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Little is known about dispersal in deep-sea ecosystems, especially for sponges, which are abundant ecosystem engineers. Understanding patterns of gene flow in deep-sea sponges is essential, especially in areas where rising pressure from anthropogenic activities makes difficult to combine management and conservation. Here, we combined population genomics and oceanographic modelling to understand how Northeast Atlantic populations (Cantabrian Sea to Norway) of the deep-sea sponge Phakellia ventilabrum are connected. The analysis comprised ddRADseq derived SNP datasets of 166 individuals collected from 57 sampling stations from 17 different areas, including two Marine Protected Areas, one Special Area of Conservation and other areas with different levels of protection. The 4,017 neutral SNPs used indicated high connectivity and panmixis amongst the majority of areas (Ireland to Norway), spanning ca. 2,500-km at depths of 99-900 m. This was likely due to the presence of strong ocean currents allowing long-distance larval transport, as supported by our migration analysis and by 3D particle tracking modelling. On the contrary, the Cantabrian Sea and Roscoff (France) samples, the southernmost areas in our study, appeared disconnected from the remaining areas, probably due to prevailing current circulation patterns and topographic features, which might be acting as barriers for gene flow. Despite this major genetic break, our results suggest that all protected areas studied are well-connected with each other. Interestingly, analysis of SNPs under selection replicated results obtained for neutral SNPs. The relatively low genetic diversity observed along the study area, though, highlights the potential fragility of this species to changing climates, which might compromise resilience to future threats.

The deep-sea is vast, remote, and largely underexplored. However, methodological advances in environmental DNA (eDNA) surveys could aid in the exploration efforts, such as using sponges as natural eDNA filters for studying fish biodiversity. In this study, we analyzed the eDNA from 116 sponge tissue samples and compared these to 18 water eDNA samples and visual surveys obtained on an Arctic seamount. Across survey methods, we revealed approximately 30% of the species presumed to inhabit this area and 11 fish species were detected via sponge derived eDNA alone. These included commercially important fish such as the Greenland halibut and Atlantic mackerel. Fish eDNA detection was highly variable across sponge samples. Highest detection rates were found in sponges with low microbial activity such as those from the class Hexactinellida. The different survey methods also detected alternate fish communities, highlighted by only one species overlap between the visual surveys and the sponge eDNA samples. Therefore, we conclude that sponge eDNA can be a useful tool for surveying deep-sea demersal fish communities and it synergises with visual surveys improving overall biodiversity assessments. Datasets such as this can form comprehensive baselines on fish biodiversity across seamounts, which in turn can inform marine management and conservation practices in the regions where such surveys are undertaken.

Little is known about dispersal in deep-sea ecosystems, especially for sponges, which are abundant ecosystem engineers. Understanding patterns of gene flow in deep-sea sponges is essential, especially in areas where rising pressure from anthropogenic activities makes difficult to combine management and conservation. Here, we combined population genomics and oceanographic modelling to understand how Northeast Atlantic populations (Cantabrian Sea to Norway) of the deep-sea sponge Phakellia ventilabrum are connected. The analysis comprised ddRADseq derived SNP datasets of 166 individuals collected from 57 sampling stations from 17 different areas, including two Marine Protected Areas, one Special Area of Conservation and other areas with different levels of protection. The 4,017 neutral SNPs used indicated high connectivity and panmixis amongst the majority of areas (Ireland to Norway), spanning ca. 2,500-km at depths of 99–900 m. This was likely due to the presence of strong ocean currents allowing long-distance larval transport, as supported by our migration analysis and by 3D particle tracking modelling. On the contrary, the Cantabrian Sea and Roscoff (France) samples, the southernmost areas in our study, appeared disconnected from the remaining areas, probably due to prevailing current circulation patterns and topographic features, which might be acting as barriers for gene flow. Despite this major genetic break, our results suggest that all protected areas studied are well-connected with each other. Interestingly, analysis of SNPs under selection replicated results obtained for neutral SNPs. The relatively low genetic diversity observed along the study area, though, highlights the potential fragility of this species to changing climates, which might compromise resilience to future threats.

