Many predator species make regular excursions from near-surface waters to the twilight (200 to 1,000 m) and midnight (1,000 to 3,000 m) zones of the deep pelagic ocean. While the occurrence of significant vertical movements into the deep ocean has evolved independently across taxonomic groups, the functional role(s) and ecological significance of these movements remain poorly understood. Here, we integrate results from satellite tagging efforts with model predictions of deep prey layers in the North Atlantic Ocean to determine whether prey distributions are correlated with vertical habitat use across 12 species of predators. Using 3D movement data for 344 individuals who traversed nearly 1.5 million km of pelagic ocean in >42,000 d, we found that nearly every tagged predator frequented the twilight zone and many made regular trips to the midnight zone. Using a predictive model, we found clear alignment of predator depth use with the expected location of deep pelagic prey for at least half of the predator species. We compared high-resolution predator data with shipboard acoustics and selected representative matches that highlight the opportunities and challenges in the analysis and synthesis of these data. While not all observed behavior was consistent with estimated prey availability at depth, our results suggest that deep pelagic biomass likely has high ecological value for a suite of commercially important predators in the open ocean. Careful consideration of the disruption to ecosystem services provided by pelagic food webs is needed before the potential costs and benefits of proceeding with extractive activities in the deep ocean can be evaluated.

Marine phytoplankton are a diverse group of photoautotrophic organisms and key mediators in the global carbon cycle. Phytoplankton physiology and biomass accumulation are closely tied to mixed layer depth, but the intracellular metabolic pathways activated in response to changes in mixed layer depth remain less explored. Here, metatranscriptomics was used to characterize the phytoplankton community response to a mixed layer shallowing (from 233 to 5 m) over the course of two days during the late spring in the Northwest Atlantic. Most phytoplankton genera downregulated core photosynthesis, carbon storage, and carbon fixation genes as the system transitioned from a deep to a shallow mixed layer and shifted towards catabolism of stored carbon supportive of rapid cell growth. In contrast, phytoplankton genera exhibited divergent transcriptional patterns for photosystem light harvesting complex genes during this transition. Active virus infection, taken as the ratio of virus to host transcripts, increased in the Bacillariophyta (diatom) phylum and decreased in the Chlorophyta (green algae) phylum upon mixed layer shallowing. A conceptual model is proposed to provide ecophysiological context for our findings, in which integrated light limitation and lower division rates during transient deep mixing are hypothesized to disrupt resource-driven, oscillating transcript levels related to photosynthesis, carbon fixation, and carbon storage. Our findings highlight shared and unique transcriptional response strategies within phytoplankton communities acclimating to the dynamic light environment associated with transient deep mixing and shallowing events during the annual North Atlantic bloom.

The straining regions of the ocean in between mesoscale eddies contain large vertical velocities that may be important in regulating phytoplankton accumulation rates. We analyze time series of variables measured by ocean Bio-Argo floats (mixed layer depths [MLDs], chlorophyll, and carbon concentrations) in conjunction with variables derived from satellite altimetry (strain rates, Lyapunov exponents, vertical velocities) to determine the evolution of mixed layer phytoplankton biomass in response to straining by the mesoscale geostrophic flow. A Lagrangian (water parcel following) framework is justified by restricting the analysis to profiles whose value of a Quasi-Planktonic Index — an index quantifying averaged distance between a float trajectory and a surface geostrophic trajectory over three consecutive time steps — is less than 5 km. Bin-averaged Lagrangian derivatives of phytoplankton biomass and chlorophyll concentration are positive for elevated strain rate and upwelling quasigeostrophic vertical velocities. Lagrangian derivatives of MLD and phytoplankton carbon averaged at straining fronts (in rotated along- and across-front coordinates) have features in common with submesoscale dynamics, including increasing phytoplankton carbon (and chlorophyll) and a shoaling mixed layer over the front. To elucidate a mechanism, we average time derivatives of modeled cell division rates, finding the pattern approximately matches the pattern of phytoplankton accumulation rates and is controlled primarily by the term modulating light stress, suggesting that frontal dynamics cause accelerations of division rates by increasing available light. Regions of increasing chlorophyll are also approximately co-located with upwelling quasigeostrophic velocity, suggesting non-Lagrangian behavior of floats causes some imprint of larger scale, more persistent mesoscale signals.

Phytoplankton form the base of marine food webs and play an important role in carbon cycling, making it important to quantify rates of biomass accumulation and loss. As phytoplankton drift with ocean currents, rates should be evaluated in a Lagrangian as opposed to an Eulerian framework. In this study, we quantify the Lagrangian (from Bio-Argo floats and surface drifters with satellite ocean colour) and Eulerian (from satellite ocean colour and altimetry) statistics of mesoscale chlorophyll and velocity by computing decorrelation time and length scales and relate the frames by scaling the material derivative of chlorophyll. Because floats profile vertically and are not perfect Lagrangian observers, we quantify the mean distance between float and surface geostrophic trajectories over the time spanned by three consecutive profiles (quasi-planktonic index, QPI) to assess how their sampling is a function of their deviations from surface motion. Lagrangian and Eulerian statistics of chlorophyll are sensitive to the filtering used to compute anomalies. Chlorophyll anomalies about a 31 d time filter reveal an approximate equivalence of Lagrangian and Eulerian tendencies, suggesting they are driven by ocean colour pixel-scale processes and sources or sinks. On the other hand, chlorophyll anomalies about a seasonal cycle have Eulerian scales similar to those of velocity, suggesting mesoscale stirring helps set distributions of biological properties, and ratios of Lagrangian to Eulerian timescales depend on the magnitude of velocity fluctuations relative to an evolution speed of the chlorophyll fields in a manner similar to earlier theoretical results for velocity scales. The results suggest that stirring by eddies largely sets Lagrangian time and length scales of chlorophyll anomalies at the mesoscale.

