Area Of Particular Environmental Interest Research Articles

Taxonomy, phylogeny, and biodiversity of Lumbrineridae (Annelida, Polychaeta) from the Central Pacific Clarion-Clipperton Zone.

The DNA taxonomy of six species of the annelid family Lumbrineridae collected from the Clarion-Clipperton Zone (CCZ) in the Central Pacific, an area of potential mining interest for polymetallic nodules, is presented. Lumbrinerids are an ecologically important and understudied annelid family within the deep sea, with many species still undescribed. This study aims to document the taxonomy and biodiversity of the CCZ using specimens collected from the UK-1, OMS, and NORI-D exploration contract areas and Areas of Particular Environmental Interest. Species were identified through a combination of morphological and molecular phylogenetic analysis. We present informal species descriptions associated with voucher specimens, accessible through the Natural History Museum (London) collections, to improve future taxonomic and biodiversity studies of this region. Five taxa in this study had no morphological or genetic matches within the literature and therefore are possibly new to science, but their suboptimal morphological preservation prevented the formalisation of new species. The most abundant taxon Lumbrineridescf.laubieri (NHM_0020) was compared with the holotype of Lumbrinerideslaubieri Miura, 1980 from the deep Northeast Atlantic. Currently no reliable morphological characters separating the Pacific and Atlantic specimens have been found and molecular data from the Atlantic specimens was not available.

Abyssal seafloor response to fresh phytodetrital input in three areas of particular environmental interest (APEIs) in the western clarion-clipperton zone (CCZ)

The abyssal seafloor (3500–6000m) remains largely unexplored but with deep-sea mining imminent, anthropogenic impacts may soon reach abyssal communities. Thus, there is a growing need for baseline studies of biodiversity, ecosystem functioning, and connectivity in both potential mining and no-mining areas across the Clarion-Clipperton Zone (CCZ), a key target region for polymetallic nodule mining. In this study, in situ pulse-chase lander experiments were conducted for 1.5 days in three no-mining areas (called Areas of Particular Environmental Interest, or APEIs) in the western CCZ, a region with a seafloor particulate organic carbon (POC) flux gradient. A decreasing trend was seen in mean seafloor respiration, macrofaunal abundance, and biomass from the more eutrophic APEI 7 to the more oligotrophic APEI 1, although this trend was not statistically significant (p = 0.18) most likely due to small samples sizes and high variability. In this study, most (96%) of the 13C-labeled processed phytodetritus was respired within 1.5 days. Experimental uptake of phytodetritus by macrofauna and bacteria was detected but was lower than in the previously studied and more eutrophic eastern CCZ over similar time scales (1.5 d). Bacteria dominated the short-term (∼1.5 d) uptake of organic carbon at the seafloor, yet macrofauna processed more organic carbon per unit biomass than previously found in the eastern CCZ (0.003 mg C m−2 d−1 and 0.5 × 10−5 mg C m−2 d−1 for the western and eastern CCZ, respectively). Our study provides important information on C-uptake and respiration rates in areas set aside from mining in the western CCZ and suggests high variability may occur in the rates of benthic Corg-cycling across the CCZ. We recommend that benthic ecosystem functions be explored across gradients of POC flux which may be a major environmental factor driving ecosystem dynamics in the CCZ.

Abyssal seafloor response to fresh phytodetrital input in three Areas of Particular Environmental Interest (APEIs) in the western Clarion-Clipperton Zone (CCZ)

Abyssal seafloor response to fresh phytodetrital input in three Areas of Particular Environmental Interest (APEIs) in the western Clarion-Clipperton Zone (CCZ)

Abyssal fauna of polymetallic nodule exploration areas, eastern Clarion-Clipperton Zone, central Pacific Ocean: Amphinomidae and Euphrosinidae (Annelida, Amphinomida).

