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Mineral-rich hardgrounds, such as ferromanganese (FeMn) crusts and phosphorites, occur on seamounts and continental margins, gaining attention for their resource potential due to their enrichment in valuable metals in some regions. This study focuses on the Southern California Borderland (SCB), an area characterized by uneven and heterogeneous topography featuring FeMn crusts, phosphorites, basalt, and sedimentary rocks that occur at varying depths and are exposed to a range of oxygen concentrations. Due to its heterogeneity, this region serves as an optimal setting for investigating the relationship between mineral-rich hardgrounds and benthic fauna. This study characterizes the density, diversity, and community composition of macrofauna (>300 μm) on hardgrounds as a function of substrate type and environment (depth and oxygen ranges). Rocks and their macrofauna were sampled quantitatively using remotely operated vehicles (ROVs) during expeditions in 2020 and 2021 at depths above, within, and below the oxygen minimum zone (OMZ). A total of 3,555 macrofauna individuals were counted and 416 different morphospecies (excluding encrusting bryozoans and hydrozoans) were identified from 82 rocks at depths between 231 and 2,688 m. Average density for SCB macrofauna was 11.08 ± 0.87 ind. 200 cm-2 and mean Shannon-Wiener diversity per rock (H[loge]) was 2.22 ± 0.07. A relationship was found between substrate type and macrofaunal communities. Phosphorite rocks had the highest H of the four substrates compared on a per-rock basis. However, when samples were pooled by substrate, FeMn crusts had the highest H and rarefaction diversity. Of all the environmental variables examined, water depth explained the largest variance in macrofaunal community composition. Macrofaunal density and diversity values were similar at sites within and outside the OMZ. This study is the first to analyze the macrofaunal communities of mineral-rich hardgrounds in the SCB, which support deep-ocean biodiversity by acting as specialized substrates for macrofaunal communities. Understanding the intricate relationships between macrofaunal assemblages and mineral-rich substrates may inform effects from environmental disruptions associated with deep-seabed mining or climate change. The findings contribute baseline information useful for effective conservation and management of the SCB and will support scientists in monitoring changes in these communities due to environmental disturbance or human impact in the future.

Aim: Shoaling of large oxygen minimum zones (OMZs) that form along eastern margins of the world's oceans can reduce habitat availability for some pelagic fishes. Our aim was to test the hypothesis that habitat compression caused by shoaling of the Pacific OMZ in tropical regions creates a boundary to the southern distribution of shortfin mako sharks (Isurus oxyrinchus) in the Eastern North Pacific Ocean. Location: Eastern North Pacific and Western North Atlantic oceans. Methods: We compared environmental conditions between areas used by satellite‐tagged mako sharks in the Eastern North Pacific, encompassing the world's largest OMZ, to those used in the Western North Atlantic where no OMZ is present. In the Pacific we quantified the effects of temperature and dissolved oxygen (DO) on depth use and tested if sharks spent less time in areas with strong habitat compression over the OMZ than expected by chance. Results: The southern distribution of sharks in the Pacific corresponded with the apex of OMZ shoaling in the North Equatorial Current. Sharks in the Atlantic occupied areas with warm surface temperatures (≥ 26°C) more often than the Pacific, and waters with these temperatures in the Atlantic had greater DO at depth. Sharks in the Pacific reduced time near the surface in warm temperatures and consistently avoided depths with low DO and spent less time in areas with strong habitat compression than expected by chance. Main Conclusions: The combination of warm surface temperatures and shoaling of the OMZ creates a soft boundary to mako shark movements in the Eastern North Pacific Ocean. The expected expansion of OMZs due to climate change could have considerable impact on future distribution of mako sharks and other pelagic fish. As such, development of species distribution models to predict the effects of climate change on pelagic fish distributions should incorporate oxygen availability. [ABSTRACT FROM AUTHOR]

Oxygen minimum zones (OMZ) are characterized by a dissolved oxygen (DO) concentration <0.5 ml L−1. In situ DO measurements suggest that the global OMZ upper limit is shallower than 60 years ago due to global warming. The benthic foraminifera assemblage, trace elements, and biological productivity and denitrification tracers have been used in paleoceanographic research, given that they indicate dysoxic and suboxic changing conditions of bottom water. Studies of benthic foraminifera assemblages in the southwestern margin of Baja California Sur are scarce. Thus, it is necessary to conduct a paleoecological analysis of the benthic foraminifera community on an approximately 1200-year timeline. The sediment core LONO09-MC02 (1200 years old) collected at 680 m depth was examined to analyze benthic foraminifera relative abundances and estimate the DO concentration in the OMZ of Baja California Sur's southwestern margin. The DO concentrations inferred from benthic foraminifera assemblages ranged between 0.09- and 0.17-ml l−1 (4.4 and 8.0 μM). This suggests that the OMZ has maintained dysoxic conditions over the past 1200 years. Moreover, the cluster and principal component analysis defined dysoxic, suboxic, and oxic assemblages. The decoupling between geochemical tracers was associated with biogeochemical cycles of the North Pacific subtropical gyre and varied in response to intermediate water circulation in the Pacific Ocean. The low estimate of dissolved oxygen concentration corresponded with minimal solar activity, favoring La Niña-like oceanographic conditions with high primary productivity. [ABSTRACT FROM AUTHOR]

Introduction: This study investigates the biogeochemical impact of Godavari River discharge (GRD) on the Bay of Bengal (BoB), focusing on the formation of an intense and shallow oxygen minimum zone (OMZ) near the river mouth during the fall season. Unlike the BoB's typical intermediate-depth OMZ, this subsurface (~40-200 m) phenomenon is attributed to the interplay of GRD-driven nutrient enrichment, coastal upwelling, enhanced productivity, and subsequent organic matter decomposition. Data and Methods: Our analysis using the Biogeochemical-Argo floats and World Ocean Atlas 2018 data reveals that a clear shoaling and intensification of the OMZ in the fall season. Further, a comparative analysis at two geographically distinct locations highlighted the pivotal role of GRD. Results, Discussion, and Implications: The location directly influenced by GRD exhibited significantly higher chlorophyll-a blooms, net primary production during the southwest monsoon, and pronounced oxygen consumption during the fall compared to the other. Our analysis suggests that GRD fuels primary productivity, leading to organic matter abundance and intense oxygen depletion in the subsurface layers, driving the observed shallow OMZ. Understanding the complex interplay between GRD, stratification, upwelling, and biogeochemical processes is crucial for predicting the impact of altered riverine inputs on coastal ecosystems, greenhouse gas emissions, and the overall health of the coastal BoB. [ABSTRACT FROM AUTHOR]

Organic-rich whale bones and wood falls occur widely in the deep sea and support diverse faunal communities, contributing to seafloor habitat diversity. Changes in community structure through succession on deep-sea bone/wood substrates are modulated by ecosystem engineers, i.e., bone-eating Osedax annelids, and wood-boring Xylophaga bivalves. Here, we use a comparative experimental approach and Ocean Networks Canada's (ONC) cabled observatory in hypoxic Barkley Canyon to study, at high temporal resolution, colonization and succession on whale-bone, Douglas fir wood, and control carbonate rock over 8.3 mo. Experimental substrates were similar in size and mounted on PVC plates near the seafloor at 890 m depth and monitored by high-definition video camera for 5-min intervals every 6-12 h over a period of 8.3 mo. A broad range of seafloor and sea-surface environmental variables were also monitored at this site over the 8.3 mo to account for environmental variability and food input. Following loss of the high-definition camera, substrates were surveyed approximately annually with lower resolution ROV video for an additional 8.5 y. We find that megafaunal abundances, species diversity, and community structure varied substantially over 8.3 mo on each substrate, with markedly different patterns on whale bones due to the development of extensive white bacterial mats. A combination of seafloor and sea surface variables explained < 35% of bone/wood community variation. Bone-eating Osedax annelids failed to colonize whale bones even after 9.2 years, and boring Xylophaga bivalves colonized the wood at much lower rates than in better oxygenated deep-sea locations. Species diversity on whale-bone and wood substrates appeared to be substantially reduced due to the absence of ecosystem engineers and the low oxygen concentrations. We hypothesize that Osedax/Xylophaga colonization, bone/wood degradation, and bone/wood community development may be limited by oxygen concentrations of 0.22 - 0.33 ml.l on the NE Pacific margin, and that OMZ expansion due to climate change will reduce whale-bone and wood degradation, and the contribution of whale falls and wood falls to beta diversity, on the NE Pacific margin. [ABSTRACT FROM AUTHOR]

Shallow-sea environmental variations in the eastern Arabian Sea (EAS) have been related to monsoon variability, poorly ventilated intermediate water, and primary productivity changes. To understand these conditions, benthic foraminiferal assemblages, oxygen isotope ratios, and total organic carbon content have been examined in sediment Core SK291/GC13 from the EAS. We aim to retrieve decadal to centennial time-scale changes in past oxygenation and paleoproductivity during the last 7400 calibrated years before the present (cal yr BP). Our proxy data suggest that the oxic benthic foraminifera species were dominant during ∼7400–6000 cal yr BP. The change from oxic to dysoxic occurred during ca. 6000–3500 cal yr BP, and transition from dysoxic to suboxic environmental conditions happened at ca. ∼3500 cal yr BP. These changes are contemporaneous with weakening of the Indian summer monsoon beginning at ∼5500 cal yr BP which culminated into a major arid phase so-called ca 4.2 ka event in the Indian subcontinent. The EAS experienced low productivity during the early phase of our studied interval (∼7400–6000 cal yr BP) and higher productivity in the middle Holocene (6000–3000 cal yr BP), as revealed by the total organic carbon and benthic assemblages. Furthermore, productivity increased in the warm period and decreased in the Little Ice Age on a decadal scale. The spectral analysis of total organic content and suboxic species indicates a 23-year Hale (solar) cycle during the last millennium, indicating a close relationship between solar activity and monsoon variability. The strengthening of the El Nino-Southern Oscillation perhaps amplified due to increased solar activity in the last millennium, which influenced the climate of the EAS. The study highlights paleo-oxygenation shifts based on benthic foraminiferal assemblages, which are crucial for understanding changes in the oxygen minimum zones (OMZs) and their potential impacts on marine ecosystems. [ABSTRACT FROM AUTHOR]