Mesoscale eddies shape the foraging ecology of predators such as marine mammals and seabirds. A growing number of animal tracking studies show that predators alter their swimming, diving, and foraging behavior within mesoscale eddies. However, little is known about how Southern Ocean eddies influence the distribution of mesopelagic micronekton (fish, squid, and crustaceans), which are major prey items of megafauna. Studies in other oceanic regions have found that eddies can influence the abundance and community composition of micronekton. Here, we analyze acoustic observations from a 14-day survey of a cyclonic mesoscale eddy, its surrounding waters, and the Polar Frontal Zone (PFZ) waters where the eddy formed. We report and interpret spatial patterns of acoustic backscatter at 18 and 75 kHz, proxies indicating combined changes in species, size, and abundance of micronekton. We find that the vertical distribution of acoustic backscatter matched the underwater light conditions characteristic of the eddy core, periphery, and surrounding waters, at scales smaller than 10 km. The median water-column integrated acoustic backscatter values in the eddy core were only half of those measured in the Sub-Antarctic Zone waters surrounding the eddy, but similar to those measured in the PFZ, where the eddy originated 27 days prior. These results suggest that, as for physical and chemical tracers, the eddy maintained its biological characteristics from its source waters creating a unique habitat compared to its surroundings.

Under most natural marine conditions, phytoplankton cells suspended in the water column are too distantly spaced for direct competition for resources (i.e., overlapping cell boundary layers) to be a routine occurrence. Accordingly, resource-based competitive exclusion should be rare. In contrast, contemporary ecosystem models typically predict an exclusion of larger phytoplankton size classes under low-nutrient conditions, an outcome interpreted as reflecting the competitive advantage of small cells having much higher nutrient 'affinities' than larger cells. Here, we develop mechanistically-focused expressions for steady-state, nutrient-limited phytoplankton growth that are consistent with the discrete, distantly-spaced cells of natural populations. These expressions, when encompassed in a phytoplankton-zooplankton model, yield sustained diversity across all size classes over the full range in nutrient concentrations observed in the ocean. In other words, our model does not exhibit resource-based competitive exclusion between size classes previously associated with size-dependent differences in nutrient 'affinities'.

Ocean eddies are coherent, rotating features that can modulate pelagic ecosystems across many trophic levels. These mesoscale features, which are ubiquitous at mid-latitudes, may increase productivity of nutrient-poor regions, accumulate prey and modulate habitat conditions in the water column. However, in nutrient-poor subtropical gyres—the largest marine biome—the role of eddies in modulating behaviour throughout the pelagic predator community remains unknown despite predictions for these gyres to expand and pelagic predators to become increasingly important for food security. Using a large-scale fishery dataset in the North Pacific Subtropical Gyre, we show a pervasive pattern of increased pelagic predator catch inside anticyclonic eddies relative to cyclones and non-eddy areas. Our results indicate that increased mesopelagic prey abundance in anticyclone cores may be attracting diverse predators, forming ecological hotspots where these predators aggregate and exhibit increased abundance. In this energetically quiescent gyre, we expect that isolated mesoscale features (and the habitat conditions in them) exhibit primacy over peripheral submesoscale dynamics in structuring the foraging opportunities of pelagic predators. Our finding that eddies influence coupling of epi- to mesopelagic communities corroborates the growing evidence that deep scattering layer organisms are vital prey for a suite of commercially important predator species and, thus, provide valuable ecosystem services.

Dissolved organic carbon (DOC) produced by primary production in the sunlit ocean can be physically transported to the mesopelagic zone. The majority of DOC exported to this zone is remineralized by heterotrophic microbes over a range of timescales. Capturing a deep convective mixing event is rare, as is observing how microbes respond in situ to the exported DOC. Here, we report ship and Argo float observations from hydrostation North Atlantic Aerosols and Marine Ecosystems Study (NAAMES) 2 Station 4 (N2S4; 47.46°N, 38.72°W), a retentive anticyclonic eddy in the subtropical region of the western North Atlantic. Changes in biogeochemistry and bacterioplankton responses were tracked as the water column mixed to approximately 230 m and restratified over the subsequent 3 days. Over this period, rapid changes in bacterioplankton production (BP) and cell abundance were observed throughout the water column. BP increased by 91% in the euphotic zone (0–100 m) and 55% in the upper mesopelagic zone (100–200 m), corresponding to 33% and 103% increases in cell abundance, respectively. Within the upper mesopelagic, BP upon the occupation of N2S4 (20 ± 4.7 nmol C L^-1 d^-1) was significantly greater than the average upper mesopelagic BP rate (2.0 ± 1.6 nmol C L^-1 h^-1) at other stations that had been stratified for longer periods of time. BP continued to increase to 31 ± 3.0 nmol C L^-1 d^-1 over the 3-day occupation of N2S4. The rapid changes in BP in the upper mesopelagic did not coincide with rapid changes in community composition, but the taxa that increased in their relative contribution included those typically observed in the epipelagic zone. We interpret the subtle but significant community structure dynamics at N2S4 to reflect how injection of labile organic matter into the upper mesopelagic zone by physical mixing supports continued growth of euphotic zone-associated bacterioplankton lineages on a timescale of days.

