Towards a scientific community consensus on designating Vulnerable Marine Ecosystems from imagery.

Management of deep-sea fisheries in areas beyond national jurisdiction by Regional Fisheries Management Organizations/Arrangements (RFMO/As) requires identification of areas with Vulnerable Marine Ecosystems (VMEs). Currently, fisheries data, including trawl and longline bycatch data, are used by many RFMO/As to inform the identification of VMEs. However, the collection of such data creates impacts and there is a need to collect non-invasive data for VME identification and monitoring purposes. Imagery data from scientific surveys satisfies this requirement, but there currently is no established framework for identifying VMEs from images. Thus, the goal of this study was to bring together a large international team to determine current VME assessment protocols and establish preliminary global consensus guidelines for identifying VMEs from images. An initial assessment showed a lack of consistency among RFMO/A regions regarding what is considered a VME indicator taxon, and hence variability in how VMEs might be defined. In certain cases, experts agreed that a VME could be identified from a single image, most often in areas of scleractinian reefs, dense octocoral gardens, multiple VME species' co-occurrence, and chemosynthetic ecosystems. A decision flow chart is presented that gives practical interpretation of the FAO criteria for single images. To further evaluate steps of the flow chart related to density, data were compiled to assess whether scientists perceived similar density thresholds across regions. The range of observed densities and the density values considered to be VMEs varied considerably by taxon, but in many cases, there was a statistical difference in what experts considered to be a VME compared to images not considered a VME. Further work is required to develop an areal extent index, to include a measure of confidence, and to increase our understanding of what levels of density and diversity correspond to key ecosystem functions for VME indicator taxa. Based on our results, the following recommendations are made: 1. There is a need to establish a global consensus on which taxa are VME indicators. 2. RFMO/As should consider adopting guidelines that use imagery surveys as an alternative (or complement) to using bycatch and trawl surveys for designating VMEs. 3. Imagery surveys should also be included in Impact Assessments. And 4. All industries that impact the seafloor, not just fisheries, should use imagery surveys to detect and identify VMEs.

Towards a revision of the bamboo corals (Octocorallia): Part 5, new genera and species of Keratoisididae from the Tasmanian deep sea.

Four new genera and six new species of deep-sea bamboo corals (Family: Keratoisididae) from the seamounts off the eastern and southern coast of Tasmania are described and illustrated. In situ high-definition images from remotely operated vehicles, compound microscopy, and Scanning Electron Microscopy (SEM) of 29 specimens collected at depths of 729 to 3950 m were used to provide data on distinguishing morphological characters. The species described include two new species in the genus Keratoisis, two species in the newly described genus Adinisis, and one species each in the new genera Tanyostea and Dokidisis. The species Keratoisis magnifica and Lepidisis solitaria are re-assigned to the newly described genus Onkoisis due to the shape and design of the polyp arrangement and sclerites, which differ considerably from that of the type species of Keratoisis.

Towards a revision of the bamboo corals (Octocorallia): Part 4, delineating the family Keratoisididae.

The systematics of bamboo corals of the Family Keratoisididae are evaluated using both DNA sequences and morphological data. Sequence data were obtained from 398 specimens, from which 77 unique haplotypes representing the mtMutS and 18S gene regions were identified. These were aligned with sequences downloaded from GenBank from an additional 12 keratoisids and 6 octocoral outgroups. Phylogenetic analyses recovered seven well-supported major clades, the most recently derived of which consists of several subclades. Each clade and subclade can be characterized by a suite of morphological characters that include axis construction, branching pattern, polyp form, and sclerite type and arrangement. This analysis also shows that keratoisid genera described >100 years ago are paraphyletic and need revision and that a large number of new genera will need to be described.

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.

Toward a revision of the bamboo corals: Part 3, deconstructing the Family Isididae.

Bamboo corals are distinguished from most other octocorals by an articulated skeleton. The nodes are proteinaceous and sclerite-free while the internodes are composed of non-scleritic calcium carbonate. This articulation of the skeleton was thought to be unique and a strong synapomorphy for the family Isididae. Our phylogeny, based on the amplification of mtMutS and 18S, shows an articulating skeleton with sclerite-free nodes has arisen independently at least five times during the evolutionary history of Octocorallia rather than being a synapomorphy characteristic of a monophyletic bamboo coral clade. The family Isididae is currently composed of four subfamilies (Circinisidinae, Isidinae, Keratoisidinae, and Mopseinae). Not only is the family polyphyletic, but our genetic analyses suggest also the subfamily Isidinae is polyphyletic based on current taxonomic classifications, and Mopseinae is not monophyletic. The type, Isis, is found outside of the well-supported Calcaxonia Pennatulacea clade where the other members of Isididae cluster. The current classification of the family Isididae does not reflect the evolutionary history of an articulated skeleton. To better reflect the evolutionary history of these taxa we propose that three of the four the subfamilies, the genus Isidoides, and genera within the subfamily Isidinae, be elevated to family level to produce a classification with five families with a bamboo-like skeleton: Chelidonisididae, Isididae, Isidoidae, Keratoisididae, and Mopseidae.

