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Epigenetic modifications are thought to be one of the molecular mechanisms involved in plastic adaptive responses to environmental variation. However, studies reporting associations between genome-wide epigenetic changes and habitat-specific variations in life history traits (e.g., lifespan, reproduction) are still scarce, likely due to the recent application of methylome resequencing methods to non-model species. In this study, we examined associations between whole genome DNA methylation and environmentally driven life history variation in 2 lineages of a marine fish, the capelin (Mallotus villosus), from North America and Europe. In both lineages, capelin harbor 2 contrasting life history tactics (demersal vs. beach-spawning). Performing whole genome and methylome sequencing, we showed that life history tactics are associated with epigenetic changes in both lineages, though the effect was stronger in European capelin. Genetic differentiation between the capelin harboring different life history tactics was negligible, but we found genome-wide methylation changes in both lineages. We identified 9,125 European and 199 North American differentially methylated regions (DMRs) due to life history. Gene ontology (GO) enrichment analysis for both lineages revealed an excess of terms related to neural function. Our results suggest that environmental variation causes important epigenetic changes that are associated with contrasting life history tactics in lineages with divergent genetic backgrounds, with variable importance of genetic variation in driving epigenetic variation. Our study emphasizes the potential role of genome-wide epigenetic variation in adaptation to environmental variation.

The USA, EU, and Canada established a trilateral working group on the ecosystem approach to ocean health and stressors, under the Atlantic Ocean Research Alliance. Recognizing the Atlantic Ocean as a shared resource and responsibility, the working group sought to advance understanding of the Atlantic Ocean and its dynamic systems to improve ocean health, enhance ocean stewardship, and promote the sustainable use and management of its resources. This included consideration of multiple ocean-use sectors such as fishing, shipping, tourism and offshore energy. The working group met for 4 years and worked through eight steps that covered the development of common language as a basis for collaboration, challenges of stakeholder engagement, review of the governance mandates, exploring the links between sectors and ecosystems effects, identifying gaps in knowledge and uptake of science, identification of tools for ecosystem based management, customary best practice for tool development and communication of key research priorities. The key findings were that ecosystem-based management enables new benefits and opportunities, and that we need to make the business case. That adequate mandates and effective tools exist for ecosystem-based management, and that ecosystem-based management urgently requires integration of human dimensions, so we must diversify the conversation. Also that stakeholders don't see their stake in ecosystem based management, so greater engagement with stakeholders and targeting of ocean literacy is required and that a sustainable future requires a sustained investment in ecosystem-based management, so long term commitment is key. This article is protected by copyright. All rights reserved.

Deep-sea sponges are important contributors to carbon and nitrogen cycling due to their large filtration capacity. Species of the suborder Astrophorina form dense sponge grounds in the North Atlantic, where they serve as prey for spongivores, but also have non-trophic interactions with commensal epi- and endobionts. At the Flemish Cap (NW Atlantic) Astrophorina sponges are present in four previously described deep-sea epifaunal assemblages: the deep-sea coral assemblage, the lower slope assemblages 1 and 2, and the deep-sea sponge assemblage. To investigate their role in trophic and non-trophic interactions at the Flemish Cap, we developed trophic-/ non-trophic interaction web models for each of the four faunal assemblages using the published literature. By excluding the sponges from the models, we estimated how many trophic, facultative and obligatory non-trophic links would be lost, and how this removal affected food-web properties (number of compartments, links, link density, and connectance). Astrophorina sponges were mostly linked via facultative non-trophic links to 60, 59, 86, and 92 compartments in the deep-sea coral, the lower slope 1 and 2, and the deep-sea sponge assemblages, respectively. Direct trophic links only existed to Echinasteridae and Pterasteridae. As removing Astrophorina sponges from the interaction webs of the different assemblages had the highest impact on food-web properties compared to removing any other fauna present, these sponges were considered “highest impact taxa”. They were also identified, along with sea pens, as “structural species”/ “habitat formers” and “foundation species” based on non-trophic interactions in the deep-sea coral and deep-sea sponge assemblages.

The European Union’s deep-sea fisheries regulations (Regulation (EU) No. 2016/2336) established obligations to manage deep-sea fisheries and to protect vulnerable marine ecosystems (VMEs). The European Commission is scheduled to complete a review of the regulations in 2021, providing an opportunity for new scientific information to be incorporated into the implementation of the regulations. Here, we summarise research outputs from the EU-funded Horizon 2020 ATLAS Project and explain their relevance to the regulation of deep-sea fisheries in EU waters. ATLAS research has increased our understanding of the distribution of VMEs and their importance in terms of ecosystem functioning. ATLAS research has also highlighted the utility of molecular techniques to understand fish population structure and the potential for habitat suitability models to help incorporate climate change into decision-making. Building on these scientific advances, we provide recommendations to help increase the effectiveness of management measures to conserve deep-sea fish stocks and protect VMEs.