BACKGROUND: Despite a rising interest in the diversity and ecology of fungi in marine environments, there are few published genomes of fungi isolated from the ocean. The basidiomycetous yeast (unicellular fungus) genus Rhodotorula are prevalent and abundant in the open ocean, and they have been isolated from a wide range of other environments. Many of these environments are nutrient poor, such as the Antarctica and the Atacama deserts, raising the question as to how Rhodotorula yeasts may have adapted their metabolic strategies to optimize survival under low nutrient conditions. In order to understand their adaptive strategies in the ocean, the genome of R. sphaerocarpa ETNP2018 was compared to that of fourteen representative Rhodotorula yeasts, isolated from a variety of environments. RESULTS: Rhodotorula sphaerocarpa ETNP2018, a strain isolated from the oligotrophic part of the eastern tropical North Pacific (ETNP) oxygen minimum zone (OMZ), hosts the smallest of the fifteen genomes and yet the number of protein-coding genes it possesses is on par with the other strains. Its genome exhibits a distinct reduction in genes dedicated to Major Facilitator Superfamily transporters as well as biosynthetic enzymes. However, its core metabolic pathways are fully conserved. Our research indicates that the selective pressures of the ETNP OMZ favor a streamlined genome with reduced overall biosynthetic potential balanced by a stable set of core metabolisms and an expansion of mechanisms for nutrient acquisition. CONCLUSIONS: In summary, this study offers insights into the adaptation of fungi to the oligotrophic ocean and provides valuable information for understanding the ecological roles of fungi in the ocean.

Ocean deoxygenation is an important consequence of climate change that poses an imminent threat to marine life and global food security. However, our understanding of the complex interactions between changes in circulation, solubility, and respiration that drive global‐scale deoxygenation is incomplete. Here, we consider idealized biogeochemical steady states in equilibrium with perpetually slower and warmer oceans constructed from climate‐model simulations of the 2090s that we hold constant in time. In contrast to simulations of the end‐of‐century transient state, our idealized states are intensely deoxygenated in the abyss, consistent with perpetually reduced ventilation and throttled Antarctic Bottom Water formation. We disentangle the effects of the deoxygenation drivers on preformed oxygen and true oxygen utilization (TOU) using the novel concept of upstream exposure time, which precisely connects TOU to oxygen utilization rates and preformed oxygen to ventilation. For our idealized steady states, deoxygenation below 2,000 m depth is due to increased TOU, driven dominantly by slower circulations that allow respiration to act roughly 2–3 times longer thereby overwhelming the effects of reduced respiration rates. Above 500 m depth, decreased respiration and slower circulation closely compensate, resulting in little expansion of upper‐ocean hypoxia. The bulk of preformed oxygen loss is driven by ventilation shifting equatorward to where warmer surface waters hold less oxygen. Warming‐driven declines in solubility account for less than 10% of the total oxygen loss. Although idealized, our analysis suggests that long‐term changes in the marine oxygen cycle could be driven dominantly by changes in circulation rather than by thermodynamics or biology. Plain Language Summary: Climate change is driving oxygen out of the ocean, threatening marine life and global food security. However, the precise contributions of the chemical, physical, and biological processes that control oxygen levels are not well known because of their complex interactions. To better understand these interactions, we consider idealized simulations of a global oxygen cycle that is fully equilibrated with a perpetually warmer and slower ocean constructed from climate‐model simulations of the 2090s but held constant in time for our analyses. Compared to typical predictions, these idealized states exhibit intense deep‐ocean deoxygenation, for which we precisely quantify the contributions from changes in solubility, respiration, and ocean circulation. We find that deep‐ocean deoxygenation is driven by the slower circulation allowing respiration to act for 2–3 times longer thereby overcoming lower respiration rates. The surface origin of oxygen shifts away from cold high‐latitude waters toward warmer waters, in which atmospheric oxygen is less soluble, further reducing oxygen levels. Warming‐driven decreases in solubility alone only account for a mere 10% of the total oxygen loss. The upper ocean remains well oxygenated because changes in respiration and circulation compensate almost perfectly. Our results highlight the central importance of circulation in controlling oxygen in the ocean. Key Points: Key drivers of deoxygenation are quantified for oxygen cycles idealized by being in equilibrium with perpetually slower oceansWidespread intense abyssal ocean deoxygenation is driven predominantly by slower circulations allowing respiration to act over longer timesMost of the reduction in preformed oxygen is driven by changes in ventilation patterns and not by warming‐driven reduced solubility [ABSTRACT FROM AUTHOR]

Vertical migrants are a diverse group of organisms, which includes crustaceans, cephalopods and mesopelagic fishes. They play an active role in the biogeochemical cycles but are in general not included in numerical models. In this study we introduce a fully coupled Earth system model that represents vertical migration and with this resolves the key components of the mesopelagic ecosystem, namely migrating zooplankton and mesopelagic fish, including their feedbacks on biogeochemical cycles. The redistribution of nutrients in the water column by vertical migration results in a reduction of the net primary production of 14%–21%, as well as in an asymmetric response in the low oxygenated waters in the tropical Pacific (an increase in the northern and a decrease in the southern oxygen minimum zone). On a global scale, we find the active transport of carbon out of the surface layer to be equivalent to ∼25% of the total export (∼30% relative to passive sinking). In the low latitudes, migration results regionally in a reduction of the shallow export by 2%–10% and an increase of the deep carbon export by 6%–15%. In our simulations, mesopelagic fish, with a biomass of 3–3.4 Gt wet weight, have a slightly larger impact on active carbon flux than migrating zooplankton. Key Points: We present a global model that represents the impact of mesopelagic fish and vertically migrating organisms on the biogeochemical cyclesThe simulated redistribution of nutrients by vertical migration results in a reduction of the global net primary production by 14%–21% compared to the simulation without vertical migrationThe active carbon export out of the surface layers by vertical migration amounts to ∼25% of the total transport (∼30% of the passive sinking) [ABSTRACT FROM AUTHOR]

IntroductionThis study investigates the biogeochemical impact of Godavari River discharge (GRD) on the Bay of Bengal (BoB), focusing on the formation of an intense and shallow oxygen minimum zone (OMZ) near the river mouth during the fall season. Unlike the BoB’s typical intermediate-depth OMZ, this subsurface (~40-200 m) phenomenon is attributed to the interplay of GRD-driven nutrient enrichment, coastal upwelling, enhanced productivity, and subsequent organic matter decomposition.Data and MethodsOur analysis using the Biogeochemical-Argo floats and World Ocean Atlas 2018 data reveals that a clear shoaling and intensification of the OMZ in the fall season. Further, a comparative analysis at two geographically distinct locations highlighted the pivotal role of GRD.Results, Discussion, and ImplicationsThe location directly influenced by GRD exhibited significantly higher chlorophyll-a blooms, net primary production during the southwest monsoon, and pronounced oxygen consumption during the fall compared to the other. Our analysis suggests that GRD fuels primary productivity, leading to organic matter abundance and intense oxygen depletion in the subsurface layers, driving the observed shallow OMZ. Understanding the complex interplay between GRD, stratification, upwelling, and biogeochemical processes is crucial for predicting the impact of altered riverine inputs on coastal ecosystems, greenhouse gas emissions, and the overall health of the coastal BoB.

Mineral-rich hardgrounds, such as ferromanganese (FeMn) crusts and phosphorites, occur on seamounts and continental margins, gaining attention for their resource potential due to their enrichment in valuable metals in some regions. This study focuses on the Southern California Borderland (SCB), an area characterized by uneven and heterogeneous topography featuring FeMn crusts, phosphorites, basalt, and sedimentary rocks that occur at varying depths and are exposed to a range of oxygen concentrations. Due to its heterogeneity, this region serves as an optimal setting for investigating the relationship between mineral-rich hardgrounds and benthic fauna. This study characterizes the density, diversity, and community composition of macrofauna (>300 μm) on hardgrounds as a function of substrate type and environment (depth and oxygen ranges). Rocks and their macrofauna were sampled quantitatively using remotely operated vehicles (ROVs) during expeditions in 2020 and 2021 at depths above, within, and below the oxygen minimum zone (OMZ). A total of 3,555 macrofauna individuals were counted and 416 different morphospecies (excluding encrusting bryozoans and hydrozoans) were identified from 82 rocks at depths between 231 and 2,688 m. Average density for SCB macrofauna was 11.08 ± 0.87 ind. 200 cm−2 and mean Shannon-Wiener diversity per rock (H′[loge]) was 2.22 ± 0.07. A relationship was found between substrate type and macrofaunal communities. Phosphorite rocks had the highest H′ of the four substrates compared on a per-rock basis. However, when samples were pooled by substrate, FeMn crusts had the highest H′ and rarefaction diversity. Of all the environmental variables examined, water depth explained the largest variance in macrofaunal community composition. Macrofaunal density and diversity values were similar at sites within and outside the OMZ. This study is the first to analyze the macrofaunal communities of mineral-rich hardgrounds in the SCB, which support deep-ocean biodiversity by acting as specialized substrates for macrofaunal communities. Understanding the intricate relationships between macrofaunal assemblages and mineral-rich substrates may inform effects from environmental disruptions associated with deep-seabed mining or climate change. The findings contribute baseline information useful for effective conservation and management of the SCB and will support scientists in monitoring changes in these communities due to environmental disturbance or human impact in the future.

