Hydrothermal vents supporting persistent plumes and microbial chemoautotrophy at Gakkel Ridge (Arctic Ocean).

Hydrothermal vents emit hot fluids enriched in energy sources for microbial life. Here, we compare the ecological and biogeochemical effects of hydrothermal venting of two recently discovered volcanic seamounts, Polaris and Aurora of the Gakkel Ridge, in the ice-covered Central Arctic Ocean. At both sites, persistent hydrothermal plumes increased up to 800 m into the deep Arctic Ocean. In the two non-buoyant plumes, rates of microbial carbon fixation were strongly elevated compared to background values of 0.5-1 µmol m-3 day-1 in the Arctic deep water, which suggests increased chemoautotrophy on vent-derived energy sources. In the Polaris plume, free sulfide and up to 360 nM hydrogen enabled microorganisms to fix up to 46 µmol inorganic carbon (IC) m-3 day-1. This energy pulse resulted in a strong increase in the relative abundance of SUP05 by 25% and Candidatus Sulfurimonas pluma by 7% of all bacteria. At Aurora, microorganisms fixed up to 35 µmol IC m-3 day-1. Here, metal sulfides limited the bioavailability of reduced sulfur species, and the putative hydrogen oxidizer Ca. S. pluma constituted 35% and SUP05 10% of all bacteria. In accordance with this data, transcriptomic analysis showed a high enrichment of hydrogenase-coding transcripts in Aurora and an enrichment of transcripts coding for sulfur oxidation in Polaris. There was neither evidence for methane consumption nor a substantial increase in the abundance of putative methanotrophs or their transcripts in either plume. Together, our results demonstrate the dominance of hydrogen and sulfide as energy sources in Arctic hydrothermal vent plumes.

Miles down for lunch: deep-sea in situ observations of Arctic finned octopods Cirroteuthis muelleri suggest pelagic-benthic feeding migration.

Deep-sea cephalopods are diverse, abundant, and poorly understood. The Cirrata are gelatinous finned octopods and among the deepest-living cephalopods ever recorded. Their natural feeding behaviour remains undocumented. During deep-sea surveys in the Arctic, we observed Cirroteuthis muelleri. Octopods were encountered with their web spread wide, motionless and drifting in the water column 500-2600 m from the seafloor. Individuals of C. muelleri were also repeatedly observed on the seafloor where they exhibited a repeated, behavioural sequence interpreted as feeding. The sequence (11-21 s) consisted of arm web spreading, enveloping and retreating. Prey capture happened during the enveloping phase and lasted 5-49 s. Numerous traces of feeding activity were also observed on the seafloor. The utilization of the water column for drifting and the deep seafloor for feeding is a novel migration behaviour for cephalopods, but known from gelatinous fishes and holothurians. By benthic feeding, the octopods benefit from the enhanced nutrient availability on the seafloor. Drifting in the water column may be an energetically efficient way of transportation while simultaneously avoiding seafloor-associated predators. In situ observations are indispensable to discover the behaviour of abundant megafauna, and the energetic coupling between the pelagic and benthic deep sea.

Habitat types and megabenthos composition from three sponge-dominated high-Arctic seamounts.

Seamounts are isolated underwater mountains stretching > 1000 m above the seafloor. They are identified as biodiversity hotspots of marine life, and host benthic assemblages that may vary on regional (among seamounts) and local (within seamounts) scales. Here, we collected seafloor imagery of three seamounts at the Langseth Ridge in the central Arctic Ocean to assess habitats and megabenthos community composition at the Central Mount (CM), the Karasik Seamount (KS), and the Northern Mount (NM). The majority of seafloor across these seamounts comprised bare rock, covered with a mixed layer of sponge spicule mats intermixed with detrital debris composed of polychaete tubes, and sand, gravel, and/or rocks. The megabenthos assemblages consisted of in total 15 invertebrate epibenthos taxa and 4 fish taxa, contributing to mean megabenthos densities of 55,745 ind. ha-1 at CM, 110,442 ind. ha-1 at KS, and 65,849 ind. ha-1 at NM. The faunal assemblages at all three seamounts were dominated by habitat-forming Tetractinellida sponges that contributed between 66% (KS) and 85% (CM) to all megabenthos. Interestingly, taxa richness did not differ at regional and local scale, whereas the megabenthos community composition did. Abiotic and biogenic factors shaping distinct habitat types played a major role in structuring of benthic communities in high-Arctic seamounts.