Little is known about dispersal in deep-sea sponges, yet understanding patterns of gene flow and connectivity is essential for their effective management. Given rising pressure from harmful anthropogenic activities, schemes that manage resource extraction whilst conserving species diversity are increasingly necessary. Here, we used ddRADseq derived SNPs to investigate the genetic diversity and connectivity for the deep-sea sponge Phakellia ventilabrum across the northeast Atlantic Ocean (from the Cantabrian Sea to Norway). The analysis of 166 individuals collected from 57 sampling stations were grouped into 17 different areas, including two MPAs, one SAC and other areas with different levels of protection. The 4,017 neutral SNPs we uncovered indicated high connectivity and panmixis amongst the majority of areas, spanning a ca. 2,500-kilometre range and depths of 99–900 m. This was likely due to the presence of strong ocean currents aiding larval transport, as supported by our migration analysis and also by 3D particle tracking modelling using information on the reproductive cycle of P. ventilabrum. We also observed significant genetic similarity between samples from the Cantabrian Sea and Roscoff (France) as compared to the remainder of the collection areas, likely arising from physical drivers such as prevailing current circulation patterns and topographic features, acting as barriers for gene flow. Despite this, our results suggest that all protected areas studied are well connected with each other. The relatively low genetic diversity observed in all areas, though, highlights the potential fragility of this species to changing climates, which might compromise resilience to future threats.

Este artículo contiene 26 páginas, 2 tablas, 14 figuras., Biogeochemical cycling of silicon (Si), largely affected by biological drivers, is pivotal to the ecological functioning of the ocean. Most knowledge regarding biological utilization of Si derives from research on phototrophic organisms circumscribed to the photic ocean (i.e., diatoms). Utilization of Si in the aphotic ocean, where heterotrophic silicifiers become relevant Si users, remains poorly investigated. Here we quantify the flux rates and stocks characterizing Si cycling across dense aggregations of the hexactinellid sponge Vazella pourtalesii established in the aphotic zone of the central Scotian Shelf, Nova Scotia, Canada. Although individual rates of silicic acid consumption were low compared to other sponge species and diatoms, the large abundance of individuals (6.5 million) over the extension of these sponge grounds (2105 km2 ) leads to massive annual silicic acid consumption, invested in producing their siliceous skeletons of biogenic silica. This sponge activity accumulates large biogenic silica stocks both in the living population and in the sediments. Skeletal pieces in sediment revealed that a good portion of biogenic silica deposited to the bottom after sponge death recycles as silicic acid before being permanently buried. This biogenic silica–silicic acid turnover, facilitated by an unconventional silicification pattern that favors delamination and dissolution of V. pourtalesii spicules, causes silicic acid enrichment at oceanographic dimensions in the bottom water of the central Scotian Shelf. Silicic acid efflux from the bottom sustains a feedback mechanism that fulfills sponge needs for silicic acid and facilitates the persistence of sponge aggregations in the long term., This research was completed by funds from the SponGES H2020 grant (BG-01-2015.2. Agreement Number 679849-2), from Fisheries and Oceans Canada SPERA and IGS projects to EK, and from MINECO PBS project (CTM2015-67221-R) to MM.

AIM: To characterize the functional diversity and selected ecological functions of marine epibenthic invertebrate communities at the ecosystem scale and to evaluate the relative contributions of environmental filtering, including bottom‐contact fishing, and competitive interactions to benthic community assembly. LOCATION: Flemish Cap, an ecosystem production unit and fishing bank in the high seas of the north‐west Atlantic Ocean. METHODS: Through the use of Hierarchical Modelling of Species Communities (HMSC), we have explored seven community response traits to the environment applied to 105 epibenthic species and evaluated the influence of such traits on the community assembly processes. Assumed bioturbation, nutrient cycling and habitat provision functions, linked to individual or a combination of biological traits, were mapped using random forest modelling. RESULTS: Functional richness within benthic communities reached an asymptote for trawl sets with roughly more than 30 species. Assemblages on top of the Flemish Cap (

Effective conservation management of deep-sea sponges, including design of appropriate marine protected areas, requires an understanding of the connectivity between populations throughout a species’ distribution. We provide the first consideration of larval connectivity among deep-sea sponge populations along the southeastern coast of North America, illustrate the influence of the Gulf Stream on dispersal, and complement published distribution models by evaluating colonization potential. Connectivity among known populations of the hexactinellid sponge Vazella pourtalesii was simulated using a 3-D biophysical dispersal model throughout its distribution from Florida, United States to Nova Scotia, Canada. We found no exchange with an estimated pelagic larval duration of 2 weeks between populations north and south of Cape Hatteras, North Carolina at surface, mid-water and seabed release depths, irrespective of month of release or application of a horizontal diffusion constant specific to cross-Gulf Stream diffusivity. The population north of Cape Hatteras and south of Cape Cod was isolated. There was some evidence that Gulf Stream eddies formed near Cape Hatteras could travel to the northwest, connecting the populations in the two sub-regions, however that would require a much longer pelagic duration than what is currently known. More likely almost all larval settlement will be in the immediate area of the adults. At sub-regional scales, connectivity was found from the Strait of Florida through to the Blake Plateau, southeastern United States, with the latter area showing potential for recruitment from more than one source population. The influence of the Charleston Bump, a shallow feature rising from the Blake Plateau, was substantial. Particles seeded just north of the Bump were transported greater distances than those seeded to the south, some of which were caught in an associated gyre, promoting retention at the seabed. To the north on the Scotian Shelf, despite weaker currents and greater distances between known occurrences, unidirectional transport was detected from Emerald Basin to the Northeast Channel between Georges and Browns Banks. These major conclusions remained consistent through simulations run with different averaging periods for the currents (decades to daily) and using two ocean model products (BNAM and GLORYS12V1).

