In situ observation of a macrourid fish at 7259†m in the Japan Trench: swimbladder buoyancy at extreme depth.

A macrourid, Coryphaenoides yaquinae sp. inc., was observed to be attracted to bait and exhibiting normal foraging behaviour during a period of 80 min within view of a baited video camera on the sea floor at 7259 m - the deepest ever observation of a fish species with a swim bladder. The buoyancy provided by an oxygen-filled swim bladder at 74.4 MPa pressure was estimated to be 0.164 N, at a theoretical energy cost of 20 kJ, 200 times less than the cost of equivalent lipid buoyancy. During normal metabolism, 192 days would be required to fill the swimbladder. At these depths, oxygen is very incompressible, so changes in volume during ascent or descent are small. However, swimbladder function is crucially dependent on a very low rate of diffusion of oxygen across the swimbladder wall. The oxygen in the swimbladder could theoretically sustain aerobic metabolism for over 1 year but is unlikely to be used as a reserve.

Large effective population size masks population genetic structure in Hirondellea amphipods within the deepest marine ecosystem, the Mariana Trench.

The examination of genetic structure in the deep-ocean hadal zone has focused on divergence between tectonic trenches to understand how environment and geography may drive species divergence and promote endemism. There has been little attempt to examine localized genetic structure within trenches, partly because of logistical challenges associated with sampling at an appropriate scale, and the large effective population sizes of species that can be sampled adequately may mask underlying genetic structure. Here we examine genetic structure in the superabundant amphipod Hirondellea gigas in the Mariana Trench at depths of 8126-10,545 m. RAD sequencing was used to identify 3182 loci containing 43,408 single nucleotide polymorphisms (SNPs) across individuals after stringent pruning of loci to prevent paralogous multicopy genomic regions being erroneously merged. Principal components analysis of SNP genotypes resolved no genetic structure between sampling locations, consistent with a signature of panmixia. However, discriminant analysis of principal components identified divergence between all sites driven by 301 outlier SNPs in 169 loci and significantly associated with latitude and depth. Functional annotation of loci identified differences between singleton loci used in analysis and paralogous loci pruned from the data set and also between outlier and nonoutlier loci, all consistent with hypotheses explaining the role of transposable elements driving genome dynamics. This study challenges the traditional perspective that highly abundant amphipods within a trench form a single panmictic population. We discuss the findings in relation to eco-evolutionary and ontogenetic processes operating in the deep sea, and highlight key challenges associated with population genetic analysis in nonmodel systems with inherent large effective population sizes and genomes.

Mercury isotopes identify near-surface marine mercury in deep-sea trench biota.

Mercury isotopic compositions of amphipods and snailfish from deep-sea trenches reveal information on the sources and transformations of mercury in the deep oceans. Evidence for methyl-mercury subjected to photochemical degradation in the photic zone is provided by odd-mass independent isotope values (Δ199Hg) in amphipods from the Kermadec Trench, which average 1.57‰ (±0.14, n = 12, SD), and amphipods from the Mariana Trench, which average 1.49‰ (±0.28, n = 13). These values are close to the average value of 1.48‰ (±0.34, n = 10) for methyl-mercury in fish that feed at ∼500-m depth in the central Pacific Ocean. Evidence for variable contributions of mercury from rainfall is provided by even-mass independent isotope values (Δ200Hg) in amphipods that average 0.03‰ (±0.02, n = 12) for the Kermadec and 0.07‰ (±0.01, n = 13) for the Mariana Trench compared to the rainfall average of 0.13 (±0.05, n = 8) in the central Pacific. Mass-dependent isotope values (δ202Hg) are elevated in amphipods from the Kermadec Trench (0.91 ±0.22‰, n = 12) compared to the Mariana Trench (0.26 ±0.23‰, n = 13), suggesting a higher level of microbial demethylation of the methyl-mercury pool before incorporation into the base of the foodweb. Our study suggests that mercury in the marine foodweb at ∼500 m, which is predominantly anthropogenic, is transported to deep-sea trenches primarily in carrion, and then incorporated into hadal (6,000-11,000-m) food webs. Anthropogenic Hg added to the surface ocean is, therefore, expected to be rapidly transported to the deepest reaches of the oceans.

