Ecology of a polymetallic nodule occurrence gradient: Implications for deep-sea mining.

Abyssal polymetallic nodule fields constitute an unusual deep-sea habitat. The mix of soft sediment and the hard substratum provided by nodules increases the complexity of these environments. Hard substrata typically support a very distinct fauna to that of seabed sediments, and its presence can play a major role in the structuring of benthic assemblages. We assessed the influence of seafloor nodule cover on the megabenthos of a marine conservation area (area of particular environmental interest 6) in the Clarion Clipperton Zone (3950-4250 m water depth) using extensive photographic surveys from an autonomous underwater vehicle. Variations in nodule cover (1-20%) appeared to exert statistically significant differences in faunal standing stocks, some biological diversity attributes, faunal composition, functional group composition, and the distribution of individual species. The standing stock of both the metazoan fauna and the giant protists (xenophyophores) doubled with a very modest initial increase in nodule cover (from 1% to 3%). Perhaps contrary to expectation, we detected little if any substantive variation in biological diversity along the nodule cover gradient. Faunal composition varied continuously along the nodule cover gradient. We discuss these results in the context of potential seabed-mining operations and the associated sustainable management and conservation plans. We note in particular that successful conservation actions will likely require the preservation of areas comprising the full range of nodule cover and not just the low cover areas that are least attractive to mining.

Megafaunal variation in the abyssal landscape of the Clarion Clipperton Zone.

The potential for imminent polymetallic nodule mining in the Clarion Clipperton Fracture Zone (CCZ) has attracted considerable scientific and public attention. This concern stems from both the extremely large seafloor areas that may be impacted by mining, and the very limited knowledge of the fauna and ecology of this region. The environmental factors regulating seafloor ecology are still very poorly understood. In this study, we focus on megafaunal ecology in the proposed conservation zone 'Area of Particular Environmental Interest 6' (study area centred 17°16'N, 122°55'W). We employ bathymetric data to objectively define three landscape types in the area (a level bottom Flat, an elevated Ridge, a depressed Trough; water depth 3950-4250 m) that are characteristic of the wider CCZ. We use direct seabed sampling to characterise the sedimentary environment in each landscape, detecting no statistically significant differences in particle size distributions or organic matter content. Additional seafloor characteristics and data on both the metazoan and xenophyophore components of the megafauna were derived by extensive photographic survey from an autonomous underwater vehicle. Image data revealed that there were statistically significant differences in seafloor cover by nodules and in the occurrence of other hard substrata habitat between landscapes. Statistically significant differences in megafauna standing stock, functional structuring, diversity, and faunal composition were detected between landscapes. The Flat and Ridge areas exhibited a significantly higher standing stock and a distinct assemblage composition compared to the Trough. Geomorphological variations, presumably regulating local bottom water flows and the occurrence of nodule and xenophyophore test substrata, between study areas may be the mechanism driving these assemblage differences. We also used these data to assess the influence of sampling unit size on the estimation of ecological parameters. We discuss these results in the contexts of regional benthic ecology and the appropriate management of potential mining activities in the CCZ and elsewhere in the deep ocean.

Seamount mining test provides evidence of ecological impacts beyond deposition.

In July 2020, Japan undertook the first deep-sea mining test of cobalt-rich crusts on the summit of Takuyo-Daigo Seamount within their exclusive economic zone (EEZ). Seabed mining regulations are currently being developed by the International Seabed Authority (ISA)1; however, a lack of experimental data has so far constrained our understanding of the associated impacts, particularly from the release of sediment plumes.2 The area of sediment re-deposition from the crust mining test was determined using modeled data and in situ observations. To investigate biological impacts, variations in seabed megafauna (animals > 1 cm) were quantified from seabed imagery collected around the excavation site before, 1 month, and 13 months after the test in areas both inside and outside (adjacent) expected deposition. Observable responses varied across community components: densities of sessile animals were similar between deposition and adjacent areas throughout the study; mobile epifauna were less abundant only in the deposition area following disturbance; and highly mobile swimmers showed reduced densities after the test in both deposition and adjacent areas following disturbance. These results stress that monitoring of highly mobile taxa may be essential to fully assess disturbance extent and magnitude. Fish may avoid areas even outside plume deposition, possibly owing to the creation of suboptimal feeding patches resulting from deposition. Our findings suggest sufficiently large (>300 × 300 m), distant, and representative control areas are essential to optimally map deep-sea mining impacts in ferromanganese crust habitats to ensure impact assessments encompass the full range of functional components in the megabenthic community (including mobile fishes) that typically inhabit seamounts.