Este artículo contiene 13 páginas, 3 figuras, 4 tablas., Deep-sea sponge grounds are vulnerable marine ecosystems, which through their benthic-pelagic coupling of nutrients, are of functional relevance to the deep-sea realm. The impact of fishing bycatch is here evaluated for the first time at a bathyal, sponge-dominated ecosystem in the high seas managed by the Northwest Atlantic Fisheries Organization. Sponge biomass surfaces created from research survey data using both random forest modeling and a gridded surface revealed 231,140 t of sponges in the area. About 65% of that biomass was protected by current fisheries closures. However, projections of trawling tracks estimated that the sponge biomass within them would be wiped out in just 1 year by the current level of fishing activity if directed on the sponges. Because these sponges filter 56,143 ± 15,047 million litres of seawater daily, consume 63.11 ± 11.83 t of organic carbon through respiration, and affect the turnover of several nitrogen nutrients, their removal would likely affect the delicate ecological equilibrium of the deep-sea benthic ecosystem. We estimated that, on Flemish Cap, the economic value associated with seawater filtration by the sponges is nearly double the market value of the fish catch. Hence, fishery closures are essential to reach sponge conservation goals as economic drivers cannot be relied upon., This research has been performed within the scope of the SponGES project, which received funding from the European Union’s Horizon 2020 research and innovation programme under grant agreement No. 679849. This document reflects only the authors’ views and the Executive Agency for Small and Medium-sized Enterprises (EASME) is not responsible for any use that may be made of the information it contains. The EU groundfish surveys in the NAFO area were co-funded by the Spanish Institute of Oceanography (IEO), the Portuguese Institute for Sea and Atmosphere (IPMA), the Spanish Institute for Marine Research Superior Council of Scientific Investigations (IIM-CSIC) and the European Union through the European Maritime and Fisheries Fund (EMFF) within the National Program of collection, management and use of data in the fisheries sector and support for scientific advice regarding the Common Fisheries Policy. AC is supported by Program Investigador (IF/00029/2014/CP1230/CT0002) from FCT. This study had also the support of Fundação para a Ciência e Tecnologia (FCT), through the strategic projects UID/MAR/04292/2013 granted to MARE. Fisheries and Oceans, Canada provided support through their International Governance Strategy project funds (Science Advice in Support of the 2020 NAFO Review of Closed Areas to Protect Vulnerable Marine Ecosystems in the NRA) to EK.

The Northwest Atlantic Fisheries Organization (NAFO) Roadmap is the general framework aimed toward implementing an Ecosystem Approach to Fisheries (EAF) in NAFO. The core principles are: a) the approach has to be objective-driven, b) it should consider long-term ecosystem sustainability, c) it must be place-based, and d) the consequences of trade-offs in managing human activities have to be explicitly defined. The Roadmap is not a fixed plan; it evolves as different components are developed, refined, and implemented. Core elements of the Roadmap include the identification of ecosystem-based management units, a hierarchical approach to define exploitation rates by considering ecosystem, multispecies, and stock level sustainability, and the integration of impacts on benthic communities. The modular design of the plan has allowed NAFO to start implementing some components (e.g. closures to bottom fishing for the protection of Vulnerable Marine Ecosystems), while work on others is still ongoing (e.g. multispecies modelling). Even though the Roadmap is far from being fully implemented, NAFO has made important progress towards EAF over the last decade. This paper summarizes the Roadmap structure and the current level of implementation, describes some of the challenges faced, and examines those still ahead.