Evidence for seasonal cycles in deep-sea fish abundances: A great migration in the deep SE Atlantic?

Animal migrations are of global ecological significance, providing mechanisms for the transport of nutrients and energy between distant locations. In much of the deep sea (>200 m water depth), the export of nutrients from the surface ocean provides a crucial but seasonally variable energy source to seafloor ecosystems. Seasonal faunal migrations have been hypothesized to occur on the deep seafloor as a result, but have not been documented. Here, we analyse a 7.5-year record of photographic data from the Deep-ocean Environmental Long-term Observatory Systems seafloor observatories to determine whether there was evidence of seasonal (intra-annual) migratory behaviours in a deep-sea fish assemblage on the West African margin and, if so, identify potential cues for the behaviour. Our findings demonstrate a correlation between intra-annual changes in demersal fish abundance at 1,400 m depth and satellite-derived estimates of primary production off the coast of Angola. Highest fish abundances were observed in late November with a smaller peak in June, occurring approximately 4 months after corresponding peaks in primary production. Observed changes in fish abundance occurred too rapidly to be explained by recruitment or mortality, and must therefore have a behavioural driver. Given the recurrent patterns observed, and the established importance of bottom-up trophic structuring in deep-sea ecosystems, we hypothesize that a large fraction of the fish assemblage may conduct seasonal migrations in this region, and propose seasonal variability in surface ocean primary production as a plausible cause. Such trophic control could lead to changes in the abundance of fishes across the seafloor by affecting secondary production of prey species and/or carrion availability for example. In summary, we present the first evidence for seasonally recurring patterns in deep-sea demersal fish abundances over a 7-year period, and demonstrate a previously unobserved level of dynamism in the deep sea, potentially mirroring the great migrations so well characterized in terrestrial systems.

As migrações dos animais são importantes para a ecologia global pois fornecem mecanismos para o transporte de nutrientes e energia entre diferentes locais. Em grande parte do oceano profundo (>200 m de profundidade), a exportação de nutrientes da superfície para os ecossistemas do fundo marinho é uma fonte de energia crucial, mas que varia entre estações. Consequentemente, calcula-se que ocorram migrações sazonais de animais no fundo marinho, mas tal nunca foi reportado. Neste estudo, nós analisamos dados fotográficos do observatório do fundo marinho DELOS colhidos ao longo de 7.5 anos, para determinar se existem indícios de comportamentos migratórios sazonais (intra-anuais) na comunidade de peixes de profundidade na costa oeste africana ao largo de Angola e, se se confirmar, tentar identificar o que desencadeia este comportamento. Os resultados obtidos mostram que há uma correlação entre as alterações intra-anuais da densidade de peixes demersais a 1,400 m de profundidade e as estimativas de produção primária obtidas por satélite. A densidade de peixes atinge o seu máximo no final de Novembro, com um pico menos acentuado em Junho, aproximadamente quatro meses após os respectivos picos de produção primária na superfície. As alterações na densidade de peixe ocorreram de uma forma tão rápida que não podem ser explicadas por recrutamento ou mortalidade, e como tal devem só podem ser geradas por uma alteração do comportamento. Dado a recorrência do padrão observado, e importância da estrutura trófica fundo-topo em ecossistemas do oceano profundo, nós colocamos a hipótese de que uma fracção grande da comunidade de peixes faz migrações sazonais nesta região, e propomos que a variação sazonal da produção primária na superfície esteja na sua origem. Este controlo trófico poderá levar a alterações na densidade de peixes no fundo marinho via, por exemplo, a produção secundária de presas e/ou disponibilidade de corpos em processo de decomposição. Resumindo, nós apresentamos aqui a primeira evidência de padrões sazonais recorrentes na densidade de peixes demersais de profundidade ao longo de um período de sete anos, e provamos existir um nível de dinamismo nunca dantes observado no oceano profundo, que poderá espelhar as grandes migrações comummente observadas em sistemas terrestres.

