Eastern Tropical South Pacific Research Articles

The Northernmost Record of a Stranded Southern Right Whale (Eubalaena australis) Calf in Ecuador’s North Coast: Notes on Stranding Causes and Distribution Expansion in the Southeastern Tropical Pacific

The Northernmost Record of a Stranded Southern Right Whale (Eubalaena australis) Calf in Ecuador’s North Coast: Notes on Stranding Causes and Distribution Expansion in the Southeastern Tropical Pacific

Millennial‐Scale Changes in Atmospheric Nitrous Oxide During the Holocene

AbstractNitrous oxide (N2O) is an important greenhouse gas which destroys the ozone in the stratosphere. Primary sources of atmospheric N2O are nitrification and denitrification in terrestrial soils and the ocean, and the main sink is photolysis in the stratosphere. Studies have mostly focused on the climate‐related response of N2O during glacial‐interglacial periods. However, its mechanism of variation during the Holocene remains unclear. We present a high‐resolution N2O record from the South Pole Ice (SPICE) core covering the Holocene epoch. The millennial‐scale N2O trend agrees with existing records. We constructed a Holocene composite consisting of the new N2O measurements in SPICE and existing records from other ice core sites. The N2O composite reveals four distinct periods of N2O variation during 11.5–10.0 ka, 10.0–6.2 ka, 6.2–2.2 ka, and 2.2–1.4 ka, including two maxima in 11.0–10.0 ka and 3.0–2.2 ka and minima in 8.8–6.2 ka and approximately 1.4 ka. Apart from these, our new high‐resolution record from SPICE shows a short‐term N2O decrease around 2.8 ka which is not observed in other records possibly due to lower sample resolution and/or higher age smoothing. Comparison of our new Holocene N2O composite with the paleo‐proxy records suggests the plausible linkage of major monsoon (Asian, North African, South and North American, and Australian‐Indonesian monsoon) and upwelling (Arabian Sea and Eastern Tropical South Pacific) regions in regulating the atmospheric N2O during the Holocene.

Quantifying N2 fixation and its contribution to export production near the Tonga-Kermadec Arc using nitrogen isotope budgets

The spatial distribution of marine di-nitrogen (N2) fixation informs our understanding of the sensitivities of this process as well as the potential for this new nitrogen (N) source to drive export production, influencing the global carbon (C) cycle and climate. Using geochemically-derived δ15N budgets, we quantified rates of N2 fixation and its importance for supporting export production at stations sampled near the southwest Pacific Tonga-Kermadec Arc. Recent observations indicate that shallow (<300 m) hydrothermal vents located along the arc provide significant dissolved iron to the euphotic zone, stimulating N2 fixation. Here we compare measurements of water column δ15NNO3+NO2 with sinking particulate δ15N collected by short-term sediment traps deployed at 170 m and 270 m at stations in close proximity to subsurface hydrothermal activity, and the δ15N of N2 fixation. Results from the δ15N budgets yield high geochemically-based N2 fixation rates (282 to 638 µmol N m-2 d-1) at stations impacted by hydrothermal activity, supporting 64 to 92% of export production in late spring. These results are consistent with contemporaneous 15N2 uptake rate estimates and molecular work describing high Trichodesmium spp. and other diazotroph abundances associated with elevated N2 fixation rates. Further, the δ15N of sinking particulate N collected at 1000 m over an annual cycle revealed sinking fluxes peaked in the summer and coincided with the lowest δ15N, while lower winter sinking fluxes had the highest δ15N, indicating isotopically distinct N sources supporting export seasonally, and aligning with observations from most other δ15N budgets in oligotrophic regions. Consequently, the significant regional N2 fixation input to the late spring/summer Western Tropical South Pacific results in the accumulation of low-δ15NNO3+NO2 in the upper thermocline that works to lower the elevated δ15NNO3+NO2 generated in the oxygen deficient zones in the Eastern Tropical South Pacific.

