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Seamounts, pinnacles, and crests are abrupt seafloor features that modify physical processes and ecological patterns. Fishers often target these local bathymetric highs, which can have high catch-per-unit-effort. Increases in the abundance of target species has been qualitatively noted around these features and promontories, however, a quantitative evaluation of local highs as preferred habitat for fishes is still lacking. Here, we used an extensive database of fish abundances (N = 2381) from mesophotic and upper bathyal depths (40-300 m) gathered over 8 yr around the Main Hawaiian Islands to evaluate the effects of macro-scale habitat categorized by combining bathymetric position index (BPI) and slope. A numerical model simulating physical ocean processes was used to test the hypothesis that local highs, herein called crests, are distinct and preferred habitat for many species due to their modified flow conditions. We show that crests host a unique, diverse fish assemblage with double the abundance relative to other habitats. Therefore, fishes are concentrated at crests rather than being uniformly distributed along the island shelf at preferred depths. These habitats are characterized by enhanced current amplitudes and convergence zones. Direct correlations between fish abundance, convergence, and current amplitude strongly suggest that flow-enhanced food availability is likely the driver of habitat-related changes in fish assemblage. These results have important implications for conservation and management. We identify a simple and informative method to classify habitat, quantitatively link bathymetry-induced flow alterations to fish abundances, and provide information for refining definitions of essential fish habitat for species that are important commercial targets worldwide.

Highlights • All known observations for Area of Particular Environmental Interest 6 presented. • Assess morphology, sediments, nodules, oceanography, biogeochemistry and ecology. • APEI-6 partially representative of nearby exploration areas yet clear differences. • Present scientific synthesis and management implications for Clarion Clipperton Zone. To protect the range of habitats, species, and ecosystem functions in the Clarion Clipperton Zone (CCZ), a region of interest for deep-sea polymetallic nodule mining in the Pacific, nine Areas of Particular Environmental Interest (APEIs) have been designated by the International Seabed Authority (ISA). The APEIs are remote, rarely visited and poorly understood. Here we present and synthesise all available observations made at APEI-6, the most north eastern APEI in the network, and assess its representativity of mining contract areas in the eastern CCZ. The two studied regions of APEI-6 have a variable morphology, typical of the CCZ, with hills, plains and occasional seamounts. The seafloor is predominantly covered by fine-grained sediments, and includes small but abundant polymetallic nodules, as well as exposed bedrock. The oceanographic parameters investigated appear broadly similar across the region although some differences in deep-water mass separation were evident between APEI-6 and some contract areas. Sediment biogeochemistry is broadly similar across the area in the parameters investigated, except for oxygen penetration depth, which reached >2 m at the study sites within APEI-6, deeper than that found at UK1 and GSR contract areas. The ecology of study sites in APEI-6 differs from that reported from UK1 and TOML-D contract areas, with differences in community composition of microbes, macrofauna, xenophyophores and metazoan megafauna. Some species were shared between areas although connectivity appears limited. We show that, from the available information, APEI-6 is partially representative of the exploration areas to the south yet is distinctly different in several key characteristics. As a result, additional APEIs may be warranted and caution may need to be taken in relying on the APEI network alone for conservation, with other management activities required to help mitigate the impacts of mining in the CCZ.

Industrial interest in deep-sea mineral extraction began decades ago, and today it is at an all-time high, accelerated by global demand for metals. Several seafloor ecosystem disturbance experiments began in the 1970s, including the Disturbance and Recolonization experiment (DISCOL) conducted in the Peru Basin in 1989. A large seafloor disturbance was created by repeatedly ploughing the seafloor over an area of ∼10.8 km2. Though a number of studies in abyssal mining regions have evaluated megafaunal biodiversity and ecosystem responses, few have included quantitative and detailed data on fishes or scavengers despite their ecological importance as top predators. We used towed camera transects (1989–1996, 2015) and baited camera data (1989–1992) to evaluate the fish community at the DISCOL site. The abyssal fish community included 16 taxa and was dominated by Ipnops meadi. Fish density was lower in ploughed habitat at 6 months and 3 years following disturbance but thereafter increased over time. Twenty-six years after disturbance there were no differences in overall total fish densities between reference and experimental areas, but the dominant fish, I. meadi, still exhibited much lower densities in ploughed habitat, likely avoiding these areas and suggesting that the fish community remains affected after decades. At the scale of industrial mining, these results could translate to population-level effects. The scavenging community was dominated by eelpouts (Pachycara spp.), hermit crabs (Probeebei mirabilis) and shrimp. The large contribution of hermit crabs appears to be unique amongst abyssal scavenger studies worldwide. The abyssal fish community at DISCOL was similar to that in the more northerly Clarion–Clipperton Zone (CCZ), though some species have only been observed at DISCOL thus far. Also, further species-level identifications are required to refine this assessment. Additional studies across the polymetallic nodule provinces of the Pacific are required to further evaluate the environmental drivers of fish density, diversity and species biogeographies. This information will be important for the development of appropriate management plans aimed at minimizing human impact from deep-sea mining.