Isolation and Characterization of Microsatellite DNA Markers in the Deep-Sea Amphipod Paralicella tenuipes by Illumina MiSeq Sequencing.

Here, we describe the development of 16 polymorphic microsatellite markers using an Illumina MiSeq sequencing approach in the deep-sea amphipod Paralicella tenuipes A total of 25 577 844 DNA sequences were filtered for microsatellite motifs of which 197 873 sequences were identified. From these sequences, 64 had sufficient flanking regions for primer design and 16 of these loci were polymorphic. Between 5 and 30 alleles were detected per locus, with an average of 13.63 alleles per locus, across a total of 120 individuals from 5 separate deep sea trenches from the Pacific Ocean. For the 16 loci, observed and expected heterozygosity values ranged from 0.116 to 0.414 and 0.422 to 0.820, respectively, with one locus displaying significant deviation from Hardy-Weinberg equilibrium. The microsatellite loci that have been isolated and described here are the first molecular markers developed for deep sea amphipods and will be invaluable for elucidating the genetic population structure and the extent of connectivity between deep ocean trenches.

Deep-sea ecosystems of the Indian Ocean >1000 m.

The Indian Ocean is the third largest of the world's oceans, accounting for ~20 % for the global marine realm. It is geomorphologically complex, hosting a wide variety of ecosystems across basins, trenches, seamounts, ridges, and fracture zones. While modern exploration has contributed significantly to our knowledge of its coastal ecosystems, deeper waters (>1000 m) remain relatively unknown despite accounting for over 90 % of its total area. This study provides the first comprehensive review of the Indian Ocean's diverse deep sea, presenting ecosystem knowledge summaries for each major seafloor feature, contextualised with the broader historical, socioeconomic, geological, and oceanographic conditions. Unsurprisingly, some ecosystems are better characterised than others, from the relatively well-surveyed Java (Sunda) Trench and hydrothermal vents of the Carlsberg, Central and Southwest Indian Ridges, to the unexplored Southeast Indian Ridge and hadal features of the western Indian Ocean. Similarly, there is a large depth discrepancy in available records with a clear bias towards shallower sampling. We identify four outstanding problems to be addressed for the advancement of deep-sea research in the Indian Ocean: 1) inconsistencies in research extent and effort over spatial scales, 2) severe lack of data over temporal scales, 3) unexplored deep pelagic environments, and 4) a need to place the Indian Ocean's deep-sea ecosystems in a global context. By synthesising and championing existing research, identifying knowledge gaps, and presenting the outstanding problems to be addressed, this review provides a platform to ensure this forgotten ocean is prioritised for deep-sea research during the UN Ocean Decade and beyond.

Submarine canyon development controlled by slope failure and oceanographic process interactions.

The southeastern Australian margin hosts a series of submarine canyons. Although the origin and evolution of canyons within the northwestern segment of the margin is relatively well studied, their quantitative morphology, interaction with longshore drift currents and slope failure remain poorly understood in the southeastern region. In this study, high-resolution bathymetry and 3D seismic reflection datasets revealed five main submarine canyons present in the central offshore Otway Basin. The canyons have V-shaped, evolving to U-shaped morphology downslope with sedimentary infill characterized by medium-high amplitudes, recognizable stratal pattern and localized chaotic seismic reflections. Analysis reveals these canyons were initiated by retrogressive slope failure on the continental slope, that once the shelf edge was reached and subsequently incised, development of the canyon heads was influenced by the shelf parallel Zeehan Current, active on the continental shelf of the region. The heads of the shelf-incised canyons preferentially migrate northwestward and were infilled by laterally accreting sedimentary packages characterized by southeast dipping seismic reflections. These indicate an early erosive phase followed by a period of deposition, a result of the prevalent eastward flowing Zeehan Current. These results have important implications for understanding of the mechanisms that control initiation and development of submarine canyons, and their morphology, both offshore southeastern Australia, and similar settings on continental margins worldwide.

Bone-eating worms from the Antarctic: the contrasting fate of whale and wood remains on the Southern Ocean seafloor.