This is a contribution in a series of taxonomic publications on benthic fauna of polymetallic nodule fields in the eastern abyssal Clarion-Clipperton Zone (CCZ). The material was collected during environmental surveys targeting exploration contract areas 'UK-1', 'OMS' and 'NORI-D', as well as an Area of Particular Environmental Interest, 'APEI-6'. The annelid families Amphinomidae and Euphrosinidae are investigated here. Taxonomic data are presented for six species from 41 CCZ-collected specimens as identified by a combination of morphological and genetic approaches; of the six species, three are here described as new, one species is likely to be new but in too poor condition to be formalised and the two others likely belong to known species. Description of three new species Euphrosinellageorgievae sp. nov., Euphrosinopsisahearni sp. nov., and Euphrosinopsishalli sp. nov. increases the number of formally described new annelid species from the targeted areas to 21 and CCZ-wide to 52. Molecular data suggest that four of the species reported here are known from CCZ only, but within CCZ they have a wide distribution. In contrast, the species identified as Bathychloeiacf.sibogae Horst, 1910 was found to have a wide distribution within the Pacific based on both morphological and molecular data, using comparative material from the abyssal South Pacific. Bathychloeiacf.balloniformis Böggemann, 2009 was found to be restricted to APEI-6 based on DNA data available from CCZ specimens only, but morphological data from other locations suggest potentially a wide abyssal distribution. The genus Euphrosinopsis was previously known only from Antarctic waters, and Euphrosinellageorgievae sp. nov. was recovered as a sister taxon to the Antarctic specimens of Euphrosinellacf.cirratoformis in our molecular phylogenetic analysis, strengthening the hypothesised link between the deep-sea and Antarctic benthic fauna.

Using habitat classification and large-scale spatial planning model to identify environmental areas: A case study in the area of the Northwest Pacific Ocean

The deep sea refers to the parts of the oceans with a depth of more than 200 m, and it comprises the largest habitat on earth with the most abundant and diverse marine life. Human activities, such as deep-sea mining, fishing, and shipping, can have severe impacts on deep-sea species and habitats. With the development of the related equipment and technology, interest in deep-sea mining continues to rise; therefore, protecting the unique and fragile deep-sea marine ecosystem has become a global focus. Given the fragility of the deep-sea habitats and the growing demand for deep-sea resources, the United Nations established the International Seabed Authority (the ISA) in 1972, which represents everyone worldwide, to manage the mineral resources in the international seabed area (also known as “the Area”) while fulfilling environmental protection obligations. The most important objective of the regional environmental management plan (REMP) in the Area initiated by the ISA is to balance deep-sea mining and environment protection, and the ISA has considered numerous types of area-based management tools (ABMTs) in the development of the REMPs (Table 1). Using the different ABMTs, several key environmental areas or sites have been identified in various regions, such as the areas of particular environmental interest (APEIs), sites in need of protection (SINPs), or areas in need of protection (AINPs). Additionally, deep-sea mining is prohibited in these sites/areas to avoid its irrecoverable impact on the environment in the Area. The designation these environmental areas or sites can improve the function and resilience of the ecosystem and avoid the unrecoverable impact of deep-sea mining. In this study, we selected the international areas of the Northwest Pacific Ocean for a case study. In order to protect representative species and habitats, such as cold-water corals and seamounts, we utilized environmental factors to classify the study area into eight types of habitats. We then used the appropriate criteria to identify candidate AINPs in two scenarios with a large-scale spatial planning model and analyzed the differences in the protected proportion of the eight types of habitats. The results indicate that identifying the candidate AINPs can reach the protection target of approximately 30% of the area, and each representative species and habitat can be protected. The results obtained in this study can provide technical references for the implementation of REMPs and the designation of AINPs in the study area.

Abyssal fauna of polymetallic nodule exploration areas, eastern Clarion-Clipperton Zone, central Pacific Ocean: Annelida: Spionidae and Poecilochaetidae

This paper represents a continuation of taxonomic publications on the benthic fauna of polymetallic nodule fields in the eastern Clarion-Clipperton Zone (CCZ) using material collected during baseline environmental survey work targeting two exploration contract areas (“UK-1” and “OMS”) and one Area of Particular Environmental Interest, “APEI-6.” Families Poecilochaetidae Hannerz, 1956 and Spionidae Grube, 1850 of the annelid suborder Spioniformia were studied here. Taxonomic data are presented for 25 species from 98 records as identified by a combination of morphological and genetic approaches. Although sub-optimal morphological condition can prevent new species being formally described, it is essential that morphological, molecular, and voucher data are made available for future surveys. Descriptions of two new species—Poecilochaetus brenkei sp. nov. and Laonice shulseae sp. nov.—increase the number of formally described new annelid species from the areas targeted in this study to 15 and CCZ-wide to 46. We also discuss the commonly reported “cosmopolitan” deep-sea spionid Aurospio dibranchiata Maciolek, 1981, which we show represents several genetically distinct species (three of these from CCZ area alone) but without reliable morphological characters to separate them. Molecular data provide evidence that 15 out of 25 species reported here have a wide distribution within the eastern CCZ and that Aurospio sp. “NHM_2186” and the known species Prionospio amarsupiata Neal & Altamira in Paterson et al. 2016 may be cosmopolitan. Lastly, the molecular data provide insights into relationships within Spioniformia, suggesting that both Poecilochaetidae and Trochochaetidae belong within Spionidae.