Shifts in the Isotopic Composition of Nitrous Oxide Between El Niño and La Niña in the Eastern Tropical South Pacific

AbstractThe El Niño‐Southern Oscillation (ENSO) is a natural climate phenomenon that alters the biogeochemical and physical dynamics of the Eastern Tropical Pacific Ocean. Its two phases, El Niño and La Niña, are characterized by decreased and increased coastal upwelling, respectively, which have cascading effects on primary productivity, organic matter supply, and ocean‐atmosphere interactions. The Eastern Tropical South Pacific oxygen minimum zone is a source of nitrous oxide (N2O), a potent greenhouse gas, to the atmosphere. Here, we present the first study to directly compare N2O sources during opposing ENSO phases using N2O isotopocule analyses. Our data show that during La Niña, N2O accumulation increased six‐fold in the upper 100 m of the water column, and N2O fluxes to the atmosphere increased up to 20‐fold. N2O isotopocule data demonstrated substantial increases in δ18O up to 60.5‰ and decreases in δ15Nβ down to −10.3‰ in the oxycline, signaling a shift in N2O cycling during La Niña compared to El Niño. During El Niño, N2O production was primarily due to ammonia‐oxidizing archaea, whereas during La Niña, N2O production by incomplete denitrification supplemented that from ammonia‐oxidation, with N2O consumption likely maintaining the high site preference values (up to 26.7‰). Ultimately, our results illustrate a strong connection between upwelling intensity, biogeochemistry, and N2O flux to the atmosphere. Additionally, they highlight the combined power of N2O isotopocule analysis and repeat measurements in the same region to constrain N2O interannual variability and cycling dynamics under different climate scenarios.

Pathways of Nitrous Oxide Production in the Eastern Tropical South Pacific Oxygen Minimum Zone

AbstractOceanic emissions of nitrous oxide (N2O) account for roughly one‐third of all natural sources to the atmosphere. Hot‐spots of N2O outgassing occur over oxygen minimum zones (OMZs), where the presence of steep oxygen gradients surrounding anoxic waters leads to enhanced N2O production from both nitrification and denitrification. However, the relative contributions from these pathways to N2O production and outgassing in these regions remains poorly constrained, in part due to shared intermediary nitrogen tracers, and the tight coupling of denitrification sources and sinks. To shed light on this problem, we embed a new, mechanistic model of the OMZ nitrogen cycle within a three‐dimensional eddy‐resolving physical‐biogeochemical model of the Eastern Tropical South Pacific (ETSP), tracking contributions from remote advection, atmospheric exchange, and local nitrification and denitrification. The model indicates that net N2O production from denitrification is approximately one order of magnitude greater than nitrification within the ETSP OMZ. However, only ∼32% of denitrification‐derived N2O production ultimately outgasses to the atmosphere in this region (contributing ∼36% of the air‐sea N2O flux on an annual basis), while the remaining is exported out of the domain. Instead, remotely produced N2O advected into the OMZ region accounts for roughly half (∼57%) of the total N2O outgassing, with smaller contributions from nitrification (∼7%). Our results suggests that, together with enhanced production by denitrification, upwelling of remotely derived N2O contributes the most to N2O outgassing over the ETSP OMZ.

Marine N2O cycling from high spatial resolution concentration, stable isotopic and isotopomer measurements along a meridional transect in the eastern Pacific Ocean

Nitrous oxide (N2O) is a potent greenhouse gas and ozone depleting substance, with the ocean accounting for about one third of global emissions. In marine environments, a significant amount of N2O is produced by biological processes in Oxygen Deficient Zones (ODZs). While recent technological advances are making surface N2O concentration more available, high temporal and spatial resolution water-column N2O concentration data are relatively scarce, limiting global N2O ocean models’ predictive capability. We present a N2O concentration, stable isotopic composition and isotopomer dataset of unprecedently large spatial coverage and depth resolution in the broader Pacific, crossing both the eastern tropical South and North Pacific Ocean ODZs collected as part of the GO-SHIP P18 repeat hydrography program in 2016/2017. We complement these data with dissolved gases (nitrogen, oxygen, argon) and nitrate isotope data to investigate the pathways controlling N2O production in relation to apparent oxygen utilization and fixed nitrogen loss. N2O yield significantly increased under low oxygen conditions near the ODZs. Keeling plot analysis revealed different N2O sources above the ODZs under different oxygen regimes. Our stable isotopic data and relationships between the N2O added by microbial processes (ΔN2O) and dissolved inorganic nitrogen (DIN) deficit confirm increased N2O production by denitrification under low oxygen conditions near the oxycline where the largest N2O accumulations were observed. The slope for δ18O-N2O versus site preference (SP, the difference between the central (α) and outer (β) N atoms in the linear N2O molecule) in the eastern tropical North Pacific ODZ was lower than expected for pure N2O reduction, likely because of the observed decrease in δ15Nβ. This trend is consistent with prior ODZ studies and attributed to concurrent production of N2O from nitrite with a low δ15N or denitrification with a SP >0‰. We estimated apparent isotope effects for N2O consumption in the ETNP ODZ of 3.6‰ for 15Nbulk, 9.4‰ for 15Nα, -2.3‰ for 15Nβ, 12.0‰ for 18O, and 11.7‰ for SP. These values were generally within ranges previously reported for previous laboratory and field experiments.