How many metazoan species live in the world's largest mineral exploration region?

The global surge in demand for metals such as cobalt and nickel has created unprecedented interest in deep-sea habitats with mineral resources. The largest area of activity is a 6 million km2 region known as the Clarion-Clipperton Zone (CCZ) in the central and eastern Pacific, regulated by the International Seabed Authority (ISA). Baseline biodiversity knowledge of the region is crucial to effective management of environmental impact from potential deep-sea mining activities, but until recently this has been almost completely lacking. The rapid growth in taxonomic outputs and data availability for the region over the last decade has allowed us to conduct the first comprehensive synthesis of CCZ benthic metazoan biodiversity for all faunal size classes. Here we present the CCZ Checklist, a biodiversity inventory of benthic metazoa vital to future assessments of environmental impacts. An estimated 92% of species identified from the CCZ are new to science (436 named species from a total of 5,578 recorded). This is likely to be an overestimate owing to synonyms in the data but is supported by analysis of recent taxonomic studies suggesting that 88% of species sampled in the region are undescribed. Species richness estimators place total CCZ metazoan benthic diversity at 6,233 (+/-82 SE) species for Chao1, and 7,620 (+/-132 SE) species for Chao2, most likely representing lower bounds of diversity in the region. Although uncertainty in estimates is high, regional syntheses become increasingly possible as comparable datasets accumulate. These will be vital to understanding ecological processes and risks of biodiversity loss.

Carbonate compensation depth drives abyssal biogeography in the northeast Pacific.

Abyssal seafloor communities cover more than 60% of Earth's surface. Despite their great size, abyssal plains extend across modest environmental gradients compared to other marine ecosystems. However, little is known about the patterns and processes regulating biodiversity or potentially delimiting biogeographical boundaries at regional scales in the abyss. Improved macroecological understanding of remote abyssal environments is urgent as threats of widespread anthropogenic disturbance grow in the deep ocean. Here, we use a new, basin-scale dataset to show the existence of clear regional zonation in abyssal communities across the 5,000 km span of the Clarion-Clipperton Zone (northeast Pacific), an area targeted for deep-sea mining. We found two pronounced biogeographic provinces, deep and shallow-abyssal, separated by a transition zone between 4,300 and 4,800 m depth. Surprisingly, species richness was maintained across this boundary by phylum-level taxonomic replacements. These regional transitions are probably related to calcium carbonate saturation boundaries as taxa dependent on calcium carbonate structures, such as shelled molluscs, appear restricted to the shallower province. Our results suggest geochemical and climatic forcing on distributions of abyssal populations over large spatial scales and provide a potential paradigm for deep-sea macroecology, opening a new basis for regional-scale biodiversity research and conservation strategies in Earth's largest biome.

Investigating the benthic megafauna in the eastern Clarion Clipperton Fracture Zone (north-east Pacific) based on distribution models predicted with random forest.

The eastern Clarion Clipperton Fracture Zone (CCZ) is a heterogeneous abyssal environment harbouring relatively low abundances of highly diverse megafauna communities. Potential future mining of polymetallic nodules threatens these benthic communities and calls for detailed spatial investigation of megafauna. Based on the predicted probability of occurrence of 68 megafauna morphotypes, a seabed area extending over 62,000 km2 was divided into three assemblages covering an eastern plain area, a deeper western plain area and an area covering both seamount and abyssal hill sites. Richness, estimated as the sum of morphotypes with a predicted probability of occurrence larger than 0.5, amounts to 15.4 of 68 morphotypes. Highest richness was predicted at seamount sites, and lowest richness in the western part of the study area. Combining the predicted probability of megafauna occurrences with bathymetric variables, two seamount habitats and two plain habitats could be defined. One of these megafauna plain habitats corresponds with contiguous nodule fields of high abundance that may be targeted for future mining, showing that prospective nodule fields have a clearly differentiated megafauna assemblage. Monitoring and management schemes, including the delineation of preservation and protection areas within contract areas, need to incorporate this geological and biological heterogeneity.

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.

Benthic megafauna of the western Clarion-Clipperton Zone, Pacific Ocean.