We report the results from the first experimental study of the fate of whale and wood remains on the Antarctic seafloor. Using a baited free-vehicle lander design, we show that whale-falls in the Antarctic are heavily infested by at least two new species of bone-eating worm, Osedax antarcticus sp. nov. and Osedax deceptionensis sp. nov. In stark contrast, wood remains are remarkably well preserved with the absence of typical wood-eating fauna such as the xylophagainid bivalves. The combined whale-fall and wood-fall experiment provides support to the hypothesis that the Antarctic circumpolar current is a barrier to the larvae of deep-water species that are broadly distributed in other ocean basins. Since humans first started exploring the Antarctic, wood has been deposited on the seafloor in the form of shipwrecks and waste; our data suggest that this anthropogenic wood may be exceptionally well preserved. Alongside the new species descriptions, we conducted a comprehensive phylogenetic analyses of Osedax, suggesting the clade is most closely related to the frenulate tubeworms, not the vestimentiferans as previous reported.

Locomotory activity and feeding strategy of the hadal munnopsid isopod Rectisura cf. herculea (Crustacea: Asellota) in the Japan Trench.

Benthic fauna in the hadal zone (6500-11,000 m) rely on maintaining sufficient locomotory activity to exploit a low, patchy and uniquely distributed food supply while exposed to high pressure, low temperatures and responding to predator-prey interactions. Very little is currently known about the locomotory capabilities of hadal fauna. In situ video footage of the isopod Rectisura cf. herculea (Birstein 1957) (Asellota, Munnopsidae) was obtained from 6945 and 7703 m deep in the Japan Trench (NW Pacific Ocean). Measurements of locomotion revealed routine walking speeds of 0.19 ± 0.04 BL s(-1) (mean ± s.d.), increasing to 0.33 ± 0.04 BL s(-1) if naturally perturbed by larger organisms. When immediately threatened by the presence of predators (decapod crustaceans), the isopods are capable of eliciting backward escape jumps and burst swimming escape responses of 2.6 ± 1.5 BL s(-1) and 4.63 ± 0.9 BL s(-1), respectively. These data suggest no significant reduction in locomotory capability despite the extreme depths in which they inhabit. These observations also revealed the isopod to be a bait-attending and aggregative species and suggest that it may not be obligatorily selecting infaunal food sources as previously thought.

No recovery of a large-scale anthropogenic sediment disturbance on the Pacific seafloor after 77 years at 6460 m depth.

Habitat restoration and recolonisation of benthic communities after physical perturbation in the deep sea has long been thought to be extremely slow. This study reports on a serendipitous opportunity to survey the current state of a large mechanical disturbance of sediments at 6460 m in the Pacific Ocean. The impact was caused 77 years ago by the sinking of the USS Johnston. The surrounding debris field had little impact on the sedimentary habitat, other than in the provision of artificial hard substrates, while the troughs that formed as the ship impacted the seafloor and slid down the slope of the Philippine Trench were still completely void of animal tracks and burrows, or any observable epifauna, and in some areas subsurface stratification was still exposed at the surface. This suggests that mechanical perturbations of sediments in the deep Pacific may remain ecologically significant for, at the very least, 100 years.

Barriers to gene flow in the deepest ocean ecosystems: Evidence from global population genomics of a cosmopolitan amphipod.

The deepest marine ecosystem, the hadal zone, hosts endemic biodiversity resulting from geographic isolation and environmental selection pressures. However, the pan-ocean distribution of some fauna challenges the concept that the hadal zone is a series of isolated island-like habitats. Whether this remains true at the population genomic level is untested. We investigated phylogeographic patterns of the amphipod, Bathycallisoma schellenbergi, from 12 hadal features across the Pacific, Atlantic, Indian, and Southern oceans and analyzed genome-wide single-nucleotide polymorphism markers and two mitochondrial regions. Despite a cosmopolitan distribution, populations were highly restricted to individual features with only limited gene flow between topographically connected features. This lack of connectivity suggests that populations are on separate evolutionary trajectories, with evidence of potential cryptic speciation at the Atacama Trench. Together, this global study demonstrates that the shallower ocean floor separating hadal features poses strong barriers to dispersal, driving genetic isolation and creating pockets of diversity to conserve.