Volcanically hosted venting with indications of ultramafic influence at Aurora hydrothermal field on Gakkel Ridge.

The Aurora hydrothermal system, Arctic Ocean, hosts active submarine venting within an extensive field of relict mineral deposits. Here we show the site is associated with a neovolcanic mound located within the Gakkel Ridge rift-valley floor, but deep-tow camera and sidescan surveys reveal the site to be ≥100 m across-unusually large for a volcanically hosted vent on a slow-spreading ridge and more comparable to tectonically hosted systems that require large time-integrated heat-fluxes to form. The hydrothermal plume emanating from Aurora exhibits much higher dissolved CH4/Mn values than typical basalt-hosted hydrothermal systems and, instead, closely resembles those of high-temperature ultramafic-influenced vents at slow-spreading ridges. We hypothesize that deep-penetrating fluid circulation may have sustained the prolonged venting evident at the Aurora hydrothermal field with a hydrothermal convection cell that can access ultramafic lithologies underlying anomalously thin ocean crust at this ultraslow spreading ridge setting. Our findings have implications for ultra-slow ridge cooling, global marine mineral distributions, and the diversity of geologic settings that can host abiotic organic synthesis - pertinent to the search for life beyond Earth.

Impact of returning scientific cruises and prolonged on-site presence on litter abundance at the deep-sea nodule fields in the Peru Basin.

Marine litter can be found along coasts, continental shelves and slopes, down into the abyss. The absence of light, low temperatures and low energy regimes characterising the deeper habitats ensure the persistence of litter over time. Therefore, manmade items within the deep sea will likely accumulate to increasing quantities. Here we report the litter abundance encountered at the Pacific abyssal nodule fields from the Peru Basin at 4150 m depth. An average density of 2.67 litter items/ha was observed. Litter composed of plastic was the most abundant followed by metal and glass. At least 58 % of the items observed could be linked to the research expeditions conducted in the area and appeared to be mostly accidental disposals from ships. The data gathered was used to address temporal trends in litter abundance as well as the impact of human on-site presence and return cruises in the context of future deep-sea mining efforts.

Making marine image data FAIR.

Underwater images are used to explore and monitor ocean habitats, generating huge datasets with unusual data characteristics that preclude traditional data management strategies. Due to the lack of universally adopted data standards, image data collected from the marine environment are increasing in heterogeneity, preventing objective comparison. The extraction of actionable information thus remains challenging, particularly for researchers not directly involved with the image data collection. Standardized formats and procedures are needed to enable sustainable image analysis and processing tools, as are solutions for image publication in long-term repositories to ascertain reuse of data. The FAIR principles (Findable, Accessible, Interoperable, Reusable) provide a framework for such data management goals. We propose the use of image FAIR Digital Objects (iFDOs) and present an infrastructure environment to create and exploit such FAIR digital objects. We show how these iFDOs can be created, validated, managed and stored, and which data associated with imagery should be curated. The goal is to reduce image management overheads while simultaneously creating visibility for image acquisition and publication efforts.

The International Bathymetric Chart of the Southern Ocean Version 2.

The Southern Ocean surrounding Antarctica is a region that is key to a range of climatic and oceanographic processes with worldwide effects, and is characterised by high biological productivity and biodiversity. Since 2013, the International Bathymetric Chart of the Southern Ocean (IBCSO) has represented the most comprehensive compilation of bathymetry for the Southern Ocean south of 60°S. Recently, the IBCSO Project has combined its efforts with the Nippon Foundation - GEBCO Seabed 2030 Project supporting the goal of mapping the world's oceans by 2030. New datasets initiated a second version of IBCSO (IBCSO v2). This version extends to 50°S (covering approximately 2.4 times the area of seafloor of the previous version) including the gateways of the Antarctic Circumpolar Current and the Antarctic circumpolar frontal systems. Due to increased (multibeam) data coverage, IBCSO v2 significantly improves the overall representation of the Southern Ocean seafloor and resolves many submarine landforms in more detail. This makes IBCSO v2 the most authoritative seafloor map of the area south of 50°S.

