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In this study, we explore the correlation of I/Ca ratios in three calcitic and one aragonitic foraminiferal species. I/Ca ratios are evaluated as possible proxies for changes in ambient redox conditions across the Peruvian oxygen minimum zone to the ambient oxygen concentrations in the habitat of the foraminiferal species studied. Cleaning and measurement methods for the determination of I/Ca ratios are tested. All species show a positive trend in their I/Ca ratios as a function of higher oxygen concentrations. The most promising species appears to be Uvigerina striata, which shows a highly statistically significant correlation between I/Ca ratios and bottom water (BW) oxygenation (I/Ca = 0.032(±0.004) × [O2]BW + 0.29(±0.03), R2 = 0.61, F = 75, P < 0.0001). Only for the aragonitic species Hoeglundina elegans is this relationship not significant. The iodine volatility in acidic solutions, the species dependency of I/Ca–[O2]BW correlations, and the individual variability of single tests, need to be accounted for when applying the I/Ca ratio as a proxy for redox conditions.

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 oceans are losing oxygen (O2), and oxygen minimum zones are expanding due to climate warming (lower O2 solubility) and eutrophication related to agriculture. This trend is challenging for most marine taxa that are not well adapted to O2 depletion. For other taxa this trend might be advantageous because they can withstand low O2 concentrations or thrive under O2-depleted or even anoxic conditions. Benthic foraminifera are a group of protists that include taxa with adaptations to partly extreme environmental conditions. Several species possess adaptations to O2 depletion that are rare amongst eukaryotes, and these species might benefit from ongoing ocean deoxygenation. In addition, since some foraminifera can calcify even under anoxic conditions, they are important archives for paleoceanographic reconstruction in O2-depleted environments. This paper reviews the current state of knowledge about foraminifera from low-O2 environments. Recent advances in our understanding of specific survival strategies of foraminifera to withstand O2 depletion are summarized and discussed. These adaptations include an anaerobic metabolism, heterotrophic denitrification, symbiosis with bacteria, kleptoplasty and dormancy and have a strong impact on their preferred microhabitat in the sediments, especially the ability of some benthic foraminiferal species to denitrify. Benthic foraminifera also differ regarding their trophic strategies, which has an additional impact on the selection of their microhabitat. For example, some species are strict herbivores that feed exclusively on fresh phytodetritus and live close to the sediment surface, while some species are non-selective detrivores that occupy intermediate to deep infaunal habitats. There is evidence that foraminifers have the capacity to undergo phagocytosis, even under anoxia, and some foraminiferal species which can withstand low-O2 conditions seem to prey on meiofauna. Also, due to their high abundances in O2-depleted environments and their metabolic adaptations, benthic foraminifera are key players in marine nutrient cycling, especially within the marine N and P cycles. This review summarizes the denitrification rates for the species that are known to denitrify and the intracellular nitrate concentrations of the species that are known to intracellularly store nitrate. Finally, equations are provided that can be used to estimate the intracellular nutrient storage and denitrification rates of foraminifera and might be integrated into biogeochemical models.

In this study we present an initial dataset of Mn/Ca and Fe/Ca ratios in tests of benthic foraminifera from the Peruvian oxygen minimum zone (OMZ) determined with SIMS. These results are a contribution to a better understanding of the proxy potential of these elemental ratios for ambient redox conditions. Foraminiferal tests are often contaminated by diagenetic coatings, like Mn rich carbonate- or Fe and Mn rich (oxyhydr)oxide coatings. Thus, it is substantial to assure that the cleaning protocols are efficient or that spots chosen for microanalyses are free of contaminants. Prior to the determination of the element/Ca ratios, the distributions of several elements (Ca, Mn, Fe, Mg, Ba, Al, Si, P and S) in tests of the shallow infaunal species Uvigerina peregrina and Bolivina spissa were mapped with an electron microprobe (EMP). To visualize the effects of cleaning protocols uncleaned and cleaned specimens were compared. The cleaning protocol included an oxidative cleaning step. An Fe rich phase was found on the inner test surface of uncleaned U. peregrina specimens. This phase was also enriched in Al, Si, P and S. A similar Fe rich phase was found at the inner test surface of B. spissa. Specimens of both species treated with oxidative cleaning show the absence of this phase. Neither in B. spissa nor in U. peregrina were any hints found for diagenetic (oxyhydr)oxide or carbonate coatings. Mn/Ca and Fe/Ca ratios of single specimens of B. spissa from different locations have been determined by secondary ion mass spectrometry (SIMS). Bulk analyses using solution ICP-MS of several samples were compared to the SIMS data. The difference between SIMS analyses and ICP-MS bulk analyses from the same sampling sites was 14.0–134.8 μmol mol−1 for the Fe/Ca and 1.68(±0.41) μmol mol−1 for the Mn/Ca ratios. This is in the same order of magnitude as the variability inside single specimens determined with SIMS at these sampling sites (1σ[Mn/Ca] = 0.35–2.07 μmol mol−1; 1σ[Fe/Ca] = 93.9–188.4 μmol mol−1). The Mn/Ca ratios in the calcite were generally relatively low (2.21–9.93 μmol mol−1) but in the same magnitude and proportional to the surrounding pore waters (1.37–6.67 μmol mol−1). However, the Fe/Ca ratios in B. spissa show a negative correlation to the concentrations in the surrounding pore waters. Lowest foraminiferal Fe/Ca ratios (87.0–101.0 μmol mol−1) were found at 465 m water depth, a location with a strong sharp Fe peak in the pore water next to the sediment surface and respectively, high Fe concentrations in the surrounding pore waters. Previous studies found no living specimens of B. spissa at this location. All these facts hint that the analysed specimens already were dead before the Fe flux started and the sampling site just recently turned anoxic due to fluctuations of the lower boundary of the OMZ near the sampling site (465 m water depth). Summarized Mn/Ca and Fe/Ca ratios are potential proxies for redox conditions, if cleaning protocols are carefully applied. The data presented here may be rated as base for the still pending detailed calibration.

