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  1. null (Ed.)
    Abstract. Oxygen-depleted regions of the global ocean are rapidly expanding, with important implications for global biogeochemical cycles. However, our ability to make projections about the future of oxygen in the ocean is limited by a lack of empirical data with which to test and constrain the behavior of global climatic and oceanographic models. We use depth-stratified plankton tows to demonstrate that some species of planktic foraminifera are adapted to life in the heart of the pelagic oxygen minimum zone (OMZ). In particular, we identify two species, Globorotaloides hexagonus and Hastigerina parapelagica, living within the eastern tropical North Pacific OMZ. The tests of the former are preserved in marine sediments and could be used to trace the extent and intensity of low-oxygen pelagic habitats in the fossil record. Additional morphometric analyses of G. hexagonus show that tests found in the lowest oxygen environments are larger, more porous, less dense, and have more chambers in the final whorl. The association of this species with the OMZ and the apparent plasticity of its test in response to ambient oxygenation invites the use of G. hexagonus tests in sediment cores as potential proxies for both the presence and intensity of overlying OMZs. 
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  2. null (Ed.)
    Climate warming is expected to intensify hypoxia in the California Current System (CCS), threatening its diverse and productive marine ecosystem. We analyzed past regional variability and future changes in the Metabolic Index (), a species-specific measure of the environment’s capacity to meet temperature-dependent organismal oxygen demand. Across the traits of diverse animals,  exhibits strong seasonal to interdecadal variations throughout the CCS, implying that resident species already experience large fluctuations in available aerobic habitat. For a key CCS species, northern anchovy, the long-term biogeographic distribution and decadal fluctuations in abundance are both highly coherent with aerobic habitat volume. Ocean warming and oxygen loss by 2100 are projected to decrease  below critical levels in 30 to 50% of anchovies’ present range, including complete loss of aerobic habitat—and thus likely extirpation—from the southern CCS. Aerobic habitat loss will vary widely across the traits of CCS taxa, disrupting ecological interactions throughout the region. 
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  3. null (Ed.)
    The capacity to extract oxygen from the environment and transport it to respiring tissues in support of metabolic demand reportedly has implications for species’ thermal tolerance, body size, diversity and biogeography. Here, we derived a quantifiable linkage between maximum and basal metabolic rate and their oxygen, temperature and size dependencies. We show that, regardless of size or temperature, the physiological capacity for oxygen supply precisely matches the maximum evolved demand at the highest persistently available oxygen pressure and this is the critical PO2 for the maximum metabolic rate, Pcrit-max. For most terrestrial and shallow-living marine species, Pcrit-max is the current atmospheric pressure, 21 kPa. Any reduction in oxygen partial pressure from current values will result in a calculable decrement in maximum metabolic performance. However, oxygen supply capacity has evolved to match demand across temperatures and body sizes and so does not constrain thermal tolerance or cause the well-known reduction in mass-specific metabolic rate with increasing body mass. The critical oxygen pressure for resting metabolic rate, typically viewed as an indicator of hypoxia tolerance, is, instead, simply a rate-specific reflection of the oxygen supply capacity. A compensatory reduction in maintenance metabolic costs in warm-adapted species constrains factorial aerobic scope and the critical PO2 to a similar range, between ∼2 and 6, across each species’ natural temperature range. The simple new relationship described here redefines many important physiological concepts and alters their ecological interpretation. 
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  4. null (Ed.)
    The Eastern Tropical North Pacific (ETNP) is a large, persistent, and intensifying oxygen minimum zone (OMZ) that accounts for almost half of the total area of global OMZs. Within the OMZ core (350–700 m depth), dissolved oxygen is typically near or below the analytical detection limit of modern sensors (10 nM). Steep oxygen gradients above and below the OMZ core lead to vertical structuring of microbial communities that also vary between particle-associated (PA) and free-living (FL) size fractions. Here, we use 16S amplicon sequencing (iTags) to analyze the diversity and distribution of prokaryotic populations between FL and PA size fractions and among the range of ambient redox conditions. The hydrographic conditions at our study area were distinct from those previously reported in the ETNP and other OMZs, such as the ETSP. Trace oxygen concentrations (0.35 mM) were present throughout the OMZ core at our sampling location. Consequently, nitrite accumulations typically reported for OMZ cores were absent as were sequences for anammox bacteria (Brocadiales genus Candidatus Scalindua), which are commonly found across oxic-anoxic boundaries in other systems. However, ammonia-oxidizing bacteria (AOB) and archaea (AOA) distributions and maximal autotrophic carbon assimilation rates (1.4 mM C d􀀀1) coincided with a pronounced ammonium concentration maximum near the top of the OMZ core. In addition, members of the genus Nitrospina, a dominant nitrite-oxidizing bacterial (NOB) clade were present suggesting that both ammonia and nitrite oxidation occur at trace oxygen concentrations. Analysis of similarity test (ANOSIM) and Non-metric Dimensional Scaling (nMDS) revealed that bacterial and archaeal phylogenetic representations were significantly different between size fractions. Based on ANOSIM and iTag profiles, composition of PA assemblages was less influenced by the prevailing depth-dependent biogeochemical regime than the FL fraction. Based on the presence of AOA, NOB and trace oxygen in the OMZ core we suggest that nitrification is an active process in the nitrogen cycle of this region of the ETNP OMZ. 