A vast icefish breeding colony discovered in the Antarctic.

A breeding colony of notothenioid icefish (Neopagetopsis ionah, Nybelin 1947) of globally unprecedented extent has been discovered in the southern Weddell Sea, Antarctica. The colony was estimated to cover at least ∼240 km2 of the eastern flank of the Filchner Trough, comprised of fish nests at a density of 0.26 nests per square meter, representing an estimated total of ∼60 million active nests and associated fish biomass of >60,000 tonnes. The majority of nests were each occupied by 1 adult fish guarding 1,735 eggs (±433 SD). Bottom water temperatures measured across the nesting colony were up to 2°C warmer than the surrounding bottom waters, indicating a spatial correlation between the modified Warm Deep Water (mWDW) upflow onto the Weddell Shelf and the active nesting area. Historical and concurrently collected seal movement data indicate that this concentrated fish biomass may be utilized by predators such as Weddell seals (Leptonychotes weddellii, Lesson 1826). Numerous degraded fish carcasses within and near the nesting colony suggest that, in death as well as life, these fish provide input for local food webs and influence local biogeochemical processing. To our knowledge, the area surveyed harbors the most spatially expansive continuous fish breeding colony discovered to date globally at any depth, as well as an exceptionally high Antarctic seafloor biomass. This discovery provides support for the establishment of a regional marine protected area in the Southern Ocean under the Convention on the Conservation of Antarctic Marine Living Resources (CCAMLR) umbrella. VIDEO ABSTRACT.

Sea-ice derived meltwater stratification slows the biological carbon pump: results from continuous observations.

The ocean moderates the world's climate through absorption of heat and carbon, but how much carbon the ocean will continue to absorb remains unknown. The North Atlantic Ocean west (Baffin Bay/Labrador Sea) and east (Fram Strait/Greenland Sea) of Greenland features the most intense absorption of anthropogenic carbon globally; the biological carbon pump (BCP) contributes substantially. As Arctic sea-ice melts, the BCP changes, impacting global climate and other critical ocean attributes (e.g. biodiversity). Full understanding requires year-round observations across a range of ice conditions. Here we present such observations: autonomously collected Eulerian continuous 24-month time-series in Fram Strait. We show that, compared to ice-unaffected conditions, sea-ice derived meltwater stratification slows the BCP by 4 months, a shift from an export to a retention system, with measurable impacts on benthic communities. This has implications for ecosystem dynamics in the future warmer Arctic where the seasonal ice zone is expected to expand.

In situ observation of sponge trails suggests common sponge locomotion in the deep central Arctic.

In 2016, the research ice-breaker Polarstern surveyed the submerged peaks of the permanently ice-covered Langseth Ridge, a tectonic feature comprising the Karasik seamount and two deeper seamount peaks, abutting the Gakkel ultra-slow spreading ridge (87°N 62°E to 85.5°N 57.4°E)1. A towed marine camera sled and a hybrid remotely operated vehicle revealed these peaks to be covered by a dense demosponge community, at first glance reminiscent of North Atlantic Geodia grounds (sensu2). Sponges were observed on top of a thick layer of spicule mat (Figure 1 and Video S1), intermixed with underlying layers of empty siboglinid tubes and bivalve shells, a substrate covering almost the entire seafloor. We observed trails of densely interwoven spicules connected directly to the underside or lower flanks of sponge individuals (Figure 1), suggesting these trails are traces of motile sponges. This is the first time abundant sponge trails have been observed in situ and attributed to sponge mobility. Given the low primary production in this permanently ice-covered region, these trails may relate to feeding behavior and/or a strategy for dispersal of juveniles. Such trails may remain visible for long periods given the regionally low sedimentation rates.

PlasPI marine cameras: Open-source, affordable camera systems for time series marine studies.