Present day oceans are well ventilated, with the exception of mid-depth oxygen minimum zones (OMZs) under high surface water productivity, regions of sluggish circulation, and restricted marginal basins. In the Mesozoic, however, entire oceanic basins transiently became dysoxic or anoxic. The Cretaceous ocean anoxic events (OAEs) were characterised by laminated organic-carbon rich shales and low-oxygen indicating trace fossils preserved in the sedimentary record. Yet assessments of the intensity and extent of Cretaceous near-bottom water oxygenation have been hampered by deep or long-term diagenesis and the evolution of marine biota serving as oxygen indicators in today's ocean. Sedimentary features similar to those found in Cretaceous strata were observed in deposits underlying Recent OMZs, where bottom-water oxygen levels, the flux of organic matter, and benthic life have been studied thoroughly. Their implications for constraining past bottom-water oxygenation are addressed in this review. We compared OMZ sediments from the Peruvian upwelling with deposits of the late Cenomanian OAE 2 from the north-west African shelf. Holocene laminated sediments are encountered at bottom-water oxygen levels of < 7 μmol kg−1 under the Peruvian upwelling and < 5 μmol kg−1 in California Borderland basins and the Pakistan Margin. Seasonal to decadal changes of sediment input are necessary to create laminae of different composition. However, bottom currents may shape similar textures that are difficult to discern from primary seasonal laminae. The millimetre-sized trace fossil Chondrites was commonly found in Cretaceous strata and Recent oxygen-depleted environments where its diameter increased with oxygen levels from 5 to 45 μmol kg−1. Chondrites has not been reported in Peruvian sediments but centimetre-sized crab burrows appeared around 10 μmol kg−1, which may indicate a minimum oxygen value for bioturbated Cretaceous strata. Organic carbon accumulation rates ranged from 0.7 and 2.8 g C cm−2 kyr−1 in laminated OAE 2 sections in Tarfaya Basin, Morocco, matching late Holocene accumulation rates of laminated Peruvian sediments under Recent oxygen levels below 5 μmol kg−1. Sediments deposited at > 10 μmol kg−1 showed an inverse exponential relationship of bottom-water oxygen levels and organic carbon accumulation depicting enhanced bioirrigation and decomposition of organic matter with increased oxygen supply. In the absence of seasonal laminations and under conditions of low burial diagenesis, this relationship may facilitate quantitative estimates of palaeo-oxygenation. Similarities and differences between Cretaceous OAEs and late Quaternary OMZs have to be further explored to improve our understanding of sedimentary systems under hypoxic conditions.

Deciphering the dynamics of dissolved oxygen in the mid-depth ocean during the last deglaciation is essential to understand the influence of climate change on modern oxygen minimum zones (OMZs). Many paleo-proxy records from the eastern Pacific Ocean indicate an extension of oxygen-depleted conditions during the deglaciation, but the degree of deoxygenation has not been quantified to date. The Peruvian OMZ, one of the largest OMZs in the world, is a key area to monitor such changes in near-bottom-water oxygenation in relation to changing climatic conditions. Here, we analysed the potential to use the composition of foraminiferal assemblages from the Peruvian OMZ as a quantitative redox proxy. A multiple regression analysis was applied to a joint dataset of living (rose-bengal-stained, fossilizable calcareous species) benthic foraminiferal distributions from the Peruvian continental margin. Bottom-water oxygen concentrations ([O2]BW) during sampling were used as the dependant variable. The correlation was significant (R2=0.82; p<0.05), indicating that the foraminiferal assemblages are rather governed by oxygen availability than by the deposition of particulate organic matter (R2=0.53; p=0.31). We applied the regression formula to three sediment cores from the northern part of the Peruvian OMZ between 3 and 8∘ S and 997 and 1250 m water depth, thereby recording oxygenation changes at the lower boundary of the Peruvian OMZ. Each core displayed a similar trend of decreasing oxygen levels since the Last Glacial Maximum (LGM). The overall [O2]BW change from the LGM and the Holocene was constrained to 30 µmol kg−1 at the lower boundary of the OMZ.