Trait-based approaches that complement taxonomy-based studies have increased in popularity among the scientific community over the last decades. The collection of biological and ecological characteristics of species (i.e., traits) provides insight into species and ecosystem vulnerability to environmental and anthropogenic changes, as well as ecosystem functioning. Here, we present the FUN Azores trait database, describe our approach, evaluate its scope, compare it to other marine trait databases, and explore the spatial distribution of its traits with “functional maps.” While most of the available trait databases to date contain essential information to understand the functional diversity of a taxonomic or functional group, our ecosystem-based approach provides a comprehensive assessment of diverse fauna (i.e., meio-, macro-, and megafauna) from benthic and pelagic environments in the Azores Marine Park; including ridges, seamounts, hydrothermal vents, and the overlying water column. We used a collaborative approach involving 30 researchers with different expertise to develop the FUN Azores database, which contains compiled data on 14 traits representing morphological, behavioral, and life history characteristics for 1,210 species across 10 phyla. The “functional maps” show a distinct distribution of the two most common size classes, suggesting different communities with different functionalities. The following traits had the best scoring coverage (i.e., >95% of the species scored): maximum body size, body form, skeleton material, feeding structure, motility, environmental position, substratum affinity, distribution, and depth range; while traits related to species behavior (e.g., sociability or aggregation tendencies) and life history (e.g., developmental mechanism) had lower scoring coverage, highlighting the need for further research to fill these knowledge gaps. We found a larger number of species in the benthic compared to the pelagic environment and differing

Trait-based approaches that complement taxonomic-based studies have increased in popularity among the scientific community over the last decades. The collection of biological and ecological characteristics of species (i.e., traits) provides insight into species and ecosystem vulnerability to environmental and anthropogenic changes, as well as ecosystem functioning. While most of the available trait databases to date contain essential information to understand the functional diversity of a taxonomic group or functional group based on size, the FUN Azores trait database has an ecosystem-based approach that provides a comprehensive assessment of diverse fauna (meio-, macro-, and megafauna) from benthic and pelagic environments in the Azores Marine Park; including ridges, seamounts, and hydrothermal vents. We used a collaborative approach involving 30 researchers with different expertise to develop the trait database; which contains compiled data on 14 traits representing morphological, behavioral, and life history characteristics for 1210 species, across 10 phyla

The Ægir Ridge System (ARS) is an ancient extinct spreading axis in the Nordic seas extending from the upper slope east of Iceland (∼550 m depth), as part of its Exclusive Economic Zone (EEZ), to a depth of ∼3,800 m in the Norwegian basin. Geomorphologically a rift valley, the ARS has a canyon-like structure that may promote increased diversity and faunal density. The main objective of this study was to characterize benthic habitats and related macro- and megabenthic communities along the ARS, and the influence of water mass variables and depth on them. During the IceAGE3 expedition (Icelandic marine Animals: Genetics and Ecology) on RV Sonne in June 2020, benthic communities of the ARS were surveyed by means of a remotely-operated vehicle (ROV) and epibenthic sledge (EBS). For this purpose, two working areas were selected, including abyssal stations in the northeast and bathyal stations in the southwest of the ARS. Video and still images of the seabed were usedtoqualitatively describebenthic habitats based on the presence of habitat-forming taxa and the physical environment. Patterns of diversity and community composition of the soft-sediment macrofauna, retrieved from the EBS, were analyzed in a semiquantitative manner. These biological data were complemented by producing high-resolution bathymetric maps using the vessel’s multi-beam echosounder system. As suspected, we were able to identify differences in species composition and number of macro- and megafaunal communities associated with a depth gradient. A biological canyon effect became evident in dense aggregates of megafaunal filter feeders and elevated macrofaunal densities. Analysis of videos and still images from the ROV transects also led to the discovery of a number ofVulnerable Marine Ecosystems (VMEs) dominated by sponges and soft corals characteristic of the Arctic region. Directions for future research encompass a more detailed, quantitative study of the megafauna and more coherent sampling over the