Imaging underwater can be particularly problematic and expensive given the harsh environmental conditions posed by salinity and for some deployments, pressure. To counter these difficulties, expensive waterproof pressure resistant housings are often used, commonly built from expensive materials such as titanium, if intended for long duration deployments. Further, environmental investigations often benefit from replicate data collection, which additionally increases study costs. In this paper we present a new camera system, developed with off the shelf and 3D printed cost effective components for use in shallow waters of <150 m depth. Integrating Raspberry Pi Zero W microcomputers with open source design files and software, it is hoped these camera systems will be of interest to the global marine and freshwater research communities.

New High-Tech Flexible Networks for the Monitoring of Deep-Sea Ecosystems.

Increasing interest in the acquisition of biotic and abiotic resources from within the deep sea (e.g., fisheries, oil-gas extraction, and mining) urgently imposes the development of novel monitoring technologies, beyond the traditional vessel-assisted, time-consuming, high-cost sampling surveys. The implementation of permanent networks of seabed and water-column-cabled (fixed) and docked mobile platforms is presently enforced, to cooperatively measure biological features and environmental (physicochemical) parameters. Video and acoustic (i.e., optoacoustic) imaging are becoming central approaches for studying benthic fauna (e.g., quantifying species presence, behavior, and trophic interactions) in a remote, continuous, and prolonged fashion. Imaging is also being complemented by in situ environmental-DNA sequencing technologies, allowing the traceability of a wide range of organisms (including prokaryotes) beyond the reach of optoacoustic tools. Here, we describe the different fixed and mobile platforms of those benthic and pelagic monitoring networks, proposing at the same time an innovative roadmap for the automated computing of hierarchical ecological information on deep-sea ecosystems (i.e., from single species' abundance and life traits to community composition, and overall biodiversity).

Depression chains in seafloor of contrasting morphology, Atacama Trench margin: a comment on Marsh et al. (2018).

This comment presents acoustic and visual data showing deep seafloor depression chains similar to those reported in Marsh et al. (R. Soc. open sci. 5: 180286), though from a different deep-sea setting. Marsh et al. present data collected during cruise JC120 from polymetallic nodule rich sites within the Clarion-Clipperton Fracture Zone (CCFZ), at water depths of between 3999 and 4258 m. Within this comment, we present data collected with equivalent acoustic and imaging devices on-board the RV Sonne (SO261-March/April 2018) from the Atacama Trench, approximately 4000 m depth, which shows comparable depression chains in the seafloor. In contrast with the CCFZ observations, our study area was wholly free of polymetallic nodules, an observation therefore weakening the 'ballast collection' by deep-sea diving mammals formation hypothesis discussed in their paper. We support their alternate hypothesis that if these features are indeed generated by deep-diving megafauna, then they are more likely the resultant traces of infauna feeding or marks made during opportunistic capture of benthic fish/cephalopods. We observed these potential prey fauna with lander and towed camera systems during the cruise, with example images of these presented here. Both the SO261 and JC120 cruises employed high-resolution sidescan systems at deployment altitudes seldom used routinely until the last few years during scientific deep-sea surveys. Given that both cruises found these depression chains in contrasting physical regions of the East Pacific, they may have a more ubiquitous distribution than at just these sites. Thus, the impacts of cetacean foraging behaviour on deep seafloor communities, and the potential relevance of these prey sources to deep-diving species, should be considered.

Underwater hyperspectral imaging as an in situ taxonomic tool for deep-sea megafauna.

Identification of benthic megafauna is commonly based on analysis of physical samples or imagery acquired by cameras mounted on underwater platforms. Physical collection of samples is difficult, particularly from the deep sea, and identification of taxonomic morphotypes from imagery depends on resolution and investigator experience. Here, we show how an Underwater Hyperspectral Imager (UHI) can be used as an alternative in situ taxonomic tool for benthic megafauna. A UHI provides a much higher spectral resolution than standard RGB imagery, allowing marine organisms to be identified based on specific optical fingerprints. A set of reference spectra from identified organisms is established and supervised classification performed to identify benthic megafauna semi-autonomously. The UHI data provide an increased detection rate for small megafauna difficult to resolve in standard RGB imagery. In addition, seafloor anomalies with distinct spectral signatures are also detectable. In the region investigated, sediment anomalies (spectral reflectance minimum at ~675 nm) unclear in RGB imagery were indicative of chlorophyll a on the seafloor. Underwater hyperspectral imaging therefore has a great potential in seafloor habitat mapping and monitoring, with areas of application ranging from shallow coastal areas to the deep sea.