The abyssal seafloor makes up three quarters of the ocean floor, and it is generally characterized as a food-limited habitat with low numbers of megafauna, particularly fishes. Baited camera observations from three abyssal seamount summits in the equatorial Pacific challenge this idea. On each of two deployments at the southernmost seamount, over 100 synaphobranchid eels (Ilyophis arx) were recorded feeding on standard bait (1 kg mackerel). This is the highest number of fishes per kg of bait ever recorded below 1000 m, including observations from large organic falls such as cetacean and shark carcasses. It is also the highest number that has ever been recorded at carrion of any kind or size at abyssal depths. We suggest an abyssal ‘seamount effect’ may be responsible, highlighting the potential importance of seamounts in structuring abyssal communities.

© 2020 The Authors The abyssal seafloor of the Clarion Clipperton Zone (CCZ) in the central Pacific has the largest known deposits of polymetallic nodules and associated benthic faunal communities with high biodiversity. The environmental factors that structure these communities, both at regional and local scales, are not well understood. In this study, seabed image surveys were used to assess distribution patterns in invertebrate and fish megafauna (>1 cm) at multiple scales in relation to key environmental factors: food supply to the seabed varying at the regional scale (hundreds of km), seabed geomorphological variations varying at the broad local scale (tens of km), and seabed nodule cover varying at the fine local scale (tens of meters). We found significant differences in megafaunal density and community composition between all study areas. Variations in faunal density did not appear to match with regional productivity gradients, although faunal density generally decreased with increasing water depth (from E to W). In contrast, geomorphology and particularly nodule cover appeared to exert strong control on local faunal abundance and community composition, but not in species richness. Local variations in faunal density and beta-diversity, particularly those driven by nodule presence (within study areas), were of comparable magnitude to those observed at a regional level (between study areas). However, regional comparisons of megabenthic assemblages showed clear shifts in dominance between taxonomic groups (perceivable even at Phylum levels) across the mid-eastern CCZ seabed, suggesting a higher regional heterogeneity than was previously thought.

Despite rapidly growing interest in deep-sea mineral exploitation, environmental research and management have focused on impacts to seafloor environments, paying little attention to pelagic ecosystems. Nonetheless, research indicates that seafloor mining will generate sediment plumes and noise at the seabed and in the water column that may have extensive ecological effects in deep midwaters (1), which can extend from an approximate depth of 200 meters to 5 kilometers. Deep midwater ecosystems represent more than 90% of the biosphere (2), contain fish biomass 100 times greater than the global annual fish catch (3), connect shallow and deep-sea ecosystems, and play key roles in carbon export (4), nutrient regeneration, and provisioning of harvestable fish stocks (5). These ecosystem services, as well as biodiversity, could be negatively affected by mining. Here we argue that deep-sea mining poses significant risks to midwater ecosystems and suggest how these risks could be evaluated more comprehensively to enable environmental resource managers and society at large to decide whether and how deep-sea mining should proceed. Midwater animal biodiversity: Squid, fish, shrimp, copepods, medusa, filter-feeding jellies, and marine worms are among the midwater creatures that could be affected by deep sea mining. Photos by E. Goetze, K. Peijnenburg, D. Perrine, Hawaii Seafood Council (B. Takenaka, J. Kaneko), S. Haddock, J. Drazen, B. Robison, DEEPEND (Dante Fenolio), and MBARI. Interest in deep-sea mining for sulfide deposits near hydrothermal vents, polymetallic nodules on the abyssal seafloor, and cobalt-rich crusts on seamounts (6) has grown substantially in the last decade. Equipment and system development are already occurring. The International Seabed Authority (ISA), the international organization created under the United Nations Convention on the Law of the Sea (UNCLOS) to manage deep-sea mining beyond national jurisdiction, is developing mineral exploitation regulations, the Mining Code. Currently, 30 ISA exploration contracts cover over 1.5 million … [↵][1]1To whom correspondence should be addressed. Email: jdrazen{at}hawaii.edu. [1]: #xref-corresp-1-1