While the study of dispersal and connectivity in the ocean typically centres on pelagic species and planktonic larval stages of benthic species, the present work explores an overlooked locomotor means in post‐settlement benthic stages that redefines their dispersal potential.Members of the echinoderm class Holothuroidea colonize a diversity of marine environments world‐wide, where they play key ecological and economical roles, making their conservation a priority. Holothuroids are commonly called sea cucumbers or sea slugs to reflect their slow movements and are assumed to disperse chiefly through pelagic larvae.The present study documents and explores their unexpected ability to actively modify their buoyancy, leading them to tumble or float at speeds orders of magnitudes faster than through benthic crawling. Two focal species representing different taxonomic orders, geographic distributions and reproductive strategies were studied over several years.Active buoyancy adjustment (ABA) was achieved through a rapid increase in water‐to‐flesh ratio by up to 740%, leading to bloating, and simultaneously detachment from the substrate. It occurred as early as 6 months post settlement in juveniles and was recorded in wild adult populations. In experimental trials, ABA was triggered by high conspecific density, decreasing salinity and increasing water turbidity. Based on field video footage, ABA‐assisted movements generated speeds of up to 90 km/day.These findings imply that displacement during planktonic larval stages may not supersede the locomotor capacity of benthic stages, challenging the notion of sedentarity. Combining the present results and anecdotal reports, ABA emerges as a generalized means of dispersal among benthic animals, with critical implications for world‐wide management and conservation of commercially and ecologically significant species., This study documents the unexpected ability of so‐called sessile or sedentary benthic marine animals (e.g. sea cucumbers) to actively modify their buoyancy to tumble or float at speeds orders of magnitudes faster than anticipated. These findings redefine the locomotor capacity of benthic animals, challenging the notion of sedentarity, with critical implications for worldwide management and conservation of commercially and ecologically significant species.

A new Tedania species (Porifera) was collect using remotely operated vehicles during the Canadian mission HUD2010-029 and the British RRS Discovery Cruise DY081, on the Orphan Seamount near the Orphan Knoll, northwest Atlantic, between 2999.88 and 3450.4 m depth. Orphan Knoll is an isolated, drowned continental fragment 550 km northeast Newfoundland in the Labrador Sea. This region is biologically rich and complex and in 2007, the regional fisheries management organization operating in the area regulated that no vessel shall engage in bottom-contact fishing activities until reviewed in 2020 with a review slated at the end of this year. Members of the genus Tedania are uncommon in the temperate northern hemisphere with only six species known previously: Tedania (Tedania) anhelans; Tedania (Tedania) pilarriosae; Tedania (Tedania) suctoria; Tedania (Tedania) urgorrii; Tedania (Tedaniopsis) gurjanovae; and Tedania (Tedaniopsis) phacellina. The particular features of the new sponge we describe are the very peculiar external morphology which is tree-like with dichotomous branching—a morphology not previously described in this subgenus; and the combination of spicules found: long styles, the typical tornotes of the subgenus and two sizes of onychaetes. Additional information is provided on other species of Tedaniopsis described from the Atlantic Ocean. Based on the characteristics reported, we propose a new species, Tedania (Tedaniopsis) rappi sp. nov. in honor of Prof. Hans Tore Rapp (1972–2020), University of Bergen, Norway, a renowned sponge taxonomist and coordinator of the Horizon 2020 SponGES project. The holotype of T. (T.) phacellina Topsent, 1912 from the Azores, the only other northern Atlantic species in the subgenus Tedaniopsis, was reviewed for comparison., SPONGES, Deep-sea Sponge Grounds Ecosystems of the North Atlantic: an integrated approach towards their preservation and sustainable exploitation, SI

The molecular mechanisms underlying intraspecific variation in life history strategies are still poorly understood, despite the importance of this question for understanding of organism’s responses to environmental variability. Theoretical work proposed that epigenetic mechanisms such as DNA methylation might regulate intraspecific variation in life history strategies, however this assumption has rarely been verified empirically in wild populations. We examined associations between genome-wide methylation changes and environmentally-driven life history variation in two lineages of a marine fish that diverged approximatively 2.5 Mya, the capelin (Mallotus villosus), from North America and Europe. In both lineages, capelin harbour two contrasted life history strategies: some are strictly semelparous, experience fast actuarial senescence, but benefit from high hatching success by spawning on demersal sites where water temperature is low and relatively stable. In contrast, others are facultative iteroparous, have slower actuarial senescence, and suffer from lower hatching success by breeding in the intertidal zone where temperature is warmer, thermohaline parameters are less stable, and egg desiccation risk is high. Performing whole genome and epigenome sequencing, we showed that these contrasted life history strategies are more likely governed by epigenetic changes than by differences in DNA sequence. While genetic differentiation between the capelin harbouring different life history strategies was negligible, we detected parallel genome-wide methylation changes across lineages. We identified 1,067 differentially methylated regions (DMRs) comprising 15,818 CpGs, with 22% of them located within 5-kb around genes comprising promotor regions. We found that all DMRs were hypermethylated in demersal-spawning individuals. This striking result suggests that lower water temperature at demersal sites leads to an overall hypermethylation of the genome determined during the epigenetic reprogramming occurring over embryonic development. Our study emphasizes that parallel epigenetics changes in lineages with divergent genetic background could have a functional role in the regulation of intraspecific life history variation.