Atmospheric marine particle concentrations impact cloud properties, which strongly impact the amount of solar radiation reflected back into space or absorbed by the ocean surface. While satellites can provide a snapshot of current conditions at the overpass time, models are necessary to simulate temporal variations in both particle and cloud properties. However, poor model accuracy limits the reliability with which these tools can be used to predict future climate. Here, we leverage the comprehensive ocean ecosystem and atmospheric aerosol–cloud dataset obtained during the third deployment of the North Atlantic Aerosols and Marine Ecosystems Study (NAAMES3). Airborne and ship-based measurements were collected in and around a cold-air outbreak during a 3 d (where d stands for day) intensive operations period from 17–19 September 2017. Cold-air outbreaks are of keen interest for model validation because they are challenging to accurately simulate, which is due, in part, to the numerous feedbacks and sub-grid-scale processes that influence aerosol and cloud evolution. The NAAMES observations are particularly valuable because the flight plans were tailored to lie along Lagrangian trajectories, making it possible to spatiotemporally connect upwind and downwind measurements with the state-of-the-art FLEXible PARTicle (FLEXPART) Lagrangian particle dispersion model and then calculate a rate of change in particle properties. Initial aerosol conditions spanning an east–west, closed-cell-to-clear-air transition region of the cold-air outbreak indicate similar particle concentrations and properties. However, despite the similarities in the aerosol fields, the cloud properties downwind of each region evolved quite differently. One trajectory carried particles through a cold-air outbreak, resulting in a decrease in accumulation mode particle concentration (–42 %) and cloud droplet concentrations, while the other remained outside of the cold-air outbreak and experienced an increase in accumulation mode particle concentrations (+62 %). The variable meteorological conditions between these two adjacent trajectories result from differences in the local sea surface temperature in the Labrador Current and surrounding waters, altering the stability of the marine atmospheric boundary layer. Further comparisons of historical satellite observations indicate that the observed pattern occurs annually in the region, making it an ideal location for future airborne Lagrangian studies tracking the evolution of aerosols and clouds over time under cold-air outbreak conditions.

Many large marine predators make excursions from surface waters to the deep ocean below 200 m. Moreover, the ability to access meso- and bathypelagic habitats has evolved independently across marine mammals, reptiles, birds, teleost fishes, and elasmobranchs. Theoretical and empirical evidence suggests a number of plausible functional hypotheses for deep-diving behavior. Developing ways to test among these hypotheses will, however, require new ways to quantify animal behavior and biophysical oceanographic processes at coherent spatiotemporal scales. Current knowledge gaps include quantifying ecological links between surface waters and mesopelagic habitats and the value of ecosystem services provided by biomass in the ocean twilight zone. Growing pressure for ocean twilight zone fisheries creates an urgent need to understand the importance of the deep pelagic ocean to large marine predators.

Seasonal shifts in phytoplankton accumulation and loss largely follow changes in mixed layer depth, but the impact of mixed layer depth on cell physiology remains unexplored. Here, we investigate the physiological state of phytoplankton populations associated with distinct bloom phases and mixing regimes in the North Atlantic. Stratification and deep mixing alter community physiology and viral production, effectively shaping accumulation rates. Communities in relatively deep, early-spring mixed layers are characterized by low levels of stress and high accumulation rates, while those in the recently shallowed mixed layers in late-spring have high levels of oxidative stress. Prolonged stratification into early autumn manifests in negative accumulation rates, along with pronounced signatures of compromised membranes, death-related protease activity, virus production, nutrient drawdown, and lipid markers indicative of nutrient stress. Positive accumulation renews during mixed layer deepening with transition into winter, concomitant with enhanced nutrient supply and lessened viral pressure.

Ocean ecosystems play a central role in the vitality of our biosphere and both influence and are influenced by the overlying atmosphere. Plankton-based compounds released from the ocean through bubble bursting and wave breaking contribute to atmospheric aerosol load and influence the formation and properties of clouds (Meskhidze et al., 2013). In parallel, ocean ecosystems are recipients of atmospheric depositions, often transported from distant sources (van de Meent et al., 2011). Ocean-atmosphere feedbacks are particularly amplified when pristine atmospheres overlay strong annual plankton cycles. The western subarctic Atlantic is one such place. The North Atlantic Aerosol and Marine Ecosystem Study (NAAMES) was a National Aeronautics and Space Administration (NASA) Earth Venture Suborbital mission focused on plankton blooms, aerosols, and clouds in this subarctic region (~40–55°N) and conducted between 2015 and 2019. The mission entailed four field campaigns targeting specific events in the annual plankton cycle and included a diversity of ship-based, airborne, and autonomous measurements (Behrenfeld et al.). At the time of writing, nearly 80 publications stem from NAAMES (21 represented within this frontiers research topic) and more are in development. This Editorial provides a brief synopsis of many of the NAAMES scientific findings.

We investigate how the near‐surface chlorophyll‐a (CHL) evolves in Gulf Stream (GS) warm‐core rings (WCRs) and cold‐core rings (CCRs) using multi‐platform satellite observations. Averaged CHL anomaly (CHLA) within the rings exhibits both positive and negative linear trends during the evolution of the WCRs while negative trends dominate in CCRs. This difference is associated with a variety of physical processes occurring during the evolution process. Meanwhile, eddy‐centric analysis reveals four spatial patterns of CHLA long‐term trends, some of which highlights the importance of rings in shaping surface CHL. Short‐term fluctuations of CHLA in WCRs and CCRs are closely correlated with mixed layer depth (MLD) and sea surface temperature anomaly (SSTA) and highlight the complex interplay between multiple mechanisms. In addition, we find higher concentration CHL in some WCRs than that in CCRs during the same season, providing an alternative view of the characteristics of the surface ecosystem in Gulf Stream rings.

