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Eastern Boundary Systems support major fisheries whose early life stages depend on upwelling production. Upwelling can be highly variable at the regional scale, with substantial repercussions for new productivity and microbial loop activity. A holistic assessment of plankton community structure is challenging due to the range in body forms and sizes of the taxa. Thus, studies that integrate the classic trophic web based on new production with the microbial loop are rare. Underwater imaging can overcome this limitation, and together with machine learning, enables fine resolution studies spanning large spatial scales. We used the In-situ Ichthyoplankton Imaging System (ISIIS) to investigate the drivers of plankton community structure in the northern California Current, sampled along the Newport Hydrographic (NH) and Trinidad Head (TR) lines, in OR and CA, respectively. The non-invasive imaging of particles and plankton (250μm –15cm) over 1644km (30 transects) in the winters and summers of 2018 and 2019 yielded 1.194 billion classified plankton images. The imaged plankton community ranged from protists, crustaceans, and gelatinous taxa to larval fishes. To assess community structure, >2000 single-taxon distribution profiles were analyzed using high resolution spatial correlations. Co-occurrences on the NH line were consistently significantly higher off-shelf while those at TR tended to be highest on-shelf. Taxa co-occurrences at TR increased significantly with upwelling strength and in 2019 TR summer co-occurrences were similar to those on the NH line. Random Forests models identified the concentrations of microbial loop taxa such as protists,Oithonacopepods, and appendicularians as important drivers of co-occurrences at NH line, while at TR, cumulative upwelling and chlorophyll a were of the highest importance. Our results indicate that the microbial loop is actively driving plankton community structure in intermittent upwelling systems such as the NH line and may induce temporal stability. Where upwelling is more continuous such as at TR, primary production may dominate patterns of community structure, obscuring the underlying role of the microbial loop. Future changes in upwelling strength are likely to disproportionately affect plankton community structure in continuous upwelling regions, while high microbial loop activity enhances community structure resilience.

Blooms of the colonial pelagic tunicate Pyrosoma atlanticum in 2014–2018 followed a marine heatwave in the eastern Pacific Ocean. Pyrosome blooms could alter pelagic food webs of the northern California Current (NCC) by accelerating the biological pump via active transport, fecal pellet production and mortality events. Although aggregations of P. atlanticum have the potential to shape marine trophic dynamics via carbon export, little is known about pyrosome vertical distribution patterns. In this study, we estimated the distribution of P. atlanticum in the NCC along transects off of Oregon (45°N and 124°W) and northern California (41°N and 124°W), USA during February and July 2018. Depth-stratified plankton tows provided volume-normalized pyrosome abundance and biovolume estimates that complemented fine-scale counts by a vertically deployed camera system. Pyrosomes were numerous offshore during February, especially off Oregon. Colonies were distributed non-uniformly in the water column with peak numbers associated with vertical gradients in environmental parameters, notably density and chl-a. Vertical distributions shifted over the 24-h period, indicative of diel vertical migration. Understanding the vertical distribution of these gelatinous grazers in the NCC gives insight to their behavior and ecological role in biologically productive temperate ecosystems as conditions become more favorable for recurring blooms.

Microbial mortality impacts the structure of food webs, carbon flow, and the interactions that create dynamic patterns of abundance across gradients in space and time in diverse ecosystems. In the oceans, estimates of microbial mortality by viruses, protists, and small zooplankton do not account fully for observations of loss, suggesting the existence of underappreciated mortality sources. We examined how ubiquitous mucous mesh feeders (i.e. gelatinous zooplankton) could contribute to microbial mortality in the open ocean. We coupled capture of live animals by blue-water diving to sequence-based approaches to measure the enrichment and selectivity of feeding by two coexisting mucous grazer taxa (pteropods and salps) on numerically dominant marine prokaryotes. We show that mucous mesh grazers consume a variety of marine prokaryotes and select between coexisting lineages and similar cell sizes. We show that Prochlorococcus may evade filtration more than other cells and that planktonic archaea are consumed by macrozooplanktonic grazers. Discovery of these feeding relationships identifies a new source of mortality for Earth's dominant marine microbes and alters our understanding of how top-down processes shape microbial community and function. This article is protected by copyright. All rights reserved.

Cnidarian jellyfish can be dominant players in the food webs of highly productive Eastern Boundary Currents (EBC). However, the trophic role of inconspicuous hydromedusae in EBCs has traditionally been overlooked. We collected mesozooplankton from five stations along two cross-shelf transects in the Northern California Current (NCC) during winter and summer of 2018–2019. We analyzed gut contents of 11 hydromedusan species and the prey community to (i) determine prey resource use by hydromedusae and (ii) determine temporal shifts in the trophic niche of hydromedusae, focusing on the two most collected species (Clytia gregaria and Eutonina indicans). Hydromedusae in the NCC fed mostly on copepods, appendicularians and invertebrate larvae. Nonmetric multidimensional scaling of hydromedusan diets showed seasonal shifts in prey resource driven by the abundant C. gregaria, which fed mostly on copepod eggs during winter and fed mostly on appendicularians and copepods during summer. Prey selectivity for copepod eggs increased during winter for C. gregaria and E. indicans. Intriguingly, theoretical ingestion rates show that both species acquire similar amounts of carbon during upwelling and nonupwelling conditions. Hydromedusae’s consistent presence and predation impact across seasons may lead to significant effects in carbon and energy transfer through the NCC food web.