Shallow-water sponges are often cited as being ‘climate change winners’ due to their resiliency against climate change effects compared to other benthic taxa. However, little is known of the impacts of climate change on deep-water sponges. The deep-water glass sponge Vazella pourtalesii is distributed off eastern North America, forming dense sponge grounds with enhanced biodiversity on the Scotian Shelf off Nova Scotia, Canada. While the strong natural environmental variability that characterizes these sponge grounds suggests this species is resilient to a changing environment, its physiological limitations remain unknown, and the impact of more persistent anthropogenic climate change on its distribution has never been assessed. We used Random Forest and generalized additive models to project the distribution of V. pourtalesii in the northwest Atlantic using environmental conditions simulated under moderate and worst-case CO2 emission scenarios. Under future (2046-2065) climate change, the suitable habitat of V. pourtalesii will increase up to 4 times its present-day size and shift into deeper waters and higher latitudes, particularly in its northern range where ocean warming will serve to improve the habitat surrounding this originally sub-tropical species. However, not all areas projected as suitable habitat in the future will realistically be populated, and the reduced likelihood of occurrence in its core habitat on the Scotian Shelf suggests that the existing Vazella sponge grounds may be negatively impacted. An effective monitoring programme will require tracking changes in the density and distribution of V. pourtalesii at the margins between core habitat and where losses and gains were projected.

The Scotian Shelf harbors unique aggregations of the glass sponge Vazella pourtalesii that provides an important habitat for benthic and pelagic fauna. Recent studies have shown that these sponge grounds have persisted in the face of strong inter-annual and multi-decadal variability in temperature and salinity. However, little is known of these environmental characteristics on hourly-seasonal time scales. This study presents the first hydrodynamic observations and associated (food) particle supply mechanisms for the Vazella sponge grounds, highlighting the influence of natural variability in environmental conditions on sponge growth and resilience. Near-bottom environmental conditions were characterized by high temporal resolution data collected with a benthic lander, deployed during a period of 10 months in the Sambro Bank Sponge Conservation Area. The lander was equipped with temperature and oxygen sensors, a current meter, a sediment trap and a video camera. In addition, water column profiles of temperature and salinity were collected in an array across the sponge grounds from high to lower sponge presence probability. Over the course of the lander deployment, temperature fluctuated between 8.8–12°C with an average of 10.6 ± 0.4°C. Dissolved oxygen concentration was on average 6.3 mg l–1, and near-bottom current speed was on average 0.12 m s–1, with peaks up to 0.47 m s–1. Semi-diurnal tidal currents promoted constant resuspension of particulate matter in the benthic boundary layer. Surface storm events episodically caused extremely turbid conditions on the seafloor that persisted for several days, with particles being resuspended to more than 13 m above the seabed. The carbon flux in the near-bottom sediment trap peaked during storm events and also after a spring bloom in April, when fresh phytodetritus was observed in the bottom boundary layer. While resuspension events can represent a major stressor for sponges, limiting their filtration capability and remobilizing them, episodes of strong currents and lateral particle transport likely play an important role in food supply and the replenishment of nutrients and oxygen. Our results contextualize human-induced threats such as bottom fishing and climate change by providing more knowledge of the natural environmental conditions under which sponge grounds persist.

Este artículo contiene 24 páginas, 11 figuras, 2 tablas., Hexactinellid sponges are common in the deep sea, but their functional integration into those ecosystems remains poorly understood. The phylogenetically related species Schaudinnia rosea and Vazella pourtalesii were herein incubated for nitrogen and phosphorous, returning markedly different nutrient fluxes. Transmission electron microscopy (TEM) revealed S. rosea to host a low abundance of extracellular microbes, while Vazella pourtalesii showed higher microbial abundance and hosted most microbes within bacteriosyncytia, a novel feature for Hexactinellida. Amplicon sequences of the microbiome corroborated large between-species differences, also between the sponges and the seawater of their habitats. Metagenome-assembled genome of the V. pourtalesii microbiota revealed genes coding for enzymes operating in nitrification, denitrification, dissimilatory nitrate reduction to ammonium, nitrogen fixation, and ammonia/ammonium assimilation. In the nitrification and denitrification pathways some enzymes were missing, but alternative bridging routes allow the microbiota to close a N cycle in the holobiont. Interconnections between aerobic and anaerobic pathways may facilitate the sponges to withstand the low-oxygen conditions of deep-sea habitats. Importantly, various N pathways coupled to generate ammonium, which, through assimilation, fosters the growth of the sponge microbiota. TEM showed that the farmed microbiota is digested by the sponge cells, becoming an internal food source. This microbial farming demands more ammonium that can be provided internally by the host sponges and some 2.6 million kg of ammonium from the seawater become annually consumed by the aggregations of V. pourtalesii. Such ammonium removal is likely impairing the development of the free-living bacterioplankton and the survival chances of other sponge species that feed on bacterioplankton. Such nutritional competitive exclusion would favor the monospecific character of the V. pourtalesii aggregations. These aggregations also affect the surrounding environment through an annual release of 27.3 million kg of nitrite and, in smaller quantities, of nitrate and phosphate. The complex metabolic integration among the microbiota and the sponge suggests that the holobiont depends critically on the correct functioning of its N-driven microbial engine. The metabolic intertwining is so delicate that it changed after moving the sponges out of their habitat for a few days, a serious warning on the conservation needs of these sponge aggregations., This research was funded mostly by the SponGES H2020 grant (BG-01-2015.2, agreement number 679849-2) awarded to HR (coordinator) and the associated international partner consortium. Much of the logistics was also funded by the Fisheries and Oceans Canada International Governance Strategy (IGS) projects awarded to EK and HR. Manuscript elaboration benefited from funding by the Dark-Si grant (MICIU-PID2019- 108627RB-I00) to MM.