In the Clarion-Clipperton Zone (CCZ) in the north-eastern Pacific Ocean ca. 30 billion tonnes of polymetallic nodules, rich in metals critical for frontier technologies, lay on the sediment surface over an area of 4–5 million square kilometres. For this reason, there is accelerating interest in deep-sea mineral mining in the CCZ. Few data exist concerning marine biodiversity in this enormous region and a solid understanding of baseline biodiversity and ecosystem function are necessary in order to inform effective management strategies, conservation and monitoring in the event that mining goes ahead in the CCZ. In this study, 10 in situ baited-camera experiments were carried out in the BGR (The Federal Institute for Geosciences and Natural Resources, Germany) licence area in the eastern CCZ to document the biodiversity of scavengers and their feeding activities. Twelve different bait-attending taxa were identified. The most common of these were macrourid, ophidiid and zoarcid fishes, and dendrobranchiate shrimp. Successional patterns regarding their time of arrival were also observed. The bait consumption rate was measured, and a mean scavenging rate of 878 g d–1 ± 113 (SE, n = 9) was calculated, with the rattail Coryphaenoides spp. responsible for the majority of the bait consumption. Polymetallic nodule size did not significantly affect community structure, diversity, or scavenging rates within the BGR licence area, but significant differences in community structure were found between the BGR area and other areas of the eastern CCZ (the Ocean Minerals of Singapore and United Kingdom 1 areas). This study increases our knowledge of deep-sea scavenging communities in an area targeted for deep-sea mining.

Seamounts are ubiquitous global features often characterized by biological hotspots of diversity, biomass, and abundance, though the mechanisms responsible are poorly understood. One controversial explanation suggests seamount-induced chlorophyll enhancements (SICE) subsidize seamount ecosystems. Using a decade of satellite chlorophyll data, we report substantial long-term chlorophyll enhancements around 17% of Pacific seamounts and 45% of shallow (

Aphyonids are poorly-known, live-bearing brotulas (Ophidiiformes, Bythitidae) that until recently were considered to be in a distinct family, Aphyonidae. A single, ca. 9.3 cm total length aphyonid observed during a remotely-operated vehicle survey in the Mariana Archipelago at 2504.2 m on Explorer Ridge (20.68152°N, 145.08750°E) is the first seen alive in its natural habitat. Collection to verify its identification was not possible, but based on observations it was a species of either Barathronus or Nybelinella. The fish swam 1–10 cm over sediment between rocks and small boulders on a 45° talus slope. Swimming speeds were consistently slow, 0.33 ± 0.15 body lengths per second, and the fish appeared to be neutrally buoyant. Although there are few other records of aphyonid-clade fishes in the Pacific away from continental margins, this observation suggests that they will be found elsewhere in the basin when appropriate methods are used to detect these small fishes in the high-relief, rugose habitats of central Pacific oceanic islands and seamounts.

The Clarion-Clipperton Zone (CCZ) is one of the richest manganese nodule provinces in the world and has recently become a focus area for manganese nodule mining interests. However, this vast area remains poorly studied and highly undersampled. In this study, the abyssal bait-attending fauna is documented for the first time using a series of baited camera deployments in various locations across the CCZ. A bait-attending community intermediate between those typical of the California margin and Hawaii was found in the larger CCZ area, generally dominated by rattail fishes, dendrobranchiate shrimp, and zoarcid and ophidiid fishes. Additionally, the western and eastern ends of the CCZ had different communities, with the western region characterized by decreased dominance of rattails and small shrimps and increased dominance of ophidiids (especially Bassozetus sp. and Barathrites iris ) and large shrimps. This trend may be related to increasing distance from the continental margin. We also test the hypothesis that bait-attending communities change across the CCZ in response to key environmental predictors, especially topography and nodule cover. Our analyses showed that higher nodule cover and elevated topography, as quantified using the benthic positioning index (BPI), increase bait-attending community diversity. Elevated topography generally had higher relative abundances, but taxa also showed differing responses to the BPI metric and bottom temperature, causing significant community compositional change over varying topography and temperatures. Larger individuals of the dominant scavenger in the CCZ, Coryphaenoides spp., were correlated with areas of higher nodule cover and with abyssal hills, suggesting these areas may be preferred habitat. Our results suggest that nodule cover is important to all levels of the benthic ecosystem and that nodule mining could have negative impacts on even the top-level predators and scavengers in the CCZ. Additionally, there is continuous change in diversity, dominance, and relative abundance across the CCZ and across gradients in bathymetric and oceanographic variables. This work increased the understanding of the biogeography of the demersal scavengers and top predators as well as the key environmental drivers of their distributions across the CCZ in order to better predict and manage the impacts of nodule mining.