Many fishes employ distinct swimming modes for routine swimming and predator escape. These steady and escape swimming modes are characterized by dramatically differing body kinematics that lead to context-adaptive differences in swimming performance. Physonect siphonophores, such as Nanomia bijuga , are colonial cnidarians that produce multiple jets for propulsion using swimming subunits called nectophores. Physonect siphonophores employ distinct routine and steady escape behaviors but–in contrast to fishes–do so using a decentralized propulsion system that allows them to alter the timing of thrust production, producing thrust either synchronously (simultaneously) for escape swimming or asynchronously (in sequence) for routine swimming. The swimming performance of these two swimming modes has not been investigated in siphonophores. In this study, we compare the performances of asynchronous and synchronous swimming in N. bijuga over a range of colony lengths (i.e., numbers of nectophores) by combining experimentally derived swimming parameters with a mechanistic swimming model. We show that synchronous swimming produces higher mean swimming speeds and greater accelerations at the expense of higher costs of transport. High speeds and accelerations during synchronous swimming aid in escaping predators, whereas low energy consumption during asynchronous swimming may benefit N. bijuga during vertical migrations over hundreds of meters depth. Our results also suggest that when designing underwater vehicles with multiple propulsors, varying the timing of thrust production could provide distinct modes directed toward speed, efficiency, or acceleration.

Suspension feeders, including ascidians (Phylum Chordata, Class Ascidiacea), experience a dilute prey field composed of extremely small particles. The filtration apparatus of ascidians is based on a mucous-mesh that is continuously secreted and ingested. The rate and metabolic cost of this mesh secretion has not been quantified to date. We used video boroscopy to quantify the mucous-mesh production rate of the solitary ascidian Herdmania momus under different food and temperature treatments. H. momus individuals with an average (±95% CI) biomass of 30.7 ± 1.1 mg and a branchial sac area of 10.3 ± 1.2 cm2 produced an average of 276 ± 33.5 cm2 of mucous-mesh h-1, corresponding to a median turnover rate of 625 ± 82 mesh d-1. Since the mean mesh mass was 2.44 ± 0.58 mg, this production rate corresponds to roughly 50 ± 8 times the individual’s biomass per day. Food concentration had no detectable effect on mesh production rate, whereas a temperature difference of ~9°C (20 vs. 29°C) moderately increased mesh production by 30-50%. The current study reveals that the feeding process of H. momus involves a high expenditure on mucous-mesh synthesis that, combined with low food availability, may limit its growth in oligotrophic waters and under changing climate regimes.

Jellyfish have provided insight into important components of animal propulsion, such as suction thrust, passive energy recapture, vortex wall effects, and the rotational mechanics of turning. These traits are critically important to jellyfish because they must propel themselves despite severe limitations on force production imposed by rudimentary cnidarian muscular structures. Consequently, jellyfish swimming can occur only by careful orchestration of fluid interactions. Yet these mechanics may be more broadly instructive because they also characterize processes shared with other animal swimmers, whose structural and neurological complexity can obscure these interactions. In comparison with other animal models, the structural simplicity, comparative energetic efficiency, and ease of use in laboratory experimentation allow jellyfish to serve as favorable test subjects for exploration of the hydrodynamic bases of animal propulsion. These same attributes also make jellyfish valuable models for insight into biomimetic or bioinspired engineeringof swimming vehicles. Here, we review advances in understanding of propulsive mechanics derived from jellyfish models as a pathway toward the application of animal mechanics to vehicle designs.

During 2016-2018, unprecedented aggregations of the colonial pelagic tunicate Pyrosoma atlanticum were observed in the Northern California Current (NCC). Pyrosomes are common in tropical and sub-tropical ocean waters, but little is known about their abundance, distribution, and trophic ecology in mid-latitude systems. To assess these factors, pyrosomes were collected during cruises in the NCC in May and August 2017. A generalized additive model (GAM) was used to identify relationships between in situ environmental variables (temperature, salinity, fluorescence) and distribution and abundance patterns of pyrosomes in May 2017. Fatty acid (FA) profiles were then characterized as diet indicators, and bulk stable isotope analysis of carbon and nitrogen was used to examine spatial variations in potential food sources and trophic level. The GAM identified sea surface temperature and surface salinity as significant variables related to pyrosome densities. The most abundant FA in the pyrosomes was docosahexanoic acid (22:6ω3), which serves in pelagic systems as a biomarker for dinoflagellates. Common FA biomarkers for bacteria, carnivory, and dinoflagellates differed by latitude, suggesting that pyrosomes have different diets over a broad latitudinal range. The δ15N values of P. atlanticum indicate that pyrosomes may be feeding at a relatively low trophic level compared to other zooplankton groups in this region. Offshore pyrosomes had lower δ13C values than those collected on the shelf, suggesting incorporation of nearshore carbon in pyrosome tissues. Previously documented rapid reproduction and growth of pyrosomes coupled with efficient feeding behavior for common NCC plankters may support their continued presence in this mid-latitude region.