As the resolution of observations and models improves, emerging evidence indicates that ocean variability on 1–200 km scales is of fundamental importance to ocean circulation, air‐sea interaction, and biogeochemistry. In many regions, salinity variability dominates over thermal effects in forming density fronts. Unfortunately, current satellite observations of sea surface salinity (SSS) only resolve scales ࣙ40 km (or larger, depending on the product). In this study we investigate small‐scale variability (ࣘ25 km) by reconstructing gridded SSS observations made by the Soil Moisture Active Passive (SMAP) satellite in the northwest Atlantic Ocean. Using altimetric geostrophic currents, we numerically advect SMAP SSS fields to produce a Lagrangian reconstruction that represents small scales. Reconstructed fields are compared to in situ salinity observations made by a ship‐board thermosalinograph, revealing a marked improvement in small‐scale salinity variability when compared to the original SMAP fields, particularly from the continental shelf to the Gulf Stream. In the Sargasso Sea, however, both SMAP and the reconstructed fields contain higher variability than is observed in situ. Enhanced small‐scale salinity variability is concentrated in two bands: a northern band aligned with the continental shelfbreak, and a southern band aligned with the Gulf Stream mean position. Seasonal differences in the small‐scale variability appear to covary with the seasonal cycle of the large‐scale SSS gradients resulting from the freshening of the coastal waters during periods of elevated river outflow.

The pelagic thresher shark Alopias pelagicus is an understudied elasmobranch harvested in commercial fisheries of the tropical Indo-Pacific. The species is endangered, overexploited throughout much of its range, and has a decreasing population trend. Relatively little is known about its movement ecology, precluding an informed recovery strategy. Here, we report the first results from an individual pelagic thresher shark outfitted with a pop-up satellite archival transmitting (PSAT) tag to assess its movement with respect to the species' physiology and trophic ecology. A 19 d deployment in the Red Sea revealed that the shark conducted normal diel vertical migration, spending the majority of the day at 200-300 m in the mesopelagic zone and the majority of the night at 50–150 m in the epipelagic zone, with the extent of these movements seemingly not constrained by temperature. In contrast, the depth distribution of the shark relative to the vertical distribution of oxygen suggested that it was avoiding hypoxic conditions below 300 m even though that is where the daytime peak of acoustic backscattering occurs in the Red Sea. Telemetry data also indicated crepuscular and daytime overlap of the shark’s vertical habitat use with distinct scattering layers of small mesopelagic fishes and nighttime overlap with nearly all mesopelagic organisms in the Red Sea as these similarly undergo nightly ascents into epipelagic waters. We identify potential depths and diel periods in which pelagic thresher sharks may be most susceptible to fishery interactions, but more expansive research efforts are needed to inform effective management.

Ocean sunfishes or molas (Molidae) are difficult to study as a result of their extensive movements and low densities in remote waters. In particular, little is known of the environmental niche separation and differences in the reproductive or movement ecology of molids in sympatry. We investigated spatiotemporal dynamics in the distribution of the common mola Mola mola, sharptail mola Masturus lanceolatus, and slender mola Ranzania laevis in the eastern North Pacific. We used observer data from a commercial fishery consisting of 85000+ longline sets spanning 24 yr, >50° in longitude, and >45° in latitude. Satellite altimetry analysis, species distribution modeling, and multivariate ordination revealed thermal niche separation, spatiotemporal segregation, and distinct community associations of the 3 molid species. Our quantitative findings suggest that the common mola is a more temperate species, while slender and sharptail mola are more (sub)tropical species, and that slender (and possibly also sharptail) mola undergo spawning migrations to the region around the Hawaiian Islands. In addition, we identified potential effects of fishing gear type on molid catch probability, an increasing trend in catch probability of a vulnerable species perhaps related to a shift in the distribution of fishing effort, and the possible presence in the fishery of a fourth molid species being misidentified as a congener, all of which are important conservation considerations for these enigmatic fishes.

Mesoscale eddies play a key role in structuring open ocean ecosystems, affecting the entire trophic web from primary producers to large pelagic predators including sharks and elephant seals. Recent advances in the tracking of pelagic predators have revealed that these animals forage in the mesopelagic and the depth and duration of their foraging dives are affected by the presence of eddies. The ways in which eddies impact the distribution of mesopelagic micronekton, however, remain largely unknown. During a multi-seasonal experiment we used a shipboard scientific echosounder transmitting at 38 kHz to observe the distribution of acoustic backscattering in the energetic mesoscale eddy field of the northwestern Atlantic. Observations were collected at 24 stations with 6 located in anticyclonic and 7 in cyclonic eddies. The sampled anticyclonic eddies are characterized by intense acoustic backscattering in the mesopelagic and changes in the intensity of acoustic backscattering layers match gradients of surface properties. Furthermore, mesopelagic daytime backscattering is positively correlated with sea level anomaly. These results suggest that anticyclonic eddies in the northwestern Atlantic impact the distribution of mesopelagic micronekton and may have the potential to locally enhance or structure spatially mesopelagic communities.