The Oceanic Biological Pump: Rapid carbon transfer to depth at Continental Margins during Winter.

The oceanic biological pump is responsible for the important transfer of CO2-C as POC "Particulate Organic Carbon" to the deep sea. It plays a decisive role in the Earth's carbon cycle and significant effort is spent to quantify its strength. In this study we used synchronized daily time-series data of surface chlorophyll-a concentrations from the NASA's MODIS satellite in combination with hourly to daily observations from sea surface buoys and from an Internet Operated Vehicle (IOV) on the seafloor within Barkley Canyon (Northeast Pacific) to investigate the importance of winter processes in the export of fresh phytodetritus. The results indicate that phytoplankton pulses during winter can be as important in POC transfer to depth as the pulses associated with spring and summer blooms. Short winter phytoplankton pulses were observed to disappear from surface waters after low-pressure systems affected the area. Pulses of chlorophyll reached the IOV, at 870 m depth on the canyon seafloor, 12-72 hours later. These observed short pulses of biological carbon production regularly observed in the region from December to March have not been considered a significant component of the biological pump when compared with the denser summer productivity blooms.

Seasonal monitoring of deep-sea megabenthos in Barkley Canyon cold seep by internet operated vehicle (IOV).

Knowledge of the processes shaping deep-sea benthic communities at seasonal scales in cold-seep environments is incomplete. Cold seeps within highly dynamic regions, such as submarine canyons, where variable current regimes may occur, are particularly understudied. Novel Internet Operated Vehicles (IOVs), such as tracked crawlers, provide new techniques for investigating these ecosystems over prolonged periods. In this study a benthic crawler connected to the NEPTUNE cabled infrastructure operated by Ocean Networks Canada was used to monitor community changes across 60 m2 of a cold-seep area of the Barkley Canyon, North East Pacific, at ~890 m depth within an Oxygen Minimum Zone (OMZ). Short video-transects were run at 4-h intervals during the first week of successive calendar months, over a 14 month period (February 14th 2013 to April 14th 2014). Within each recorded transect video megafauna abundances were computed and changes in environmental conditions concurrently measured. The responses of fauna to environmental conditions as a proxy of seasonality were assessed through analysis of abundances in a total of 438 video-transects (over 92 h of total footage). 7698 fauna individuals from 6 phyla (Cnidaria, Ctenophora, Arthropoda, Echinodermata, Mollusca, and Chordata) were logged and patterns in abundances of the 7 most abundant taxa (i.e. rockfish Sebastidae, sablefish Anoplopoma fimbria, hagfish Eptatretus stoutii, buccinids (Buccinoidea), undefined small crabs, ctenophores Bolinopsis infundibulum, and Scyphomedusa Poralia rufescens) were identified. Patterns in the reproductive behaviour of the grooved tanner crab (Chionnecetes tanneri) were also indicated. Temporal variations in biodiversity and abundance in megabenthic fauna was significantly influenced by variabilities in flow velocity flow direction (up or down canyon), dissolved oxygen concentration and month of study. Also reported here for the first time are transient mass aggregations of grooved tanner crabs through these depths of the canyon system, in early spring and likely linked to the crab's reproductive cycle.

Resilience of benthic deep-sea fauna to mining activities.