The Arabian Sea (AS) hosts the world's thickest and most intense oxygen minimum zone (OMZ), and previous studies have documented a dramatic decline of dissolved oxygen (DO) in the northeastern AS in recent decades. In this study, using the recently released data from Biogeochemical‐Argo floats, we found a surprising trend of recovery in deoxygenation within the core region of the OMZ in the AS (ASOMZ) since 2013. The average DO concentration increased by approximately threefold, from ∼0.63 μM in 2013 to ∼1.68 μM in 2022, and the thickness of the ASOMZ decreased by 13%. We find that the weakening of Oman upwelling resulting from the weakening of the summer monsoon is the main driver of oxygenation in the ASOMZ. In addition, the reduction of primary production linked to warming‐driven stratification reinforces deoxygenation recovery at depth. Plain Language Summary: The deoxygenation of the ocean is one of the most important changes occurring in the marine environment, impacting marine biodiversity, primary production, and carbon and nitrogen biogeochemical cycles. Ocean deoxygenation can cause the expansion of naturally occurring low dissolved oxygen (DO) water bodies known as OMZs. As the world's thickest and most intense oxygen minimum zone (OMZ), the Arabian Sea (AS) has undergone a dramatic decrease in DO, particularly in the northeastern AS in recent decades. Based on an unprecedented collection of DO from biogechemical‐Argo floats over the past decade, we found a surprising trend of recovery in deoxygenation within the core region of the OMZ in the AS (ASOMZ) since 2013. The average DO concentration in the ASOMZ increased by almost thrice, and the thickness decreased by 13%. Weakening of the summer monsoon over the AS and strengthening of stratification induced by global warming are possible causes for the recovery from deoxygenation. OMZs have major impacts on global carbon and nitrogen cycles. Our findings not only overturn the previous view of the deoxygenation trend in the ASOMZ but also provide valuable insights for projection model studies of global OMZs. Key Points: Deoxygenation in the oxygen minimum zone in the Arabian Sea has been recovered during the past decadeThe decrease in Oman upwelling due to the weakening of the summer monsoon is likely the main mechanism for the increase in dissolved oxygenThe strengthening of stratification in the AS leading to a decrease in production also contributed to the recovery of deoxygenation [ABSTRACT FROM AUTHOR]

Nitrous oxide (N2O) is a potent greenhouse gas and a major cause of ozone depletion. One-third of atmospheric N2O originates in aquatic environments. Reduction of N2O to dinitrogen gas (N2) requires the nitrous oxide reductase enzyme, which is encoded by the gene nosZ. Organisms that contain nosZ are the only known biological sinks of N2O and are found in diverse genera and a wide range of environments. The two clades of nosZ (Clade I and II) contain great diversity, making it challenging to study the population structure and distribution of nosZ containing organisms in the environment. A database of over 11,000 nosZ sequences was compiled from NCBI (representing diverse aquatic environments) and unpublished sequences and metagenomes (primarily from oxygen minimum zones, OMZs, where N2O levels are often elevated). Sequences were clustered into archetypes based on DNA and amino acid sequence identity and their clade, phylogeny, and environmental source were determined. Further analysis of the source and environmental distribution of the sequences showed strong habitat separation between clades and phylogeny. Although there are more Clade I nosZ genes in the compilation, Clade II is more diverse phylogenetically and has a wider distribution across environmental sources. On the other hand, Clade I nosZ genes are predominately found within marine sediment and are primarily from the phylum Pseudonomonadota. The majority of the sequences analyzed from marine OMZs represented distinct phylotypes between dierent OMZs showing that the nosZ gene displays regional and environmental separation. This study expands the known diversity of nosZ genes and provides a clearer picture of how the clades and phylogeny of nosZ organisms are distributed across diverse environments [ABSTRACT FROM AUTHOR]

The diversity of organisms inhabiting deep-sea ecosystems, such as seamounts, has hitherto remained under-studied. In this study, we report on the faunal diversity in the water column adjacent to the summit and periphery of two seamounts (SMS2 and SMS3) and an oxygen minimum zone site located away from the seamounts in the southeast Arabian Sea. Environmental DNA (eDNA) in the water column was metabarcoded using the Cytochrome Oxidase C subunit I (COI) gene marker and Oxford Nanopore sequencing technology. Hydrographic conditions revealed that the summits of the seamounts intersect with the core oxygen minimum zone between depths of 300 and 600 m. Comparisons of COI gene sequences with those in available databases, MIDORI and BOLD, indicated the existence of a diverse group of novel organisms in the study area. Crustaceans dominated (75–95%) in the summit and periphery of the SMS2 and the OMZ site, while Cnidaria (56–63%) and Chordata (55%), respectively, dominated the summit and periphery of SMS3. Overall, the current study highlights the broad diversity of organisms living in the water column around the seamounts and underscores the potential of eDNA for exploring them. [ABSTRACT FROM AUTHOR]

The semi-permanent coastal upwelling system within the Humboldt Current System (HCS) off northern Chile provides nutrient-rich, oxygen-poor water to the euphotic zone that contributes to high levels of primary production and restricts the vertical distribution of most zooplankton species according to their tolerance to hypoxia within this highly heterogeneous environment. The day/night vertical distributions of two calanoid copepods, Eucalanus inermis and Nannocalanus minor, were studied concerning the presence of oxygen-poor water associated with the Oxygen minimum zone (OMZ) at a coastal and an offshore station off northern Chile (~20°S), by stratified sampling from the surface to a depth of 600 m. At the coastal station, most of the population of both species remained above the OMZ, whereas at the offshore station, most populations remained at the edge of the OMZ. Both species exhibited reverse diel vertical migration between day and night and were positively correlated with dissolved oxygen and chlorophyll-a, suggesting that both species may prefer oxygenated waters and avoid the OMZ. However, E. inermis and N. minor exhibited distinct behavior, with E. inermis displaying significant variations among stations and strata and N. minor showing differences only among strata, particularly in the upper strata where it was more aggregated. The results helped us understand the behavior and distribution of zooplankton related to OMZ and how they may change as ocean temperature increases worldwide. [ABSTRACT FROM AUTHOR]

Marine dinitrogen (N2) fixation fuels primary production and thereby influences the Earth's climate. Yet, its geographical distribution and controlling environmental parameters remain debatable. We measured N2 fixation rates from the two spatially and physicochemically contrasting regions of the Arabian Sea during the winter monsoon: (a) the colder and nutrient‐rich waters in the northern region owing to winter convection and (b) the warmer and nutrient‐poor waters in the southern region unaffected by winter convection. We found higher N2 fixation rates at the surface of northern region due to convective mixing driven supply of phosphate (intuitively iron also) from the underlying suboxic waters. N2 fixation was favored by high nutrient concentrations in the euphotic waters, whereas remained unaffected by nutrient availability in the aphotic waters. We conclude that diazotrophs dwelling in the euphotic zone chose phosphate over fixed nitrogen‐poor waters. However, we found that among oligotrophic waters, anticyclonic eddy extremes the barrier of fixed nitrogen supply, and thereby, elevates N2 fixation. While the Arabian Sea loses about 20%–40% of the global ocean fixed nitrogen, we estimate that N2 fixation in the Arabian Sea offsets only up to 42% of its fixed nitrogen‐loss by denitrification, but this offset could be higher if diazotrophic activity is further examined up to the deeper depths of the Arabian Sea. Plain Language Summary: Phytoplankton facilitate the removal of greenhouse carbon dioxide gas from the atmosphere. However, phytoplankton's carbon dioxide assimilation ability is largely controlled by the reactive forms of nitrogen (N, such as ammonium, nitrate and nitrite). In contrast, the N‐limited environments often create a suitable niche for diazotrophs—organisms genetically proficient to convert dinitrogen (N2) gas to a reactive form of N. Previous research has hypothesized close spatial coupling between N2 fixation and N‐loss processes because the latter result in an N‐deficit, that is, a conducive condition for diazotrophs. Here, we measured N2 fixation rates in and above the N‐loss dominated waters of the Arabian Sea. The highest N2 fixation occurred near the coast of the Arabian Sea. We found relatively high N2 fixation rates above the waters with the highest N‐loss owing to convective mixing driven supply of nutrients such as phosphate and probably iron. N2 fixation rates below the sunlit layer were consistent but low. Our study showed that physical processes substantially influence N2 fixation by affecting nutrients supply. Key Points: Winter convection elevates N2 fixation through the enhanced supply of nutrients from the suboxic waters to the surface waters of the Arabian SeaDiazotrophy offsets up to 42% of the Arabian Sea's denitrification deficitWinter convective mixing strengthens the long‐held hypothesis of coupling between N‐gain and ‐loss processes [ABSTRACT FROM AUTHOR]