Report on hydrozoans (Cnidaria), excluding Stylasteridae, from the Emperor Seamounts, western North Pacific Ocean.

Fourteen species of hydroids, collected during August 2019 by ROV SuBastian of the Schmidt Ocean Institute, are reported from the Emperor Seamount chain in the western North Pacific Ocean. Two others, Candelabrum sp. and Eudendrium sp., were observed only on videos taken by the ROV. From collections and video observations, eight species of hydroids were found at Jingu Seamount, three at Yomei, Nintoku, and Annei seamounts, and one at Koko Seamount and Hess Rise. At Suiko and Godaigo seamounts, hydroids were seen in videos but they could not be identified. Latebrahydra schulzei, an endobiotic associate of the hexactinellid sponge Walteria flemmingii Schulze, 1886 from Annei Seamount and Hess Rise, is described as a new genus and species tentatively attributed to Hydractiniidae L. Agassiz, 1862. Another new species, Hydractinia galeai, is described from Jingu Seamount. Among its distinctive characters is a zooid termed a sellectozooid, likely serving in both food capture and defence. Hydroids examined from Yomei, Nintoku, and Jingu seamounts are elements of a cold-water fauna occurring in the North Pacific Boreal Bathyal province, while those of Annei and Koko seamounts, and Hess Rise, are part of the biota of the Central North Pacific Bathyal province. Hydroids identified as Bouillonia sp., from Nintoku Seamount, represent the first record of this predominantly deep water tubulariid genus in the North Pacific Ocean. Bonneviella superba Nutting, 1915, from Jingu Seamount, is reported for the first time outside the Aleutian Islands. Bonneviella cf. gracilis Fraser, 1939, known elsewhere only from Dease Strait in the western Canadian Arctic, was also collected on Jingu. In addition to hydroids, medusae of Ptychogastria polaris Allman, 1878 were observed on videos from Nintoku, Jingu, Annei, and Koko seamounts at depths between 2423-1422 m. An unidentified siphonophore was observed near bottom at 2282 m on Nintoku Seamount.

Toward a revision of the bamboo corals: Part 1, species in the Muricellisidinae (Octocorallia: Isididae).

The subfamily Muricellisidinae was erected by Kükenthal in 1915 and placed in the family Isididae in order to accommodate an unusual species collected in Sagami Bay, Japan. In 1931, Thomson and Dean added a second species collected in Indonesia during the Siboga Expedition. The holotypes of both species have been re-examined. Muricellisis echinata was found to be an anthothelid living on the axis of a keratoisid bamboo coral and M. cervicornis is a melithaeid. Both species are redescribed.

Characterization of deep-sea benthic invertebrate megafauna of the Galapagos Islands.

The deep sea represents the largest and least explored biome on the planet. Despite the iconic status of the Galapagos Islands and being considered one of the most pristine locations on earth, the deep-sea benthic ecosystems of the archipelago are virtually unexplored in comparison to their shallow-water counterparts. In 2015, we embarked on a multi-disciplinary scientific expedition to conduct the first systematic characterization of deep-sea benthic invertebrate communities of the Galapagos, across a range of habitats. We explored seven sites to depths of over 3,300 m using a two-part Remotely Operated Vehicle (ROV) system aboard the E/V Nautilus, and collected 90 biological specimens that were preserved and sent to experts around the world for analysis. Of those, 30 taxa were determined to be undescribed and new to science, including members of five new genera (2 sponges and 3 cnidarians). We also systematically analysed image frame grabs from over 85 h of ROV footage to investigate patterns of species diversity and document the presence of a range of underwater communities between depths of 290 and 3,373 m, including cold-water coral communities, extensive glass sponge and octocoral gardens, and soft-sediment faunal communities. This characterization of Galapagos deep-sea benthic invertebrate megafauna across a range of ecosystems represents a first step to study future changes that may result from anthropogenic impacts to the planet's climate and oceans, and informed the creation of fully protected deep-water areas in the Galapagos Marine Reserve that may help preserve these unique communities in our changing planet.

Four new species of Magelona (Annelida: Magelonidae) from Easter Island, Guam and Hawaii.