With increasing demand for mineral resources, extraction of polymetallic sulphides at hydrothermal vents, cobalt-rich ferromanganese crusts at seamounts, and polymetallic nodules on abyssal plains may be imminent. Here, we shortly introduce ecosystem characteristics of mining areas, report on recent mining developments, and identify potential stress and disturbances created by mining. We analyze species' potential resistance to future mining and perform meta-analyses on population density and diversity recovery after disturbances most similar to mining: volcanic eruptions at vents, fisheries on seamounts, and experiments that mimic nodule mining on abyssal plains. We report wide variation in recovery rates among taxa, size, and mobility of fauna. While densities and diversities of some taxa can recover to or even exceed pre-disturbance levels, community composition remains affected after decades. The loss of hard substrata or alteration of substrata composition may cause substantial community shifts that persist over geological timescales at mined sites.

Association of deep-sea incirrate octopods with manganese crusts and nodule fields in the Pacific Ocean.

Incirrate octopods (those without fins) are among the larger megafauna inhabiting the benthic environments of all oceans, commonly in water depths down to about 3,000 m. They are known to protect and brood their eggs until the juveniles hatch, but to date there is little published information on octopod deep-sea life cycles and distribution. For this study, three manganese-crust and nodule-abundant regions of the deep Pacific were examined by remote operated-vehicle and towed camera surveys carried out between 2011 and 2016. Here, we report that the depth range of incirrate octopods can now be extended to at least 4,290 m. Octopods (twenty-nine individuals from two distinct species) were observed on the deep Ka'ena and Necker Ridges of the Hawaiian Archipelago, and in a nodule-abundant region of the Peru Basin. Two octopods were observed to be brooding clutches of eggs that were laid on stalks of dead sponges attached to nodules at depths exceeding 4,000 m. This is the first time such a specific mineral-biota association has been observed for incirrate octopods. Both broods consisted of approximately 30 large (2.0-2.7 cm) eggs. Given the low annual water temperature of 1.5oC, it is likely that egg development, and hence brooding, takes years [1]. Stalked-sponge fauna in the Peru Basin require the presence of manganese nodules as a substrate, and near total collapse of such sponge populations was observed following the experimental removal of nodules within the DISCOL (DISturbance and COLonisation) area of the Peru Basin [2]. Stalked fauna are also abundant on the hard substrates of the Hawaiian archipelago. The brooding behavior of the octopods we observed suggests that, like the sponges, they may also be susceptible to habitat loss following the removal of nodule fields and crusts by commercial exploitation.

High-Frequency Patterns in the Abundance of Benthic Species near a Cold-Seep - An Internet Operated Vehicle Application.

Three benthic megafaunal species (i.e. sablefish Anoplopoma fimbria; pacific hagfish Eptatretus stoutii and a group of juvenile crabs) were tested for diel behavioral patterns at the methane hydrates site of Barkley Canyon (890 m depth), off Vancouver Island (BC, Canada). Fluctuations of animal counts in linear video-transects conducted with the Internet Operated Deep-Sea Crawler "Wally" in June, July and December of 2013, were used as proxy of population activity rhythms. Count time series and environmental parameters were analyzed under the hypothesis that the environmental conditioning of activity rhythms depends on the life habits of particular species (i.e. movement type and trophic level). Non-linear least squares modeling of biological time series revealed significant diel periods for sablefish in summer and for hagfish and crabs in December. Combined cross-correlation and redundancy (RDA) analyses showed strong relationships among environmental fluctuations and detected megafauna. In particular, sablefish presence during summer months was related to flow magnitude, while the activity of pacific hagfish and juvenile crabs in December correlated with change in chemical parameters (i.e. chlorophyll and oxygen concentrations, respectively). Waveform analyses of animal counts and environmental variables confirmed the phase delay during the 24 h cycle. The timing of detection of sablefish occurred under low flow velocities, a possible behavioral adaptation to the general hypoxic conditions. The proposed effect of chlorophyll concentrations on hagfish counts highlights the potential role of phytodetritus as an alternative food source for this opportunistic feeder. The juvenile crabs seemed to display a cryptic behavior, possibly to avoid predation, though this was suppressed when oxygen levels were at a minimum. Our results highlight the potential advantages such mobile observation platforms offer in multiparametric deep-sea monitoring in terms of both spatial and temporal resolution and add to the vastly understudied field of diel rhythms of deep-sea megafauna.