from the continental shelf of Spitsbergen through the Eurasian Basin of the Arctic Ocean. Cores taken by a multiple corer were raised from 50 stations with water depths between 94 and 4427 m, from areas with moderate primary production values to areas that are among the least productive ones in the world. We believe, that in the Arctic Ocean the vertical distribution of living foraminifers is determined by the restricted availability of food. Live foraminiferal faunas are dominated by potentially infaunal species or epifaunal species. Species confined to the infaunal microhabitat are absent in Arctic sediments that we examined, and predominantly infaunal living species are nowhere dominant. In general, an infaunal mode of life is restricted to the seasonally ice-free areas and thus to areas with at least moderate primary production during the summer period. Under the permanent ice cover living species are usually restricted to the top centimeter of the sediment surface, even though some are able to dwell deeper in the sediment under ice-free conditions.

Fifty short sediment cores collected with a multiple corer and five box cores from the central Arctic Ocean were analysed to study the ecology and distribution of benthic foraminifers. To work out living faunal associations, standing stock and diversity, separate analyses of living (Rose Bengal stained) and dead foraminifers were carried out for the sediment surface. The size fractions between 63 and 125 μm and >125 μm were counted separately to allow comparison with former Arctic studies and with studies from the adjacent Norwegian–Greenland Sea, Barents Sea and the North Atlantic Ocean. Benthic foraminiferal associations are mainly controlled by the availability of food, and competition for food, while water mass characteristics, bottom current activity, substrate composition, and water depth are of minor importance. Off Spitsbergen in seasonally ice-free areas, high primary production rates are reflected by high standing stocks, high diversities, and foraminiferal associations (>125 μm) that are similar to those of the Norwegian–Greenland Sea. Generally, in seasonally ice-free areas standing stock and diversity increase with increasing food supply. In the central Arctic Ocean, the oligotrophic permanently ice-covered areas are dominated by epibenthic species. The limited food availability is reflected by very low standing stocks and low diversities. Most of these foraminiferal associations do not correspond to those of the Norwegian–Greenland Sea. The dominant associations include simple agglutinated species such as Sorosphaerae, Placopsilinellae, Komokiacea and Aschemonellae, as well as small calcareous species such as Stetsonia horvathi and Epistominella arctica. Those of the foraminiferal species that usually thrive under seasonally ice-free conditions in middle bathyal to lower bathyal water depth are found under permanently ice-covered conditions in water depths about 1000 m shallower, if present at all.

[1] To investigate the use of benthic foraminifera as a means to document ancient methane release, we determined the stable isotopic composition of tests of live (Rose Bengal stained) and dead specimens of epibenthic Fontbotia wuellerstorfi, preferentially used in paleoceanographic reconstructions, and of endobenthic high-latitude Cassidulina neoteretis and Cassidulina reniforme from a cold methane-venting seep off northern Norway. We collected foraminiferal tests from three push cores and nine multiple cores obtained with a remotely operated vehicle and a video-guided multiple corer, respectively. All sampled sites except one control site are situated at the Hakon Mosby mud volcano (HMMV) on the Barents Sea continental slope in 1250 m water depth. At the HMMV in areas densely populated by pogonophoran tube worms, δ13C values of cytoplasm-containing epibenthic F. wuellerstorfi are by up to 4.4‰ lower than at control site, thus representing the lowest values hitherto reported for this species. Live C. neoteretis and C. reniforme reach δ13C values of −7.5 and −5.5‰ Vienna Pee Dee Belemnite (VPDB), respectively, whereas δ13C values of their empty tests are higher by 4‰ and 3‰. However, δ13C values of empty tests are never lower than those of stained specimens, although they are still lower than empty tests from the control site. This indicates that authigenic calcite precipitates at or below the sediment surface are not significantly influencing the stable isotopic composition of foraminiferal shells. The comparatively high δ13C results rather from upward convection of pore water and fluid mud during active methane venting phases at these sites. These processes mingle tests just recently calcified with older ones secreted at intermittent times of less or no methane discharge. Since cytoplasm-containing specimens of suspension feeder F. wuellerstorfi are almost exclusively found attached to pogonophores, which protrude up to 3 cm above the sediment, and δ13C values of bottom-water-dissolved inorganic carbon (DIC) are not significantly depleted, we conclude that low test δ13C values of F. wuellerstorfi are the result of incorporation of heavily 13C-depleted methanotrophic biomass that these specimens feed on rather than because of low bottom water δ13CDIC. Alternatively, the pogonophores, which are rooted at depth in the upper sediment column, may serve as a conduit for depleted δ13CDIC that ultimately influences the calcification process of F. wuellerstorfi attached to the pogonophoran tube well above the sediment/water interface. The lowest δ13C of live specimens of the endobenthic C. neoteretis and C. reniforme are within the range of pore water δ13CDIC values, which exceed those that could be due to organic matter decomposition, and thus, in fact, document active methane release in the sediment.

