SUMMARY: New data force us to raise previous estimates of oceanic denitrification. Our revised estimate of ~ 450 Tg N yr -1 (Tg = 10 12 g) produces an oceanic fixed N budget with a large deficit ) that can be explained only by positing an ocean that has deviated far from a steady-state, the need for a major upwards revision of fixed N inputs, particularly nitrogen fixation, or both. Oceanic denitrification can be significantly altered by small re-distributions of carbon and dissolved oxygen. Since fixed N is a limiting nutrient, uncompensated changes in denitrification affect the ocean's ability to sequester atmospheric CO 2 via the "biological pump". We have also had to modify our concepts of the oceanic N 2 O regime to take better account of the extremely high N 2 O saturations that can arise in productive, low oxygen waters. Recent results from the western Indian Shelf during a period when hypoxic, suboxic and anoxic waters were present produced a maximum surface N 2 O saturation of > 8000%, a likely consequence of "stop and go" denitrification. The sensitivity of N 2 O production and consumption to small changes in the oceanic dissolved oxygen distribution and to the "spin-up" phase of denitrification suggests that the oceanic source term for N 2 O could change rapidly.

Eutrophication of surface waters and hypoxia in bottom waters has been increasing in many coastal areas, leading to very large depletions of marine life in the affected regions. These areas of high surface productivity and low bottom-water oxygen concentration are caused by increasing runoff of nutrients from land. Although the local ecological and socio-economic effects have received much attention, the potential contribution of increasing hypoxia to global-change phenomena is unknown. Here we report the intensification of one of the largest low-oxygen zones in the ocean, which develops naturally over the western Indian continental shelf during late summer and autumn. We also report the highest accumulations yet observed of hydrogen sulphide (H2S) and nitrous oxide (N2O) in open coastal waters. Increased N2O production is probably caused by the addition of anthropogenic nitrate and its subsequent denitrification, which is favoured by hypoxic conditions. We suggest that a global expansion of hypoxic zones may lead to an increase in marine production and emission of N2O, which, as a potent greenhouse gas, could contribute significantly to the accumulation of radiatively active trace gases in the atmosphere.

Abstract. Coastal hypoxia (defined here as <1.42 ml L −1 ; 62.5 µM; 2 mg L −1 , approx. 30% oxygen saturation) develops seasonally in many estuaries, fjords, and along open coasts as a result of natural upwelling or from anthropogenic eutrophication induced by riverine nutrient inputs. Permanent hypoxia occurs naturally in some isolated seas and marine basins as well as in open slope oxygen minimum zones. Responses of benthos to hypoxia depend on the duration, predictability, and intensity of oxygen depletion and on whether H 2 S is formed. Under suboxic conditions, large mats of filamentous sulfide oxidizing bacteria cover the seabed and consume sulfide. They are hypothesized to provide a detoxified microhabitat for eukaryotic benthic communities. Calcareous foraminiferans and nematodes are particularly tolerant of low oxygen concentrations and may attain high densities and dominance, often in association with microbial mats. When oxygen is sufficient to support metazoans, small, soft-bodied invertebrates (typically annelids), often with short generation times and elaborate branchial structures, predominate. Large taxa are more sensitive than small taxa to hypoxia. Crustaceans and echinoderms are typically more sensitive to hypoxia, with lower oxygen thresholds, than annelids, sipunculans, molluscs and cnidarians.

The anaerobic oxidation of ammonium (anammox) contributes significantly to the global loss of fixed nitrogen and is carried out by a deep branching monophyletic group of bacteria within the phylum Planctomycetes. Various studies have implicated anammox to be the most important process responsible for the nitrogen loss in the marine oxygen minimum zones (OMZs) with a low diversity of marine anammox bacteria. This comprehensive study investigated the anammox bacteria in the suboxic zone of the Black Sea and in three major OMZs (off Namibia, Peru and in the Arabian Sea). The diversity and population composition of anammox bacteria were investigated by both, the 16S rRNA gene sequences and the 16S-23S rRNA internal transcribed spacer (ITS). Our results showed that the anammox bacterial sequences of the investigated samples were all closely related to the Candidatus Scalindua genus. However, a greater microdiversity of marine anammox bacteria than previously assumed was observed. Both phylogenetic markers supported the classification of all sequences in two distinct anammox bacterial phylotypes: Candidatus Scalindua clades 1 and 2. Scalindua 1 could be further divided into four distinct clusters, all comprised of sequences from either the Namibian or the Peruvian OMZ. Scalindua 2 consisted of sequences from the Arabian Sea and the Peruvian OMZ and included one previously published 16S rRNA gene sequence from Lake Tanganyika and one from South China Sea sediment (97.9-99.4% sequence identity). This cluster showed only

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