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.

A Contrast of Criteria for Special Places Important for Biodiversity Outcomes

This paper contrasts seven spatial biodiversity conservation area designations by six different bodies: Other Effective Area-Based Conservation Measures (OECMs), and the Ecologically and Biologically Significant Areas (EBSAs) of the Convention on Biological Diversity (CBD); the Vulnerable Marine Ecosystems (VMEs) of the Food And Agriculture Organization (FAO); the Key Biodiversity Areas (KBAs) under criteria developed by the IUCN; the Areas of Particular Environmental Interest (APEIs) of the International Seabed Authority (ISA); the Particularly Sensitive Sea Areas (PSSAs) of the International Maritime Organization (IMO); and the Locally Managed Marine Areas (LMMAs) used by small island States in the Pacific Ocean; on five themes: biological and ecological features, functions served by areas receiving these labels, governance, threats and pressures, and other considerations. The seven different labels for such areas were generally similar in the biologically and ecological criteria to be met, and the functions typically served by these areas. Differences among the labels increased when considering governance, threat and pressures, and other considerations. Implications of these similarities and differences for policy development and outcomes are discussed. Performance reviews of the various labels under these themes could provide insight into both the effectiveness of the provisions in the Agreements and Decisions and how evidence is acquired and used to inform their application, allowing improvements to each approach to learn from experiences with other labels.

Regional Variation in Communities of Demersal Fishes and Scavengers Across the CCZ and Pacific Ocean

We synthesize and analyze data from visual transecting approaches and baited camera studies to evaluate fish and invertebrate scavenger communities across the Clarion-Clipperton Zone (CCZ), an area of intense deep-sea mining interest, and neighboring areas of the abyssal Pacific. In abyssal regions including the CCZ most of the top predators are large mobile fishes and crustaceans, and the majority of these are also opportunistic scavengers. Top predators can exert important ecosystem influences and they can be susceptible to sustained anthropogenic disturbances, necessitating their study in the CCZ mining region. In total 157 baited camera deployments from 3 mining exploration license areas, 4 APEIs (Areas of Particular Environmental Interest – one type of no mining zone) and 4 other areas in the Pacific (Hawaii, California, New Zealand and Guam) and 122 visual transects from 7 exploration license areas, 4 no mining zones, and the Peru Basin (DISCOL area) were examined. Many taxa were observed in both sampling techniques but visual transects viewed few fishes overall. Fish and scavenger communities and diversity varied across the CCZ, significantly for baited camera data with a parallel but insignificant pattern for visual transects suggesting that even for these highly mobile species, not all regions of the CCZ are equivalent and the CCZ cannot be managed as one homogenous region. Further CCZ communities were different than communities elsewhere in the abyssal Pacific. The regional variations in community composition are largely the result of varying abundances of species rather than species presence/absence given that most, but not all, of the fishes and scavengers observed have very large ranges. On a more local scale, seamounts had a significantly different scavenger community than neighboring abyssal plains and thus contribute to regional diversity. Visual transect data revealed a similar but insignificant pattern due to low sample sizes. Given the coarse spatial resolution of sampling of fish and scavenger communities in the CCZ, it is not possible to evaluate if no mining zones (APEIs) adequately represent these communities nor where, or if, any biogeographic boundaries exist in the CCZ region. It is possible to conclude that a network of APEIs that covers the spectrum of available habitats at regional and more local scales will be key to conserving fish and scavenger biodiversity.

Diversity of Deep-Sea Scale-Worms (Annelida, Polynoidae) in the Clarion-Clipperton Fracture Zone

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 km2 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.