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  5. null (Ed.)
    Increasing deoxygenation (loss of oxygen) of the ocean, including expansion of oxygen minimum zones (OMZs), is a potentially important consequence of global warming. We examined present-day variability of vertical distributions of 23 calanoid copepod species in the Eastern Tropical North Pacific (ETNP) living in locations with different water column oxygen profiles and OMZ intensity (lowest oxygen concentration and its vertical extent in a profile). Copepods and hydrographic data were collected in vertically stratified day and night MOCNESS (Multiple Opening/Closing Net and Environmental Sensing System) tows (0–1000 m) during four cruises over a decade (2007– 2017) that sampled four ETNP locations: Costa Rica Dome, Tehuantepec Bowl, and two oceanic sites further north (21– 22 N) off Mexico. The sites had different vertical oxygen profiles: some with a shallow mixed layer, abrupt thermocline, and extensive very low oxygen OMZ core; and others with a more gradual vertical development of the OMZ (broad mixed layer and upper oxycline zone) and a less extensive OMZ core where oxygen was not as low. Calanoid copepod species (including examples from the genera Eucalanus, Pleuromamma, and Lucicutia) demonstrated different distributional strategies (implying different physiological characteristics) associated with this variability. We identified sets of species that (1) changed their vertical distributions and depth of maximum abundance associated with the depth and intensity of the OMZ and its oxycline inflection points; (2) shifted their depth of diapause; (3) adjusted their diel vertical migration, especially the nighttime upper depth; or (4) expanded or contracted their depth range within the mixed layer and upper part of the thermocline in association with the thickness of the aerobic epipelagic zone (habitat compression concept). These distribution depths changed by tens to hundreds of meters depending on the species, oxygen profile, and phenomenon. For example, at the lower oxycline, the depth of maximum abundance for Lucicutia hulsemannae shifted from  600 to  800 m, and the depth of diapause for Eucalanus inermis shifted from  500 to  775 m, in an expanded OMZ compared to a thinner OMZ, but remained at similar low oxygen levels in both situations. These species or life stages are examples of “hypoxiphilic” taxa. For the migrating copepod Pleuromamma abdominalis, its nighttime depth was shallow ( 20 m) when the aerobic mixed layer was thin and the low-oxygen OMZ broad, but it was much deeper ( 100 m) when the mixed layer and higher oxygen extended deeper; daytime depth in both situations was  300 m. Because temperature decreased with depth, these distributional depth shifts had metabolic implications. The upper ocean to mesopelagic depth range encompasses a complex interwoven ecosystem characterized by intricate relationships among its inhabitants and their environment. It is a critically important zone for oceanic biogeochemical and export processes and hosts key food web components for commercial fisheries. Among the zooplankton, there will likely be winners and losers with increasing ocean deoxygenation as species cope with environmental change. Changes in individual copepod species abundances, vertical distributions, and life history strategies may create potential perturbations to these intricate food webs and processes. Present-day variability provides a window into future scenarios and potential effects of deoxygenation. 
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  6. The Wire Flyer towed vehicle is a new platform able to collect high-resolution water column sections. The vehicle is motivated by a desire to effectively capture spatial structures at the submesoscale. The vehicle fills a niche that is not achieved by other existing towed and repeat profiling systems. The Wire Flyer profiles up and down along a ship-towed cable autonomously using controllable wings for propulsion. At ship speeds between 2 and 5 kt (1.02–2.55ms21), the vehicle is able to profile over prescribed depth bands down to 1000 m. The vehicle carries sensors for conductivity, temperature, depth, oxygen, turbidity, chlorophyll, pH, and oxidation reduction potential. During normal operations the vehicle is typically commanded to cover vertical regions between 300 and 400min height with profiles that repeat at kilometer spacing. The vertical profiling speed can be user specified up to 150mmin21. The high-density sampling capability at depths below the upper few hundred meters makes the vehicle distinct from other systems. During operations an acoustic modem is used to communicate with the vehicle to provide status information, data samples, and the ability to modify the sampling pattern. This paper provides an overview of the vehicle system, describes its operation, and presents results from several cruises. 