Few studies have thus far explored the microbiomes of glass sponges (Hexactinellida). The present study seeks to elucidate the composition of the microbiota associated with the glass sponge Vazella pourtalesii and the functional strategies of the main symbionts. We combined microscopic approaches with metagenome-guided microbial genome reconstruction and amplicon community profiling towards this goal. Microscopic imaging revealed that the host and microbial cells appeared within dense biomass patches that are presumably syncytial tissue aggregates. Based on abundances in amplicon libraries and metagenomic data, SAR324 bacteria, Crenarchaeota, Patescibacteria and Nanoarchaeota were identified as abundant members of the V. pourtalesii microbiome and their genomic potentials were thus analyzed in detail. A general pattern emerged in that the V. pourtalesii symbionts had very small genome sizes in the range of 0.5-2.2 Mb and low GC contents, even below those of seawater relatives. Based on functional analyses of metagenome-assembled genomes (MAGs), we propose two major microbial strategies: the “givers”, namely Crenarchaeota and SAR324, heterotrophs and facultative anaerobes, produce and partly secrete all required amino acids and vitamins. The “takers”, Nanoarchaeota and Patescibacteria, are anaerobes with reduced genomes that tap into the microbial community for resources, e.g., lipids and DNA, likely using pili-like structures. We posit that the existence of microbial cells in sponge syncytia together with the low-oxygen conditions in the seawater environment are factors that shape the unique compositional and functional properties of the microbial community associated with V. pourtalesii.ImportanceWe investigated the microbial community of V. pourtalesii that forms globally unique, monospecific sponge grounds under low-oxygen conditions on the Scotian Shelf, where it plays a key role for its vulnerable ecosystem. The microbial community was found to be concentrated within biomass patches and is dominated by small cells (

Establishment of adequate conservation areas represents a challenging but crucial task in the conservation of genetic diversity and biological variability. Anthropogenic pressures on marine ecosystems and organisms are steadily increasing. Whether and to what extent these pressures influence marine genetic biodiversity is only starting to be revealed. Using 16S rRNA gene amplicon sequencing, we analysed the microbial community structure of 33 individuals of the habitat-forming glass sponge Vazella pourtalesii, as well as reference seawater, sediment, and biofilm samples. We assessed how two anthropogenic impacts, i.e. habitat destruction by trawling and artificial substrate provision (moorings made of composite plastic), correspond with in situ V. pourtalesii microbiome variability. In addition, we evaluated the role of two bottom fishery closures in preserving sponge-associated microbial diversity on the Scotian Shelf, Canada. Our results illustrate that V. pourtalesii sponges collected from pristine sites within fishery closures contained distinct and taxonomically largely novel microbial communities. At the trawled site we recorded significant quantitative differences in distinct microbial phyla, such as a reduction in Nitrospinae in sponges and environmental references. Individuals of V. pourtalesii growing on the mooring were significantly enriched in Bacteroidetes, Verrucomicrobia and Cyanobacteria in comparison to sponge individuals growing on the natural seabed. Due to a concomitant enrichment of these taxa in the mooring biofilm, we propose that biofilms on artificial substrates may ‘prime’ sponge-associated microbial communities when small sponges settle on such substrates. These observations likely have relevant management implications when considering the increase of artificial substrates in the marine environment, e.g., marine litter, off-shore wind parks, and petroleum platforms.

Correction to:Scientific Reportshttps://doi.org/10.1038/s41598-019-52250-1, published online 01 Novenber 2019 The original version of this Article contained errors in the Abstract. “Sponge biomass surfaces created from research survey data using both random forest modeling and a gridded surface revealed 231,140 t of sponges in the area. About 65% of that biomass was protected by current fisheries closures.” now reads: “Sponge biomass surface created from research survey data using random forest modeling revealed 231,136 t of sponges in the area. About 42% of that biomass was protected by current fisheries closures.” This error has now been corrected in the PDF and HTML versions of the Article.

Midwater trawl surveys were conducted during 2007–10 at meso- and bathypelagic depths in and near The Gully, a large submarine canyon off Nova Scotia that is also a Marine Protected Area. The fish assemblage in the canyon was highly diverse but 20 species together comprised more than 90% of the catch by number and 80% by weight. The most numerous was the gonostomatid Cyclothone microdon while the myctophid Benthosema glaciale was next in number and first in weight. Most of those principal species would be expected in catches taken in oceanic waters beyond the shelf break. Only the bottom-spawning Melanostigma atlanticum was, within the surveyed area, distinctively a species of the canyon. Multivariate analyses showed that the primary variations in the assemblage were aligned with drivers that act in open ocean: depth, water mass and both diel and seasonal cycles. However, the effect of the canyon was evident in an up-canyon decline in the catches of most species. We hypothesize that the oceanic species are passively carried into The Gully by the known inflow and are there exposed to intense predation, depleting their numbers. We estimate that biomass flux as sufficient to support the Marine Protected Area's signature species: northern bottlenose whales (Hyperoodon ampullatus).

