Description: Highlights • Deep-sea mineral exploration and exploitation licenses have been issued recently. • Mining will modify the abiotic and biotic environment. • At directly mined sites, species are removed and cannot resist disturbance. • Recovery is highly variable in distinct ecosystems and among benthic taxa. • Community changes may persist over geological time-scales at directly mined sites. Abstract 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.

Description: Polynoidae contains ~900 species within 18 subfamilies, some of them restricted to the deep sea. Macellicephalinae is the most diverse among these deep-sea subfamilies. In the abyssal Equatorial Pacific Ocean, the biodiversity of benthic communities is at stake in the Clarion-Clipperton Fracture Zone (CCFZ) owing to increased industrial interest in polymetallic nodules. The records of polychaetes in this region are scarce. Data gathered during the JPI Oceans cruise SO239 made a significant contribution to fill this gap, with five different localities sampled between 4000 and 5000 m depth. Benthic samples collected using an epibenthic sledge or a remotely operated vehicle resulted in a large collection of polynoids. The aims of this study are as follows: (1) to describe new species of deep-sea polynoids using morphology and molecular data (COI, 16S and 18S); and (2) to evaluate the monophyly of Macellicephalinae. Based on molecular and morphological phylogenetic analyses, ten subfamilies are synonymized with Macellicephalinae in order to create a homogeneous clade determined by the absence of lateral antennae. Within this clade, the Anantennata clade was well supported, being determined by the absence of a median antenna. Furthermore, 17 new species and four new genera are described, highlighting the high diversity hidden in the deep. A taxonomic key for the 37 valid genera of the subfamily Macellicephalinae is provided.

Description: Commercial-scale mining for polymetallic nodules could have a major impact on the deepsea environment, but the effects of these mining activities on deep-sea ecosystems are very poorly known. The first commercial test mining for polymetallic nodules was carried out in 1970. Since then a number of small-scale commercial test mining or scientific disturbance studies have been carried out. Here we evaluate changes in faunal densities and diversity of benthic communities measured in response to these 11 simulated or test nodule mining disturbances using meta-analysis techniques. We find that impacts are often severe immediately after mining, with major negative changes in density and diversity of most groups occurring. However, in some cases, the mobile fauna and small-sized fauna experienced less negative impacts over the longer term. At seven sites in the Pacific, multiple surveys assessed recovery in fauna over periods of up to 26 years. Almost all studies show some recovery in faunal density and diversity for meiofauna and mobile megafauna, often within one year. However, very few faunal groups return to baseline or control conditions after two decades. The effects of polymetallic nodule mining are likely to be long term. Our analyses show considerable negative biological effects of seafloor nodule mining, even at the small scale of test mining experiments, although there is variation in sensitivity amongst organisms of different sizes and functional groups, which have important implications for ecosystem responses. Unfortunately, many past studies have limitations that reduce their effectiveness in determining responses. We provide recommendations to improve future mining impact test studies. Further research to assess the effects of test-mining activities will inform ways to improve mining practices and guide effective environmental management of mining activities.

Description: Polymetallic nodule mining at abyssal depths in the Clarion Clipperton Fracture Zone (Eastern Central Pacific) will impact one of the most remote and least known environments on Earth. Since vast areas are being targeted by concession holders for future mining, large-scale effects of these activities are expected. Hence, insight into the fauna associated with nodules is crucial to support effective environmental management. In this study video surveys were used to compare the epifauna from sites with contrasting nodule coverage in four license areas. Results showed that epifaunal densities are more than two times higher at dense nodule coverage (〉25 versus ≤10 individuals per 100 m2), and that taxa such as alcyonacean and antipatharian corals are virtually absent from nodule-free areas. Furthermore, surveys conducted along tracks from trawling or experimental mining simulations up to 37 years old, suggest that the removal of epifauna is almost complete and that its full recovery is slow. By highlighting the importance of nodules for the epifaunal biodiversity of this abyssal area, we urge for cautious consideration of the criteria for determining future preservation zones.