Of the more than 150 ctenophore species, the oceanic ctenophore Eurhamphaea vexilligera is notable for its bright orange-yellow ink, secreted from numerous small vesicles that line its substomodeal comb rows. To date, in situ observations by scuba divers have proved the most fruitful method of observing these animals' natural behavior. We present the results of one such contemporary scuba-based observation of E. vexilligera, conducted in the Gulf Stream waters off the coast of Florida, using high-resolution photography and video. Utilizing underwater camera systems purpose built for filming gelatinous zooplankton, we observed E. vexilligera ink release and swimming behavior in situ. From these data, we describe the timeline and mechanics of E. vexilligera ink release in detail, as well as the animal's different swimming behaviors and resulting ink dispersal patterns. We also describe a rolling swimming behavior, accompanied and possibly facilitated by a characteristic change in overall body shape. These observations provide further insight into the behavioral ecology of this distinctive ctenophore and may serve as the foundation for future kinematic studies.

Pulsatile jet propulsion is a common swimming mode used by a diverse array of aquatic taxa from chordates to cnidarians. This mode of locomotion has interested both biologists and engineers for over a century. A central issue to understanding the important features of jet-propelling animals is to determine how the animal interacts with the surrounding fluid. Much of our knowledge of aquatic jet propulsion has come from simple theoretical approximations of both propulsive and resistive forces. Although these models and basic kinematic measurements have contributed greatly, they alone cannot provide the detailed information needed for a comprehensive, mechanistic overview of how jet propulsion functions across multiple taxa, size scales and through development. However, more recently, novel experimental tools such as high-speed 2D and 3D particle image velocimetry have permitted detailed quantification of the fluid dynamics of aquatic jet propulsion. Here, we provide a comparative analysis of a variety of parameters such as efficiency, kinematics and jet parameters, and review how they can aid our understanding of the principles of aquatic jet propulsion. Research on disparate taxa allows comparison of the similarities and differences between them and contributes to a more robust understanding of aquatic jet propulsion.

International audience; Salps are gelatinous planktonic suspension feeders that filter large volumes of water in the food-dilute open ocean. Their life cycle allows periodic exponential growth and population blooms. Dense swarms of salps have a high grazing impact that can deplete the photic zone of phytoplankton and export huge quantities of organic matter to the deep sea. Previous studies described their feeding manner as mostly nonselective, with larger particles retained at higher efficiencies than small particles. To examine salp diets, we used direct in situ sampling (InEx method) of undisturbed solitary Salpa maxima. Aggregates ("chains") of Salpa fusiformis and Thalia democratica were studied using in situ incubations. Our findings suggest that in situ feeding rates are higher than previously reported and that cell removal is size independent with $ 1 μm picoeukaryotes preferentially removed over both larger eukaryotes and smaller bacteria. The prey : predator size ratios we measured (1 : 10 4-1 : 10 5) are an order of magnitude smaller than previously reported values and to the best of our knowledge, are the smallest values reported so far for any planktonic suspension feeders. Despite differences among the three species studied, they had similar prey preferences with no correlation between salp body length and prey size. Our findings shed new light on prey : predator relationships in planktonic systems-in particular, that factors other than size influence filtration efficiency-and suggest that in situ techniques should be devised and applied for the study of suspension feeding in the ocean.

Pyrosomes are widely distributed pelagic tunicates that have the potential to reshape marine food webs when they bloom. However, their grazing preferences and interactions with the background microbial community are poorly understood. This is the first study of the marine microorganisms associated with pyrosomes undertaken to improve the understanding of pyrosome biology, the impact of pyrosome blooms on marine microbial systems, and microbial symbioses with marine animals. The diversity, relative abundance, and taxonomy of pyrosome-associated microorganisms were compared to seawater during a Pyrosoma atlanticum bloom in the Northern California Current System using high-throughput sequencing of the 16S rRNA gene, microscopy, and flow cytometry. We found that pyrosomes harbor a microbiome distinct from the surrounding seawater, which was dominated by a few novel taxa. In addition to the dominant taxa, numerous more rare pyrosome-specific microbial taxa were recovered. Multiple bioluminescent taxa were present in pyrosomes, which may be a source of the iconic pyrosome luminescence. We also discovered free-living marine microorganisms in association with pyrosomes, suggesting that pyrosome feeding impacts all microbial size classes but preferentially removes larger eukaryotic taxa. This study demonstrates that microbial symbionts and microbial prey are central to pyrosome biology. In addition to pyrosome impacts on higher trophic level marine food webs, the work suggests that pyrosomes also alter marine food webs at the microbial level through feeding and seeding of the marine microbial communities with their symbionts. Future efforts to predict pyrosome blooms, and account for their ecosystem impacts, should consider pyrosome interactions with marine microbial communities.

An abundance of swimming animals have converged upon a common swimming strategy using multiple propulsors coordinated as metachronal waves. The shared kinematics suggest that even morphologically and systematically diverse animals use similar fluid dynamic relationships to generate swimming thrust. We quantified the kinematics and hydrodynamics of a diverse group of small swimming animals who use multiple propulsors, e.g. limbs or ctenes, which move with antiplectic metachronal waves to generate thrust. Here we show that even at these relatively small scales the bending movements of limbs and ctenes conform to the patterns observed for much larger swimming animals. We show that, like other swimming animals, the propulsors of these metachronal swimmers rely on generating negative pressure along their surfaces to generate forward thrust (i.e., suction thrust). Relying on negative pressure, as opposed to high pushing pressure, facilitates metachronal waves and enables these swimmers to exploit readily produced hydrodynamic structures. Understanding the role of negative pressure fields in metachronal swimmers may provide clues about the hydrodynamic traits shared by swimming and flying animals.