During the Arctic Expedition ARK VIII/3 (August to September 1991) with RV Polarstern, the macrofauna, meiofauna, Foraminifera, bacteria were sampled and sediment chemistry was determined at 13 stations along a transect from the Barents Sea slope across the deep Arctic Eurasian Basins towards the Lomonosov Ridge. Water depths ranged from 258 to 4427 m. In general, higher values for all benthic compartments as well as total organic carbon (TOC) and total hydrolysable amino acids (THAA) were recorded for the Barents Sea slope than for the deeper stations in the basins and the ridge slopes. The only significant correlation found was between macrofaunal abundance and depth. Bacterial and all faunal abundances as well as bacterial and macrofaunal biomass decreased significantly with increasing latitude. Although correlations between food items such as TOC and THAA and the fauna were weak, significant relationships between the bacterial and faunal size-classes reflected a distinct food chain typical of oligotrophic systems. The smallest compartments -bacteria, meiofauna and Foraminifera- were more abundant than the macrofauna in the central Arctic Ocean. Macrofauna biomass dominated the biomass on the Barents Sea shelf and slope and on the Lomonosov Ridge, but bacterial biomass was equally or even more important on the Gakkel Ridge and in the deep basins. The results reveal the Eurasian Basin as one of the most oligotrophic regions in the World Ocean. Although primary production is low, recent foraminiferal investigations have revealed that benthic communities in the central Arctic Ocean are driven by the sedimentation of fresh organic material. Lateral transport of organic material from the Siberian shelf may provide additional food. The various benthic compartments compete either for fresh organic matter or for refractory material that is transferred to higher levels of the food chain by bacterial mineralisation.

We reconstructed late Quaternary deep (3000–4100 m) and intermediate depth (1000–2500 m) paleoceanographic history of the Eurasian Basin, Arctic Ocean from ostracode assemblages in cores from the Lomonosov Ridge, Gakkel Ridge, Yermak Plateau, Morris Jesup Rise, and Amundsen and Makarov Basins obtained during the 1991 Polarstern cruise. Modern assemblages on ridges and plateaus between 1000 and 1500 m are characterized by abundant, relatively species-rich benthic ostracode assemblages, in part, reflecting the influence of high organic productivity and inflowing Atlantic water. In contrast, deep Arctic Eurasian basin assemblages have low abundance and low diversity and are dominated by Krithe and Cytheropteron reflecting faunal exchange with the Greenland Sea via the Fram Strait. Major faunal changes occurred in the Arctic during the last glacial/interglacial transition and the Holocene. Low-abundance, low-diversity assemblages from the Lomonosov and Gakkel Ridges in the Eurasian Basin from the last glacial period have modern analogs in cold, low-salinity, low-nutrient Greenland Sea deep water; glacial assemblages from the deep Nansen and Amundsen Basins have modern analogs in the deep Canada Basin. During Termination 1 at intermediate depths, diversity and abundance increased coincident with increased biogenic sediment, reflecting increased organic productivity, reduced sea-ice, and enhanced inflowing North Atlantic water. During deglaciation deep Nansen Basin assemblages were similar to those living today in the deep Greenland Sea, perhaps reflecting deepwater exchange via the Fram Strait. In the central Arctic, early Holocene faunas indicate weaker North Atlantic water inflow at middepths immediately following Termination 1, about 8500–7000 year B.P., followed by a period of strong Canada Basin water overflow across the Lomonosov Ridge into the Morris Jesup Rise area and central Arctic Ocean. Modern perennial sea-ice cover evolved over the last 4000–5000 years. Late Quaternary faunal changes reflect benthic habitat changes most likely caused by changes in the import of cold, deepwater of Greenland Sea origin and warmer and middepth Atlantic water to the Eurasian Basin through the Fram Strait, and export of Arctic Ocean deepwater.