Description: In the abyssal equatorial Pacific Ocean, most of the seafloor of the Clarion-Clipperton Fracture Zone (CCFZ), a 6 million km2 polymetallic nodule province, has been preempted for future mining. In light of the large environmental footprint that mining would leave and given the diversity and the vulnerability of the abyssal fauna, the International Seabed Authority has implemented a regional management plan that includes the creation of nine Areas of Particular Environmental Interest (APEIs) located at the periphery of the CCFZ. The scientific principles for the design of the APEIs were based on the best – albeit very limited – knowledge of the area. The fauna and habitats in the APEIs are unknown, as are species' ranges and the extent of biodiversity across the CCFZ. As part of the Joint Programming Initiative Healthy and Productive Seas and Oceans (JPI Oceans) pilot action “Ecological aspects of deep-sea mining”, the SO239 cruise provided data to improve species inventories, determine species ranges, identify the drivers of beta diversity patterns and assess the representativeness of an APEI. Four exploration contract areas and an APEI (APEI no. 3) were sampled along a gradient of sea surface primary productivity that spanned a distance of 1440 km in the eastern CCFZ. Between three and eight quantitative box cores (0.25 m2; 0–10 cm) were sampled in each study area, resulting in a large collection of polychaetes that were morphologically and molecularly (cytochrome c oxidase subunit I and 16S genes) analyzed. A total of 275 polychaete morphospecies were identified. Only one morphospecies was shared among all five study areas and 49 % were singletons. The patterns in community structure and composition were mainly attributed to variations in organic carbon fluxes to the seafloor at the regional scale and nodule density at the local scale, thus supporting the main assumptions underlying the design of the APEIs. However, the APEI no. 3, which is located in an oligotrophic province and separated from the CCFZ by the Clarion Fracture Zone, showed the lowest densities, lowest diversity, and a very low and distant independent similarity in community composition compared to the contract areas, thus questioning the representativeness and the appropriateness of APEI no. 3 to meet its purpose of diversity preservation. Among the four exploration contracts, which belong to a mesotrophic province, the distance decay of similarity provided a species turnover of 0.04 species km−1, an average species range of 25 km and an extrapolated richness of up to 240 000 polychaete species in the CCFZ. By contrast, nonparametric estimators of diversity predict a regional richness of up to 498 species. Both estimates are biased by the high frequency of singletons in the dataset, which likely result from under-sampling and merely reflect our level of uncertainty. The assessment of potential risks and scales of biodiversity loss due to nodule mining thus requires an appropriate inventory of species richness in the CCFZ.

Description: The polymetallic nodules lying on the seafloor of the Clarion-Clipperton Fracture Zone (CCFZ) represent over 30 billion metric tons of manganese. A single mining operation has potential to directly impact approximately 200 km 2 of the seabed per year. Yet, the biodiversity and functioning of the bentho-demersal ecosystem in the CCFZ remain poorly understood. Recent studies indicate a high species diversity in a food-poor environment, although the area remains poorly sampled. Undersampling is aggravated by a combination of low densities of fauna and high habitat heterogeneity at multiple spatial scales. This study examines the Polynoidae, a diverse family of mobile polychaetes. Sampling with an epibenthic sledge and a remotely operated vehicle was performed during the cruise SO239 within the eastern CCFZ. Five areas under the influence of a sea surface productivity gradient were visited. Specimens were identified using morphology and DNA: (i) to provide a more comprehensive account of polynoid diversity within the CCFZ, (ii) to infer factors potentially driving alpha and beta diversity, and (iii) to test the hypothesis that epibenthic polychaetes have low species turnover and large species range. Patterns of species turnover across the eastern CCFZ were correlated with organic carbon fluxes to the seafloor but there was also a differentiation in the composition of assemblages north and south of the Clarion fracture. In contrast to the previous studies, patterns of alpha taxonomic and phylogenetic diversity both suggest that polynoid assemblages are the most diverse at Area of Particular Environmental Interest no. 3, the most oligotrophic study site, located north of the Clarion fracture. Without ruling out the possibility of sampling bias, the main hypothesis explaining such high diversity is the diversification of polynoid subfamily Macellicephalinae, in response to oligotrophy. We propose that macellicephalins evolved under extremely low food supply conditions through adoption of a semi-pelagic mode of life, which enabled them to colonise new niches at the benthic boundary layer and foster their radiation at great depths.