It has been well documented that animals (and machines) swimming or flying near a solid boundary get a boost in performance1-6. This ground effect is often modeled as an interaction between a mirrored pair of vortices represented by a true vortex and an opposite sign ‘virtual vortex’ on the other side of the wall7. However, most animals do not swim near solid surfaces and thus near body vortex-vortex interactions in open-water swimmers have been poorly investigated. In this study we examine the most energetically efficient metazoan swimmer known to date, the jellyfishAurelia aurita, to elucidate the role that vortex interactions can play in animals that swim away from solid boundaries. We used high speed video tracking, laser-based digital particle image velocimetry (dPIV) and an algorithm for extracting pressure fields from flow velocity vectors to quantify swimming performance and the effect of near body vortex-vortex interactions. Here we show that a vortex ring (stopping vortex), created underneath the animal during the previous swim cycle, is critical for increasing propulsive performance. This well positioned stopping vortex acts in the same way as a virtual vortex during wall-effect performance enhancement, by helping converge fluid at the underside of the propulsive surface and generating significantly higher pressures which result in greater thrust. These findings advocate that jellyfish can generate a wall-effect boost in open water by creating what amounts to a ‘virtual wall’ between two real, opposite sign vortex rings. This explains the significant propulsive advantage jellyfish possess over other metazoans and represents important implications for bio-engineered propulsion systems.

We present the Aquatic Symbiosis Genomics Project, a global collaboration to generate high quality genome sequences for a wide range of eukaryotes and their microbial symbionts. Launched under the Symbiosis in Aquatic Systems Initiative of the Gordon and Betty Moore Foundation, the ASG Project brings together researchers from across the globe who hope to use these reference genomes to augment and extend their analyses of the dynamics, mechanisms and environmental importance of symbiosis. Applying large-scale, high-throughput sequencing and assembly technologies, the ASG collaboration will assemble and annotate the genomes of 500 symbiotic organisms – both the “hosts” and the microbial symbionts with which they associate. These data will be released openly to benefit all who work on symbiosis, from conservation geneticists to those interested in the origin of the eukaryotic cell.

Oceanic ecosystems are dominated by minute microorganisms that play a major role in food webs and biogeochemical cycles 1 . Many microorganisms thrive in the dilute environment due to their capacity to locate, attach to, and use patches of nutrients and organic matter 2,3 . We propose that some free-living planktonic bacteria have traded their ability to stick to nutrient-rich organic particles for a non-stick cell surface that helps them evade predation by mucous filter feeders. We used a combination of in situ sampling techniques and next-generation sequencing to study the biological filtration of microorganisms at the phylotype level. Our data indicate that some marine bacteria, most notably the highly abundant Pelagibacter ubique and most other members of the SAR 11 clade of the Alphaproteobacteria, can evade filtration by slipping through the mucous nets of both pelagic and benthic tunicates. While 0.3 µm polystyrene beads and other similarly-sized bacteria were efficiently filtered, SAR11 members were not captured. Reversed-phase chromatography revealed that most SAR11 bacteria have a much less hydrophobic cell surface than that of other planktonic bacteria. Our data call for a reconsideration of the role of surface properties in biological filtration and predator-prey interactions in aquatic systems. In situ sampling reveals that members of the SAR11 clade show significantly lower retention by mucous filter feeders, and that this is probably due to their reduced hydrophobic cell surface, suggesting that cell surface properties are important factors in predator–prey interactions.

Appendicularians are ubiquitous marine grazers that use tangential filtration to collect micron and submicron prey. The food-concentrating filter (FCF) is the primary determinant of appendicularian prey selectivity, but the precise means by which it concentrates and conveys particles to the pharyngeal filter remain poorly understood. We used high-speed videography to examine the mechanics of the FCF of Oikopleura dioica with high resolution to better understand how filter structure, hydrodynamics, and animal behavior affect feeding selectivity. We observed that filtration occurs through the process of serial adhesion and detachment of particles, and in this respect, differs from an industrial tangential filtration system. We demonstrate how filter elasticity and the hydrodynamics of pulsatile flow caused by the kinematics of the animal's tail contribute to the filter's self-cleaning abilities. We present experimental observations showing that smaller particles (3 μm) adhere significantly more to the filter than larger ones (10 μm), but that, overall adhesion is quite low for an array of particle sizes (3–20 μm). This study provides a mechanistic basis for predicting the effects of appendicularian grazing on particle size spectra in the upper ocean.

Despite its delicate morphology, the lobate ctenophore Mnemiopsis leidyi thrives in coastal ecosystems as an influential zooplankton predator. Coastal ecosystems are often characterized as energetic systems with high levels of natural turbulence in the water column. To understand how natural wind-driven turbulence affects the feeding ecology of M. leidyi, we used a combination of approaches to quantify how naturally and laboratory generated turbulence affects the behavior, feeding processes and feeding impact of M. leidyi. Experiments using laboratory generated turbulence demonstrated that turbulence can reduce M. leidyi feeding rates on copepods and Artemia nauplii by > 50%. However, detailed feeding data from the field, collected during highly variable surface conditions, showed that wind-driven turbulence did not affect the feeding rates or prey selection of M. leidyi. Additional laboratory experiments and field observations suggest that the feeding process of M. leidyi is resilient to wind-driven turbulence because M. leidyi shows a behavioral response to turbulence by moving deeper in the water column. Seeking refuge in deeper waters enables M. leidyi to maintain high feeding rates even under high turbulence conditions generated by wind driven mixing. As a result, M. leidyi exerted a consistently high predatory impact on prey populations during highly variable and often energetic wind-driven mixing conditions. This resilience adds to our understanding of how M. leidyi can thrive in a wide spectrum of environments around the world. The limits to this resilience also set boundaries to its range expansion into novel areas.