Vulnerable marine ecosystems (VMEs) are particularly susceptible to bottom-fishing activity as they are easily disturbed and slow to recover. A data-driven approach was developed to provide management options for the protection of VMEs under the European Union “deep-sea access regulations.” A total of two options within two scenarios were developed. The first scenario defined VME closure areas without consideration of fishing activity. Option 1 proposed closures for the protection of VME habitats and likely habitat, while Option 2 also included areas where four types of VME geophysical elements were present. The second scenario additionally considered fishing. This scenario used VME biomass—fishing intensity relationships to identify a threshold where effort of mobile bottom-contact gears was low and unlikely to have caused significant adverse impacts. Achieving a high level of VME protection requires the creation of many closures (> 100), made up of many small (∼50 km2) and fewer larger closures (> 1000 km2). The greatest protection of VMEs will affect approximately 9% of the mobile fleet fishing effort, while closure scenarios that avoid highly fished areas reduce this to around 4–6%. The framework allows managers to choose the level of risk-aversion they wish to apply in protecting VMEs by comparing alternative strategies.

In the deep ocean symbioses between microbes and invertebrates are emerging as key drivers of ecosystem health and services. We present a large-scale analysis of microbial diversity in deep-sea sponges (Porifera) from scales of sponge individuals to ocean basins, covering 52 locations, 1077 host individuals translating into 169 sponge species (including understudied glass sponges), and 469 reference samples, collected anew during 21 ship-based expeditions. We demonstrate the impacts of the sponge microbial abundance status, geographic distance, sponge phylogeny, and the physicalbiogeochemical environment as drivers of microbiome composition, in descending order of relevance. Our study further discloses that fundamental concepts of sponge microbiology apply robustly to sponges from the deep-sea across distances of >10,000 km. Deep-sea sponge microbiomes are less complex, yet more heterogeneous, than their shallow-water counterparts. Our analysis underscores the uniqueness of each deep-sea sponge ground based on which we provide critical knowledge for conservation of these vulnerable ecosystems.