Microbiomes of Hadal Fishes across Trench Habitats Contain Similar Taxa and Known Piezophiles.

Hadal snailfishes are the deepest-living fishes in the ocean, inhabiting trenches from depths of ∼6,000 to 8,000 m. While the microbial communities in trench environments have begun to be characterized, the microbes associated with hadal megafauna remain relatively unknown. Here, we describe the gut microbiomes of two hadal snailfishes, Pseudoliparis swirei (Mariana Trench) and Notoliparis kermadecensis (Kermadec Trench), using 16S rRNA gene amplicon sequencing. We contextualize these microbiomes with comparisons to the abyssal macrourid Coryphaenoides yaquinae and the continental shelf-dwelling snailfish Careproctus melanurus. The microbial communities of the hadal snailfishes were distinct from their shallower counterparts and were dominated by the same sequences related to the Mycoplasmataceae and Desulfovibrionaceae. These shared taxa indicate that symbiont lineages have remained similar to the ancestral symbiont since their geographic separation or that they are dispersed between geographically distant trenches and subsequently colonize specific hosts. The abyssal and hadal fishes contained sequences related to known, cultured piezophiles, microbes that grow optimally under high hydrostatic pressure, including Psychromonas, Moritella, and Shewanella. These taxa are adept at colonizing nutrient-rich environments present in the deep ocean, such as on particles and in the guts of hosts, and we hypothesize they could make a dietary contribution to deep-sea fishes by degrading chitin and producing fatty acids. We characterize the gut microbiota within some of the deepest fishes to provide new insight into the diversity and distribution of host-associated microbial taxa and the potential of these animals, and the microbes they harbor, for understanding adaptation to deep-sea habitats. IMPORTANCE Hadal trenches, characterized by high hydrostatic pressures and low temperatures, are one of the most extreme environments on our planet. By examining the microbiome of abyssal and hadal fishes, we provide insight into the diversity and distribution of host-associated life at great depth. Our findings show that there are similar microbial populations in fishes geographically separated by thousands of miles, reflecting strong selection for specific microbial lineages. Only a few psychropiezophilic taxa, which do not reflect the diversity of microbial life at great depth, have been successfully isolated in the laboratory. Our examination of deep-sea fish microbiomes shows that typical high-pressure culturing methodologies, which have largely remained unchanged since the pioneering work of Claude ZoBell in the 1950s, may simulate the chemical environment found in animal guts and helps explain why the same deep-sea genera are consistently isolated.

Trench connection.

'Trench Connection' was the first international symposium focusing primarily on the hadal zone (depths greater than 6000 m). It was held at the University of Tokyo's Atmosphere and Ocean Research Institute in November 2010. The symposium was successful in having attracted an international collective of scientists and engineers to discuss the latest developments in the exploration and understanding of the deepest environments on Earth. The symposium sessions were categorized into three themes: (i) new deep-submergence technology; (ii) trench ecology and evolution; and (iii) the physical environment. Recent technological developments have overcome the challenges of accessing the extreme depths, which have in turn prompted an international renewed interest in researching physical and biological aspects of the hadal ecosystems. This bringing together of international participants from different disciplines led to healthy discussions throughout the symposium, providing potential opportunities and realizations of where the future of unravelling hadal ecology lies. Hadal science is still at relatively rudimentary levels compared with those of shallower marine environments; however, it became apparent at the symposium that it is now an ever-expanding scientific field.

Eurythenes atacamensis sp. nov. (Crustacea: Amphipoda) exhibits ontogenetic vertical stratification across abyssal and hadal depths in the Atacama Trench, eastern South Pacific Ocean.