Coordination of multiple propulsors can provide performance benefits in swimming organisms. Siphonophores are marine colonial organisms that orchestrate the motion of multiple swimming zooids for effective swimming. However, the kinematics at the level of individual swimming zooids (nectophores) have not been examined in detail. We used high speed, high resolution microvideography and particle image velocimetry (PIV) of the physonect siphonophore,Nanomia bijuga, to study the motion of the nectophores and the associated fluid motion during jetting and refilling. The integration of nectophore and velum kinematics allow for a high-speed (maximum ~1 m s−1), narrow (1-2 mm) jet and rapid refill as well as a 1:1 ratio of jetting to refill time. Overall swimming performance is enhanced by velocity gradients produced in the nectophore during refill, which lead to a high pressure region that produces forward thrust. Generating thrust during both the jet and refill phases augments the distance travelled by 17% over theoretical animals, which generate thrust only during the jet phase. The details of velum kinematics and associated fluid mechanics elucidate how siphonophores effectively navigate three-dimensional space and could be applied to exit flow parameters in multijet underwater vehicles.Summary statement:Colonial siphonophores produce high speed jets and generate forward thrust during refill using a flexible velum to achieve effective propulsion.

It has been well documented that animals (and machines) swimming or flying near a solid boundary get a boost in performance. This ground effect is often modelled as an interaction between a mirrored pair of vortices represented by a true vortex and an opposite sign ‘virtual vortex’ on the other side of the wall. However, most animals do not swim near solid surfaces and thus near body vortex–vortex interactions in open-water swimmers have been poorly investigated. In this study, we examine the most energetically efficient metazoan swimmer known to date, the jellyfishAurelia aurita, to elucidate the role that vortex interactions can play in animals that swim away from solid boundaries. We used high-speed video tracking, laser-based digital particle image velocimetry (dPIV) and an algorithm for extracting pressure fields from flow velocity vectors to quantify swimming performance and the effect of near body vortex–vortex interactions. Here, we show that a vortex ring (stopping vortex), created underneath the animal during the previous swim cycle, is critical for increasing propulsive performance. This well-positioned stopping vortex acts in the same way as a virtual vortex during wall-effect performance enhancement, by helping converge fluid at the underside of the propulsive surface and generating significantly higher pressures which result in greater thrust. These findings advocate that jellyfish can generate a wall-effect boost in open water by creating what amounts to a ‘virtual wall’ between two real, opposite sign vortex rings. This explains the significant propulsive advantage jellyfish possess over other metazoans and represents important implications for bio-engineered propulsion systems.

Obelia spp. are cnidarian hydromedusae with a cosmopolitan distribution but very little is known about their feeding. The small size of Obelia (bell diameter ∼ 1 mm, tentacle width ∼ 0.05 mm) suggests that feeding occurs in a viscous regime characterized by thick boundary layers. During feeding observations with a natural prey assemblage the majority of prey were captured at the tentacle tips during the contraction phase. Swimming kinematics from high speed videography confirmed that swimming was a low Re number process (Re

Aggregations of the neustonic hydrozoan Velella velella occur periodically in the northern California Current. Despite the regular occurrence of notable bloom events in this productive upwelling zone, little is known about their trophic ecology. We used gut content and stable isotope analyses (SIA) to elucidate V. velella prey selectivity and trophic niche to address their potential impacts on the marine ecosystem. The dominant prey items ingested by V. velella colonies were non-motile prey including cladocerans and northern anchovy (Engraulis mordax) eggs, though copepods were also common in gut contents. Removal rates of northern anchovy eggs could be magnified in bloom years and in areas of high spawning biomass. Stable isotope analysis revealed differences in isotopic niche width and overlap among V. velella based on latitudinal gradients and to a lesser extent on V. velella size and demonstrates the need for continued work to fully understand the trophic ecology of this unique neustonic organism.

International audience; Mucous-mesh grazers (pelagic tunicates and thecosome pteropods) are common in oceanic waters and efficiently capture, consume and repackage particles many orders of magnitude smaller than themselves. They feed using an adhesive mucous mesh to capture prey particles from ambient seawater. Historically, their grazing process has been characterized as non-selective, depending only on the size of the prey particle and the pore dimensions of the mesh. The purpose of this review is to reverse this assumption by reviewing recent evidence that shows mucous-mesh feeding can be selective. We focus on large planktonic microphages as a model of selective mucus feeding because of their important roles in the ocean food web: as bac-terivores, prey for higher trophic levels, and exporters of carbon via mucous aggregates, faecal pellets and jelly-falls. We identify important functional variations in the filter mechanics and hydrodynamics of different taxa. We review evidence that shows this feeding strategy depends not only on the particle size and dimensions of the mesh pores, but also on particle shape and surface properties , filter mechanics, hydrodynamics and grazer behaviour. As many of these organisms remain critically understudied, we conclude by suggesting priorities for future research.