Changes in airborne high spectral resolution lidar (HSRL) measurements of scattering, depolarization, and attenuation coincided with a shift in phytoplankton community composition across an anticyclonic eddy in the North Atlantic. We normalized the total depolarization ratio (δ) by the particulate backscattering coefficient (b_bp) to account for the covariance in δ and b_bp that has been attributed to multiple scattering. A 15% increase in δ/b_bp inside the eddy coincided with decreased phytoplankton biomass and a shift to smaller and more elongated phytoplankton cells. Taxonomic changes (reduced dinoflagellate relative abundance inside the eddy) were also observed. The δ signal is thus potentially most sensitive to changes in phytoplankton shape because neither the observed change in the particle size distribution (PSD) nor refractive index (assuming average refractive indices) are consistent with previous theoretical modeling results. We additionally calculated chlorophyll-a (Chl) concentrations from measurements of the diffuse light attenuation coefficient (K_d) and divided by b_bp to evaluate another optical metric of phytoplankton community composition (Chl:b_bp), which decreased by more than a factor of two inside the eddy. This case study demonstrates that the HSRL is able to detect changes in phytoplankton community composition. High spectral resolution lidar measurements reveal complex structures in both the vertical and horizontal distribution of phytoplankton in the mixed layer providing a valuable new tool to support other remote sensing techniques for studying mixed layer dynamics. Our results identify fronts at the periphery of mesoscale eddies as locations of abrupt changes in near-surface optical properties.

We examine the effects of Southern Ocean eddies on phytoplankton cell division rates in a global, multiyear, eddy‐resolving, 3‐D ocean simulation of the Community Earth System Model. We first identify and track eddies in the simulation and validate their distribution and demographics against observed eddy trajectory characteristics. Next, we examine how simulated cyclones and anticyclones differentially modify iron, light, and ultimately population‐specific cell division rates. We use an eddy‐centric, depth‐averaged framework to explicitly examine the dynamics of the phytoplankton population across the entire water column within an eddy. We find that population‐averaged iron availability is elevated in anticyclones throughout the year. The dominant mechanism responsible for vertically transporting iron from depth in anticyclones is eddy‐induced Ekman upwelling. During winter, in regions with deep climatological mixed layer depths, anticyclones also induce anomalously deep mixed layer depths, which further supply new iron from depth via an increased upward mixing flux. However, this additional contribution comes at the price of deteriorating light availability as biomass is distributed deeper in the water column. Therefore, even though population‐averaged specific division rates are elevated in Southern Ocean anticyclones throughout most of the year, in the winter, severe light stress can dominate relieved iron stress and lead to depressed division rates in some anticyclones, particularly in the deep mixing South Pacific Antarctic Circumpolar Current. The opposite is true in cyclones, which exhibit a consistently symmetric physical and biogeochemical response relative to anticyclones.

We examine various contributions to the vertical velocity field within large mesoscale eddies by analyzing multiple solutions to an idealized numerical model of a representative anticyclonic warm core Gulf Stream ring. Initial conditions are constructed to reproduce the observed density and nutrient profiles collected during the Warm Core Rings Program. The contributions to vertical fluxes diagnosed from the numerical simulations are compared against a divergence-based, semidiagnostic equation and a generalized omega equation to better understand the dynamics of the vertical velocity field. Frictional decay alone is found to be ineffective in raising isopycnals and transporting nutrients to the upper ocean. With representative wind forcing, the magnitude of vorticity gradient–induced Ekman pumping is not necessarily larger than the current-induced counterpart on a time scale relevant to ecosystem response. Under realistic forcing conditions, strain deformation can perturb the ring to be noncircular and induce vertical velocities much larger than the Ekman vertical velocities. Nutrient budget diagnosis, together with analysis of the relative magnitudes of the various types of vertical fluxes, allows us to describe the time-scale dependence of nutrient delivery. At time scales that are relevant to individual phytoplankton (from hours to days), the magnitudes of nutrient flux by Ekman velocities and deformation-induced velocities are comparable. Over the life span of a typical warm core ring, which can span multiple seasons, surface current–induced Ekman pumping is the most effective mechanism in upper-ocean nutrient enrichment because of its persistence in the center of anticyclones regardless of the direction of the wind forcing.

We examine the structure and drivers of anomalous phytoplankton biomass in Southern Ocean eddies tracked in a global, multiyear, eddy‐resolving, 3‐D ocean simulation of the Community Earth System Model. We examine how simulated anticyclones and cyclones differentially modify phytoplankton biomass concentrations, growth rates, and physical transport. On average, cyclones induce negative division rate anomalies that drive negative net population growth rate anomalies, reduce dilution across shallower mixed layers, and advect biomass anomalously downward via eddy‐induced Ekman pumping. The opposite is true in anticyclones. Lateral transport is dominated by eddy stirring rather than eddy trapping. The net effect on anomalous biomass can exceed 10–20% of background levels at the regional scale, consistent with observations. Moreover, we find a strong seasonality in the sign and magnitude of regional anomalies and the processes that drive them. The most dramatic seasonal cycle is found in the South Pacific Antarctic Circumpolar Current, where physical and biological processes dominate at different times, modifying biomass in different directions throughout the year. Here, in cyclones, during winter, anomalously shallow mixed layer depths first drive positive surface biomass anomalies via reduced dilution, and later drive positive depth‐integrated biomass anomalies via reduced light limitation. During spring, reduced iron availability and elevated grazing rates suppress net population growth rates and drive the largest annual negative surface and depth‐integrated biomass anomalies. During summer and fall, lateral stirring and eddy‐induced Ekman pumping create small negative surface anomalies but positive depth‐integrated anomalies. The same mechanisms drive biomass anomalies in the opposite direction in anticyclones.