Diving deeper: Mesopelagic fish biomass estimates comparison using two different models

A growing population on a planet with limited resources demands finding new sources of protein. Hence, fisheries are turning their perspectives towards mesopelagic fish, which have, so far, remained relatively unexploited and poorly studied. Large uncertainties are associated with regards to their biomass, turn-over rates, susceptibility to environmental forcing and ecological and biogeochemical role. Models are useful to disentangle sources of uncertainties and to understand the impact of different processes on the biomass. In this study, we employed two food-web models – OSMOSE and the model by Anderson et al. (2019, or A2019) – coupled to a regional physical–biogeochemical model to simulate mesopelagic fish in the Eastern Tropical South Pacific ocean. The model by A2019 produced the largest biomass estimate, 26 to 130% higher than OSMOSE depending on the mortality parameters used. However, OSMOSE was calibrated to match observations in the coastal region off Peru and its temporal variability is affected by an explicit life cycle and food web. In contrast, the model by A2019 is more convenient to perform uncertainty analysis and it can be easily coupled to a biogeochemical model to estimate mesopelagic fish biomass. However, it is based on a flow analysis that had been previously applied to estimate global biomass of mesopelagic fish but has never been calibrated for the Eastern Tropical South Pacific. Furthermore, it assumes a steady-state in the energy transfer between primary production and mesopelagic fish, which may be an oversimplification for this highly dynamic system. OSMOSE is convenient to understand the interactions of the ecosystem and how including different life stages affects the model response. The combined strengths of both models allow us to study mesopelagic fish from a holistic perspective, taking into account energy fluxes and biomass uncertainties based on primary production, as well as complex ecological interactions.

Subsurface Thermohaline Biases in the Southern Tropical Pacific and the Roles of Wind Stress and Precipitation

Abstract Thermohaline structure and time evolution in the subsurface ocean play a critical role in climate variability and predictability. They are still poorly represented in ocean and climate models. Here, the characteristics of subsurface thermohaline biases in the southern tropical Pacific and their causes are investigated through CMIP-based analyses and model-based experiments. There exists a pronounced subsurface cold bias at 200-m depth over the southern tropical Pacific in CMIP6 simulations with an ensemble mean of about −4°C and an extreme close to −10°C. This cold bias is accompanied by a fresh subsurface bias of about −0.9 psu in the ensemble mean (−1.9 psu minimum). Similar subsurface thermohaline biases also exist in CMIP5 outputs, indicating that reduction of these biases remains a long-standing challenge for model developments. To understand the causes of these biases, attribution analyses and POP2-based sensitivity experiments are performed. It is found that the subsurface thermohaline biases are attributed to the model deficiencies in simulating wind stress curl and precipitation in the southern tropical Pacific. By conducting CESM2-based coupled experiments, a warm SST bias in the southeastern tropical Pacific is found to be responsible for the poor simulations in wind stress curl and precipitation. The consequences of these biases are also analyzed. The subsurface thermohaline biases cause the density field to increase substantially along 10°S, flattening the zonal isopycnal surface and reducing equatorward interior transport. In addition, the anomalously cold and fresh subsurface signals in the southern tropical Pacific are seen to propagate to the equator, leading to an overall spurious cooling in the equatorial subsurface. Significance Statement Subsurface biases severely degrade the credibility of climate models in their predictions and projections; hence, it is important to understand the causes of these subsurface biases. Our study analyzes the characteristics of subsurface thermohaline biases in the southern tropical Pacific and investigates their causes. A pronounced subsurface cold bias is found over the southern tropical Pacific, accompanied by an obvious subsurface fresh bias. By performing attribution analyses and numerical experiments, it is found that the subsurface thermohaline biases are attributed to the model deficiencies in simulating wind stress and precipitation, which are further attributed to the warm SST bias in the southeastern tropical Pacific. These results provide a guide for improving climate model performances.

Associations between picocyanobacterial ecotypes and cyanophage host genes across ocean basins and depth.