© The Author(s), 2020. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Howell, K. L., Hilario, A., Allcock, A. L., Bailey, D. M., Baker, M., Clark, M. R., Colaco, A., Copley, J., Cordes, E. E., Danovaro, R., Dissanayake, A., Escobar, E., Esquete, P., Gallagher, A. J., Gates, A. R., Gaudron, S. M., German, C. R., Gjerde, K. M., Higgs, N. D., Le Bris, N., Levin, L. A., Manea, E., McClain, C., Menot, L., Mestre, N. C., Metaxas, A., Milligan, R. J., Muthumbi, A. W. N., Narayanaswamy, B. E., Ramalho, S. P., Ramirez-Llodra, E., Robson, L. M., Rogers, A. D., Sellanes, J., Sigwart, J. D., Sink, K., Snelgrove, P. V. R., Stefanoudis, P., V., Sumida, P. Y., Taylor, M. L., Thurber, A. R., Vieira, R. P., Watanabe, H. K., Woodall, L. C., & Xavier, J. R. A blueprint for an inclusive, global deep-sea ocean decade field program. Frontiers in Marine Science, 7, (2020): 584861, doi:10.3389/fmars.2020.584861., The ocean plays a crucial role in the functioning of the Earth System and in the provision of vital goods and services. The United Nations (UN) declared 2021–2030 as the UN Decade of Ocean Science for Sustainable Development. The Roadmap for the Ocean Decade aims to achieve six critical societal outcomes (SOs) by 2030, through the pursuit of four objectives (Os). It specifically recognizes the scarcity of biological data for deep-sea biomes, and challenges the global scientific community to conduct research to advance understanding of deep-sea ecosystems to inform sustainable management. In this paper, we map four key scientific questions identified by the academic community to the Ocean Decade SOs: (i) What is the diversity of life in the deep ocean? (ii) How are populations and habitats connected? (iii) What is the role of living organisms in ecosystem function and service provision? and (iv) How do species, communities, and ecosystems respond to disturbance? We then consider the design of a global-scale program to address these questions by reviewing key drivers of ecological pattern and process. We recommend using the following criteria to stratify a global survey design: biogeographic region, depth, horizontal distance, substrate type, high and low climate hazard, fished/unfished, near/far from sources of pollution, licensed/protected from industry activities. We consider both spatial and temporal surveys, and emphasize new biological data collection that prioritizes southern and polar latitudes, deeper (> 2000 m) depths, and midwater environments. We provide guidance on observational, experimental, and monitoring needs for different benthic and pelagic ecosystems. We then review recent efforts to standardize biological data and specimen collection and archiving, making “sampling design to knowledge application” recommendations in the context of a new global program. We also review and comment on needs, and recommend actions, to develop capacity in deep-sea res, Development of this paper was supported by funding from the Scientific Committee on Oceanic Research (SCOR) awarded to KH and AH as working group 159 co-chairs. KH, BN, and KS are supported by the UKRI funded One Ocean Hub NE/S008950/1. AH work is supported by the CESAM (UIDP/50017/2020 + 1432 UIDB/50017/2020) that is funded by Fundação para a Ciência e a Tecnologia (FCT)/MCTES through national funds. AA is supported by Science Foundation Ireland and the Marine Institute under the Investigators Program Grant Number SFI/15/IA/3100 co-funded under the European Regional Development Fund 2014–2020. AC is supported through the FunAzores -ACORES 01-0145-FEDER-000123 grant and by FCT through strategic project UID/05634/2020 and FCT and Direção-Geral de Politica do Mar (DGPM) through the project Mining2/2017/005. PE is funded by national funds (OE), through FCT in the scope of the framework contract foreseen in the numbers 4, 5 and 6 of the article 23, of the Decree-Law 57/2016, of August 29, changed by Law 57/2017, of July 19. SG research is supported by CNRS funds. CG is supported by an Independent Study Award and the Investment in Science Fund at WHOI. KG gratefully acknowledges support from Synchronicity Earth. LL is funded by the NOAA Office of Ocean Exploration and Research (NA19OAR0110305) and the US National Science Foundation (OCE 1634172). NM is supported by FCT and DGPM, through the project Mining2/2017/001 and the FCT grants CEECIND/00526/2017, UIDB/00350/2020 + UIDP/00350/2020. SR is funded by the FCTgrant CEECIND/00758/2017. JS is supported by ANID FONDECYT #1181153 and ANID Millennium Science Initiative Program #NC120030. JX research is funded by the European Union’s Horizon 2020 research and innovation program through the SponGES project (grant agreement no. 679849) and further supported by national funds through FCT within the scope of UIDB/04423/2020 and UIDP/04423/2020. The Natural Sciences and Engineering Council of Canada supports AM and PVRS. MB and th

Lithistid sponges are globally distributed in temperate and sub-tropical areas, constituting an important component of deep-sea benthic communities where they form structurally complex and vulnerable marine ecosystems (VMEs). In this study, we assess the diversity and investigate the spatial and bathymetric distribution of the lithistid sponges of the Azores archipelago (North Atlantic) based on historical records and examination of samples accidentally collected during deep-sea longline fishing operations in the region. Eleven lithistid species are recognized to occur in the Azores, including Leiodermatium tuba, recently described from material collected in several Northeast Atlantic seamounts that is hereby reported for the first time to the archipelago. We provide molecular barcodes (mtDNA COI and rRNA 28S) for seven of these species, including Discodermia ramifera, Macandrewia azorica, and Exsuperantia archipelagus, for which the Azores constitutes the type locality. We further discuss the phylogenetic and biogeographic affinities of the Azorean lithistids in the context of the Porifera classification, and the wider Northeast Atlantic upper bathyal fauna. Our study also warrants the addition of some lithistid species to the list of VME indicators for the Northeast Atlantic in support of the sustainable management and conservation of these species and habitats, as well as the ecological functions they deliver.