The North Atlantic phytoplankton spring bloom is the pinnacle in an annual cycle that is driven by physical, chemical, and biological seasonality. Despite its important contributions to the global carbon cycle, transitions in plankton community composition between the winter and spring have been scarcely examined in the North Atlantic. Phytoplankton composition in early winter was compared with latitudinal transects that captured the subsequent spring bloom climax. Amplicon sequence variants (ASVs), imaging flow cytometry, and flow-cytometry provided a synoptic view of phytoplankton diversity. Phytoplankton communities were not uniform across the sites studied, but rather mapped with apparent fidelity onto subpolar- and subtropical-influenced water masses of the North Atlantic. At most stations, cells < 20-μm diameter were the main contributors to phytoplankton biomass. Winter phytoplankton communities were dominated by cyanobacteria and pico-phytoeukaryotes. These transitioned to more diverse and dynamic spring communities in which pico- and nano-phytoeukaryotes, including many prasinophyte algae, dominated. Diatoms, which are often assumed to be the dominant phytoplankton in blooms, were contributors but not the major component of biomass. We show that diverse, small phytoplankton taxa are unexpectedly common in the western North Atlantic and that regional influences play a large role in modulating community transitions during the seasonal progression of blooms.

During the North Atlantic Aerosols and Marine Ecosystems Study in the western North Atlantic, float‐based profiles of fluorescent dissolved organic matter and backscattering exhibited distinct spike layers at ~ 300 m. The locations of the spikes were at depths similar or shallower to where a ship‐based scientific echo sounder identified layers of acoustic backscatter, an Underwater Vision Profiler detected elevated concentration of zooplankton, and mesopelagic fish were sampled by a mesopelagic net tow. The collocation of spike layers in bio‐optical properties with mesopelagic organisms suggests that some can be detected with float‐based bio‐optical sensors. This opens the door to the investigation of such aggregations/layers in observations collected by the global biogeochemical‐Argo array allowing the detection of mesopelagic organisms in remote locations of the open ocean under‐sampled by traditional methods.

Results from our 2017 cruise to the Santa Barbara Channel illustrate the value that student leadership training can bring to ocean science. The Across the Channel: Investigating Diel Dynamics (ACIDD) mission, conducted from December 16 to 22, 2017, aboard R/V Sally Ride, was led by two PhD students as co-principal investigators and chief scientists (authors Bisson and Baetge). The 21-​member science team was composed almost entirely of our graduate student peers at the University of California, Santa Barbara (UCSB), as well as three artists. As an integrated team, we conceived, adapted, and executed research cruise plans and developed far-​reaching connections with the public based on our coupled artistic-​oceanographic pursuit.

Every night across the world's oceans, numerous marine animals arrive at the surface of the ocean to feed on plankton after an upward migration of hundreds of metres. Just before sunrise, this migration is reversed and the animals return to their daytime residence in the dark mesopelagic zone (at a depth of 200–1,000 m). This daily excursion, referred to as diel vertical migration (DVM), is thought of primarily as an adaptation to avoid visual predators in the sunlit surface layer and was first recorded using ship-net hauls nearly 200 years ago. Nowadays, DVMs are routinely recorded by ship-mounted acoustic systems (for example, acoustic Doppler current profilers). These data show that night-time arrival and departure times are highly conserved across ocean regions and that daytime descent depths increase with water clarity, indicating that animals have faster swimming speeds in clearer waters. However, after decades of acoustic measurements, vast ocean areas remain unsampled and places for which data are available typically provide information for only a few months, resulting in an incomplete understanding of DVMs. Addressing this issue is important, because DVMs have a crucial role in global ocean biogeochemistry. Night-time feeding at the surface and daytime metabolism of this food at depth provide an efficient pathway for carbon and nutrient export.

Here we use observations from a satellite-mounted light-detection-and-ranging (lidar) instrument to describe global distributions of an optical signal from DVM animals that arrive in the surface ocean at night. Our findings reveal that these animals generally constitute a greater fraction of total plankton abundance in the clear subtropical gyres, consistent with the idea that the avoidance of visual predators is an important life strategy in these regions. Total DVM biomass, on the other hand, is higher in more productive regions in which the availability of food is increased. Furthermore, the 10-year satellite record reveals significant temporal trends in DVM biomass and correlated variations in DVM biomass and surface productivity. These results provide a detailed view of DVM activities globally and a path for refining the quantification of their biogeochemical importance.

Mesoscale eddies are critical components of the ocean’s "internal weather" system. Mixing and stirring by eddies exerts significant control on biogeochemical fluxes in the open ocean, and eddies may trap distinctive plankton communities that remain coherent for months and can be transported hundreds to thousands of kilometers. Debate regarding how and why predators use fronts and eddies, for example as a migratory cue, enhanced forage opportunities, or preferred thermal habitat, has been ongoing since the 1950s. The influence of eddies on the behavior of large pelagic fishes, however, remains largely unexplored. Here, we reconstruct movements of a pelagic predator, the blue shark (Prionace glauca), in the Gulf Stream region using electronic tags, earth-observing satellites, and data-assimilating ocean forecasting models. Based on >2,000 tracking days and nearly 500,000 high-resolution time series measurements collected by 15 instrumented individuals, we show that blue sharks seek out the interiors of anticyclonic eddies where they dive deep while foraging. Our observations counter the existing paradigm that anticyclonic eddies are unproductive ocean "deserts" and suggest anomalously warm temperatures in these features connect surface-oriented predators to the most abundant fish community on the planet in the mesopelagic. These results also shed light on the ecosystem services provided by mesopelagic prey. Careful consideration will be needed before biomass extraction from the ocean twilight zone to avoid interrupting a key link between planktonic production and top predators. Moreover, robust associations between targeted fish species and oceanographic features increase the prospects for effective dynamic ocean management.