Biogeography and Connectivity Across Habitat Types and Geographical Scales in Pacific Abyssal Scavenging Amphipods

Recently, there has been a resurgent interest in the exploration of deep-sea mineral deposits, particularly polymetallic nodules in the Clarion-Clipperton Zone (CCZ), central Pacific. Accurate environmental impact assessment is critical to the effective management of a new industry and depends on a sound understanding of species taxonomy, biogeography, and connectivity across a range of scales. Connectivity is a particularly important parameter in determining ecosystem resilience, as it helps to define the ability of a system to recover post-impact. Scavenging amphipods in the superfamilies Alicelloidea Lowry and De Broyer, 2008 and Lysianassoidea Dana, 1849 contribute to a unique and abundant scavenging community in abyssal ecosystems. They are relatively easy to sample and in recent years have become the target of several molecular and taxonomic studies, but are poorly studied in the CCZ. Here, a molecular approach is used to identify and delimit species, and to investigate evolutionary relationships of scavenging amphipods from both abyssal plain and deep (>3000 m) seamount habitats in three APEIs (Areas of Particular Environmental Interest, i.e., designated conservation areas) in the western CCZ. A total of 17 different morphospecies of scavenging amphipods were identified, which include at least 30 genetic species delimited by a fragment of the cytochrome c oxidase subunit I (COI) barcode gene. The scavenging communities sampled in the western CCZ included the most common species (Abyssorchomene gerulicorbis (Shulenberger and Barnard, 1976), A. chevreuxi (Stebbing, 1906), Paralicella caperesca Shulenberger and Barnard, 1976, and P. tenuipes Chevreux, 1908) reported for other ocean basins. Only four morphospecies, representing five genetic species, were shared between APEIs 1, 4, and 7. The two abyssal plain sites at APEIs 4 and 7 were dominated by two and three of the most common scavenging species, respectively, while the APEI 1 seamount site was dominated by two species potentially new to science that appeared to be endemic to the site. The presence of common species in all sites and high genetic diversity, yet little geographic structuring, indicate connectivity over evolutionary time scales between the areas, which span about 1500 km. Similar to recent studies, the differences in amphipod assemblages found between the seamount and abyssal sites suggest that ecological conditions on seamounts generate distinct community compositions.

Environment, ecology, and potential effectiveness of an area protected from deep-sea mining (Clarion Clipperton Zone, abyssal Pacific)

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.

Megafaunal Ecology of the Western Clarion Clipperton Zone

The Clarion Clipperton Zone (CCZ) is a vast area of the central Pacific Ocean where the abyssal seabed is a focus for future polymetallic nodule mining. Broad-scale environmental gradients occur east-to-west across the CCZ seabed, including organic matter supply and nodule abundance, factors that influence benthic faunal community structure and function. A network of protected areas across the CCZ, called Areas of Particular Environmental Interest (APEIs), has been designated to cover this variation. Most previous studies of the benthic environment and megafaunal communities have focussed on the eastern CCZ, leaving the impact of these large-scale gradients unexamined and the network design untested. Seamounts are a further source of heterogeneity in the region. We examined the benthic megafaunal ecology of three APEIs in the western CCZ, spanning a range of environmental conditions. We used a combination of seabed photography and direct sampling to assess the environment and megafauna on the soft sediment habitats on the abyssal plain in three APEIs, and seamounts in two of those APEIs. We found that environmental conditions on abyssal plains differed between the three APEIs in terms of water depth, nodule abundance and coverage, sediment particle size distribution, and estimated organic matter flux. Megafauna were low density and high diversity, with few common morphotypes between sites and many morphotypes being observed only once. Xenophyophores dominated the assemblages. The density and diversity of invertebrates were greater at the sites with lower organic matter inputs and greater nodule abundance. Seamounts in the same APEIs were nodule-free and had coarser sediments than on the plain. Invertebrate megafaunal diversity was lower on the seamounts than on the plains, and most morphotypes recorded on the seamounts were only found on seamounts. Low morphotype overlap also suggests little connectivity between APEIs, and between seamounts and adjacent abyssal plains. Our results provide the first evaluation of the seabed habitats and megafaunal ecology in the western CCZ, highlighting environmental gradients that influence benthic communities, and are important for evaluating the design of the network of protected areas.