Vulnerable marine ecosystems (VMEs) are particularly susceptible to bottom-fishing activity as they are easily disturbed and slow to recover. A data-driven approach was developed to provide management options for the protection of VMEs under the European Union “deep-sea access regulations.” A total of two options within two scenarios were developed. The first scenario defined VME closure areas without consideration of fishing activity. Option 1 proposed closures for the protection of VME habitats and likely habitat, while Option 2 also included areas where four types of VME geophysical elements were present. The second scenario additionally considered fishing. This scenario used VME biomass—fishing intensity relationships to identify a threshold where effort of mobile bottom-contact gears was low and unlikely to have caused significant adverse impacts. Achieving a high level of VME protection requires the creation of many closures (> 100), made up of many small (∼50 km2) and fewer larger closures (> 1000 km2). The greatest protection of VMEs will affect approximately 9% of the mobile fleet fishing effort, while closure scenarios that avoid highly fished areas reduce this to around 4–6%. The framework allows managers to choose the level of risk-aversion they wish to apply in protecting VMEs by comparing alternative strategies.

Sponges are amongst the most difficult benthic taxa to properly identify, which has led to a prevalence of cryptic species in several sponge genera, especially in those with simple skeletons. This is particularly true for sponges living in remote or hardly accessible environments, such as the deep-sea, as the inaccessibility of their habitat and the lack of accurate descriptions usually leads to misclassifications. However, species can also remain hidden even when they belong to genera that have particularly characteristic features. In these cases, researchers inevitably pay attention to these peculiar features, sometimes disregarding small differences in the other “typical” spicules. The genus Melonanchora Carter, 1874, is among those well suited for a revision, as their representatives possess a unique type of spicule (spherancorae). After a thorough review of the material available for this genus from several institutions, four new species of Melonanchora, M. tumultuosa sp. nov., M. insulsa sp. nov., M. intermedia sp. nov. and M. maeli sp. nov. are formally described from different localities across the Atlanto-Mediterranean region. Additionally, all Melonanchora from the Okhotsk Sea and nearby areas are reassigned to other genera; Melonanchora kobjakovae is transferred to Myxilla (Burtonanchora) while two new genera, Hanstoreia gen. nov. and Arhythmata gen. nov. are created to accommodate Melonanchora globogilva and Melonanchora tetradedritifera, respectively. Hanstoreia gen. nov. is closest to Melonanchora, whereas Arhythmata gen. nov., is closer to Stelodoryx, which is most likely polyphyletic and in need of revision.

Sponges are amongst the most difficult benthic taxa to properly identify, which has led to a prevalence of cryptic species in several sponge genera, especially in those with simple skeletons. This is particularly true for sponges living in remote or hardly accessible environments, such as the deep-sea, as the inaccessibility of their habitat and the lack of accurate descriptions usually leads to misclassifications. However, species can also remain hidden even when they belong to genera that have particularly characteristic features. In these cases, researchers inevitably pay attention to these peculiar features, sometimes disregarding small differences in the other “typical” spicules. The genus Melonanchora Carter, 1874, is among those well suited for a revision, as their representatives possess a unique type of spicule (spherancorae). After a thorough review of the material available for this genus from several institutions, four new species of Melonanchora, M. tumultuosa sp. nov., M. insulsa sp. nov., M. intermedia sp. nov. and M. maeli sp. nov. are formally described from different localities across the Atlanto-Mediterranean region. Additionally, all Melonanchora from the Okhotsk Sea and nearby areas are reassigned to other genera; Melonanchora kobjakovae is transferred to Myxilla (Burtonanchora) while two new genera, Hanstoreia gen. nov. and Arhythmata gen. nov. are created to accommodate Melonanchora globogilva and Melonanchora tetradedritifera, respectively. Hanstoreia gen. nov. is closest to Melonanchora, whereas Arhythmata gen. nov., is closer to Stelodoryx, which is most likely polyphyletic and in need of revision.