Cyanophages, viruses that infect cyanobacteria, are globally abundant in the ocean's euphotic zone and are a potentially important cause of mortality for marine picocyanobacteria. Viral host genes are thought to increase viral fitness by either increasing numbers of genes for synthesizing nucleotides for virus replication, or by mitigating direct stresses imposed by the environment. The encoding of host genes in viral genomes through horizontal gene transfer is a form of evolution that links viruses, hosts, and the environment. We previously examined depth profiles of the proportion of cyanophage containing various host genes in the Eastern Tropical North Pacific Oxygen Deficient Zone (ODZ) and at the subtropical North Atlantic (BATS). However, cyanophage host genes have not been previously examined in environmental depth profiles across the oceans. We examined geographical and depth distributions of picocyanobacterial ecotypes, cyanophage, and their viral-host genes across ocean basins including the North Atlantic, Mediterranean Sea, North Pacific, South Pacific, and Eastern Tropical North and South Pacific ODZs using phylogenetic metagenomic read placement. We determined the proportion of myo and podo-cyanophage containing a range of host genes by comparing to cyanophage single copy core gene terminase (terL). With this large dataset (22 stations), network analysis identified statistical links between 12 of the 14 cyanophage host genes examined here with their picocyanobacteria host ecotypes. Picyanobacterial ecotypes, and the composition and proportion of cyanophage host genes, shifted dramatically and predictably with depth. For most of the cyanophage host genes examined here, we found that the composition of host ecotypes predicted the proportion of viral host genes harbored by the cyanophage community. Terminase is too conserved to illuminate the myo-cyanophage community structure. Cyanophage cobS was present in almost all myo-cyanophage and did not vary in proportion with depth. We used the composition of cobS phylotypes to track changes in myo-cyanophage composition. Picocyanobacteria ecotypes shift with changes in light, temperature, and oxygen and many common cyanophage host genes shift concomitantly. However, cyanophage phosphate transporter gene pstS appeared to instead vary with ocean basin and was most abundant in low phosphate regions. Abundances of cyanophage host genes related to nutrient acquisition may diverge from host ecotype constraints as the same host can live in varying nutrient concentrations. Myo-cyanophage community in the anoxic ODZ had reduced diversity. By comparison to the oxic ocean, we can see which cyanophage host genes are especially abundant (nirA, nirC, and purS) or not abundant (myo psbA) in ODZs, highlighting both the stability of conditions in the ODZ and the importance of nitrite as an N source to ODZ endemic LLV Prochlorococcus.

Moisture Source for the Precipitation of Tropical Cyclones over the Pacific Ocean through a Lagrangian Approach

Abstract Tropical cyclones (TCs) are an important component of the hydrological cycle at tropical latitudes. In this study, we investigated the origin of precipitation associated with TCs formed from 1980 to 2018 over the Pacific Ocean in three subbasins: the western North Pacific Ocean (WNP), central and east Pacific Ocean (NEPAC), and South Pacific Ocean (SPO) basins. The analysis was performed throughout the TC lifetime during genesis, when they reached the lifetime maximum intensity (LMI), and the dissipation stage. The backward trajectories of all precipitant atmospheric parcels residing over the TC locations from the global outputs of the Lagrangian Flexible Particle (FLEXPART) dispersion model fed by the ERA-Interim dataset were used to identify moisture sources. The South and East China Seas and the western tropical North Pacific Ocean were identified as the principal moisture sources in the WNP basin, while the atmospheric moisture that precipitated mainly came from the eastern tropical North and South Pacific Ocean in the NEPAC basin, followed by the Caribbean Sea. Meanwhile, the Coral Sea, western tropical South Pacific Ocean, and northern Australia are the origins of the moisture in the SPO. The mean moisture uptake per TC was higher during the hurricane category than during any other stage in each basin.

Subtropical clouds key to Southern Ocean teleconnections to the tropical Pacific

Excessive precipitation over the southeastern tropical Pacific is a major common bias that persists through generations of global climate models. While recent studies suggest an overly warm Southern Ocean as the cause, models disagree on the quantitative importance of this remote mechanism in light of ocean circulation feedback. Here, using a multimodel experiment in which the Southern Ocean is radiatively cooled, we show a teleconnection from the Southern Ocean to the tropical Pacific that is mediated by a shortwave subtropical cloud feedback. Cooling the Southern Ocean preferentially cools the southeastern tropical Pacific, thereby shifting the eastern tropical Pacific rainbelt northward with the reduced precipitation bias. Regional cloud locking experiments confirm that the teleconnection efficiency depends on subtropical stratocumulus cloud feedback. This subtropical cloud feedback is too weak in most climate models, suggesting that teleconnections from the Southern Ocean to the tropical Pacific are stronger than widely thought.