Quantifying past ocean nitrate concentrations is crucial for understanding the global nitrogen cycle. Here, we reconstruct deglacial bottom-water nitrate concentrations ([NO3-]BW) in the oxygen-deficient zones of the Sea of Okhotsk, the Gulf of California, the Mexican Margin, and the Gulf of Guayaquil. Using the pore density of denitrifying benthic foraminifera as a nitrate proxy, differences in [NO3-]BW are observed at the study sites spanning the Last Glacial Maximum to the Holocene. Changes in water-column denitrification, water-mass ventilation, primary productivity, and sea surface temperatures may account for nitrate differences at the study sites. The [NO3-]BW in the Sea of Okhotsk, the Gulf of California, and the Gulf of Guayaquil are influenced by the intermediate water masses while, the [NO3-]BW at the Mexican Margin is likely influenced by deglacial changes in the Pacific Deep Water. The comparison of past and present [NO3-] shows that the modern Gulf of Guayaquil and the Gulf of California currently have stronger oxygen-deficient zones with higher denitrification than during the Last Glacial Maximum. In contrast, the modern Mexican Margin and the Sea of Okhotsk may have higher oxygen than during the Last Glacial Maximum, indicated by low modern denitrification.

In the vicinity of hydrothermal vent fields, unique habitats are observed that are influenced by hydrothermal fluids. Benthic foraminifera can be part of the communities found around these hydrothermal vent fields. They can form suitable indicators for different marine environmental conditions because their tests are often well preserved in the sediments. In this work, living (rose-bengal-stained) and dead benthic foraminifera from six sediment core tops were investigated with increasing distance from the Rainbow hydrothermal vent field at the Mid-Atlantic Ridge. The proximal-to-distal transect was sampled starting at ∼ 200 m distance from the active vents and followed the vent plume for ∼ 41 km. The biodiversity and abundance of benthic foraminifera tended to increase with distance from the Rainbow hydrothermal vent field, and there was a significant difference between the living and dead assemblage. The Shannon–Wiener diversity was lower at 0.2–1 km distance from the vent field with 2.3 (living) and 2.8 (dead) and showed higher constant values of 3.0 to 3.3 (living) and ∼ 3.6 (dead) from a distance of ∼ 2 km onwards. The population density of living benthic foraminifera showed a similar pattern to the biodiversity, while the density of empty foraminiferal tests was subject to strong fluctuations. Differences in the species composition between live and dead assemblages indicate environmental fluctuations, which could be triggered by seasonal nutrient pulses or brief contacts of the vent plume with the sediment. Species composition changed with distance from the black smokers. While miliolids dominated sediments closer to the vent field, hyaline perforate and agglutinated species constituted the major parts of the assemblages at greater distances. Thus, miliolids may be better adapted to the environment potentially influenced by hydrothermal vent fluids than the hyaline and agglutinated species. Specifically, miliolids seem to tolerate elevated porewater concentrations of copper, cobalt, and zinc and are possibly influenced by intrusions of acidic hydrothermal fluids. This result is in contrast to studies from other venting sites with acidic fluids, where agglutinated species dominate. High biodiversity and abundances of benthic foraminifera suggest that a diverse benthic ecosystem occurs below the distal Rainbow vent plume.

Many benthic organisms show aggregated distribution patterns due to the spatial heterogeneity of niches or food availability. In particular, high-abundance patches of benthic foraminifera have been reported that extend from centimetres to metres in diameter in salt marshes or shallow waters. The dimensions of spatial variations of shelf or deep-sea foraminiferal abundances have not yet been identified. Therefore, we studied the distribution of Globobulimina turgida dwelling in the 0–3 cm surface sediment at 118m water depth in the Alsbäck Deep, Gullmar Fjord, Sweden. Standing stock data from 58 randomly replicated samples depicted a log-normal distribution of G. turgida with weak evidence for an aggregated distribution on a decimetre scale. A model simulation with different patch sizes, outlines, and impedances yielded no significant correlation with the observed variability of G. turgida standing stocks. Instead, a perfect match with a random log-normal distribution of population densities was obtained. The data–model comparison revealed that foraminiferal populations in the Gullmar Fjord were not moulded by any underlying spatial structure beyond 10 cm diameter. Log-normal population densities also characterise data from contiguous, gridded, or random sample replicates reported in the literature. Here, a centimetre-scale heterogeneity was found and interpreted to be a result of asexual reproduction events and restricted mobility of juveniles. Standing stocks of G. turgida from the Alsbäck Deep temporal data series from 1994 to 2021 showed two distinct cohorts of samples of either high or low densities. These cohorts are considered to represent two distinct ecological settings: hypoxic and well-ventilated conditions in the Gullmar Fjord. Environmental forcing is therefore considered to impact the population structure of benthic foraminifera rather than their reproduction dynamics.

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