Jellyfish disruption of fisheries has been described in some coastal systems, but few thorough investigations have been conducted. To ascertain the economic impact and trend of jellyfish blooms in the Northern California Current (NCC), we mailed surveys to resident commercial shrimpers, salmon trollers, rockfish (blue, black), and groundfish fishers (n = 872). We asked fishers to estimate the damages caused by jellyfish—including costs of relocating to avoid blooms, lost fishing time, time lost to bycatch sorting, fish depreciation, and gear damage. Of the total respondents (n = 111), 67% reported that jellyfish reduce their seasonal revenue, but the degree of impact ranged considerably by fishery and location. Highest jellyfish nuisance corresponded to regions with the most salmon trolling effort. Using the mean revenue losses provided by respondents, we estimate that the combined economic impact of jellyfish on Oregon's salmon and pink shrimp fishers was over $650 000 in peak jellyfish season (June–September) in 2012. Fishers reported that jellyfish biomass varies annually, but most respondents (51%) reported observing no appreciable change in jellyfish populations in the last 5 years. Since economic impact analyses have been conducted primarily in areas with anomalous, high-density blooms, data from the NCC, which is not known to be experiencing increases in jellyfish abundance, provides baseline information on the socio-economic impact of jellyfish blooms in this region. In addition, the finding that jellyfish impact hook and line fisheries—not solely net fisheries—has implications for many other regions where fishers employ this gear type.

The influential predatory role of the lobate comb jellyfish Mnemiopsis leidyi has largely been attributed to the generation of a hydrodynamically silent feeding current to entrain and initiate high encounter rates with prey. However, for high encounter rates to translate to high ingestion rates, M. leidyi must effectively capture the entrained prey. To investigate the capture mechanisms, we quantified fluid interactions between the lobes using particle image velocimetry (PIV) and quantified, using 3-dimensional videography, the outcome of encounter events with slow swimming Artemia prey. The auricles, which produce the feeding current of M. leidyi, were the primary encounter structures, first contacting 59% of the prey in the feeding current. Upon detection, the auricles manipulated the Artemia to initiate captures on the tentillae, which are coated with sticky cells (colloblasts). Using this mechanism of sensory-scanning to capture prey entrained in the feeding current, M. leidyi employs a similar foraging strategy to that of feeding-current foraging copepods. As such, M. leidyi has a higher capture efficiency than do medusae, contributing to the greater predatory effect of M. leidyi in both its endemic and invasive ecosystems.

In the version of this Letter originally published, the authors incorrectly stated that primers 28F-519R were reported in ref. 54 to underestimate the abundance of SAR11 in the ocean. This statement has now been amended in all versions of the Letter.

Planktonic organisms are exposed to turbulent water motion that affects the fundamental activities of swimming and feeding. The goal of this study was to measure the influence of realistic turbulence levels on (i) swimming behavior and (ii) fluid interactions during feeding by the lobate ctenophore, Mnemiopsis leidyi, a highly successful suspension-feeding predator. A laboratory turbulence generator produced turbulence (e = 0.5–1.4 × 10^(−6) W kg^(−1)) representative of a field site in Woods Hole, MA, USA. Compared with still water, M. leidyi avoided regions in the experimental vessel where turbulence was greatest (e = 1.1–1.4 × 10^(−6) W kg^(−1)) by increasing its swimming speeds and accelerations. Both laboratory and in situ particle image velocimetry data demonstrated that feeding currents of M. leidyi were eroded by ambient fluid motions. Despite this, the overall flux to the feeding structures remained constant due to higher swimming speeds in turbulent conditions. Instantaneous shear deformation rates produced by background turbulence were higher than those produced by ctenophore feeding currents and frequently exceeded the published escape thresholds of copepod prey. Feeding current erosion and fluid mechanical signal noise within turbulent flows affect the mechanics of predator–prey interactions during suspension feeding by the ctenophore M. leidyi.

Gelatinous zooplankton exhibit a wide range of propulsive swimming modes. One of the most energetically efficient is the rowing behaviour exhibited by many species of schyphomedusae, which employ vortex interactions to achieve this result. Ctenophores (comb jellies) typically use a slow swimming, cilia-based mode of propulsion. However, species within the genusOcyropsishave developed an additional propulsive strategy of rowing the lobes, which are normally used for feeding, in order to rapidly escape from predators. In this study, we used high-speed digital particle image velocimetry to examine the kinematics and fluid dynamics of this rarely studied propulsive mechanism. This mechanism allowsOcyropsisto achieve size-adjusted speeds that are nearly double those of other large gelatinous swimmers. The investigation of the fluid dynamic basis of this escape mode reveals novel vortex interactions that have not previously been described for other biological propulsion systems. The arrangement of vortices during escape swimming produces a similar configuration and impact as that of the well-studied ‘vortex rebound’ phenomenon which occurs when a vortex ring approaches a solid wall. These results extend our understanding of how animals use vortex–vortex interactions and provide important insights that can inform the bioinspired engineering of propulsion systems.