Organic-rich whale bones and wood falls occur widely in the deep sea and support diverse faunal communities, contributing to seafloor habitat diversity. Changes in community structure through succession on deep-sea bone/wood substrates are modulated by ecosystem engineers, i.e., bone-eating Osedax annelids, and wood-boring Xylophaga bivalves. Here, we use a comparative experimental approach and Ocean Networks Canada’s (ONC) cabled observatory in hypoxic Barkley Canyon to study, at high temporal resolution, colonization and succession on whale-bone, Douglas fir wood, and control carbonate rock over 8.3 mo. Experimental substrates were similar in size and mounted on PVC plates near the seafloor at 890 m depth and monitored by high-definition video camera for 5-min intervals every 6-12 h over a period of 8.3 mo. A broad range of seafloor and sea-surface environmental variables were also monitored at this site over the 8.3 mo to account for environmental variability and food input. Following loss of the high-definition camera, substrates were surveyed approximately annually with lower resolution ROV video for an additional 8.5 y. We find that megafaunal abundances, species diversity, and community structure varied substantially over 8.3 mo on each substrate, with markedly different patterns on whale bones due to the development of extensive white bacterial mats. A combination of seafloor and sea surface variables explained < 35% of bone/wood community variation. Bone-eating Osedax annelids failed to colonize whale bones even after 9.2 years, and boring Xylophaga bivalves colonized the wood at much lower rates than in better oxygenated deep-sea locations. Species diversity on whale-bone and wood substrates appeared to be substantially reduced due to the absence of ecosystem engineers and the low oxygen concentrations. We hypothesize that Osedax/Xylophaga colonization, bone/wood degradation, and bone/wood community development may be limited by oxygen concentrations of 0.22 - 0.33 ml.l on the NE Pacific margin, and that OMZ expansion due to climate change will reduce whale-bone and wood degradation, and the contribution of whale falls and wood falls to beta diversity, on the NE Pacific margin.

This study reports on the fine-scale movements of swordfish (Xiphias gladius) tagged off of the Antofagasta region of northern Chile, where harpoon fishing operations date back more than seven millenia. The study was conducted within a highly productive swordfish foraging area where movements and habitat use remain poorly characterized. Swordfish were outfitted with both data storage and satellite-linked archival tags scheduled for short (<30 days) and longer-term (>100 days) data collection. All tags were deployed on basking swordfish using harpoon-based methods across two field seasons in March 2019 and April 2022 around oceanic fronts and offshore seamounts. Fine-scale depth and temperature data were recorded over 145 days from four adult swordfish (100-200 kg). Swordfish exhibited regionally unique diurnal behaviors with greater average daytime depths than those recorded at night (78.0 ± 66.5 and 12.7 ± 4.9 m, respectively). The average daytime depths were shallower than those previously documented for other Pacific regions, including Chile's offshore waters. Daytime basking was observed on 76% of the daily records (110 out of 145 days) and accounted for approximately 12% of the collective daylight hours. Findings align with previous swordfish tagging studies and suggest that depth distribution patterns vary considerably depending on regional oceanographic conditions. Movement data are compared with previous studies, and we discussed relative to regional fishing operations. [ABSTRACT FROM AUTHOR]

Continuous understanding of the ongoing ocean acidification (OA) is essential for predicting the future impact of OA on marine ecosystems. Here we report the results of open ocean time-series measurements (19 cruises) of seawater pH in total hydrogen ion scale (pHT) and associated parameters in the Arabian Sea (AS) and the Bay of Bengal (BoB). During southwest monsoon (SWM), the pHT within the 30 to 100 m water column shows the maximum difference between the two basins with BoB pHT being lower (up to ~0.39 units) than AS which could be due to freshwater influx from rivers, mixed layer dynamics, and cold-core eddies. However, during Spring inter-monsoon (SIM), the pHT of BoB follows the trend of AS. A contrasting finding is that the lowest pHT occurs at 350 to 500 m in the BoB while it is ~1000 m in the AS. The pHT within the 150 to 1500 m layer of these two basins shows lower values by 0.03 (±0.02) in the BoB as compared to the AS. The possible reasons for the low pHT within the BoB oxygen minimum zone (OMZ) could be due to intrusion of western Pacific water in the BoB, freshwater influx from rivers, variations in OMZ of the two basins, higher temperature (~2°C) within the OMZ of the AS, and denitrification in the AS. The pHT in both the basins (500 to 1000 m) is lower than in the North Atlantic and higher than in the North Pacific waters; however, the pHT in the 200 to 500 m is lower in the BoB than in all these basins. This study highlights the under-saturation of calcium carbonate at very shallow depths (~ 100 m) in the BoB, indicating that the plankton in the BoB are facing a major risk from OA compared to the AS and need further investigation. [ABSTRACT FROM AUTHOR]

The distribution of dissolved oxygen (DO) affects the survival of marine organisms. In order to explore the temporal and spatial variation of sea surface DO and oxycline in the Oxygen Minimum Zone (OMZ) of Bay of Bengal (BoB), we investigated the vertical DO distribution from 0 to 200 m in the central BoB using Bio-Argo and remote sensing data from 2013 to 2017. The results showed that the salinity stratification caused by the input of a large amount of runoff and rainfall induced the significant change of the DO in the central BoB, and DO decreased significantly at ~40 m, and dropped to less than 20 μmol·kg-1 within 200 m. The sea surface DO was negatively correlated with the sea surface temperature (r=-0.75). The oxycline depth was significantly correlated with the 23 isotherm and sea level anoma ℃ ly (r=0.93 and 0.81). The surface DO concentration was mainly affected by the surface temperature, and the oxycline depth was mainly controlled by the mesoscale eddies in the OMZ of the BoB. [ABSTRACT FROM AUTHOR]

Abstract The Arabian Sea (AS) hosts the world's thickest and most intense oxygen minimum zone (OMZ), and previous studies have documented a dramatic decline of dissolved oxygen (DO) in the northeastern AS in recent decades. In this study, using the recently released data from Biogeochemical‐Argo floats, we found a surprising trend of recovery in deoxygenation within the core region of the OMZ in the AS (ASOMZ) since 2013. The average DO concentration increased by approximately threefold, from ∼0.63 μM in 2013 to ∼1.68 μM in 2022, and the thickness of the ASOMZ decreased by 13%. We find that the weakening of Oman upwelling resulting from the weakening of the summer monsoon is the main driver of oxygenation in the ASOMZ. In addition, the reduction of primary production linked to warming‐driven stratification reinforces deoxygenation recovery at depth.

Nitrous oxide (N2O) is a potent greenhouse gas and a major cause of ozone depletion. One-third of atmospheric N2O originates in aquatic environments. Reduction of N2O to dinitrogen gas (N2) requires the nitrous oxide reductase enzyme, which is encoded by the gene nosZ. Organisms that contain nosZ are the only known biological sinks of N2O and are found in diverse genera and a wide range of environments. The two clades of nosZ (Clade I and II) contain great diversity, making it challenging to study the population structure and distribution of nosZ containing organisms in the environment. A database of over 11,000 nosZ sequences was compiled from NCBI (representing diverse aquatic environments) and unpublished sequences and metagenomes (primarily from oxygen minimum zones, OMZs, where N2O levels are often elevated). Sequences were clustered into archetypes based on DNA and amino acid sequence identity and their clade, phylogeny, and environmental source were determined. Further analysis of the source and environmental distribution of the sequences showed strong habitat separation between clades and phylogeny. Although there are more Clade I nosZ genes in the compilation, Clade II is more diverse phylogenetically and has a wider distribution across environmental sources. On the other hand, Clade I nosZ genes are predominately found within marine sediment and are primarily from the phylum Pseudonomonadota. The majority of the sequences analyzed from marine OMZs represented distinct phylotypes between different OMZs showing that the nosZ gene displays regional and environmental separation. This study expands the known diversity of nosZ genes and provides a clearer picture of how the clades and phylogeny of nosZ organisms are distributed across diverse environments.

Abstract Background Despite a rising interest in the diversity and ecology of fungi in marine environments, there are few published genomes of fungi isolated from the ocean. The basidiomycetous yeast (unicellular fungus) genus Rhodotorula are prevalent and abundant in the open ocean, and they have been isolated from a wide range of other environments. Many of these environments are nutrient poor, such as the Antarctica and the Atacama deserts, raising the question as to how Rhodotorula yeasts may have adapted their metabolic strategies to optimize survival under low nutrient conditions. In order to understand their adaptive strategies in the ocean, the genome of R. sphaerocarpa ETNP2018 was compared to that of fourteen representative Rhodotorula yeasts, isolated from a variety of environments. Results Rhodotorula sphaerocarpa ETNP2018, a strain isolated from the oligotrophic part of the eastern tropical North Pacific (ETNP) oxygen minimum zone (OMZ), hosts the smallest of the fifteen genomes and yet the number of protein-coding genes it possesses is on par with the other strains. Its genome exhibits a distinct reduction in genes dedicated to Major Facilitator Superfamily transporters as well as biosynthetic enzymes. However, its core metabolic pathways are fully conserved. Our research indicates that the selective pressures of the ETNP OMZ favor a streamlined genome with reduced overall biosynthetic potential balanced by a stable set of core metabolisms and an expansion of mechanisms for nutrient acquisition. Conclusions In summary, this study offers insights into the adaptation of fungi to the oligotrophic ocean and provides valuable information for understanding the ecological roles of fungi in the ocean.