Description: Imaging is increasingly used to capture information on the marine environment thanks to the improvements in imaging equipment, devices for carrying cameras and data storage in recent years. In that context, biologists, geologists, computer specialists and end-users must gather to discuss the methods and procedures for optimising the quality and quantity of data collected from images. The 4 th Marine Imaging Workshop was organised from 3-6 October 2022 in Brest (France) in a hybrid mode. More than a hundred participants were welcomed in person and about 80 people attended the online sessions. The workshop was organised in a single plenary session of presentations followed by discussion sessions. These were based on dynamic polls and open questions that allowed recording of the imaging community’s current and future ideas. In addition, a whole day was dedicated to practical sessions on image analysis, data standardisation and communication tools. The format of this edition allowed the participation of a wider community, including lower-income countries, early career scientists, all working on laboratory, benthic and pelagic imaging. This article summarises the topics addressed during the workshop, particularly the outcomes of the discussion sessions for future reference and to make the workshop results available to the open public.

Description: The increasing demand for metals is pushing forward the progress of deep‐sea mining industry. The abyss between the Clarion and Clipperton Fracture Zones (CCFZ), a region holding a higher concentration of minerals than land deposits, is the most targeted area for the exploration of polymetallic nodules worldwide, which may likely disturb the seafloor across large areas and over many years. Effects from nodule extraction cause acute biodiversity loss of organisms inhabiting sediments and polymetallic nodules. Attention to deep‐sea ecosystems and their services has to be considered before mining starts but the lack of basic scientific knowledge on the methodologies for the ecological surveys of fauna in the context of deep‐sea mining impacts is still scarce. We review the methodology to sample, process and investigate metazoan infauna both inhabiting sediments and nodules dwelling on these polymetallic‐nodule areas. We suggest effective procedures for sampling designs, devices and methods involving gear types, sediment processing, morphological and genetic identification including metabarcoding and proteomic fingerprinting, the assessment of biomass, functional traits, fatty acids, and stable isotope studies within the CCFZ based on both first‐hand experiences and literature. We recommend multi‐ and boxcorers for the quantitative assessments of meio‐ and macrofauna, respectively. The assessment of biodiversity at species level should be focused and/or the combination of morphological with metabarcoding or proteomic fingerprinting techniques. We highlight that biomass, functional traits, and trophic markers may provide critical insights for biodiversity assessments and how statistical modeling facilitates predicting patterns spatially across point‐source data and is essential for conservation management.

Description: 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.

Description: © The Authors, 2010. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Biogeosciences 7 (2010): 2851-2899, doi:10.5194/bg-7-2851-2010.