In plankton ecology, biological and physical dynamics are coupled, structuring how plankton interact with their environment and other organisms. This interdisciplinary field has progressed considerably over the recent past, due in large part to advances in technology that have improved our ability to observe plankton and their fluid environment simultaneously across multiple scales. Recent research has demonstrated that fluid flow interacting with plankton behavior can drive many planktonic processes and spatial patterns. Moreover, evidence now suggests that plankton behavior can significantly affect ocean physics. Biophysical processes relevant to plankton ecology span a range of scales; for example, microscale turbulence influences planktonic growth and grazing at millimeter scales, whereas features such as fronts and eddies can shape larger-scale plankton distributions. Most research in this field focuses on specific processes and thus is limited to a narrow range of spatial scales. However, biophysical interactions are intimately connected across scales, since processes at a given scale can have implications at much larger and smaller scales; thus, a cross-scale perspective on how biological and physical dynamics interact is essential for a comprehensive understanding of the field. Here, we present a review of biophysical interactions in the plankton across multiple scales, emphasizing new findings over recent decades and highlighting opportunities for cross-scale comparisons. By investigating feedbacks and interactions between processes at different scales, we aim to build cross-scale intuition about biophysical planktonic processes and provide insights for future directions in the field.

Autores: Robert Condon ... [et al.]. -- 10 páginas, 5 figuras, During the past several decades, high numbers of gelatinous zooplankton species have been reported in many estuarine and coastal ecosystems. Coupled with media-driven public perception, a paradigm has evolved in which the global ocean ecosystems are thought to be heading toward being dominated by “nuisance” jellyfish. We question this current paradigm by presenting a broad overview of gelatinous zooplankton in a historical context to develop the hypothesis that population changes reflect the human-mediated alteration of global ocean ecosystems. To this end, we synthesize information related to the evolutionary context of contemporary gelatinous zooplankton blooms, the human frame of reference for changes in gelatinous zooplankton populations, and whether sufficient data are available to have established the paradigm. We conclude that the current paradigm in which it is believed that there has been a global increase in gelatinous zooplankton is unsubstantiated, and we develop a strategy for addressing the critical questions about long-term, human-related changes in the sea as they relate to gelatinous zooplankton blooms., La financiación del National Center for Ecological Analysis and Synthesis y del Jellyfish Working Group proviene de la Nacional Science Foundation Grant no. DEB-94-21535, de la Universidad de California en Santa Bárbara, y desde el Estado de California. Michael N. Dawson fue apoyado en parte por la Nacional Science Foundation Grant no. DEB-07-17071.

Zooplankton swimming near the substratum experience boundary layer flow that is characterized by steep velocity gradients and turbulence. How do small swimming organisms navigate flows at this interface to forage and interact with mates? To address this question, we collected field measurements of the swimming behavior of the marine ostracod Paravargula trifax near complex living substrata, which were exposed to two conditions: slow “ambient flow” and faster “experimental flow.” Ostracod trajectories and background flow were recorded simultaneously using a self-contained underwater velocimetry apparatus (SCUVA). Particle image velocimetry (DPIV) produced instantaneous velocity vector fields in which the ostracods were swimming. Mean velocities, local shear stresses, turbulence intensity, and boundary shear velocity (u*) were greater in the experimental flow treatment. In slow ambient flow (urms = 0.39 ± 0.13 [mean ± SD] cm s − 1), ostracod swimming tracks were more tortuous and swimming angles corrected for background flow were randomly distributed compared with tracks in faster flow (urms = 3.49 ± 0.50 cm s − 1), indicating decreased maneuverability in rapidly flowing, turbulent water. Modeled, passive neutrally buoyant particles moved at substantially slower speeds, and their tracks were less tortuous than those of the ostracods, thus illustrating the importance of behavior as well as environmental flow in determining ostracod trajectories. Frequencies of encounters by ostracods with the benthos and with other ostracods were not different between treatments. However, in the experimental flow treatment, interactions with other ostracods occurred more frequently in the boundary layer than in the free stream, suggesting that microhabitats in the boundary layer may allow for enhanced mating encounters.

Weekly abundances of 2 salp populations (Salpa fusiformis and Thalia democratica) were determined for the period from 1967 to 1990 at a coastal station in the Western Mediterranean using a discrete abundance scale. A previous analysis modeled these data as a Markov chain with ordered states and found a significant effect of stratification on the Markov transition probabilities. An alternative analysis combining an underlying population model and a model of the observation process found qualitatively similar results, with population growth rate significantly related to stratification. However, by explicitly separating the population model from the observation model, the alternative approach has advantages including a closer connection to the underlying biology and the capability to predict salp abundance.

Salps have higher filtration rates than most other holoplankton, and are capable of packaging and exporting primary production from surface waters. A method of kinematic analysis was employed to accurately measure salp feeding rates. The data were then used to explain how diverse body morphologies and swimming motions among species and lifecycle stages influence salp feeding performance. We selected five species, representing a range of morphologies and swimming styles, and used digitized outlines from video frames to measure body-shape change during a pulse cycle. Time-varying body volume was then calculated from the digitized salp outlines to estimate the amount of fluid passing through the filtering mesh. This non-invasive method produced higher feeding rates than other methods and revealed that body volume, pulse frequency and degree of contraction are important factors for determining volume filtered. Each species possessed a unique combination of these three characteristics that resulted in comparable filtration (range: 0.44–15.33 ml s−1) and normalized filtration rates (range: 0.21–1.27 s−1) across species. The convergence of different species with diverse morphologies on similar normalized filtration suggests a tendency towards a flow optimum.