Niche partitioning among closely related, sympatric species is a fundamental concept in ecology, and its mechanisms are of broad interest for understanding ecosystem functioning and predicting the impacts of human-driven environmental change. However, identifying mechanisms by which top marine predators partition available resources has been especially challenging given the difficulty of quantifying resource use of large pelagic animals. In the eastern tropical Pacific (ETP), three large, highly mobile and ecologically similar pelagic predators (blue marlin (Makaira nigricans), black marlin (Istiompax indica) and sailfish (Istiophorus platypterus)) coexist in a vertically compressed habitat. To evaluate each species' ecological niche, we leveraged a decade of recreational fisheries data, multi-year satellite tracking with high-resolution dive data, and stable isotope analysis. Fishery interaction and telemetry-based three-dimensional seasonal utilization distributions suggested high spatial and temporal overlap among species; however, seasonal and diel variability in diving behaviour produced spatial partitioning, leading to low trophic overlap among species. Expanding oxygen minimum zones will reduce the available vertical habitat within predator guilds, likely leading to increases in interspecific competition. Thus, understanding the mechanisms of habitat partitioning among predators in the vertically compressed ETP can provide insight into how predators in other ocean regions may respond to vertically limited habitats. [ABSTRACT FROM AUTHOR]

Tropical pelagic predators are exploited by fisheries and their movements are influenced by factors including prey availability, temperature, and dissolved oxygen levels. As the biophysical parameters vary greatly within the range of circumtropical species, local studies are needed to define those species' habitat preference and model possible behavioral responses under different climate change scenarios. Here, we tagged yellowfin tuna Thunnus albacares in the Galápagos Marine Reserve and tracked the horizontal and vertical movements of eight individuals for 4–97 days. The tuna traveled a mean of 13.6 km day−1 horizontally and dispersed throughout the archipelago and in offshore waters inside the Galápagos Marine Reserve and in the surrounding Ecuadorian exclusive economic zone. Vertically, they traveled a mean of 2 km day−1, although high‐resolution data from a recovered tag suggested that transmitted data underestimated their vertical movement by a factor of 5.5. The tracked yellowfin tuna spent most of their time near the surface, with an overall mean swimming depth of 24.3 ± 46.6 m, and stayed shallower at night (11.1 ± 16.3 m) than during the day (37.7 ± 60.9 m), but on occasion dived to cold, oxygen‐poor waters below 200 m. Deep dives were commonly made during the day with a mean recovery period of 51 min between exposures to modeled oxygen‐limiting conditions <1.5 mL L−1, presumably to re‐oxygenate. The depth and frequency of dives were likely limited by dissolved oxygen levels, as oxygen‐depleted conditions reach shallow depths in this region. The main habitat of tracked yellowfin tunas was in the shallow mixed layer, which may leave them vulnerable to fishing. Vertical expansion of low‐oxygen waters under future climate change scenarios may further compress their habitat, increasing their vulnerability to surface fishing gear. [ABSTRACT FROM AUTHOR]

The tectonic closure of the Central American Seaway (CAS) during the mid‐Miocene to mid‐Pliocene (∼16–3 Ma BP) is thought of as a key interval for the onset of the present‐day tropical Pacific oxygen minimum zone (OMZ). In this study we investigate the impact of an open CAS on the ocean circulation and the OMZ in the tropical Pacific. We perform a series of sensitivity experiments with the Kiel Climate Model, where we vary the CAS sill depth from shallow to deep. We find that the eastern tropical Pacific OMZ was less developed during the period of an open CAS. This is driven mainly by an enhanced eastward subsurface current that facilitated an increased oxygen supply from the western tropical Pacific. In addition, a small decrease in net marine primary production and subsequent weaker export of particulate organic carbon induced less subsurface oxygen consumption in the Eastern Equatorial Pacific. Plain Language Summary: Oxygen plays a fundamental role for aerobic life and cycling of biogeochemical properties in the ocean. When low rates of oxygen supply combine with high rates of biological oxygen consumption, oxygen minimum zones (OMZs) are generally formed in the ocean. Today the world's largest OMZ is found in the eastern tropical Pacific where the constriction and final closure of the Central American Seaway (CAS) from 16 to 3 Ma BP created specific characteristics of ocean circulation and biogeochemistry. Using simulations with a climate and a marine biogeochemistry model we investigate whether the CAS closure led to the emergence of today's tropical Pacific OMZ. We find that the eastern tropical Pacific OMZ was less developed and oxygen levels were overall higher in the subsurface Eastern Equatorial Pacific (EEP) when the CAS was open. This was driven mainly by an enhanced eastward subsurface current from the northwestern tropical Pacific. In addition, a small decrease in net marine primary production induced less subsurface oxygen consumption from remineralization of particulate organic matter in the EEP. Key Points: An open Central American Seaway promotes higher oxygen concentrations in the subsurface Pacific waters, with strongest changes in the EEPAn increased eastward subsurface flow in the northeastern tropical Pacific transports young, oxygen‐rich waters in from the western PacificThe improved EEP ventilation is a result of the ocean current redirection in the North and South Pacific during open Central American Seaway [ABSTRACT FROM AUTHOR]

Macrobenthic samples were collected offshore Namibia on board R/V Mirabilis during the 3rd RGNO training and the National Marine Information and Research Centre's (NatMIRC's) plankton survey from 13th May to 17th May 2016. Two transects, Cape Frio (20º S) and Walvis Bay (23º S), hosted three stations each, while the third transect, Luderitz (26º S), hosted only one station. From the results, three oxygen zones were identified, namely Microxic (<0.1 ml l-1), Dysoxic (0.1-1.0 ml l-1), and Oxic (>1.0 ml l-1). A total of 20 Macrobenthic taxa were identified; Nematoda, Polychaeta, Cumacea, and Oligochaeta were the most dominant taxa and recorded abundances in all oxygen zones. Eighteen genera of macro-Nematoda were identified; Desmolaimus and Paracomesoma dominated in all oxygen zones, Metoncholaimus recorded higher abundance in the Dysoxic stations, and no abundance in the Oxic stations, and the opposite was observed for Halanonchus and Dorylaimopsis. H' Diversity values for both the general Macrofauna and Macro- Nematoda were higher in the Oxic stations and lower in both the Dysoxic and Microxic stations, while an opposite trend was observed for Dominance values. Density values were lower in Microxic stations and higher in Dysoxic stations, while at the Oxic stations, the density values fell in between the two hypoxic zones. [ABSTRACT FROM AUTHOR]

Physiological traits of five key fish species (Cape hake Merluccius spp., pelagic goby Sufflogobius bibarbatus, Cape horse mackerel Trachurus capensis, European anchovy Engraulis encrasicolus and sardine Sardinops sagax) from the northern Benguela upwelling system (NBUS) were compared during their larval stage by measurements of enzymatic activities of key metabolic enzymes (citrate synthase [CS] and pyruvate kinase [PK]). Two distinct age classes (early larvae: 8‒14 days old; late larvae: 22‒29 days old) for each species and from two areas were compared: Terrace Bay (20° S), the main spawning ground of Cape horse mackerel and anchovy, and Kunene (17° S), near the Angola–Benguela frontal zone, where warm and hypoxic water masses intrude into the NBUS. The results revealed significantly higher CS activity in both larval age classes in Cape horse mackerel, anchovy and sardine compared with Cape hake and pelagic goby. Pelagic goby and Cape horse mackerel had significantly higher PK activity compared with Cape hake, anchovy and sardine, apparent in both larval age classes and both areas. Results for anaerobic metabolism indicate higher capacity for pelagic goby and Cape horse mackerel to recover from oxygen debt built up in prey capture and predator escape behaviour and a higher potential for hypoxia tolerance when compared with Cape hake, anchovy and sardine. These results suggest higher survival probability for pelagic goby and Cape horse mackerel compared with the other species under conditions of a shoaling oxygen minimum zone and may explain their current dominance in the NBUS. [ABSTRACT FROM AUTHOR]

Climate change is expected to increase the strength of ocean Oxygen Deficient Zones (ODZs), but we lack a detailed understanding of the temporal or spatial variability of these ODZs. A 50‐year time series in the Eastern Tropical North Pacific (ETNP) ODZ revealed that it has strengthened by 30% from 1994 to 2019. We subdivided the ODZ into a core and a deep layer based on potential density and revealed that different processes control the magnitude of fixed nitrogen loss between these regions. We postulate that the depth of the upper ETNP ODZ water mass, the 13°C Water, influences the organic carbon supply to the core ODZ and therefore its strength. We correlated the maximum fixed nitrogen loss in the core ODZ with a nearby sedimentary nitrogen isotope record and found that this recent increase in the magnitude of fixed nitrogen loss occurred only a few times over the last 1,200 years. Using this correlation, we derived the first confidence interval for the natural variability of the maximum fixed nitrogen loss within the ETNP ODZ, which has a range of 3.3 μmol kg−1 (p = 0.01). While the current increase is only comparable to two previous events, it is within the confidence interval for natural variability (p = 0.03). The deep ODZ also strengthened from 2016 to 2019 by approximately 30%, but this increase occurred more rapidly than the core ODZ, and this dramatic increase was not observed over the rest of the 40 years. Climate‐driven intensification could lead to unprecedented changes in the ETNP ODZ within the next decade. Plain Language Summary: Naturally occurring ocean Oxygen Deficient Zones (ODZs) play a significant role in regulating ocean nutrient availability and ecosystem structure. Climate change is expected to strengthen these ODZs. A 50‐year time series in the world's largest ODZ revealed that it has strengthened by 30% from 1994 to 2019. Based on sedimentary records, this rapid increase has only occurred twice over the last 1,200 years. Although this sedimentary record suggests there is high but not extremely certain likelihood that climate change caused this recent intensification, continued strengthening at these rates could lead to unprecedented changes within the next decade. Key Points: The nitrogen deficit in the largest Oxygen Deficient Zone (ODZ) has increased over 50 years, indicating a 30% increase in ODZ strengthThe maximum fixed N loss of the core ODZ (σθ 26.2–26.8 kg m−3) is significantly different (p = 0.03) from its natural variabilityThough the core ODZ intensification is significant, we cannot yet claim with virtual certainty (p = 0.01) that climate change caused it [ABSTRACT FROM AUTHOR]