The shortbill spearfish (Tetrapturus angustirostris) is an understudied, istiophorid billfish primarily encountered as bycatch in pelagic commercial fisheries of the Indo-Pacific. The species is listed as data-deficient, and little is known of its biology, ecology, and population structure or status. We assessed the species' movement ecology and thermal niche with telemetry data from the first shortbill spearfishes ever outfitted with pop-up satellite archival transmitting tags (n = 3 with successfully transmitted data). Short (4–15 day) deployments offshore of the Island of Hawai'i revealed that spearfish primarily occupied the mixed layer, spending >90% of each 24-hr period between the surface and 100 m in water temperatures between 24–26°C. These individuals consistently exhibited vertical activity at night regardless of the prevailing lunar phase. Nocturnal movements throughout the mixed layer may enable shortbill spearfish to forage on mesopelagic species undergoing diel vertical migration and reduce trophic niche overlap with primarily diurnal, pelagic species. The narrow thermal distribution of shortbill spearfish in this study, almost exclusively within 2°C of sea surface temperature, suggests that they are more stenothermal than extra-generic istiophorid species.

The Mediterranean spearfish (Tetrapturus belone) is one of the least‐studied istiophorid billfishes, with little known of its biology, ecology, and behavior. To assess the species' movement and thermal niche, we analyzed telemetry data from, to our knowledge, the first and only Mediterranean spearfish ever outfitted with a pop‐up satellite archival transmitting tag. Throughout a 29‐day deployment during July and August 2015, the fish travelled in Italian waters of the Tyrrhenian and Ligurian Seas, spending on average 93% of each 24‐hr period above 30 m and exhibiting a diel activity pattern comprised of daytime vertical movement and nighttime near‐surface residency. The preferred thermal niche was 26–28°C, but the spearfish experienced temperatures as low as 14.2°C during descents. Vertical distribution was limited throughout the deployment with more time spent at depth in areas where the thermocline was comparatively deeper and weaker, consistent with habitat compression experienced by other billfishes.

Sea Surface Temperature (SST) modifies the turbulent mixing, drag, and pressure gradients within the marine atmospheric boundary layer that accelerate near‐surface flow from cool to warm SST and decelerate the flow from warm to cool SST. This phenomenon is well documented on scales of 100–1,000 km (the oceanic mesoscale); however, the nature of this air–sea coupling at scales on the order of 1–10 km (the submesoscale) remains unknown. The Advanced Spaceborne Thermal Emission and Reflection Radiometer can be used to study submesoscale phenomena because the high‐resolution infrared and near‐infrared images can used to estimate both SST and wind speed. Observations of dramatic temperature and wind gradients along the Gulf Stream landward edge are used to examine the surface wind response to submesoscale fronts in SST. Our analysis indicates that SST‐induced wind speed perturbations are observed at the scales of order 1–10 km, significantly smaller than previously suggested.

Mesoscale eddies, energetic vortices covering nearly a third of the ocean surface at any one time, modulate the spatial and temporal evolution of the mixed layer. We present a global analysis of concurrent satellite observations of mesoscale eddies with hydrographic profiles by autonomous Argo floats, revealing rich geographic and seasonal variability in the influence of eddies on mixed layer depth. Anticyclones deepen the mixed layer depth, whereas cyclones thin it, with the magnitude of these eddy‐induced mixed layer depth anomalies being largest in winter. Eddy‐centric composite averages reveal that the largest anomalies occur at the eddy center and decrease with distance from the center. Furthermore, the extent to which eddies modulate mixed layer depth is linearly related to the sea surface height amplitude of the eddies. Finally, large eddy‐mediated mixed layer depth anomalies are more common in anticyclones when compared to cyclones. We present candidate mechanisms for this observed asymmetry.

Chlorophyll rings (CRs) are defined as elevated chlorophyll along eddy peripheries and have been observed in anticyclonic oceanic eddies occasionally. This study presents observations of CRs around both anticyclonic and cyclonic eddies from a large observational data set. An innovative algorithm is developed to identify CRs from satellite observations of sea level anomalies and near-surface chlorophyll concentration in the North Pacific Ocean between 2003 and 2010. The results show that only 1% of mesoscale eddies are associated with CRs, which implies the CRs are not ubiquitous. We propose two potential generation mechanisms for CRs: horizontal advection and wind-current interaction. The former dominates the formation of about two-thirds of the CRs. The CRs associated with both cyclones and anticyclones represents an important contribution to better understanding of mesoscale physical/biological coupled phenomena.

Southern Ocean mesoscale eddies play an important role in ocean circulation and biogeochemical cycling, but their biological characteristics have not been well quantified at the basin scale. To address this, we combined a 15‐year tracked eddy data set with satellite observations of ocean color, sea surface temperature, and autonomous profiling floats to quantify the surface and subsurface properties of eddies. Anomalies of surface temperature and chlorophyll were examined in eddy‐centric composite averages constructed from thousands of eddies. Normalized surface chlorophyll anomalies (chl_norm) vary seasonally and geographically. Cyclones typically show positive chl_norm, while anticyclones have negative chl_norm. The sign of chl_norm reverses during late summer and autumn for eddies between the Subtropical and Polar Fronts. The reversal is most obvious in the Indian sector, and we attribute this to a combination of eddy stirring (deformation of surface gradients by the rotational velocity of an eddy) and deeper winter mixing in anticyclones. Both chl_norm and sea surface temperature anomalies transition from dipole structures north of the Subtropical Front to monopole structures south of the Subantarctic Front. Sea surface temperature and chl_norm composites provide evidence for eddy trapping (transporting of anomalies) and eddy stirring. This research provides a basin‐scale study of surface chlorophyll in Southern Ocean eddies and reveals counterintuitive biogeochemical signals.