Several collections of magelonids from Easter Island, Guam in the Mariana Islands and Hawaii were examined. Five magelonids are fully described and illustrated: Magelona anuheone sp. nov. (Easter Island), M. cf. symmetrica Mortimer Mackie, 2006 (Guam), M. alexandrae sp. nov. (Hawaii), M. cinthyae sp. nov. (Hawaii) and M. paulolanai sp. nov. (Hawaii and Guam). Magelona anuheone sp. nov. has a distinctly longer than wide prostomium with inflated and well-developed prostomial horns, similar to M. montera from the Indian Ocean. Magelona alexandrae sp. nov. belongs to Magelona species with crenulated upper margin of thoracic postchaetal lamellae and also well-developed prostomial horns. Magelona cinthyae sp. nov. is unique among its congeners in that several collected adult individuals presented two or more eyespots in the prostomium, a feature only previously recorded to pelagic larval stages. Magelona paulolanai sp. nov. belongs to a group of species with rudimentary prostomial horns, simple capillaries on chaetiger 9 and tridentate abdominal hooks. All species are illustrated and compared to their morphologically similar congeners.

Abyssal fauna of the UK-1 polymetallic nodule exploration area, Clarion-Clipperton Zone, central Pacific Ocean: Cnidaria.

BACKGROUND: We present data from a DNA taxonomy register of the abyssal Cnidaria collected as part of the Abyssal Baseline (ABYSSLINE) environmental survey cruise 'AB01' to the UK Seabed Resources Ltd (UKSRL) polymetallic-nodule exploration area 'UK-1' in the eastern Clarion-Clipperton Zone (CCZ), central Pacific Ocean abyssal plain. This is the second paper in a series to provide regional taxonomic data for a region that is undergoing intense deep-sea mineral exploration for high-grade polymetallic nodules. Data were collected from the UK-1 exploration area following the methods described in Glover et al. (2015b). NEW INFORMATION: Morphological and genetic data are presented for 10 species and 18 records identified by a combination of morphological and genetic data, including molecular phylogenetic analyses. These included 2 primnoid octocorals, 2 isidid octocorals, 1 anemone, 4 hydroids (including 2 pelagic siphonophores accidentally caught) and a scyphozoan jellyfish (in the benthic stage of the life cycle). Two taxa matched previously published genetic sequences (pelagic siphonophores), two taxa matched published morphological descriptions (abyssal primnoids described from the same locality in 2015) and the remaining 6 taxa are potentially new species, for which we make the raw data, imagery and vouchers available for future taxonomic study. We have used a precautionary approach in taxon assignments to avoid over-estimating species ranges. The Clarion-Clipperton Zone is a region undergoing intense exploration for potential deep-sea mineral extraction. We present these data to facilitate future taxonomic and environmental impact study by making both data and voucher materials available through curated and accessible biological collections. For some of the specimens we also provide image data collected at the seabed by ROV, wich may facilitate more accurate taxon designation in coming ROV or AUV surveys.

Platycuma bamberconfabulor sp. nov. (Crustacea: Cumacea: Nannastacidae) from Antarctica, with a note on the gut of Platycuma.

Platycuma bamberconfabulor sp. nov. is based on a specimen from the Ross Sea. The species can be distinguished from the other species in the genus through the combination of a laterally expanded and somewhat dorsoventrally flattened carapace, anterior margin of the carapace excavate in dorsal view, uropod peduncles less than twice as long as pleonite 6, and subequal pleonites 4 and 5.

Environmental influences on the Indo-Pacific octocoral Isis hippuris Linnaeus 1758 (Alcyonacea: Isididae): genetic fixation or phenotypic plasticity?

As conspicuous modular components of benthic marine habitats, gorgonian (sea fan) octocorals have perplexed taxonomists for centuries through their shear diversity, particularly throughout the Indo-Pacific. Phenotypic incongruence within and between seemingly unitary lineages across contrasting environments can provide the raw material to investigate processes of disruptive selection. Two distinct phenotypes of the Isidid Isis hippurisLinnaeus, 1758 partition between differing reef environments: long-branched bushy colonies on degraded reefs, and short-branched multi/planar colonies on healthy reefs within the Wakatobi Marine National Park (WMNP), Indonesia. Multivariate analyses reveal phenotypic traits between morphotypes were likely integrated primarily at the colony level with increased polyp density and consistently smaller sclerite dimensions at the degraded site. Sediment load and turbidity, hence light availability, primarily influenced phenotypic differences between the two sites. This distinct morphological dissimilarity between the two sites is a reliable indicator of reef health; selection primarily acting on colony morphology, porosity through branching structure, as well as sclerite diversity and size. ITS2 sequence and predicted RNA secondary structure further revealed intraspecific variation between I. hippuris morphotypes relative to such environments (ΦST = 0.7683, P < 0.001). This evidence suggests-but does not confirm-that I. hippuris morphotypes within the WMNP are two separate species; however, to what extent and taxonomic assignment requires further investigation across its full geographic distribution. Incongruence between colonies present in the WMNP with tenuously described Isis alternatives (Isis reticulataNutting, 1910, Isis minorbrachyblastaZou, Huang & Wang, 1991), questions the validity of such assignments. Furthermore, phylogenetic analyses confirm early taxonomic suggestion that the characteristic jointed axis of the Isididae is in fact a convergent trait. Thus the polyphyletic nature of the Isididae lies in its type species I. hippuris, being unrelated to the rest of its family members.