Bacterial and Archaeal Communities in Polymetallic Nodules, Sediments, and Bottom Waters of the Abyssal Clarion-Clipperton Zone: Emerging Patterns and Future Monitoring Considerations

Bacteria and archaea are key contributors to deep-sea biogeochemical cycles and food webs. The disruptions these microbial communities may experience during and following polymetallic nodule mining in the Clarion-Clipperton Zone (CCZ) of the North Pacific Ocean could therefore have broad ecological effects. Our goals in this synthesis are to characterize the current understanding of biodiversity and biogeography of bacteria and archaea in the CCZ and to identify gaps in the baseline data and sampling approaches, prior to the onset of mining in the region. This is part of a large effort to compile biogeographic patterns in the CCZ, and to assess the representivity of no-mining Areas of Particular Environmental Interest, across a range of taxa. Here, we review published studies and an additional new dataset focused on 16S ribosomal RNA (rRNA) gene amplicon characterization of abyssal bacterial and archaeal communities, particularly focused on spatial patterns. Deep-sea habitats (nodules, sediments, and bottom seawater) each hosted significantly different microbial communities. An east-vs.-west CCZ regional distinction was present in nodule communities, although the magnitude was small and likely not detectable without a high-resolution analysis. Within habitats, spatial variability was driven by differences in relative abundances of taxa, rather than by abundant taxon turnover. Our results further support observations that nodules in the CCZ have distinct archaeal communities from those in more productive surrounding regions, with higher relative abundances of presumed chemolithoautotrophic Nitrosopumilaceae suggesting possible trophic effects of nodule removal. Collectively, these results indicate that bacteria and archaea in the CCZ display previously undetected, subtle, regional-scale biogeography. However, the currently available microbial community surveys are spatially limited and suffer from sampling and analytical differences that frequently confound inter-comparison; making definitive management decisions from such a limited dataset could be problematic. We suggest a number of future research priorities and sampling recommendations that may help to alleviate dataset incompatibilities and to address challenges posed by rapidly advancing DNA sequencing technology for monitoring bacterial and archaeal biodiversity in the CCZ. Most critically, we advocate for selection of a standardized 16S rRNA gene amplification approach for use in the anticipated large-scale, contractor-driven biodiversity monitoring in the region.

Patterns of Macrofaunal Biodiversity Across the Clarion-Clipperton Zone: An Area Targeted for Seabed Mining

Macrofauna are an abundant and diverse component of abyssal benthic communities and are likely to be heavily impacted by polymetallic nodule mining in the Clarion-Clipperton Zone (CCZ). In 2012, the International Seabed Authority (ISA) used available benthic biodiversity data and environmental proxies to establish nine no-mining areas, called Areas of Particular Environmental Interest (APEIs) in the CCZ. The APEIs were intended as a representative system of protected areas to safeguard biodiversity and ecosystem function across the region from mining impacts. Since 2012, a number of research programs have collected additional ecological baseline data from the CCZ. We assemble and analyze macrofaunal biodiversity data sets from eight studies, focusing on three dominant taxa (Polychaeta, Tanaidacea, and Isopoda), and encompassing 477 box-core samples to address the following questions: (1) How do macrofaunal abundance, biodiversity, and community structure vary across the CCZ, and what are the potential ecological drivers? (2) How representative are APEIs of the nearest contractor areas? (3) How broadly do macrofaunal species range across the CCZ region? and (4) What scientific gaps hinder our understanding of macrofaunal biodiversity and biogeography in the CCZ? Our analyses led us to hypothesize that sampling efficiencies vary across macrofaunal data sets from the CCZ, making quantitative comparisons between studies challenging. Nonetheless, we found that macrofaunal abundance and diversity varied substantially across the CCZ, likely due in part to variations in particulate organic carbon (POC) flux and nodule abundance. Most macrofaunal species were collected only as singletons or doubletons, with additional species still accumulating rapidly at all sites, and with most collected species appearing to be new to science. Thus, macrofaunal diversity remains poorly sampled and described across the CCZ, especially within APEIs, where a total of nine box cores have been taken across three APEIs. Some common macrofaunal species ranged over 600–3000 km, while other locally abundant species were collected across ≤ 200 km. The vast majority of macrofaunal species are rare, have been collected only at single sites, and may have restricted ranges. Major impediments to understanding baseline conditions of macrofaunal biodiversity across the CCZ include: (1) limited taxonomic description and/or barcoding of the diverse macrofauna, (2) inadequate sampling in most of the CCZ, especially within APEIs, and (3) lack of consistent sampling protocols and efficiencies.