Eurythenes S.I. Smith in Scudder, 1882 (Crustacea: Amphipoda) are prevalent scavengers of the benthopelagic community from bathyal to hadal depths. While a well-studied genus, molecular systematic studies have uncovered cryptic speciation and multiple undescribed lineages. Here, we apply an integrative taxonomic approach and describe the tenth species, Eurythenes atacamensis sp. nov., based on specimens from the 2018 Atacamex and RV Sonne SO261 Expeditions to the southern sector of the Peru-Chile Trench, the Atacama Trench (24-⁠21°S). Eurythenes atacamensis sp. nov. is a large species, max. observed length 83.2 mm, possesses diagnostic features, including a short gnathopod 1 palm and a chelate gnathopod 2 palm, and a distinct genetic lineage based on a 16S rRNA and COI phylogeny. This species is a dominant bait-attending fauna with an extensive bathymetric range, spanning from 4974 to 8081 m. The RV Sonne SO261 specimens were recovered along a 10-station transect from abyssal to hadal depths and further examined for demographic and bathymetric-related patterns. Ontogenetic vertical stratification was evident across the trench axis, with only juveniles present at abyssal depths (4974-6025 m). Total length-depth analysis revealed that the size of females was unrelated to depth, whereas juveniles followed a sigmoidal relationship with a step-up in size at depths >7200 m. Thus, these bathymetric trends suggest that juveniles and females employ differing ecological strategies in subduction trench environments. This study highlights that even dominant and ecologically important species are still being discovered within the abyssal and hadal environments. Continued systematic expeditions will lead to an improved understanding of the eco-evolutionary drivers of speciation in the world's largest ecosystem.

Exploration of the Puerto Rico Trench in the mid-twentieth century: Today's significance and relevance.

The Puerto Rico Trench is a deep oceanic subduction zone that runs parallel with the northern coasts of Puerto Rico and the Dominican Republic. It is the deepest place in the Atlantic Ocean with a maximum depth of approximately 8400 m. Discovered by the HMS Challenger Expedition in 1875, the depth of the trench increased multiple times in the ensuing 100 years with the onset of sonar usage. It is perhaps unique among the world's deep trenches in that a series of unrelated but equally pioneering expeditions captured the true biological and geological characteristics of one of the deepest places in the world, observations that are still highly relevant today. Multiple deep water trawling campaigns and surveys using drop cameras and exploratory dives in a deep diving submersible provided great insight into the morphology of the trench, the types of habitat within the trench, the substrate, the food supply, and the diversity of species that inhabit these extraordinary depths. Many of these accounts are obscure and disparate, yet combined bear a remarkable similarity to recent work in other trenches. These unique and insightful accounts are collated and retold here alongside recent and comparable findings to contextualise these discoveries, prevent them from being forgotten, and keep the efforts of those involved to remain relevant as we continue to explore the deepest places of the world's oceans.

New species of Eurythenes from hadal depths of the Mariana Trench, Pacific Ocean (Crustacea: Amphipoda).

Eurythenes S. I. Smith in Scudder, 1882 are one of the largest scavenging deep-sea amphipods (max. 154 mm) and are found in every ocean across an extensive bathymetric range from the shallow polar waters to hadal depths. Recent systematic studies of the genus have illuminated a cryptic species complex and highlighted the benefits of using a combination of morphological and molecular identification approaches. In this study, we present the ninth species, Eurythenes plasticus sp. nov., which was recovered using baited traps between the depths 6010 and 6949 m in the Mariana Trench (Northwest Pacific Ocean) in 2014. This new Eurythenes species was found to have distinct morphological characteristics and be a well-supported clade based on sequence variation at two mitochondrial regions (16S rDNA and COI). While this species is new to science and lives in the remote hadal zone, it is not exempt from the impacts of anthropogenic pollution. Indeed, one individual was found to have a microplastic fibre, 83.74% similar to polyethylene terephthalate (PET), in its hindgut. As this species has a bathymetric range spanning from abyssal to hadal depths in the Central Pacific Ocean basin, it offers further insights into the biogeography of Eurythenes.

Whole genome sequencing of four bacterial strains from South Shetland Trench revealing biosynthetic and environmental adaptation gene clusters.