A new genus of bamboo coral (Octocorallia: Isididae) from the Bahamas.

A bamboo coral collected during a deep-sea expedition to the Bahamas in 2009 proved to have a unique combination of features for a member of the bamboo coral subfamily Keratoisidinae: the structure and shape of the polyps, the sclerites consisting entirely of rods, some of which extend the length of the polyp, the delicateness of the branches with solid internodes, and the deep funnel construction of the peristomal region into which the tentacles can contract. The specimens are described as a new genus and species.

Biotic and human vulnerability to projected changes in ocean biogeochemistry over the 21st century.

Ongoing greenhouse gas emissions can modify climate processes and induce shifts in ocean temperature, pH, oxygen concentration, and productivity, which in turn could alter biological and social systems. Here, we provide a synoptic global assessment of the simultaneous changes in future ocean biogeochemical variables over marine biota and their broader implications for people. We analyzed modern Earth System Models forced by greenhouse gas concentration pathways until 2100 and showed that the entire world's ocean surface will be simultaneously impacted by varying intensities of ocean warming, acidification, oxygen depletion, or shortfalls in productivity. In contrast, only a small fraction of the world's ocean surface, mostly in polar regions, will experience increased oxygenation and productivity, while almost nowhere will there be ocean cooling or pH elevation. We compiled the global distribution of 32 marine habitats and biodiversity hotspots and found that they would all experience simultaneous exposure to changes in multiple biogeochemical variables. This superposition highlights the high risk for synergistic ecosystem responses, the suite of physiological adaptations needed to cope with future climate change, and the potential for reorganization of global biodiversity patterns. If co-occurring biogeochemical changes influence the delivery of ocean goods and services, then they could also have a considerable effect on human welfare. Approximately 470 to 870 million of the poorest people in the world rely heavily on the ocean for food, jobs, and revenues and live in countries that will be most affected by simultaneous changes in ocean biogeochemistry. These results highlight the high risk of degradation of marine ecosystems and associated human hardship expected in a future following current trends in anthropogenic greenhouse gas emissions.

The World's largest known Gorgonian.

Gorgonians in the deep sea can be much larger than their shallow water counterparts, but there are only a few scattered measurements. We have estimated the size of a chrysogorgiid gorgonian, Iridogorgia magnispiralis Watling 2007, observed from a submersible on Twin Banks in the Northwest Hawaiian Islands, as being 5.7 m tall.

Deep-sea origin and in-situ diversification of chrysogorgiid octocorals.

The diversity, ubiquity and prevalence in deep waters of the octocoral family Chrysogorgiidae Verrill, 1883 make it noteworthy as a model system to study radiation and diversification in the deep sea. Here we provide the first comprehensive phylogenetic analysis of the Chrysogorgiidae, and compare phylogeny and depth distribution. Phylogenetic relationships among 10 of 14 currently-described Chrysogorgiidae genera were inferred based on mitochondrial (mtMutS, cox1) and nuclear (18S) markers. Bathymetric distribution was estimated from multiple sources, including museum records, a literature review, and our own sampling records (985 stations, 2345 specimens). Genetic analyses suggest that the Chrysogorgiidae as currently described is a polyphyletic family. Shallow-water genera, and two of eight deep-water genera, appear more closely related to other octocoral families than to the remainder of the monophyletic, deep-water chrysogorgiid genera. Monophyletic chrysogorgiids are composed of strictly (Iridogorgia Verrill, 1883, Metallogorgia Versluys, 1902, Radicipes Stearns, 1883, Pseudochrysogorgia Pante & France, 2010) and predominantly (Chrysogorgia Duchassaing & Michelotti, 1864) deep-sea genera that diversified in situ. This group is sister to gold corals (Primnoidae Milne Edwards, 1857) and deep-sea bamboo corals (Keratoisidinae Gray, 1870), whose diversity also peaks in the deep sea. Nine species of Chrysogorgia that were described from depths shallower than 200 m, and mtMutS haplotypes sequenced from specimens sampled as shallow as 101 m, suggest a shallow-water emergence of some Chrysogorgia species.