Iron and manganese accumulation within the Eastern Tropical North Pacific oxygen deficient zone

The Eastern Tropical North Pacific (ETNP) contains the largest oxygen deficient zone (ODZ) in the modern ocean. We determined dissolved concentrations of Fe, Fe(II), and Mn from three cruises in the region. Similar to other reported ODZs, Fe(II) was highest in the depth range associated with the secondary nitrite maximum. The main source of this feature is likely lateral advection of water overlying reducing shelf sediments within a narrow density range centered on the potential density anomaly of 26.5 kg/m3. This density horizon is similar to the Eastern Tropical South Pacific (ETSP) and reflects the intersection of the same density range with a large fraction of the continental shelf bottom waters. We also observed subsurface maxima of dissolved Mn in this density range, in contrast to the ETSP. Deep waters were enriched in Fe within the ETNP, analogous to other eastern boundary upwelling systems as well as the Arabian Sea. We argue that in these systems, reducing conditions on the shelf and overlying water column facilitate a robust shelf to basin shuttle of Fe, moving Fe from the coastal margin to deep plumes. Mn is also transported offshore in the core of the ODZ, and the relationship between Fe(II), Mn, and nitrite is remarkably similar between the ETNP, ETSP, and Arabian Sea. The exception is that Mn supply from the Peruvian shelf is less pronounced than in the other two ODZs, potentially reflecting the absence of large rivers in the Peruvian system.

Variability of Precipitation Recycling and Moisture Sources over the Colombian Pacific Region: A Precipitationshed Approach

This study assessed the precipitation recycling and moisture sources in the Colombian Pacific region between 1980–2017, based on the monitoring of moisture in the atmosphere through the Eulerian Water Accounting Model-2 layer (WAM2 layer) and the delimitation of the area contributing to terrestrial and oceanic moisture in the region is performed using the “precipitationshed” approach. The results indicate a unimodal precipitation recycling ratio for the North and Central Pacific and Patía-Mira regions, with the highest percentages between March and April, reaching 30% and 34%, respectively, and the lowest between September and October (between 19% and 21%). Moreover, monthly changes in the circulation of the region promote a remarkable variability of the sources that contribute to the precipitation of the study area and the spatial dynamics of the precipitationshed. From December to April, the main contributions come from continental sources in eastern Colombia and Venezuela, the tropical North Atlantic, and the Caribbean Sea, a period of high activity of the Orinoco Low-Level jet. In September, the moisture source region is located over the Pacific Ocean, where a southwesterly cross-equatorial circulation predominates, converging in western Colombia, known as the Choco Jet (CJ), decreasing the continental contribution. An intensified Caribbean Low-Level Jet inhibits moisture sources from the north between June and August, strengthening a southerly cross-equatorial flow from the Amazon River basin and the southeastern tropical Pacific. The March–April (September–October) season of higher (lower) recycling of continental precipitation is related to the weakening (strengthening) of the CJ in the first (second) half of the year, which decreases (increases) the contribution of moisture from the Pacific Ocean to the region, increasing (decreasing) the influence of land-based sources in the study area.

Carbon Assimilation by the Picoplanktonic Community Inhabiting the Secondary Chlorophyll Maximum of the Anoxic Marine Zones of the Eastern Tropical North and South Pacific

Anoxic marine zones (AMZs) constitute pelagic systems distinguished from the oxygen minimum zones (OMZs) by the complete absence of detectable oxygen and the accumulation of nitrite in mid-waters. At the top of the oxygen-depleted layer and below the oxycline, nutrients are abundant; light intensity is very much reduced (<1% of incident light) and a secondary chlorophyll maximum (SCM) is developed. The shoaling of the oxygen-depleted layer, product of the AMZ expansion, could enhance this SCM, which has little-known biogeochemical effects. Here, we show that the SCM is contributing a measurable signal in the particulate organic carbon (POC), enough to alter the δ13CPOC in the top of the oxygen-depleted layer. This data showed significant differences among stations with and without the development of a SCM, being 3.0‰ heavier when a SCM is developed, and indicating photosynthetic activity and/or remineralization in the top of the AMZ. More depleted δ13CPOC values were also found when no SCM was present indicating stronger chemoautotrophic activity, potentially driven by anammox and sulfur-oxidizing bacteria activity. Assimilation rate data show that when sufficient light and Prochlorococcus are present, photosynthesis exceeds chemoautotrophic carbon fixation, and can exceed heterotrophic assimilation of glucose or acetate. However, in the majority of the stations, assimilation rates of both glucose and acetate exceeded carbon fixation rates under light stimulation, suggesting that often the SCM is still a net heterotrophic system.