This correction notice addresses errors in affiliations in the article "Seasonal intensification of oxygen minimum zone: linking Godavari River discharge to fall hypoxia in the Bay of Bengal." The corrections involve the affiliations of two authors, with the correct affiliations being in Goa, India, and Thuwal, Saudi Arabia. The authors acknowledge the errors and clarify that they do not impact the scientific conclusions of the article. The original article has been updated to reflect these corrections. [Extracted from the article]

The diversity of organisms inhabiting deep-sea ecosystems, such as seamounts, has hitherto remained under-studied. In this study, we report on the faunal diversity in the water column adjacent to the summit and periphery of two seamounts (SMS2 and SMS3) and an oxygen minimum zone site located away from the seamounts in the southeast Arabian Sea. Environmental DNA (eDNA) in the water column was metabarcoded using the Cytochrome Oxidase C subunit I (COI) gene marker and Oxford Nanopore sequencing technology. Hydrographic conditions revealed that the summits of the seamounts intersect with the core oxygen minimum zone between depths of 300 and 600 m. Comparisons of COI gene sequences with those in available databases, MIDORI and BOLD, indicated the existence of a diverse group of novel organisms in the study area. Crustaceans dominated (75–95%) in the summit and periphery of the SMS2 and the OMZ site, while Cnidaria (56–63%) and Chordata (55%), respectively, dominated the summit and periphery of SMS3. Overall, the current study highlights the broad diversity of organisms living in the water column around the seamounts and underscores the potential of eDNA for exploring them.

Abstract The tectonic closure of the Central American Seaway (CAS) during the mid‐Miocene to mid‐Pliocene (∼16–3 Ma BP) is thought of as a key interval for the onset of the present‐day tropical Pacific oxygen minimum zone (OMZ). In this study we investigate the impact of an open CAS on the ocean circulation and the OMZ in the tropical Pacific. We perform a series of sensitivity experiments with the Kiel Climate Model, where we vary the CAS sill depth from shallow to deep. We find that the eastern tropical Pacific OMZ was less developed during the period of an open CAS. This is driven mainly by an enhanced eastward subsurface current that facilitated an increased oxygen supply from the western tropical Pacific. In addition, a small decrease in net marine primary production and subsequent weaker export of particulate organic carbon induced less subsurface oxygen consumption in the Eastern Equatorial Pacific.

Biological dinitrogen (N2) fixation is meagerly explored in the Bay of Bengal (Bay). Stratified, warm, oligotrophic (but relatively high iron and phosphate) and oxygen minimum zone (OMZ) waters of the Bay might be a niche for diazotrophs. Therefore, we conducted N2 fixation rate measurements during the spring inter‐monsoon in the euphotic zone, the OMZ and below the OMZ down to 1,500 m depth near the coastal and in the central Bay. We further assessed primary production and cyanobacterial community composition along with their potential environmental controlling parameters. N2 fixation rates in the euphotic zone were low (0.02–0.38 nmol N L−1 d−1) and their contribution to primary production was small (<2%). Despite conducive conditions for diazotrophy in the Bay, the reason for the relatively low euphotic zone and OMZ N2 fixation rates remained unclear and enigmatic. Interestingly, significantly higher N2 fixation rates occurred below the OMZ (>600 m depth), ranging from 0.06 to 0.11 nmol N L−1 d−1 where oxygen concentrations ranged between 0.5 and 1.6 mL L−1, rather than within the OMZ where rates ranged from 0.02 to 0.08 nmol N L−1 d−1 and oxygen concentrations were ≤0.5 mL L−1. Euphotic zone N2 fixation showed seasonality in the Bay with increasing rates from spring to summer season, perhaps owing to increasing Fe flux as the summer monsoon approaches its peak. Plain Language Summary: The growth of marine phytoplankton is primarily limited by the reactive forms of nitrogen. Dinitrogen‐fixing organisms (termed "diazotrophs") supplement this need by providing a natural fertilizer—ammonium. Diazotrophs prefer phosphate and iron‐rich, but reactive nitrogen‐deficit waters. Therefore, the oligotrophic Bay of Bengal (the northeastern Indian Ocean) with excess phosphate relative to reactive nitrogen and high iron input could be a potential niche for diazotrophy. However, our results indicate that N2 fixation rates were low and supported less than 2% of organic matter formation in the Bay of Bengal during the spring inter‐monsoon. Surprisingly, N2 fixation rates were higher below the oxygen minimum zone than within it. Thus, our study provides additional evidence that N2 fixation is feasible (low but persistent) in dark marine places with abundant oxygen in conjunction with surface ocean. Key Points: Despite most prerequisites available for diazotrophic activity, N2 fixation rates are low in the Bay of BengalThe seasonality in N2 fixation in the Bay of Bengal is attributed to the Fe flux through dust depositionHigher N2 fixation rates occurred below rather than within the oxygen minimum zone [ABSTRACT FROM AUTHOR]

Pelagic predators must contend with low prey densities that are irregularly distributed and dynamic in space and time. Based on satellite imagery and telemetry data, many pelagic predators will concentrate horizontal movements on ephemeral surface fronts—gradients between water masses—because of enhanced local productivity and increased forage fish densities.Vertical fronts (e.g. thermoclines, oxyclines) can be spatially and temporally persistent, and aggregate lower trophic level and diel vertically migrating organisms due to sharp changes in temperature, water density or available oxygen. Thus, vertical fronts represent a stable and potentially energy rich habitat feature for diving pelagic predators but remain little explored in their capacity to enhance foraging opportunities.Here, we use a novel suite of high‐resolution biologging data, including in situ derived oxygen saturation and video, to document how two top predators in the pelagic ecosystem exploit the vertical fronts created by the oxygen minimum zone of the eastern tropical Pacific.Prey search behaviour was dependent on dive shape, and significantly increased near the thermocline and hypoxic boundary for blue marlin Makaira nigricans and sailfish Istiophorus platypterus, respectively. Further, we identify a behaviour not yet reported for pelagic predators, whereby the predator repeatedly dives below the thermocline and hypoxic boundary (and by extension, below the prey). We hypothesize this behaviour is used to ambush prey concentrated at the boundaries from below.We describe how habitat fronts created by low oxygen environments can influence pelagic ecosystems, which will become increasingly important to understand in the context of global change and expanding oxygen minimum zones. We anticipate that our findings are shared among many pelagic predators where strong vertical fronts occur, and additional high‐resolution tagging is warranted to confirm this. [ABSTRACT FROM AUTHOR]

The deep ocean, one of the largest ecosystems on earth, is dominated by microorganisms that are keystones in the regulation of biogeochemical cycles. However, the evolutionary pathways underlying the specific adaptations required (e.g., high pressure and low temperature) by this unique niche remain understudied. Here, we analyzed the first representatives belonging to the order Acidimicrobiales, a group of marine planktonic Actinobacteriota, that specifically inhabits the aphotic zone of the oceanic water column (>200 m). Compared with their epipelagic counterparts, deep-sea representatives showed the same evolution in genome architecture with higher GC content, longer intergenic spaces as well as higher nitrogen (N-ARSC) and lower carbon (C-ARSC) content in encoded amino acid residue side chains consistent with the higher nitrogen concentration and lower carbon concentration in deep waters compared to the photic zone. Metagenomic recruitment showed distribution patterns that allowed the description of different ecogenomic units within the three deep water-associated genera defined by our phylogenomic analyses (UBA3125, S20-B6 and UBA9410). The entire genus UBA3125 was found exclusively associated with oxygen minimum zones linked to the acquisition of genes involved in denitrification. Genomospecies of genus S20-B6 recruited in samples from both mesopelagic (200-1,000 m) and bathypelagic (1000-4,000 m) zones, including polar regions. Diversity in the genus UBA9410 was higher, with genomospecies widely distributed in temperate zones, others in polar regions, and the only genomospecies associated with abyssal zones (>4,000 m). At the functional level, groups beyond the epipelagic zone have a more complex transcriptional regulation including in their genomes a unique WhiB paralog. In addition, they showed higher metabolic potential for organic carbon and carbohydrate degradation as well as the ability to accumulate glycogen as a source of carbon and energy. This could compensate for energy metabolism in the absence of rhodopsins, which is only present in genomes associated with the photic zone. The abundance in deep samples of cytochrome P450 monooxygenases associated with the genomes of this order suggests an important role in remineralization of recalcitrant compounds throughout the water column. [ABSTRACT FROM AUTHOR]