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  7. Oxygen minimum zones (OMZs), large midwater regions of very low oxygen, are expected to expand as a result of climate change. While oxygen is known to be important in structuring midwater ecosystems, a precise and mechanistic understanding of the effects of oxygen on zooplankton is lacking. Zooplankton are important components of midwater food webs and biogeochemical cycles. Here, we show that, in the eastern tropical North Pacific OMZ, previously undescribed submesoscale oxygen variability has a direct effect on the distribution of many major zooplankton groups. Despite extraordinary hypoxia tolerance, many zooplankton live near their physiological limits and respond to slight (≤1%) changes in oxygen. Ocean oxygen loss (deoxygenation) may, thus, elicit major unanticipated changes to midwater ecosystem structure and function. 
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  8. The pelagic red crab, Pleuroncodes planipes, is abundant throughout the Eastern Tropical Pacific in both benthic and pelagic environments to depths of several hundred meters. The oxygen minimum zones in this region reaches oxygen levels as low as 0.1 kPa at depths within the crabs vertical range. Crabs maintain aerobic metabolism to a critical PO2 of ~0.27 ± 0.2 kPa (10 °C), in part by increasing ventilation as oxygen declines. At subcritical oxygen levels, they enhance anaerobic ATP production slightly as indicated by modest increases in lactate levels. However, hypoxia tolerance is primarily mediated via a pronounced suppression of aerobic metabolism (~70%). Metabolic suppression is achieved, primarily, via reduced protein synthesis, which is a major sink for metabolic energy. Posttranslational modifications on histone H3 suggest a condensed chromatin state and, hence, decreased transcription. Under hypoxia, p-H3S10, Ac-H3K9, Ac-H3K14 were 39, 68, and 36% of control values, respectively. We also report a net decrease in protein translation. In particular, eEF2 activity is reduced due to a ~5-fold increase in inhibitory phosphorylation and a significant decrease in protein level. Elevated heat shock proteins suggest that, despite impressive tolerance, the cellular stress response is triggered during hypoxia. We discuss the implications for pelagic ecology and biogeochemical cycles. 
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  9. The effects of regional variations in oxygen and temperature levels with depth were assessed for the metabolism and hypoxia tolerance of dominant euphausiid species. The physiological strategies employed by these species facilitate prediction of changing vertical distributions with expanding oxygen minimum zones and inform estimates of the contribution of vertically migrating species to biogeochemical cycles. The migrating species from the Eastern Tropical Pacific (ETP), Euphausia eximia and Nematoscelis gracilis, tolerate a Partial Pressure (PO2) of 0.8 kPa at 10 8C (15 mM O2) for at least 12 h without mortality, while the California Current species, Nematoscelis difficilis, is incapable of surviving even 2.4 kPa PO2 (32 mM O2) for more than 3 h at that temperature. Euphausia diomedeae from the Red Sea migrates into an intermediate oxygen minimum zone, but one in which the temperature at depth remains near 22 8C. Euphausia diomedeae survived 1.6 kPa PO2 (22 mM O2) at 228C for the duration of six hour respiration experiments. Critical oxygen partial pressures were estimated for each species, and, for E. eximia, measured via oxygen consumption (2.1 kPa, 10 8C, n¼2) and lactate accumulation (1.1 kPa, 10 8C). A primary mechanism facilitating low oxygen tolerance is an ability to dramatically reduce energy expenditure during daytime forays into low oxygen waters. The ETP and Red Sea species reduced aerobic metabolism by more than 50% during exposure to hypoxia. Anaerobic glycolytic energy production, as indicated by whole-animal lactate accumulation, contributed only modestly to the energy deficit. Thus, the total metabolic rate was suppressed by 49–64%. Metabolic suppression during diel migrations to depth reduces the metabolic contribution of these species to vertical carbon and nitrogen flux (i.e., the biological pump) by an equivalent amount. Growing evidence suggests that metabolic suppression is a widespread strategy among migrating zooplankton in oxygen minimum zones and may have important implications for the economy and ecology of the oceans. The interacting effects of oxygen and temperature on the metabolism of oceanic species facilitate predictions of changing vertical distribution with climate change. 
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