Estimating Mass Flux From Size‐Fractionated Filtered Particles: Insights Into Controls on Sinking Velocities and Mass Fluxes in Recent U.S. GEOTRACES Cruises

AbstractWe compile full ocean‐depth size‐fractionated (1–51 and >51 μm) particle concentration and composition of suspended particulate matter from three recent U.S. GEOTRACES cruises, and exploit detailed information of particle characteristics measured to give insights into controls on sinking velocity and mass flux. Our model integrates the concept of fractal scaling into Stokes' Law by incorporating one of two porosity‐size power law relationships that result in fractal dimensions of 1.4 and 2.1. The medians of pump‐derived total (>1 μm) mass flux in the upper 100 m of gyre stations are 285.1, 609.2, and 99.3 mg/m2/d in the North Atlantic, Eastern Tropical South Pacific, and Western Arctic Ocean cruises, respectively. In this data set, variations in particle concentration were generally more important than sinking velocity in controlling variations in mass flux. We examine different terms in a Stokes' Law model to explore how variations in particle and water column characteristics from these three cruises affect mass flux. The decomposition of different aspects of the Stokes' relationship sheds light on the lowest total mass flux of the three cruises in the Western Arctic, which could be explained by the Arctic having the lowest particle concentrations as well as the lowest sinking velocities due to having the smallest particle sizes and the most viscous water. This work shows the importance of both particle characteristics and size distribution for mass fluxes, and similar methods can be applied to existing and future size‐fractionated filtered particulate measurements to improve our understanding of the biological pump elsewhere.

An analysis of protists in Pacific oxygen deficient zones: implications for Prochlorococcus and N2 -producing bacteria.

Ocean oxygen deficient zones (ODZs) host 30%-50% of marine N2 production. Cyanobacteria photosynthesizing in the ODZ create a secondary chlorophyll maximum and provide organic matter to N2 -producing bacteria. This chlorophyll maximum is thought to occur due to reduced grazing in anoxic waters. We first examine ODZ protists with long amplicon reads. We then use non-primer-based methods to examine the composition and relative abundance of protists in metagenomes from the Eastern Tropical North and South Pacific ODZs and compare these data to the oxic Hawaii Ocean Time-series (HOT) in the North Pacific. We identify and quantify protists in proportion to the total microbial community. From metagenomic data, we see a large drop in abundance of fungi and protists such as choanoflagellates, radiolarians, cercozoa and ciliates in the ODZs but not in the oxic mesopelagic at HOT. Diplonemid euglenozoa were the only protists that increased in the ODZ. Dinoflagellates and foraminifera reads were also present in the ODZ though less abundant compared to oxic waters. Denitrification has been found in foraminifera but not yet in dinoflagellates. DNA techniques cannot separate dinoflagellate cells and cysts. Metagenomic analysis found taxonomic groups missed by amplicon sequencing and identified trends in abundance.

Iron(II) in the world's oxygen deficient zones

Oxygen deficient zones (ODZs) are regions in the marine water column where the concentration of oxygen is very low or zero and respiration occurs using nitrate as the terminal electron acceptor. Under such conditions, iron (Fe) would be predicted to be in the +3 valence state, and its geochemistry would be largely unaffected by the absence of oxygen. In contrast, observations show that Fe(II) is a major and sometimes dominant oxidation state in ODZs, even in the presence of a substantial amount of excess nitrate. Fe(II) plays an important role in Fe transport in ODZs. They are generally enriched in Fe throughout the regions where nitrate reduction is occurring. Here, it is argued that Fe(II) maxima arise as a result of processes common to the Eastern Tropical North and South Pacific as well as the Arabian Sea. Recent work has revealed that lateral advection of Fe(II) from reducing sediments is perhaps the major source and it appears that physical processes that are still poorly understood act over a surprisingly narrow density range. Tracers including iodine and 228Ra have been extremely useful in constraining these processes. The findings reported here have important implications for Fe transport from the continental margin to the ocean interior, and for microbes involved in the nitrogen cycle that have high Fe requirements.