Sediments of the continental shelf anoxic zones of the Benguela upwelling system (BUS) and the Peruvian upwelling system are present-day hotspots of trace element accumulation. However, contribution of the lithogenic trace element fraction and early diagenetic transformation processes are poorly constrained. The identification of source and accumulation mechanisms is necessary for the validation of trace elements as proxies for productivity and redox cycling, notably in highly dynamic upwelling systems such as the BUS. Here, we analyzed redox- and biosensitive elements (U, Ba, Ni), lithogenic tracers (Al, Ti, Zr), total organic carbon (TOC) and P in eleven short sediment cores from the Namibian shelf (ca. 22–25°S) as well as grain size fractionated hinterland samples from the Tsauchab Valley at Sossusvlei (∼25°S). These samples help to constrain the lithogenic trace element contributions, a prerequisite for a realistic interpretation of marine authigenic trace element data. Our findings corroborate previous findings that Zr is a sensitive tracer of the coarse fraction in marine sediment for dust input and/or sediment reworking. Hence, the combined analysis of the coarse fraction (Zr), organic matter, and P enrichments (TOC/P) differentiates between a calm, organic matter-rich central shelf environment and a more energetic phosphorite-rich southern shelf environment. A significant correlation of U with P in cores from the southern shelf corroborates previous findings of U incorporation into apatite, the initial formation of which requires oscillating oxic to sulfidic redox conditions. Non-apatite associated U dominates in the cores from the central shelf, where it is only enriched in the buried, anoxic parts of the sediment. This also supports oscillating oxic to sulfidic redox conditions in surface sediment, where U may be recycled. Hence, U accumulates either (i) in apatite under oscillating oxic to sulfidic conditions or (ii) in buried anoxic sediment sections. This strongly questions the use of U as indicator of suboxic conditions but rather anoxic conditions. While lithogenic fractions of U were negligible, the lithogenic fractions of Ba and Ni were found to be elevated. The calculation of authigenic Ba when accounting for lithogenic Ba from the Sossusvlei fine fractions gives more realistic authigenic Ba values. The high Ba enrichment in the BUS is interpreted as uptake of Ba by diatoms and/or nucleation of barite in P-rich diatom remains, as suggested previously. Given the shallow water depth of the cores, barite formation must already take place in subsurface waters (<50 m water depth) as opposed to previous suggesting greater water depths. The lithogenic background of Ni strongly varies from 23°S to 25°S based on Sossusvlei data and the literature. When authigenic Ni contents are calculated accordingly and plotted versus TOC, more systematic trends are seen (compared to total Ni versus TOC). Essentially all cores show an increase in the authigenic Ni/TOC ratio with sediment depth suggesting better preservation and retention of Ni as compared to TOC consistent with previous observations in sediments from the Peruvian upwelling system. This trend corroborates the use of Ni as a productivity indicator for upwelling sediments. [ABSTRACT FROM AUTHOR]

Observations suggest that the tropical Pacific Ocean has lost oxygen since the 1960s leading to the expansion of its oxygen minimum zone (OMZ). Attribution to anthropogenic forcing is, however, difficult because of limited data availability and the large natural variability introduced by the Pacific Decadal Oscillation (PDO). Here, we evaluate the PDO influence on oxygen dynamics and OMZ extent using observations and hindcast simulations from two global ocean circulation models (NEMO‐PISCES, MOM6‐COBALT). In both models, the tropical Pacific oxygen content decreases by about 30 Tmol.decade−1 and the OMZ volume expands by 1.3 × 105 km3.decade−1 during PDO positive phases, while variations of similar magnitude but opposite sign are simulated during negative phases. Changes in equatorial advective oxygen supply, partially offset by biological demand, control the oxygen response to PDO. Observations which cover 39% of the tropical Pacific volume only partially capture spatio‐temporal variability, hindering the separation of anthropogenic trend from natural variations. Plain Language Summary: Human activities cause oxygen loss in the ocean, which leads to the expansion of areas with very low oxygen concentrations located in the tropics called oxygen minimum zones (OMZ). Understanding the dynamics of OMZs is crucial because they produce greenhouse gasses and are unsuitable for the life of most large marine organisms. Quantifying the response of OMZs is however complicated by natural variability that superimposes on human‐induced changes. In the Pacific Ocean, one of the strongest natural variability phenomena is the Pacific Decadal Oscillation. We used data and numerical models to assess the magnitude of oxygen changes caused by this natural phenomena in the tropical Pacific Ocean, and show that they are comparable to that of human‐induced oxygen changes. We highlight that more oxygen data is needed to accurately separate natural variations from human‐induced changes, and that a fraction of the oxygen loss attributed to human activities in prior work could in fact be due to natural variability. Key Points: Pacific Decadal Oscillation (PDO) modulates tropical Pacific oxygen content and oxygen minimum zone volume on decadal time scalesThe PDO‐induced variations are of the same order of magnitude as the anthropogenic deoxygenation signalCurrently available data are too sparse to resolve and isolate the PDO‐induced and anthropogenic signals [ABSTRACT FROM AUTHOR]

The oxygen minimum zone (OMZ) is an essential feature along the Pacific margin, extensively studied for its influence on benthic fauna, acquiring relevance in coastal zones due to its impact on aquaculture species, marine-managed areas, and marine-protected areas. The most evident temporal variability beyond seasonal timescales observed for the OMZ was related to inter-annual variability associated with El Niño, when warmer and more oxygenated waters arrive at shallower zones and the OMZ becomes deeper. However, the impact of oxygen-deficient water that appears sporadically over the shelf is not yet understood. This study provides an integrated view of the oceanographic and climatic conditions behind bottom oxygen conditions along the Chilean continental margin, considering the temporal variability in the South East (SE) Pacific during the last ~2000 years. We organized the information on redox-sensitive metals and δ15N from sedimentary cores obtained from shelf zones from 23°S to 36°S at water depths lower than 100 m. The results demonstrate the variability in the upper part of the OMZ over the shelf, which seems to respond to climatic oscillations. The element distribution indicated in particular a more intense OMZ was developed over the shelf before 1400 AD, followed by more oxygenated bottom conditions afterward, except between 1925 and 1970 off 36°S when the OMZ was intensified, a condition that was not observed at the northern sites. This period mostly coincided with a low Pacific Decadal Oscillation amplitude and reduced El Niño Southern Oscillation (ENSO) activity. In the last 50 years, the proxy data suggests less intense suboxic environment along the Chilean margin. This weakening of the upper boundary of the OMZ in the last decades is interpreted as resulting from the accumulative effect of ventilation processes at different time scales.

Oxygen minimum zones (OMZs) are extended oceanic regions for which dissolved oxygen concentration is extremely low. They are suspected to be expanding in response to global warming. However, currently, the mechanisms by which OMZ varies in response to climate variability are still uncertain. Here, the variability of the subtropical OMZ off central Chile of a regional coupled physical–biogeochemical regional model simulation was analyzed for the period 2000–2008, noting that its fluctuations were significant despite the relatively weak amplitude of the El Niño/Southern Oscillation (ENSO). In particular, the interannual variability in the OMZ volume (OMZVOL, defined as the volume with dissolved oxygen concentration (DO) ≤ 45μM) was approximately 38% larger than that of the seasonal cycle, with maximum and minimum anomalies of OMZVOL taking place during two cold La Niña (LN) years (2001 and 2007). The model analyses further reveal that these anomalies resulted from a combined effect of changes in (1) the oxygen-poor waters poleward transport by the Peru–Chile undercurrent (PCUC), (2) the intensity of quasi-zonal jets influencing the transport of water to and from the OMZ, and (3) the zonal DO transport related to mesoscale eddy activity. Specifically, the interannual variability of the PCUC modulated primarily the DO contents of the OMZ core [(DO) ≤ 20μM] and secondarily the OMZVOL, while cross-shore DO transport by the zonal jets and the eddy fluxes played a major role in ventilating and shaping the offshore extent of the OMZ. When the OMZVOL was maximum (minimum), the PCUC transport was slightly increased (reduced), which was associated with a reduction (increase) in the ventilation of the OMZ through negative (positive) anomalies of zonal advection and DO eddy fluxes. Our results demonstrate that significant natural interannual variability in the subtropical OMZ off Chile originates from the interplay between oceanic equatorial teleconnection (PCUC transport) and local non-linear dynamics (the zonal jets and mesoscale eddies).

In Arabian Sea (AS), land-locked northern boundary and strong seasonal productivity lead to the formation of one of the most intense open ocean Oxygen Minimum Zones (OMZs). Presence of this perennial OMZ has significant consequences on adjacent coastal fisheries and ecosystem. Simulations from CMIP5 suggest significant weakening of both monsoonal winds and productivity under high emission scenario. But the fate of AS OMZ in this scenario - whether it will expand or shrink - still remains elusive, mainly due to poor representation of extent and strength of AS OMZ in CMIP5 present-day simulations. To address this, we analyze the distribution of O2 in AS from a subset of three contrasted CMIP5 simulations, and complemented with a set of regional downscaled model experiments which we forced at surface and open boundaries using information from those three CMIP5 models. We tested two regional downscaling approaches - with and without correction of CMIP5 biases with respect to observations. Using a set of sensitivity experiments, we disentangle the contributions of local (atmospheric) forcing vs. remote (at the lateral boundaries) forcing in driving the future projected O2 changes. While CMIP5 projects either shrinking or expansion of the AS OMZ depending on the model, our downscaling experiments consistently project a shrinking of AS OMZ. We show that projected O2 changes in OMZ layer are affected by both local and remote processes. In the southern AS, the main response to climate change is oxygenation that originates from the boundaries, and hence downscalled and CMIP5 model responses are similar. In contrast, in northern AS, downscaling yields a substantial reduction in O2 projection discrepancies because of a minimal influence of remote forcing there leading to a stronger sensitivity to improved local physics and improved model representation of present-day conditions. We find that when corrected for present-day biases, projected deoxygenation in the northern AS is shallower. Our findings indicate the importance of downscaling of global models in regions where local forcing is dominant, and the need for correcting global model biases with respect to observations to reduce uncertainties in future O2 projections.