Whole genome sequences of four bacterial strains Dietzia maris SST1, Pseudomonas zhaodongensis SST2, Pseudomonas sp. SST3 and Halomonas sulfidaeris SST4, recovered from the South Shetland Trench sediment in Antarctica were analyzed using Ion Torrent sequencing technology. The respective sizes of their genomes (3.88, 4.99, 5.60 and 4.25 Mb) and GC contents (70.0, 60.3, 59.9 and 53.8%) are in agreement with these values of other strains of the species. The bacterial strains displayed promising antimicrobial activity against a number of pathogenic bacterial and fungal species. Whole genomes have been assembled and biosynthetic gene clusters (BGCs) have been identified using the antibiotics and Secondary Metabolite Analysis Shell (antiSMASH) web platform. Comparative analysis of the genome sequences revealed that the strains host abundant BGCs encoding for terpenes, siderophores, arylpolyene, bacteriocins, and lassopeptides. Furthermore, the key stress-related genes were identified and their distribution provided an insight into how these isolates adapt to key marine environmental conditions. This comprehensive study is a contribution to understanding the nature of life on the deep-sea environments.

Ecological variables for developing a global deep-ocean monitoring and conservation strategy.

The deep sea (>200 m depth) encompasses >95% of the world's ocean volume and represents the largest and least explored biome on Earth (<0.0001% of ocean surface), yet is increasingly under threat from multiple direct and indirect anthropogenic pressures. Our ability to preserve both benthic and pelagic deep-sea ecosystems depends upon effective ecosystem-based management strategies and monitoring based on widely agreed deep-sea ecological variables. Here, we identify a set of deep-sea essential ecological variables among five scientific areas of the deep ocean: (1) biodiversity; (2) ecosystem functions; (3) impacts and risk assessment; (4) climate change, adaptation and evolution; and (5) ecosystem conservation. Conducting an expert elicitation (1,155 deep-sea scientists consulted and 112 respondents), our analysis indicates a wide consensus amongst deep-sea experts that monitoring should prioritize large organisms (that is, macro- and megafauna) living in deep waters and in benthic habitats, whereas monitoring of ecosystem functioning should focus on trophic structure and biomass production. Habitat degradation and recovery rates are identified as crucial features for monitoring deep-sea ecosystem health, while global climate change will likely shift bathymetric distributions and cause local extinction in deep-sea species. Finally, deep-sea conservation efforts should focus primarily on vulnerable marine ecosystems and habitat-forming species. Deep-sea observation efforts that prioritize these variables will help to support the implementation of effective management strategies on a global scale.

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.

Bathyal and abyssal demersal bait-attending fauna of the Eastern Mediterranean Sea.

Baited cameras were deployed over a depth range of 532-5111 m in the Ionian Sea to characterise the large mobile fauna. The planned installation of a neutrino telescope also offers the potential for biological observatories. The current study was intended to aid observatory placement. At increasing depths, sediment was observed to become more uniform and animal burrows and tracks reduced. A total of 10 species of deep-sea fishes were identified from images; four elasmobranchs, which were not recorded deeper than 1841 m, and six teleosts. At depths > 3000 m, including Calypso Deep, the deepest point in the Mediterranean, only one fish species was observed; the Mediterranean grenadier, Coryphaenoides mediterraneus (3400-5111 m), extending this species' maximum recorded depth to 5111 m. Four species of decapod crustacea could be identified from images. The dressed deep-sea shrimp, Acanthephyra eximia (1346-5111 m) was the only invertebrate recorded at abyssal depths, including the deepest point. A faunal change was detected at ~ 1000 m depth. Incorporating other studies from the Eastern Mediterranean identified additional faunal boundaries at ~ 1500 m and ~ 2500 m. The time from landing the observation equipment to the arrival of the first fish increased exponentially with depth at a slower rate to that observed in the Atlantic Ocean. The estimated density of bait-attending deep-sea fish was, therefore, significantly impoverished compared to the Atlantic Ocean at equivalent depth. Barriers to colonisation, low resource input, and high temperature at depth relative to the Atlantic Ocean are probable causes of the impoverished fauna.