Description: The deep sea, the largest biome on Earth, has a series of characteristics that make this environment both distinct from other marine and land ecosystems and unique for the entire planet. This review describes these patterns and processes, from geological settings to biological processes, biodiversity and biogeographical patterns. It concludes with a brief discussion of current threats from anthropogenic activities to deep-sea habitats and their fauna. Investigations of deep-sea habitats and their fauna began in the late 19th century. In the intervening years, technological developments and stimulating discoveries have promoted deep-sea research and changed our way of understanding life on the planet. Nevertheless, the deep sea is still mostly unknown and current discovery rates of both habitats and species remain high. The geological, physical and geochemical settings of the deep-sea floor and the water column form a series of different habitats with unique characteristics that support specific faunal communities. Since 1840, 28 new habitats/ecosystems have been discovered from the shelf break to the deep trenches and discoveries of new habitats are still happening in the early 21st century. However, for most of these habitats the global area covered is unknown or has been only very roughly estimated; an even smaller – indeed, minimal – proportion has actually been sampled and investigated. We currently perceive most of the deep-sea ecosystems as heterotrophic, depending ultimately on the flux on organic matter produced in the overlying surface ocean through photosynthesis. The resulting strong food limitation thus shapes deep-sea biota and communities, with exceptions only in reducing ecosystems such as inter alia hydrothermal vents or cold seeps. Here, chemoautolithotrophic bacteria play the role of primary producers fuelled by chemical energy sources rather than sunlight. Other ecosystems, such as seamounts, canyons or cold-water corals have an increased productivity through specific physical processes, such as topographic modification of currents and enhanced transport of particles and detrital matter. Because of its unique abiotic attributes, the deep sea hosts a specialized fauna. Although there are no phyla unique to deep waters, at lower taxonomic levels the composition of the fauna is distinct from that found in the upper ocean. Amongst other characteristic patterns, deep-sea species may exhibit either gigantism or dwarfism, related to the decrease in food availability with depth. Food limitation on the seafloor and water column is also reflected in the trophic structure of heterotrophic deep-sea communities, which are adapted to low energy availability. In most of these heterotrophic habitats, the dominant megafauna is composed of detritivores, while filter feeders are abundant in habitats with hard substrata (e.g. mid-ocean ridges, seamounts, canyon walls and coral reefs). Chemoautotrophy through symbiotic relationships is dominant in reducing habitats. Deep-sea biodiversity is among of the highest on the planet, mainly composed of macro and meiofauna, with high evenness. This is true for most of the continental margins and abyssal plains with hot spots of diversity such as seamounts or cold-water corals. However, in some ecosystems with particularly "extreme" physicochemical processes (e.g. hydrothermal vents), biodiversity is low but abundance and biomass are high and the communities are dominated by a few species. Two large-scale diversity patterns have been discussed for deep-sea benthic communities. First, a unimodal relationship between diversity and depth is observed, with a peak at intermediate depths (2000–3000 m), although this is not universal and particular abiotic processes can modify the trend. Secondly, a poleward trend of decreasing diversity has been discussed, but this remains controversial and studies with larger and more robust data sets are needed. Because of the paucity in our knowledge of habitat coverage and species composition, biogeographic studies are mostly based on regional data or on specific taxonomic groups. Recently, global biogeographic provinces for the pelagic and benthic deep ocean have been described, using environmental and, where data were available, taxonomic information. This classification described 30 pelagic provinces and 38 benthic provinces divided into 4 depth ranges, as well as 10 hydrothermal vent provinces. One of the major issues faced by deep-sea biodiversity and biogeographical studies is related to the high number of species new to science that are collected regularly, together with the slow description rates for these new species. Taxonomic coordination at the global scale is particularly difficult, but is essential if we are to analyse large diversity and biogeographic trends. Because of their remoteness, anthropogenic impacts on deep-sea ecosystems have not been addressed very thoroughly until recently. The depletion of biological and mineral resources on land and in shallow waters, coupled with technological developments, are promoting the increased interest in services provided by deep-water resources. Although often largely unknown, evidence for the effects of human activities in deep-water ecosystems – such as deep-sea mining, hydrocarbon exploration and exploitation, fishing, dumping and littering – is already accumulating. Because of our limited knowledge of deep-sea biodiversity and ecosystem functioning and because of the specific life-history adaptations of many deep-sea species (e.g. slow growth and delayed maturity), it is essential that the scientific community works closely with industry, conservation organisations and policy makers to develop robust and efficient conservation and management options.

Description: This paper has been written under the umbrella of the Census of Marine Life synthesis initiative SYNDEEP, supported by the Alfred P. Sloan Foundation, Fondation Total and EuroCoML, which are gratefully acknowledged. ERLL is funded by the CoML-ChEss programme (A. P. Sloan Foundation) and Fondation Total. CRG acknowledges support from the CoMLChEss programme. LAL acknowledges support from the National Science Foundation and the CoML-COMARGE and ChEss programmes. DPT acknowledges funding from the CoML-FMAP programme. MV acknowledges the CoML-MAR-ECO programme (Sloan Foundation and NOAA).