Environmental Heterogeneity Throughout the Clarion-Clipperton Zone and the Potential Representativity of the APEI Network

Environmental variables such as food supply, nodule abundance, sediment characteristics, and water chemistry may influence abyssal seafloor communities and ecosystem functions at scales from meters to thousands of kilometers. Thus, knowledge of environmental variables is necessary to understand drivers of organismal distributions and community structure, and for selection of proxies for regional variations in community structure, biodiversity, and ecosystem functions. In October 2019, the Deep CCZ Biodiversity Synthesis Workshop was conducted to (i) compile recent seafloor ecosystem data from the Clarion-Clipperton Zone (CCZ), (ii) synthesize patterns of seafloor biodiversity, ecosystem functions, and potential environmental drivers across the CCZ, and (iii) assess the representativity of no-mining areas (Areas of Particular Environmental Interest, APEIs) for subregions and areas in the CCZ targeted for polymetallic nodule mining. Here we provide a compilation and summary of water column and seafloor environmental data throughout the CCZ used in the Synthesis Workshop and in many of the papers in this special volume. Bottom-water variables were relatively homogenous throughout the region while nodule abundance, sediment characteristics, seafloor topography, and particulate organic carbon flux varied across CCZ subregions and between some individual subregions and their corresponding APEIs. This suggests that additional APEIs may be needed to protect the full range of habitats and biodiversity within the CCZ.

DNA barcoding and cryptic diversity of deep-sea scavenging amphipods in the Clarion-Clipperton Zone (Eastern Equatorial Pacific)

The Clarion-Clipperton Zone (CCZ), located in the abyssal equatorial Pacific, has been subject to intensive international exploration for polymetallic nodule mining over the last four decades. Many studies have investigated the potential effects of mining on deep-sea ecosystems and highlighted the importance of defining environmental baseline conditions occurring at potential mining sites. However, current information on biodiversity and species distributions in the CCZ is still scarce and hampers the ability to effectively manage and reduce the potential impacts of mining activities. As part of the regulatory regimes adopted by the International Seabed Authority, concession holders are required to conduct an environmental impact assessment and gather baseline data on biodiversity and community structure in relation to their license areas. In the present study, we used an integrative molecular and morphological approach to assess species richness and genetic variation of deep-sea scavenging amphipods collected in two nodule-mining exploration areas (UK-1 and OMS-1 areas) and one Area of Particular Environmental Interest (APEI-6) in the eastern part of the CCZ. We analyzed the DNA sequences of the cytochrome c oxidase subunit I gene of 645 specimens belonging to ten distinct morphospecies. Molecular data uncover potential cryptic diversity in two investigated species, morphologically identified as Paralicella caperesca Shulenberger & Barnard, 1976 and Valettietta cf. anacantha (Birstein & Vinogradov, 1963). Our study highlights the importance of using molecular tools in conjunction with traditional morphological methods for modern biodiversity assessment studies, particularly to evaluate morphologically similar individuals and incomplete specimens. The results of this study can help determine species identity and ranges, information which can feed into environmental management.

Species boundaries and phylogeographic patterns in new species ofNannoniscus(Janiroidea: Nannoniscidae) from the equatorial Pacific nodule province inferred from mtDNA and morphology

AbstractSpatial patterns of genetic variation (based on COI and 16S mtDNA) for morphologically similar species in the isopod genus Nannoniscus G.O. Sars. 1870 were examined that occur broadly across the Clarion Clipperton Fracture Zone (CCZ). Samples were obtained from five different licence areas as well as an Area of Particular Environmental Interest (APEI-6) with sites located at various distances (a few to several hundred kilometres) from one another. Applying three different species delimitation (SD) methods (sGMYC, mPTP and ABGD) of the molecular data, we could distinguish between four and 12 different molecular taxonomic operational units (MOTUs). Morphological analyses could confirm five distinct phenotypic clades that represent species new to science and are described here: Nannoniscus brenkei sp. nov., Nannoniscus hilario sp. nov., Nannoniscus magdae sp. nov., Nannoniscus menoti sp. nov. and Nannoniscus pedro sp. nov. Despite the assumed limited dispersal capacity of Nannoniscus species, we found haplotypes of two species to be geographically widespread (up to > 1400 km apart), as opposed to several divergent clades occurring in close vicinity or even sympatry. Geographic distance appeared to explain the phylogeographic structure of Nannoniscus species to some extent, although oceanographic features and level of environmental heterogeneity were probably equally important.