The Antarctic Circumpolar Current has highly energetic mesoscale phenomena, but their impacts on phytoplankton biomass, productivity, and biogeochemical cycling are not understood well. We analyze satellite observations and an eddy‐rich ocean model to show that they drive chlorophyll anomalies of opposite sign in winter versus summer. In winter, deeper mixed layers in positive sea surface height (SSH) anomalies reduce light availability, leading to anomalously low chlorophyll concentrations. In summer with abundant light, however, positive SSH anomalies show elevated chlorophyll concentration due to higher iron level, and an iron budget analysis reveals that anomalously strong vertical mixing enhances iron supply to the mixed layer. Features with negative SSH anomalies exhibit the opposite tendencies: higher chlorophyll concentration in winter and lower in summer. Our results suggest that mesoscale modulation of iron supply, light availability, and vertical mixing plays an important role in causing systematic variations in primary productivity over the seasonal cycle.

Satellite-tracking of mature white sharks (Carcharodon carcharias) has revealed open-ocean movements spanning months and covering tens of thousands of kilometers. But how are the energetic demands of these active apex predators met as they leave coastal areas with relatively high prey abundance to swim across the open ocean through waters often characterized as biological deserts? Here we investigate mesoscale oceanographic variability encountered by two white sharks as they moved through the Gulf Stream region and Sargasso Sea in the North Atlantic Ocean. In the vicinity of the Gulf Stream, the two mature female white sharks exhibited extensive use of the interiors of clockwise-rotating anticyclonic eddies, characterized by positive (warm) temperature anomalies. One tagged white shark was also equipped with an archival tag that indicated this individual made frequent dives to nearly 1,000 m in anticyclones, where it was presumably foraging on mesopelagic prey. We propose that warm temperature anomalies in anticyclones make prey more accessible and energetically profitable to adult white sharks in the Gulf Stream region by reducing the physiological costs of thermoregulation in cold water. The results presented here provide valuable new insight into open ocean habitat use by mature, female white sharks that may be applicable to other large pelagic predators.

Horizontal and vertical motions associated with coherent mesoscale structures, including eddies and meanders, are responsible for significant global transports of many properties, including heat and mass. Mesoscale vertical fluxes also influence upper ocean biological productivity by mediating the supply of nutrients into the euphotic layer, with potential impacts on the global carbon cycle. The Brazil-Malvinas Confluence (BMC) is a western boundary current region in the South Atlantic with intense mesoscale activity. This region has an active role in the genesis and transformation of water masses and thus is a critical component of the Atlantic meridional overturning circulation. The collision between the Malvinas and Brazil Currents over the Patagonian shelf/slope creates an energetic front that translates offshore to form a vigorous eddy field. Recent improvements in gridded altimetric sea level anomaly fields allow us to track BMC mesoscale eddies with high spatial and temporal resolutions using an automated eddy tracker. We characterize the eddies across fourteen 5° × 5° subregions. Eddy-centric composites of tracers and geostrophic currents diagnosed from a global reanalysis of surface and in situ data reveal substantial subregional heterogeneity. The in situ data are also used to compute the evolving quasi-geostrophic vertical velocity (QG-ω) associated with each instantaneous eddy instance. The QG-ω eddy composites have the expected dipole patterns of alternating upwelling/downwelling, however, the magnitude and sign of azimuthally averaged vertical velocity varies among subregions. Maximum eddy values are found near fronts and sharp topographic gradients. In comparison with regional eddy composites, subregional composites provide refined information about mesoscale eddy heterogeneity.

Highlights

• Eddies and meanders of the Gulf Stream entrain and trap water with distinct near-surface chlorophyll concentration.

• Following formation, chlorophyll in anticyclonic eddies is observed to increase.

• The positive chlorophyll trend in anticyclonic eddies is reproduced in a simulation including eddy-induced Ekman pumping.

Marine animals, such as turtles, seabirds and pelagic fishes, are observed to travel and congregate around eddies in the open ocean. Mesoscale eddies, large swirling ocean vortices with radius scales of approximately 50–100 km, provide environmental variability that can structure these populations. In this study, we investigate the use of mesoscale eddies by 24 individual juvenile loggerhead sea turtles (Caretta caretta) in the Brazil-Malvinas Confluence region. The influence of eddies on turtles is assessed by collocating the turtle trajectories to the tracks of mesoscale eddies identified in maps of sea level anomaly. Juvenile loggerhead sea turtles are significantly more likely to be located in the interiors of anticyclones in this region. The distribution of surface drifters in eddy interiors reveals no significant association with the interiors of cyclones or anticyclones, suggesting higher prevalence of turtles in anticyclones is a result of their behavior. In the southern portion of the Brazil-Malvinas Confluence region, turtle swimming speed is significantly slower in the interiors of anticyclones, when compared to the periphery, suggesting that these turtles are possibly feeding on prey items associated with anomalously low near-surface chlorophyll concentrations observed in those features.