The International Seabed Authority (ISA) is developing regulations to control the future exploitation of deep-sea mineral resources including sulphide deposits near hydrothermal vents, polymetallic nodules on the abyssal seafloor, and cobalt crusts on seamounts. Under the UN Convention on the Law of the Sea the ISA is required to adopt are taking measures to ensure the effective protection of the marine environment from harmful effects arising from mining-related activities. Contractors are required to generate environmental baselines and assess the potential environmental consequences of deep seafloor mining. Understandably, nearly all environmental research has focused on the seafloor where the most direct mining effects will occur. However, sediment plumes and other impacts (e.g., noise) from seafloor mining are likely to be extensive in the water column. Sediment plumes created on the seafloor will affect the benthic boundary layer which extends 10s to 100s of meters above the seafloor. Separation or dewatering of ore from sediment and seawater aboard ships will require discharge of a dewatering plume at some depth in the water column. It is important to consider the potential impacts of mining on the ocean’s midwaters (depths from ~200 m to the seafloor) because they provide vital ecosystem services and harbor substantial biodiversity. The better known epipelagic or sunlit surface ocean provisions the rest of the water column through primary production and export flux (This was not the focus at this workshop as the subject was considered too large and surface discharges are unlikely). It is also home to a diverse community of organisms including commercially important fishes such as tunas, billfish, and cephalopods that contribute to the economies of many countries. The mesopelagic or twilight zone (200-1000 m) is dimly lit and home to very diverse and abundant communities of organisms. Mesopelagic plankton and small nekton form the forage base for many deep-diving marine mammals and commercially harvested epipelagic species. Furthermore, detritus from the epipelagic zone falls through the mesopelagic where it is either recycled, providing the vital process of nutrient regeneration, or sinks to greater depths sequestering carbon from short-term atmospheric cycles. The waters below the mesopelagic down to the seafloor (both the bathypelagic and abyssopelagic) are very poorly characterized but are likely large reservoirs of novel biodiversity and link the surface and benthic ecosystems. Great strides have been made in understanding the biodiversity and ecosystem function of the ocean’s midwaters, but large regions, including those containing many exploration license areas and the greater depths where mining plumes will occur, remain very poorly studied. It is clear that pelagic communities are distinct from those on the seafloor and in the benthic boundary layer. They are often sampled with different instrumentation. The fauna have relatively large biogeographic ranges and they are more apt to mix freely across stakeholder boundaries, reference areas and other spatial management zones. Pelagic organisms live in a three-dimensional habitat and their food webs and populations are vertically connected by daily or lifetime migrations and the sinking flux of detritus from the epipelagic. The fauna do not normally encounter hard surfaces, making them fragile, and difficult to capture and maintain for sensitivity or toxicity studies. Despite some existing general knowledge, ecological baselines for midwater communities and ecosystems that likely will be impacted by mining have not been documented. There is an urgent need to conduct more research and evaluate the midwater biota (microbes to fishes) in regions where mining is likely to occur. Deep-sea mining activities may affect midwater organisms in a number of ways, but it is still unclear at what scale perturbations may occur. The sediment plumes both from collectors on the seafloor and from midwater discharge will have a host of negative consequences. They may cause respiratory distress from clogged gills or respiratory surfaces. Suspension feeders, such as copepods, polychaetes, salps, and appendicularians, that filter small particles from the water and form an important basal group of the food web, may suffer from dilution of their food by inorganic sediments and/or clogging of their fragile mucous filter nets. Small particles may settle on gelatinous plankton causing buoyancy issues. Metals, including toxic elements that will enter the food web, will be released from pore waters and crushed ore materials. Sediment plumes will also absorb light and change backscatter properties, reducing visual communication and bioluminescent signaling that are very important for prey capture and reproduction in midwater animals. Noise from mining activities may alter the behaviors of marine mammals and other animals. Small particles have high surface area to volume ratios, high pelagic persistence and dispersal and as a result greater potential to result in pelagic impacts. All of these potential effects will result in mortality, migration (both horizontal and vertical), decreased fitness, and shifts in community composition. Depending on the scale and duration of these effects, there could be reduction in provisioning to commercial fish species, delivery of toxic metals to pelagic food webs and hence human seafood supply, and alterations to carbon transport and nutrient regeneration services. After four days of presentations and discussions, the workshop participants came to several conclusions and synthesized recommendations. 1. Assuming no discharge in the epipelagic zone, it is essential to minimize mining effects in the mesopelagic zone because of links to our human seafood supply as well as other ecosystem services provided by the mesopelagic fauna. This minimization could be accomplished by delivering dewatering discharge well below the mesopelagic/bathypelagic transition (below ~1000 m depth). 2. Research should be promoted by the ISA and other bodies to study the bathypelagic and abyssopelagic zones (from ~1000 m depths to just above the seafloor). It is likely that both collector plumes and dewatering plumes will be created in the bathypelagic, yet this zone is extremely understudied and contains major unknowns for evaluating mining impacts. 3. Management objectives, regulations and management actions need to prevent the creation of a persistent regional scale “haze” (enhanced suspended particle concentrations) in pelagic midwaters. Such a haze would very likely cause chronic harm to deep midwater ecosystem biodiversity, structure and function. 4. Effort is needed to craft suitable standards, thresholds, and indicators of harmful environmental effects that are appropriate to pelagic ecosystems. In particular, suspension feeders are very important ecologically and are likely to be very sensitive to sediment plumes. They are a high priority for study. 5. Particularly noisy mining activities such as ore grinding at seamounts and hydrothermal vents is of concern to deep diving marine mammals and other species. One way to minimize sound impacts would be to minimize activities in the sound-fixing-and-ranging (SOFAR) channel (typically at depths of ~1000 m) which transmits sounds over very long distances. 6. A Lagrangian (drifting) perspective is needed in monitoring and management because the pelagic ecosystem is not a fixed habitat and mining effects are likely to cross spatial management boundaries. For example, potential broad-scale impacts to pelagic ecosystems should be considered in the deliberations over preservation reference zones, the choice of stations for environmental baseline and monitoring studies and other area-based management and conservation measures. 7. Much more modeling and empirical study of realistic mining sediment plumes is needed. Plume models will help evaluate the spatial and temporal extent of pelagic (as well as benthic) ecosystem effects and help to assess risks from different technologies and mining scenarios. Plume modeling should include realistic mining scenarios (including duration) and assess the spatial-temporal scales over which particle concentrations exceed baseline levels and interfere with light transmission to elucidate potential stresses on communities and ecosystem services. Models should include both near and far field-phases, incorporating realistic near field parameters of plume generation, flocculation, particle sinking, and other processes. It is important to note that some inputs to these models such as physical oceanographic parameters are lacking and should be acquired in the near-term. Plume models need to be complemented by studies to understand effects on biological components by certain particle sizes and concentrations.

