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Abstract. Benthic foraminiferal tests are widely used for paleoceanographic reconstructions from a range of different environments with varying dissolved oxygen concentrations in the bottom water. There is ample evidence that foraminifera can live in anoxic sediments. For some species, this is explained by a switch to facultative anaerobic metabolism (i.e. denitrification). Here we show for the first time that adult specimens of three benthic foraminiferal species are not only able to survive, but are also able to calcify under anoxic conditions, at various depths in the sediment, and with or without nitrates. In fact, several specimens of Ammonia tepida (1–4%), Bulimina marginata (8–24%) and Cassidulina laevigata (16–23%) were able to calcify at different redox fronts of sediment cores, under laboratory conditions. This demonstrates ongoing metabolic processes, even in micro-environments where denitrification is not possible. Earlier observations suggest that the disappearance of foraminiferal communities after prolonged anoxia is not due to instantaneous or strongly increased adult mortality. Here we show that it cannot be explained by an inhibition of growth through chamber addition either. Our observations of ongoing calcification under anoxic conditions mean that geochemical proxy data obtained from benthic foraminifera in settings experiencing intermittent anoxia have to be reconsidered. The analysis of whole single specimens or of their successive chambers may provide essential information about short-term environmental variability and/or the causes of anoxia.

Abstract. We present a new rapid and accurate protocol to simultaneously sample benthic living foraminifera in two dimensions in a centimetre-scale vertical grid and dissolved iron and phosphorus in two dimensions at high resolution (200 μm). Such an approach appears crucial for the study of foraminiferal ecology in highly dynamic and heterogeneous sedimentary systems, where dissolved iron shows a strong variability at the centimetre scale. On the studied intertidal mudflat of the Loire estuary, foraminiferal faunas are dominated by Ammonia tepida, which accounts for 92 % of the living (CellTracker Green(CTG)-labelled) assemblage. The vertical distribution shows a maximum density in the oxygenated 0–0.4 cm surface layer. A sharp decrease is observed in the next 2 cm, followed by a second, well-defined maximum in the suboxic sediment layer (3–8 cm depth). The presented method yields new information concerning the 2-D distribution of living A. tepida in suboxic layers. First, the identification of recent burrows by visual observation of the sediment cross section and the burrowing activity as deduced from the dissolved iron spatial distribution show no direct relation to the distribution of A. tepida at the centimetre scale. This lack of relation appears contradictory to previous studies (Aller and Aller, 1986; Berkeley et al., 2007). Next, the heterogeneity of A. tepida in the 3–8 cm depth layer was quantified by means of Moran's index to identify the scale of parameters controlling the A. tepida distribution. The results reveal horizontal patches with a characteristic length of 1–2 cm. These patches correspond to areas enriched in dissolved iron likely generated by anaerobic degradation of labile organic matter. These results suggest that the routine application of our new sampling strategy could yield important new insights about foraminiferal life strategies, improving our understanding of the role of these organisms in coastal marine ecosystems.

Abstract. Oxygen and nitrate availabilities impact the marine nitrogen cycle at a range of spatial and temporal scales. Here, we demonstrate the impact of denitrifying foraminifera on the nitrogen cycle at two oxygen and nitrate contrasting stations in a fjord environment (Gullmar Fjord, Sweden). Denitrification by benthic foraminifera was determined through the combination of specific density counting per microhabitat and specific nitrate respiration rates obtained through incubation experiments using N2O microsensors. Benthic nitrate removal was calculated from submillimeter chemical gradients extracted from 2D porewater images of the porewater nitrate concentration. These were acquired by combining the DET technique (diffusive equilibrium in thin film) with chemical colorimetry and hyperspectral imagery. Sediments with high nitrate concentrations in the porewater and oxygenated overlying water were dominated by the non-indigenous species (NIS) Nonionella sp. T1. Denitrification by this species could account for 50 %–100 % of the nitrate loss estimated from the nitrate gradients. In contrast sediments below hypoxic bottom waters had low inventories of porewater nitrate, and denitrifying foraminifera were rare. Their contribution to benthic nitrate removal was negligible (< 5 %). Our study showed that benthic foraminifera can be a major contributor to nitrogen mitigation in oxic coastal ecosystems and should be included in ecological and diagenetic models aiming to understand biogeochemical cycles coupled to nitrogen.

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