Zooplankton mortality effects on the plankton community of the northern Humboldt Current System: sensitivity of a regional biogeochemical model

Abstract. Small pelagic fish off the coast of Peru in the eastern tropical South Pacific (ETSP) support around 10 % of global fish catches. Their stocks fluctuate interannually due to environmental variability which can be exacerbated by fishing pressure. Because these fish are planktivorous, any change in fish abundance may directly affect the plankton and the biogeochemical system. To investigate the potential effects of variability in small pelagic fish populations on lower trophic levels, we used a coupled physical–biogeochemical model to build scenarios for the ETSP and compare these against an already-published reference simulation. The scenarios mimic changes in fish predation by either increasing or decreasing mortality of the model's large and small zooplankton compartments. The results revealed that large zooplankton was the main driver of the response of the community. Its concentration increased under low mortality conditions, and its prey, small zooplankton and large phytoplankton, decreased. The response was opposite, but weaker, in the high mortality scenarios. This asymmetric behaviour can be explained by the different ecological roles of large, omnivorous zooplankton and small zooplankton, which in the model is strictly herbivorous. The response of small zooplankton depended on the antagonistic effects of mortality changes as well as on the grazing pressure by large zooplankton. The results of this study provide a first insight into how the plankton ecosystem might respond if variations in fish populations were modelled explicitly.

The cyanobacterium Prochlorococcus has divergent light-harvesting antennae and may have evolved in a low-oxygen ocean

Marine picocyanobacteria of the genus Prochlorococcus are the most abundant photosynthetic organisms in the modern ocean, where they exert a profound influence on elemental cycling and energy flow. The use of transmembrane chlorophyll complexes instead of phycobilisomes as light-harvesting antennae is considered a defining attribute of Prochlorococcus Its ecology and evolution are understood in terms of light, temperature, and nutrients. Here, we report single-cell genomic information on previously uncharacterized phylogenetic lineages of this genus from nutrient-rich anoxic waters of the eastern tropical North and South Pacific Ocean. The most basal lineages exhibit optical and genotypic properties of phycobilisome-containing cyanobacteria, indicating that the characteristic light-harvesting antenna of the group is not an ancestral attribute. Additionally, we found that all the indigenous lineages analyzed encode genes for pigment biosynthesis under oxygen-limited conditions, a trait shared with other freshwater and coastal marine cyanobacteria. Our findings thus suggest that Prochlorococcus diverged from other cyanobacteria under low-oxygen conditions before transitioning from phycobilisomes to transmembrane chlorophyll complexes and may have contributed to the oxidation of the ancient ocean.

Nitrification and Nitrous Oxide Production in the Offshore Waters of the Eastern Tropical South Pacific

AbstractMarine oxygen deficient zones are dynamic areas of microbial nitrogen cycling. Nitrification, the microbial oxidation of ammonia to nitrate, plays multiple roles in the biogeochemistry of these regions, including production of the greenhouse gas nitrous oxide (N2O). We present here the results of two oceanographic cruises investigating nitrification, nitrifying microorganisms, and N2O production and distribution from the offshore waters of the Eastern Tropical South Pacific. On each cruise, high‐resolution measurements of ammonium ([NH4+]), nitrite ([NO2−]), and N2O were combined with 15N tracer‐based determination of ammonia oxidation, nitrite oxidation, nitrate reduction, and N2O production rates. Depth‐integrated inventories of NH4+ and NO2− were positively correlated with one another and with depth‐integrated primary production. Depth‐integrated ammonia oxidation rates were correlated with sinking particulate organic nitrogen flux but not with primary production; ammonia oxidation rates were undetectable in trap‐collected sinking particulate material. Nitrite oxidation rates exceeded ammonia oxidation rates at most mesopelagic depths. We found positive correlations between archaeal amoA genes and ammonia oxidation rates and between Nitrospina‐like 16S rRNA genes and nitrite oxidation rates. N2O concentrations in the upper oxycline reached values of >140 nM, even at the western extent of the cruise track, supporting air‐sea fluxes of up to 1.71 μmol m−2 day−1. Our results suggest that a source of NO2− other than ammonia oxidation may fuel high rates of nitrite oxidation in the offshore Eastern Tropical South Pacific and that air‐sea fluxes of N2O from this region may be higher than previously estimated.