There is growing interest in mining polymetallic nodules from the abyssal Clarion-Clipperton Zone (CCZ) in the tropical Pacific Ocean. Despite having been the focus of environmental studies for decades, the benthic megafauna of the CCZ remain poorly known. To predict and manage the environmental impacts of mining in the CCZ, baseline knowledge of the megafauna is essential. The ABYSSLINE Project has conducted benthic biological baseline surveys in the UK Seabed Resources Ltd polymetallic-nodule exploration contract area (UK-1). Prior to ABYSSLINE research cruises in 2013 and 2015, no biological studies had been done in this area of the eastern CCZ. Using a Remotely Operated Vehicle and Autonomous Underwater Vehicle (as well as several other pieces of equipment), the megafauna within the UK Seabed Resources Ltd exploration contract area (UK-1) and at a site ~250 km east of the UK-1 area were surveyed, allowing us to make the first estimates of megafaunal morphospecies richness from the imagery collected. Here, we present an atlas of the abyssal annelid, arthropod, bryozoan, chordate, ctenophore and molluscan megafauna observed and collected during the ABYSSLINE cruises to the UK-1 polymetallic-nodule exploration contract area in the CCZ. There appear to be at least 55 distinct morphospecies (8 Annelida, 12 Arthropoda, 4 Bryozoa, 22 Chordata, 5 Ctenophora, and 4 Mollusca) identified mostly by morphology but also using molecular barcoding for a limited number of animals that were collected. This atlas will aid the synthesis of megafaunal presence/absence data collected by contractors, scientists and other stakeholders undertaking work in the CCZ, ultimately helping to decipher the biogeography of the megafauna in this threatened habitat.