Background: Lithistid demosponges, also known as rock sponges, are a polyphyletic group of sponges which are widely distributed. In the Northeast Atlantic (NEA), 17 species are known and the current knowledge on their distribution is mainly restricted to the Macaronesian islands. In the Mediterranean Sea, 14 species are recorded and generally found in marine caves. Methods: Lithistids were sampled in nine NEA seamounts during the scientific expeditions Seamount 1 (1987) and Seamount 2 (1993) organized by the MNHN of Paris. Collected specimens were identified through the analyses of external and internal morphological characters using light and scanning electron microscopy, and compared with material from various museum collections as well as literature records. Results: A total of 68 specimens were analysed and attributed to 17 species across two orders, seven families, and seven genera, representing new records of distribution. Ten of these species are new to science, viz. Neoschrammeniella inaequalis sp. nov., N. piserai sp. nov., N. pomponiae sp. nov., Discodermia arbor sp. nov., D. kellyae sp. nov., Macandrewia schusterae sp. nov., M. minima sp. nov., Exsuperantia levii sp. nov., Leiodermatium tuba sp. nov. and Siphonidium elongatus sp. nov., and are here described and illustrated. New bathymetric records were also found for D. ramifera, D. verrucosa and M. robusta. The Meteor seamount group has a higher species richness (15 species) compared to the Lusitanian seamount group (six species). The majority of the species had their distribution restricted to one seamount, and ten are only known from a single locality, but this can be a result of sample bias. Discussion: The number of species shared between the seamounts and the Macaronesian islands is very reduced. The same pattern repeats between the NEA and Mediterranean Sea. This study demonstrates that NEA seamounts are ecosystems with a higher diversity of lithistids than previously thought, increasing the numbe

The United Nations Decade of Ocean Science for Sustainable Development presents an exceptional opportunity to effect positive change in ocean use. We outline what is required of the deep-sea research community to achieve these ambitious objectives. The health of the global ocean, on which society depends, is in decline. The importance of sustainable use to ocean health has long been recognized1, yet the United Nations (UN) First World Ocean Assessment2 from 2017 highlighted increasing ocean pressures from accelerated expansion of human activities, including climate change. These pressures affect all ocean regions, from the coast to the deep sea3. In response to this concern, and to align with several international policy commitments, the UN General Assembly proclaimed 2021–2030 the Decade of Ocean Science for Sustainable Development4. The roadmap for this Ocean Decade recognizes the deep sea as a frontier of science and discovery, and calls for research to advance understanding of deep-sea ecosystems, their functions, vulnerabilities and services to society. Published in March 2020, the draft implementation plan for the Ocean Decade guides the design and implementation of ‘Actions’ that underpin the desired move from ‘the ocean we have’ to ‘the ocean we want’ (Fig. 1). The draft plan calls on the scientific community to develop Actions, at programme, project, activity or contribution levels, to help deliver on four key objectives: (1) increase capacity to generate, understand, manage and use ocean knowledge; (2) identify and generate required ocean data, information and knowledge; (3) build comprehensive understanding of the ocean and its governance systems; and (4) increase the use of ocean knowledge.

Deep-sea sponge-dominated communities are complex habitats considered hotspots of biodiversity and ecosystem functioning. They are classified as Vulnerable Marine Ecosystem and are listed as threatened or declining as a result of anthropogenic activities. Yet, studies into the distribution, community structure and composition of these habitats are scarce, hampering the development of appropriate management measures to ensure their conservation. In this study we describe a diverse benthic community, dominated by a lithistid sponge, found in two geomorphological features of important conservation status —Le Danois Bank and El Corbiro Canyon— of the Cantabrian Sea. Based on the analyses of visual transects using a photogrammetric towed vehicle and samples collected by rock dredge, we characterize the habitat and the associated community in detail. This deep-sea sponge aggregation was found on bedrock. It is dominated by one lithistid sponge, Neoschrammeniella aff. bowerbankii (0.2 ind./m2) and further composed of various sponge species as well as of other benthic invertebrates such as cnidarians, bryozoans and crustaceans. Using a non-invasive methodology (SfM – Structure from Motion) and empirical relationships of individuals size and biomass/volume obtained in laboratory for N. aff. bowerbankii, we were able to estimate a total biomass of 41 kg and volume of 39 l of this species in the surveyed area. This approach allows a fine tune methodology for estimating biomass and volume by image-based-observed area avoiding destructive techniques for this species.