There is a growing interest in the exploitation of deep-sea mineral deposits, particularly on the abyssal seafloor of the central Pacific Clarion-Clipperton Zone (CCZ), which is rich in polymetallic nodules. In order to effectively manage potential exploitation activities, a thorough understanding of the biodiversity, community structure, species ranges, connectivity, and ecosystem functions across a range of scales is needed. The benthic megafauna plays an important role in the functioning of deep-sea ecosystems and represents an important component of the biodiversity. While megafaunal surveys using video and still images have provided insight into CCZ biodiversity, the collection of faunal samples is needed to confirm species identifications to accurately estimate species richness and species ranges, but faunal collections are very rarely carried out. Using a Remotely Operated Vehicle, 55 specimens of benthic megafauna were collected from seamounts and abyssal plains in three Areas of Particular Environmental Interest (APEI 1, APEI 4, and APEI 7) at 3100-5100 m depth in the western CCZ. Using both morphological and molecular evidence, 48 different morphotypes belonging to five phyla were found, only nine referrable to known species, and 39 species potentially new to science. This work highlights the need for detailed taxonomic studies incorporating genetic data, not only within the CCZ, but in other bathyal, abyssal, and hadal regions, as representative genetic reference libraries that could facilitate the generation of species inventories.

On the impact of Citizen Science-derived data quality on deep learning based classification in marine images.

The evaluation of large amounts of digital image data is of growing importance for biology, including for the exploration and monitoring of marine habitats. However, only a tiny percentage of the image data collected is evaluated by marine biologists who manually interpret and annotate the image contents, which can be slow and laborious. In order to overcome the bottleneck in image annotation, two strategies are increasingly proposed: "citizen science" and "machine learning". In this study, we investigated how the combination of citizen science, to detect objects, and machine learning, to classify megafauna, could be used to automate annotation of underwater images. For this purpose, multiple large data sets of citizen science annotations with different degrees of common errors and inaccuracies observed in citizen science data were simulated by modifying "gold standard" annotations done by an experienced marine biologist. The parameters of the simulation were determined on the basis of two citizen science experiments. It allowed us to analyze the relationship between the outcome of a citizen science study and the quality of the classifications of a deep learning megafauna classifier. The results show great potential for combining citizen science with machine learning, provided that the participants are informed precisely about the annotation protocol. Inaccuracies in the position of the annotation had the most substantial influence on the classification accuracy, whereas the size of the marking and false positive detections had a smaller influence.

Biological effects 26 years after simulated deep-sea mining.

The potential for imminent abyssal polymetallic nodule exploitation has raised considerable scientific attention. The interface between the targeted nodule resource and sediment in this unusual mosaic habitat promotes the development of some of the most biologically diverse communities in the abyss. However, the ecology of these remote ecosystems is still poorly understood, so it is unclear to what extent and timescale these ecosystems will be affected by, and could recover from, mining disturbance. Using data inferred from seafloor photo-mosaics, we show that the effects of simulated mining impacts, induced during the "DISturbance and reCOLonization experiment" (DISCOL) conducted in 1989, were still evident in the megabenthos of the Peru Basin after 26 years. Suspension-feeder presence remained significantly reduced in disturbed areas, while deposit-feeders showed no diminished presence in disturbed areas, for the first time since the experiment began. Nevertheless, we found significantly lower heterogeneity diversity in disturbed areas and markedly distinct faunal compositions along different disturbance levels. If the results of this experiment at DISCOL can be extrapolated to the Clarion-Clipperton Zone, the impacts of polymetallic nodule mining there may be greater than expected, and could potentially lead to an irreversible loss of some ecosystem functions, especially in directly disturbed areas.

Abundance and morphology of Paleodictyon nodosum, observed at the Clarion-Clipperton Zone.

Paleodictyon is an important trace fossil characterised by a regular hexagonal structure and typical of ancient deep-ocean habitats as far back as the Ordovician. It is represented in modern deep-sea settings by Paleodictyon nodosum, known from the Mid-Atlantic Ridge, the South Atlantic, and off eastern Australia. Here we report the occurrence of P. nodosum in the Clarion Clipperton Zone (CCZ), abyssal equatorial Pacific, an area characterised by polymetallic nodule fields. At the study site within the International Seabed Authority northeastern Area of Particular Environmental Interest (APEI-6), P. nodosum appeared as a compact, regular pattern of small circular openings on the seafloor, each pattern interpreted as reflecting the activity of an individual organism. The patterns had a mean size (maximum dimension) of 45 mm ± 16 mm SD (n = 841) and occurred at a density of 0.33 individuals m-2. Most (82%) were interrupted by nodules, but those that were not displayed both regular (59%) and irregular (41%) forms, the former having equal numbers of rows along the three axes (6 x 6 x 6 and 8 x 8 x 8). In both size and morphology, our Paleodictyon traces were more similar to the Australian examples than to those from the Mid-Atlantic Ridge.

Biological responses to disturbance from simulated deep-sea polymetallic nodule mining.

Commercial-scale mining for polymetallic nodules could have a major impact on the deep-sea 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.