Physonect siphonophores are colonial cnidarians that are pervasive predators in many neritic and oceanic ecosystems. Physonects employ multiple, clonal medusan individuals, termed nectophores, to propel an aggregate colony. Here we show that developmental differences between clonal nectophores of the physonect Nanomia bijuga produce a division of labour in thrust and torque production that controls direction and magnitude of whole-colony swimming. Although smaller and less powerful, the position of young nectophores near the apex of the nectosome allows them to dominate torque production for turning, whereas older, larger and more powerful individuals near the base of the nectosome contribute predominantly to forward thrust production. The patterns we describe offer insight into the biomechanical success of an ecologically important and widespread colonial animal group, but, more broadly, provide basic physical understanding of a natural solution to multi-engine organization that may contribute to the expanding field of underwater-distributed propulsion vehicle design., Physonect siphonophores are highly mobile jellyfish with complex colonial organization. Here, Costello et al. show that division of labour among developmental stages controls the direction and propulsion of the colony, with older individuals providing thrust and younger individuals providing torque.

Marine microbes exhibit highly varied, often non-spherical shapes that have functional significance for essential processes, including nutrient acquisition and sinking rates. There is a surprising absence of data, however, on how cell shape affects grazing, which is crucial for predicting the fate of oceanic carbon. We used synthetic spherical and prolate spheroid microbeads to isolate the effect of particle length-to-width ratios on grazing and fate in the ocean. Here we show that the shape of microbe-sized particles affects predation by the appendicularian Oikopleura dioica, a globally abundant marine grazer. Using incubation experiments, we demonstrate that shape affects how particles are retained in the house and that the minimum particle diameter is the key variable determining how particles are ingested. High-speed videography revealed the mechanism behind these results: microbe-sized spheroids oriented with the long axis parallel to fluid streamlines, matching the speed and tortuosity of spheres of equivalent width. Our results suggest that the minimum particle diameter determines how elongated prey interact with the feeding-filters of appendicularians, which may help to explain the prevalence of ellipsoidal cells in the ocean, since a cell’s increased surface-to-volume ratio does not always increase predation. We provide the first evidence that grazing by appendicularians can cause non-uniform export of different shaped particles, thereby influencing particle fate.

Salps are marine invertebrates comprising multiple jet-propelled swimming units during a colonial life-cycle stage. Using theory, we show that asynchronous swimming with multiple pulsed jets yields substantial hydrodynamic benefit due to the production of steady swimming velocities, which limit drag. Laboratory comparisons of swimming kinematics of aggregate salps (Salpa fusiformisandWeelia cylindrica) using high-speed video supported that asynchronous swimming by aggregates results in a smoother velocity profile and showed that this smoother velocity profile is the result of uncoordinated, asynchronous swimming by individual zooids.In situflow visualizations ofW. cylindricaswimming wakes revealed that another consequence of asynchronous swimming is that fluid interactions between jet wakes are minimized. Although the advantages of multi-jet propulsion have been mentioned elsewhere, this is the first time that the theory has been quantified and the role of asynchronous swimming verified using experimental data from the laboratory and the field.

William H. Graham et al., © The Ecological Society of America. Jellyfish are usually perceived as harmful to humans and are seen as >pests>. This negative perception has hindered knowledge regarding their value in terms of ecosystem services. As humans increasingly modify and interact with coastal ecosystems, it is important to evaluate the benefits and costs of jellyfish, given that jellyfish bloom size, frequency, duration, and extent are apparently increasing in some regions of the world. Here we explore those benefits and costs as categorized by regulating, supporting, cultural, and provisioning ecosystem services. A geographical perspective of human vulnerability to jellyfish over four categories of human well-being (health care, food, energy, and freshwater production) is also discussed in the context of thresholds and trade-offs to enable social adaptation. Whereas beneficial services provided by jellyfish likely scale linearly with biomass (perhaps peaking at a saturation point), non-linear thresholds exist for negative impacts to ecosystem services. We suggest that costly adaptive strategies will outpace the beneficial services if jellyfish populations continue to increase in the future., Funding for the National Center for Ecological Analysis and Synthesis comes from National Science Foundation Grant DEB-94-21535, the University of California at Santa Barbara, and the State of California

Duarte, Carlos M. et al. -- 7 pages, 3 figures, 2 tables, Jellyfish (Cnidaria, Scyphozoa) blooms appear to be increasing in both intensity and frequency in many coastal areas worldwide, due to multiple hypothesized anthropogenic stressors. Here, we propose that the proliferation of artificial structures – associated with (1) the exponential growth in shipping, aquaculture, and other coastal industries, and (2) coastal protection (collectively, “ocean sprawl”) – provides habitat for jellyfish polyps and may be an important driver of the global increase in jellyfish blooms. However, the habitat of the benthic polyps that commonly result in coastal jellyfish blooms has remained elusive, limiting our understanding of the drivers of these blooms. Support for the hypothesized role of ocean sprawl in promoting jellyfish blooms is provided by observations and experimental evidence demonstrating that jellyfish larvae settle in large numbers on artificial structures in coastal waters and develop into dense concentrations of jellyfish-producing polyps, This research is a contribution to the Global Expansion of Jellyfish Blooms: Magnitude, Causes and Consequences Working Group, supported by the National Center for Ecological Analysis and Synthesis (NCEAS is supported by the National Science Foundation [grant #DEB-94-21535], the University of California at Santa Barbara, and the State of California) and the Evaluation of Ecosystem Impacts of Global Change in Mediterranean Ecosystems (MEDEICG) project, funded by the Spanish National Plan of I+D (CTM2009-07013). The C quinquecirrha experiments were conducted by DB and JM and supported by National Oceanic and Atmospheric Administration grant #NA09NOS4780214 to DB. We thank K Davies for artistic work