The deep sea is the largest biome on Earth but the least explored. Our knowledge of it comes from scattered sources spanning different spatial and temporal scales. Implementation of marine policies like the European Union’s Marine Strategy Framework Directive (MSFD) and support for Blue Growth in the deep sea are therefore hindered by lack of data. Integrated assessments of environmental status require tools to work with different and disaggregated datasets (e.g. density of deep-sea habitat-forming species, body-size distribution of commercial fishes, intensity of bottom trawling) across spatial and temporal scales. A feasibility study was conducted as part of the four-year ATLAS project to assess the effectiveness of the open-access Nested Environmental status Assessment Tool (NEAT) to assess deep-sea environmental status. We worked at nine selected study areas in the North Atlantic focusing on five MSFD descriptors (D1-Biodiversity, D3-Commercial fish and shellfish, D4-Food webs, D6-Seafloor integrity, D10-Marine litter). The objectives of the present study were to i) explore and propose indicators that could be used in the assessment of deep-sea environmental status, ii) evaluate the performance of NEAT in the deep sea, and iii) identify challenges and opportunities for the assessment of deep-sea status. Based on data availability, data quality and expert judgement, in total 24 indicators (one for D1, one for D3, seven for D4, 13 for D6, two for D10) were used in the assessment of the nine study areas, their habitats and ecosystem components. NEAT analyses revealed differences among the study areas for their environmental status ranging from “poor” to “high”. Overall, the NEAT results were in moderate to complete agreement with expert judgement, previous assessments, scientific literature on human-pressure gradients and expected management outcomes. We suggest that the assessment of deep-sea environmental status should take place at habitat and ecosystem level (rather than at species level) and at relatively large spatial scales, in comparison to shallow-water areas. Limited knowledge across space (e.g. distribution of habitat-forming species) and the scarcity of long-term data sets limit our knowledge about natural variability and human impacts in the deep sea preventing a more systematic assessment of habitat and ecosystem components in the deep sea. However, stronger cross-sectoral collaborations, the use of novel technologies and open data-sharing platforms will be critical for establishing environmental baseline indicator values in the deep sea that will contribute to the science base supporting the implementation of marine policies and stimulating Blue Growth., Sí

The deep sea plays a critical role in global climate regulation through uptake and storage of heat and carbon dioxide. However, this regulating service causes warming, acidification and deoxygenation of deep waters, leading to decreased food availability at the seafloor. These changes and their projections are likely to affect productivity, biodiversity and distributions of deep-sea fauna, thereby compromising key ecosystem services. Understanding how climate change can lead to shifts in deep-sea species distributions is critically important in developing management measures. We used environmental niche modelling along with the best available species occurrence data and environmental parameters to model habitat suitability for key cold-water coral and commercially important deep-sea fish species under present-day (1951–2000) environmental conditions and to project changes under severe, high emissions future (2081–2100) climate projections (RCP8.5 scenario) for the North Atlantic Ocean. Our models projected a decrease of 28%–100% in suitable habitat for cold-water corals and a shift in suitable habitat for deep-sea fishes of 2.0°–9.9° towards higher latitudes. The largest reductions in suitable habitat were projected for the scleractinian coral Lophelia pertusa and the octocoral Paragorgia arborea, with declines of at least 79% and 99% respectively. We projected the expansion of suitable habitat by 2100 only for the fishes Helicolenus dactylopterus and Sebastes mentella (20%–30%), mostly through northern latitudinal range expansion. Our results projected limited climate refugia locations in the North Atlantic by 2100 for scleractinian corals (30%–42% of present-day suitable habitat), even smaller refugia locations for the octocorals Acanella arbuscula and Acanthogorgia armata (6%–14%), and almost no refugia for P. arborea. Our results emphasize the need to understand how anticipated climate change will affect the distribution of deep-sea species including commercially important fishes and foundation species, and highlight the importance of identifying and preserving climate refugia for a range of area-based planning and management tools., ATLAS, A Trans-AtLantic Assessment and deep-water ecosystem-based Spatial management plan for Europe, SI