The deep ocean, one of the largest ecosystems on earth, is dominated by microorganisms that are keystones in the regulation of biogeochemical cycles. However, the evolutionary pathways underlying the specific adaptations required (e.g., high pressure and low temperature) by this unique niche remain understudied. Here, we analyzed the first representatives belonging to the order Acidimicrobiales, a group of marine planktonic Actinobacteriota, that specifically inhabits the aphotic zone of the oceanic water column (>200 m). Compared with their epipelagic counterparts, deep-sea representatives showed the same evolution in genome architecture with higher GC content, longer intergenic spaces as well as higher nitrogen (N-ARSC) and lower carbon (C-ARSC) content in encoded amino acid residue side chains consistent with the higher nitrogen concentration and lower carbon concentration in deep waters compared to the photic zone. Metagenomic recruitment showed distribution patterns that allowed the description of different ecogenomic units within the three deep water-associated genera defined by our phylogenomic analyses (UBA3125, S20-B6 and UBA9410). The entire genus UBA3125 was found exclusively associated with oxygen minimum zones linked to the acquisition of genes involved in denitrification. Genomospecies of genus S20-B6 recruited in samples from both mesopelagic (200–1,000 m) and bathypelagic (1000–4,000 m) zones, including polar regions. Diversity in the genus UBA9410 was higher, with genomospecies widely distributed in temperate zones, others in polar regions, and the only genomospecies associated with abyssal zones (>4,000 m). At the functional level, groups beyond the epipelagic zone have a more complex transcriptional regulation including in their genomes a unique WhiB paralog. In addition, they showed higher metabolic potential for organic carbon and carbohydrate degradation as well as the ability to accumulate glycogen as a source of carbon and energy. This could compensate for energy metabolism in the absence of rhodopsins, which is only present in genomes associated with the photic zone. The abundance in deep samples of cytochrome P450 monooxygenases associated with the genomes of this order suggests an important role in remineralization of recalcitrant compounds throughout the water column.

Abstract Observations suggest that the tropical Pacific Ocean has lost oxygen since the 1960s leading to the expansion of its oxygen minimum zone (OMZ). Attribution to anthropogenic forcing is, however, difficult because of limited data availability and the large natural variability introduced by the Pacific Decadal Oscillation (PDO). Here, we evaluate the PDO influence on oxygen dynamics and OMZ extent using observations and hindcast simulations from two global ocean circulation models (NEMO‐PISCES, MOM6‐COBALT). In both models, the tropical Pacific oxygen content decreases by about 30 Tmol.decade−1 and the OMZ volume expands by 1.3 × 105 km3.decade−1 during PDO positive phases, while variations of similar magnitude but opposite sign are simulated during negative phases. Changes in equatorial advective oxygen supply, partially offset by biological demand, control the oxygen response to PDO. Observations which cover 39% of the tropical Pacific volume only partially capture spatio‐temporal variability, hindering the separation of anthropogenic trend from natural variations.

Productive oxygen minimum zones are regions dominated by heterotrophic denitrification fueled by sinking organic matter. Microbial redox-sensitive transformations therein result in the loss and overall geochemical deficit in inorganic fixed nitrogen in the water column, thereby impacting global climate in terms of nutrient equilibrium and greenhouse gases. Here, geochemical data are combined with metagenomes, metatranscriptomes, and stable-isotope probing incubations from the water column and subseafloor of the Benguela upwelling system. The taxonomic composition of 16S rRNA genes and relative expression of functional marker genes are used to explore metabolic activities by nitrifiers and denitrifiers under decreased stratification and increased lateral ventilation in Namibian coastal waters. Active planktonic nitrifiers were affiliated with Candidatus Nitrosopumilus and Candidatus Nitrosopelagicus among Archaea, and Nitrospina, Nitrosomonas, Nitrosococcus, and Nitrospira among Bacteria. Concurrent evidence from taxonomic and functional marker genes shows that populations of Nitrososphaeria and Nitrospinota were highly active under dysoxic conditions, coupling ammonia and nitrite oxidation with respiratory nitrite reduction, but minor metabolic activity toward mixotrophic use of simple nitrogen compounds. Although active reduction of nitric oxide to nitrous oxide by Nitrospirota, Gammaproteobacteria, and Desulfobacterota was tractable in bottom waters, the produced nitrous oxide was apparently scavenged at the ocean surface by Bacteroidota. Planctomycetota involved in anaerobic ammonia oxidation were identified in dysoxic waters and their underlying sediments, but were not found to be metabolically active due to limited availability of nitrite. Consistent with water column geochemical profiles, metatranscriptomic data demonstrate that nitrifier denitrification is fueled by fixed and organic nitrogen dissolved in dysoxic waters, and prevails over canonical denitrification and anaerobic oxidation of ammonia when the Namibian coastal waters and sediment–water interface on the shelf are ventilated by lateral currents during austral winter. [ABSTRACT FROM AUTHOR]

The widespread Capitanian (late Guadalupian) marine anoxia/euxinia has long been regarded as a key driver of the end-Guadalupian (middle Permian) biotic crisis. However, the cause of this marine anoxia is debated, particularly regarding the influence of the Emeishan large igneous province (ELIP). To investigate the contribution of the ELIP to marine anoxia and the possible causal mechanisms, we undertook a conodont biostratigraphic and geochemical study of the middle Permian Maokou Formation in a platform-to-trough transect in the Upper Yangtze region, South China. Our results show that the depositional facies of the Maokou Formation changed from a carbonate ramp to an intra-platform trough within the Jinogondolella (J.) altudaensis zone in the northwestern Yangtze region, which can be attributed to the initial activity of the ELIP. Mantle-derived Sr inputs in the initial and main stages of the ELIP led to two decreases in 87Sr/86Sr during the Capitanian, in the J. shannoni – J. altudaensis and J. prexuanhanensis – J. xuanhanensis zones. The elevated 87Sr/86Sr values during the late Capitanian may have been due to enhanced continental weathering caused by rapid climate warming in response to subaerial eruptions of the ELIP. The deep-water anoxia–euxinia expanded during the middle Capitanian, as indicated by increased Mo EF /U EF and V/(V + Ni) values, along with the disappearance of burrows and appearance of small pyrite framboids in the J. altudaensis zone in the Cheng 20 well. However, shallow-water anoxia occurred during the late Capitanian (i.e., J. prexuanhanensis – J. xuanhanensis zone), as evidenced by positive Ce anomalies and losses of aerobic benthic species in the Erya section. Importantly, marine anoxia and negative δ13C carb excursions occurred synchronously, but earlier in deep water than in shallow water, potentially indicating an expansion of the oxygen minimum zone (OMZ). The deep-water anoxia corresponded to a decrease in 87Sr/86Sr ratios and the appearance of an intra-platform trough in the J. altudaensis zone, whereas the shallow-water anoxia in the late Capitanian coincided with elevated 87Sr/86Sr ratios. This suggests that the initial activity of the ELIP promoted the development of the OMZ in deep waters during the middle Capitanian, while the subaerial eruptions of the ELIP drove climate warming that led to the expansion of the OMZ into shallow-water platforms during the late Capitanian. • The ELIP triggered the Capitanian intra-platform trough in the N.W. Yangtze region. • 87Sr/86Sr evolution indicates ELIP-induced mantle activities and climate change. • The ELIP eruptions successively caused deep- and shallow-water anoxia. [ABSTRACT FROM AUTHOR]

Productive oxygen minimum zones are regions dominated by heterotrophic denitrification fueled by sinking organic matter. Microbial redox-sensitive transformations therein result in the loss and overall geochemical deficit in inorganic fixed nitrogen in the water column, thereby impacting global climate in terms of nutrient equilibrium and greenhouse gases. Here, geochemical data are combined with metagenomes, metatranscriptomes, and stable-isotope probing incubations from the water column and subseafloor of the Benguela upwelling system. The taxonomic composition of 16S rRNA genes and relative expression of functional marker genes are used to explore metabolic activities by nitrifiers and denitrifiers under decreased stratification and increased lateral ventilation in Namibian coastal waters. Active planktonic nitrifiers were affiliated with Candidatus Nitrosopumilus and Candidatus Nitrosopelagicus among Archaea, and Nitrospina, Nitrosomonas, Nitrosococcus, and Nitrospira among Bacteria. Concurrent evidence from taxonomic and functional marker genes shows that populations of Nitrososphaeria and Nitrospinota were highly active under dysoxic conditions, coupling ammonia and nitrite oxidation with respiratory nitrite reduction, but minor metabolic activity toward mixotrophic use of simple nitrogen compounds. Although active reduction of nitric oxide to nitrous oxide by Nitrospirota, Gammaproteobacteria, and Desulfobacterota was tractable in bottom waters, the produced nitrous oxide was apparently scavenged at the ocean surface by Bacteroidota. Planctomycetota involved in anaerobic ammonia oxidation were identified in dysoxic waters and their underlying sediments, but were not found to be metabolically active due to limited availability of nitrite. Consistent with water column geochemical profiles, metatranscriptomic data demonstrate that nitrifier denitrification is fueled by fixed and organic nitrogen dissolved in dysoxic waters, and prevails over canonical denitrification and anaerobic oxidation of ammonia when the Namibian coastal waters and sediment–water interface on the shelf are ventilated by lateral currents during austral winter.