Adult life strategy affects distribution patterns in abyssal isopods – implications for conservation in Pacific nodule areas

Abstract. With increasing pressure to extract minerals from the deep-sea bed, understanding the ecological and evolutionary processes that limit the spatial distribution of species is critical to assessing ecosystem resilience to mining impacts. The aim of our study is to gain a better knowledge about the abyssal isopod crustacean fauna of the central Pacific manganese nodule province (Clarion–Clipperton Fracture Zone, CCZ). In total, we examined 22 epibenthic sledge (EBS) samples taken at five abyssal areas located in the central northern Pacific including four contracting areas and one Area of Particular Environmental Interest (APEI3). Additional samples come from the DISturbance and reCOLonization experiment (DISCOL) area situated in the Peru Basin, southeastern Pacific. Using an integrative approach that combined morphological and genetic methods with species delimitation analyses (SDs) we assessed patterns of species range size, diversity, and community composition for four different isopod families (Munnopsidae Lilljeborg, 1864; Desmosomatidae Sars, 1897; Haploniscidae Hansen, 1916; and Macrostylidae Hansen, 1916) displaying different dispersal capacities as adults. Isopods are brooders, so their distribution and connectivity cannot be explained by larval dispersal but rather by adult locomotion. In particular, our objectives were to (1) identify potential differences in the distributional ranges of isopod families relative to their locomotory potential and to (2) evaluate the representativeness of the APEI for the preservation of regional biodiversity in the CCZ following mining disturbances. From 619 specimens, our SD analysis could distinguish 170 species, most of which were new to science (94.1 %). We found that increased locomotory ability correlated with higher species diversity with 9 species of Macrostylidae, 23 of Haploniscidae, 52 of Desmosomatidae, and 86 of Munnopsidae. This is supported by family-level rarefaction analyses. As expected, we found the largest species ranges in the families with swimming abilities, with a maximum recorded species range of 5245 and 4480 km in Munnopsidae and Desmosomatidae, respectively. The less motile Haploniscidae and Macrostylidae had maximal species ranges of 1391 and 1440 km, respectively. Overall, rarefaction analyses indicated that species richness did not vary much between areas, but the real number of species was still not sufficiently sampled. This is also indicated by the large proportion of singletons (40.5 %) found in this study. The investigated contractor areas in the CCZ were more similar in species composition and had a higher proportion of shared species between each other than the closely located APEI3 and the distantly located DISCOL area. In fact, the DISCOL area, located in the Peru Basin, had more species in common with the core CCZ areas than APEI3. In this regard, APEI3 does not appear to be representative as serving as a reservoir for the fauna of the investigated contractor areas, at least for isopods, as it has a different species composition. Certainly, more data from other APEIs, as well as preservation reference zones within contractor areas, are urgently needed in order to assess their potential as resources of recolonization of impacted seabed.

Using Habitat Classification to Assess Representativity of a Protected Area Network in a Large, Data-Poor Area Targeted for Deep-Sea Mining

Extractive activities in the ocean are expanding into the vast, poorly studied deep sea, with the consequence that environmental management decisions must be made for data-poor seafloor regions. Habitat classification can support marine spatial planning and inform decision-making processes in such areas. We present a regional, top–down, broad-scale, seafloor-habitat classification for the Clarion-Clipperton Fracture Zone (CCZ), an area targeted for future polymetallic nodule mining in abyssal waters in the equatorial Pacific Ocean. Our classification uses non-hierarchical, k-medoids clustering to combine environmental correlates of faunal distributions in the region. The classification uses topographic variables, particulate organic carbon flux to the seafloor, and is the first to use nodule abundance as a habitat variable. Twenty-four habitat classes are identified, with large expanses of abyssal plain and smaller classes with varying topography, food supply, and substrata. We then assess habitat representativity of the current network of protected areas (called Areas of Particular Environmental Interest) in the CCZ. Several habitat classes with high nodule abundance are common in mining exploration claims, but currently receive little to no protection in APEIs. There are several large unmanaged areas containing high nodule abundance on the periphery of the CCZ, as well as smaller unmanaged areas within the central CCZ, that could be considered for protection from mining to improve habitat representativity and safeguard regional biodiversity.