Aim: we examined the relationships between bathymetry, latitude and energy and the diversity of marine benthic invertebrates across wide environmental ranges of Canada's three oceans. Location: Canadian Pacific, Arctic and Atlantic Oceans from the intertidal zone to upper bathyal depths, encompassing 13 marine ecoregions. Methods: we compiled 35 benthic datasets that encompass 3,337 taxa (70% identified to species and 21% to genus) from 13,172 samples spanning 6,117 sites. Partitioning the analyses by different gear types, ecoregions or sites, we used Hill numbers to examine spatial patterns in alpha-diversity. We used resampling and extrapolation to standardized sampling effort and examined the effects of depth, latitude, chemical energy (export particulate organic carbon [POC] flux), thermal energy (bottom temperature) and seasonality of primary production on the benthic biodiversity. Results: the Canadian Arctic harboured the highest benthic diversity (e.g. epifauna and common and dominant infauna species), whereas the lowest diversity was found in the Atlantic. The Puget Trough (Pacific), Beaufort Sea, Arctic Archipelago, Hudson Bay, Northern Labrador and Southern Grand Bank (Atlantic) were the "hotspots" of diversity among the ecoregions. The infauna and epifauna both exhibited hump-shaped diversity-depth relationships, with peak diversity near shelf breaks; latitude (positively) predicted infaunal diversity, albeit weakly. Food supply, as inferred from primary production and depth, was more important than thermal energy in controlling diversity patterns. Limitations with respect to calculating POC flux in coastal (e.g. terrestrial runoff) and ice-covered regions or biological interactions may explain the negative POC flux-infaunal diversity relationship. Main conclusions: we show previously unreported diversity hotspots in the Canadian Arctic and in other ecoregions. Our analyses reveal potential controlling mechanisms of large-scale benthic biodiversity patterns in Canada's three oceans, which are inconsistent with the prevailing view of seafloor energy-diversity relationships. These results provide insightful information for conservation that can help to implement further MPA networks., Ministry of Science and Technology of Taiwan; Natural Sciences and Engineering Research Council of Canada (NSERC)

Circulation patterns in the North Atlantic Ocean have changed and re-organized multiple times over millions of years, influencing the biodiversity, distribution, and connectivity patterns of deep-sea species and ecosystems. In this study, we review the effects of the water mass properties (temperature, salinity, food supply, carbonate chemistry, and oxygen) on deep-sea benthic megafauna (from species to community level) and discussed in future scenarios of climate change. We focus on the key oceanic controls on deep-sea megafauna biodiversity and biogeography patterns. We place particular attention on cold-water corals and sponges, as these are ecosystem-engineering organisms that constitute vulnerable marine ecosystems (VME) with high associated biodiversity. Besides documenting the current state of the knowledge on this topic, a future scenario for water mass properties in the deep North Atlantic basin was predicted. The pace and severity of climate change in the deep-sea will vary across regions. However, predicted water mass properties showed that all regions in the North Atlantic will be exposed to multiple stressors by 2100, experiencing at least one critical change in water temperature (+2 ◦C), organic carbon fluxes (reduced up to 50%), ocean acidification (pH reduced up to 0.3), aragonite saturation horizon (shoaling above 1000 m) and/or reduction in dissolved oxygen (>5%). The northernmost regions of the North Atlantic will suffer the greatest impacts. Warmer and more acidic oceans will drastically reduce the suitable habitat for ecosystem-engineers, with severe consequences such as declines in population densities, even compromising their long-term survival, loss of biodiversity and reduced biogeographic distribution that might compromise connectivity at large scales. These effects can be aggravated by reductions in carbon fluxes, particularly in areas where food availability is already limited. Declines in benthic biomass and biodiversity will diminish ecosystem services such as habitat provision, nutrient cycling, etc. This study shows that the deep-sea VME affected by contemporary anthropogenic impacts and with the ongoing climate change impacts are unlikely to withstand additional pressures from more intrusive human activities. This study serves also as a warning to protect these ecosystems through regulations and by tempering the ongoing socio-political drivers for increasing exploitation of marine resources., Sí

Many of the marine policy frameworks developed to protect biodiversity in deep-sea areas, including areas beyond national jurisdiction (ABNJ), include indicators to assess policy objectives. These frameworks often have specific guidance on how the indicators should be applied and interpreted. Selection of indicators is an important process and those with strong scientific underpinnings are more likely to produce the expected outcomes. We reviewed three policy and assessment frameworks which include ABNJ regions or were developed specifically for ABNJ: (1) Oslo and Paris Convention for the Protection of the Marine Environment of the North-East Atlantic (OSPAR) ecosystem assessments, (2) the frameworks adopted to implement the UN General Assembly (UNGA) sustainable fisheries resolutions for the management of bottom fisheries to prevent Significant Adverse Impacts on vulnerable marine ecosystems, and (3) the Aichi Biodiversity Targets adopted by Parties to the Convention on Biological Diversity (CBD). We examined whether an assessment approach based on evaluation of Good Environmental Status (GES) under the European Union's Marine Strategy Framework Directive (MSFD), could be applied to ABNJ. We examined each MSFD descriptor for its applicability to deep-sea habitats considering the work of two European projects concluding that the MSFD could be applied to ABNJ to support OSPAR, UNGA and CBD policy objectives towards a common approach to the assessment of the status of deep-sea ecosystems in ABNJ. In achieving this we also introduce readers outside of Europe to the work conducted within the MSFD.