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Recent advances in technology have enabled an unprecedented development of underwater research, extending from near shore to the deepest regions of the globe. However, monitoring of biodiversity is not fully implemented in political agendas and biological observations in the deep ocean have been even more limited in space and time. The Foresight Workshop on Advances in Ocean Biological Observations: a sustained system for deep-ocean meroplankton was convened to to foster advances in the knowledge on deep-ocean invertebrate larval distributions and improve our understanding of fundamental deep-ocean ecological processes such as connectivity and resilience of benthic communities to natural and human-induced disturbance. This Meroplankton Observations Workshop had two specific goals: 1) review the state-of-the-art instrumentation available for meroplankton observations; 2) develop a strategy to implement technological innovations for in-situ meroplankton observation. Presentations and discussions are summarised in this report covering: i) key challenges and priorities for advancing the knowledge of deep-sea larval diversity and distribution: ii) recent developments in technology and future needs for plankton observation, iii) data integration and oceanographic modelling; iv) synergies and added value of a sustained observation system for meroplankton; v) steps for developing a sustained observation system for deep-ocean meroplankton and plans to maximise collaborative opportunities.

Understanding spatio-temporal variability of trace elements fingerprints (TEF) in bivalve shells is paramount to determine the discrimination power of this analytical approach and secure traceability along supply chains. Spatio-temporal variability of TEF was assessed in cockle (Cerastoderma edule) shells using inductively coupled plasma-mass spectrometry (ICP-MS). Four elemental ratios (Mg/Ca, Mn/Ca, Sr/Ca and Ba/Ca) were measured from the shells of specimens originating from eight different ecosystems along the Portuguese coast, as well as from four different areas, within one of them, over two consecutive years (2013 and 2014). TEF varied significantly in the shells of bivalves originating from the eight ecosystems surveyed in the present study. Linear discriminant function analyses assigned sampled cockles to each of the eight ecosystems with an average accuracy of 90%. Elemental ratios also displayed significant differences between the two consecutive years in the four areas monitored in the same ecosystem. Overall, while TEF displayed by cockle shells can be successfully used to trace their geographic origin, a periodical verification of TEF (>6 months and

Five chrysopetalid species are reported from samples collected at bathyal depths in three NE Atlantic regions: the Bay of Biscay, the Horseshoe Abyssal Plain and the Gulf of Cadiz. Arichlidon reyssi (Katzmann et al., 1974), Dysponetus caecus (Langerhans, 1880) and D. profundus Böggemann, 2009 are free-living forms found mainly on biogenic substrates (e.g., coral and sunken wood). A brief description and taxonomical remarks are given for each of these species and their geographical distributions and habitat records were updated accordingly. Natsushima bifurcata Miura & Laubier, 1990 and Craseoschema thyasiricola gen. et sp. nov. are symbionts inhabiting the mantle cavity of chemosynthesis-based bivalves known from four mud volcanoes from the Gulf of Cadiz. Craseoschema thyasiricola gen. et sp. nov. was found inside a thyasirid bivalve and presents mixed morphological characteristics of free-living and symbiotic forms within Calamyzinae Hartmann-Schröder, 1971. A full description of the new species is given together with DNA sequences of the genes COI, 16S and H3 that were used in a phylogenetic analysis to indicate the position of the new genus within the family, Financial support was provided by CESAM (UID/AMB/50017/2019) and by FCT/MCTES (Fundação para a Ciência e a Tecnologia /Ministério de Ciência, Tecnologia e Ensino Superior) through national funds. AR was supported by a postdoctoral grant (SFRH/BPD/112408/2015) from FCT co-financed by ESF (European Science Foundation) and MEC (Ministério da Fundação e Ciência). CR and LG were supported by postdoctoral grants, SFRH/BPD/107805/2015 and SFRH/ BPD/96142/2013, respectively. This study is a contribution of the “Macroevolutionary Transitions in Syllidae” Project (CGL2015-63593-P), supported by MINECO/FEDER (Ministério de Economia y Competitividad/Fondo Europeo de Desarrollo Regional) and European Union funds (PI: MTA)

Bottom-trawling fisheries operating in Portugal (West Iberian Margin) impose one of the largest footprints per unit of biomass landed in European waters at depths greater than 200 m, affecting the seafloor integrity and the associated benthic fauna. To investigate how trawling pressure is affecting the macrofaunal assemblages, we compared the standing stock (abundance and biomass), community structure and taxonomical and trophic diversity in areas subjected to varying trawling pressure along the SW Portuguese upper slope, between 200 and 600 m. In addition to trawling pressure, several environmental variables, namely depth, grain size and organic matter, were correlated with the biological component, which suggest that the longstanding trawling pressure presents cumulative effects to the habitat heterogeneity known to characterise the West Iberian Margin fauna. Furthermore, our results showed a depletion of macro-infaunal abundances in both the fishing ground and the adjacent area (up to 3 times lower), when compared to the area not trawled. The observed decrease in abundance with increasing trawling pressure was also associated with a loss of species and trophic richness, but univariate diversity indices related with community structure (i.e. Shannon-Wiener index, Pielou's evenness) failed to detect consistent differences across areas. Also observed was a decrease in the number of taxa – trophic guilds combinations of the core assemblage (i.e. characteristic, dominant or frequent taxa) with increasing trawling pressure. We suggest that, in disturbed sediments, the lower functional redundancy resulting from the loss of species within most feeding guilds increases the vulnerability of trophic interactions and therefore of the whole assemblage to further increases in natural and anthropogenic disturbance or their synergistic effects.

Upper Miocene heterozoan carbonates crop out extensively in a NE–SW-trending belt (42 km long and 1.5–8 km wide) along the so-called El Alcor topographic high, from Carmona to Dos Hermanas (Seville, S Spain). These carbonates formed at the southern active margin of the Guadalquivir Basin, the foreland basin of the Betic Cordillera. They change to marls basinward (NE) and to sands landward (SE and SW). Therefore, carbonate production was constrained to a limited area in an otherwise siliciclastic shelf. The carbonates (up to 40 m thick) overlie a gradually coarsening-upward succession of marls followed by silts and sandstones. The carbonate sequence can be divided into three subunits corresponding, from bottom to top, to lowstand, transgressive, and highstand system tract deposits. The lower subunit, exhibiting extensive trough cross-bedding, is interpreted as a shallow-water bar deposit. The intermediate subunit onlaps underlying sediments and was deposited in deeper, low-turbulence conditions. The upper subunit deposits accumulated in a well-oxygenated outer platform based on benthic foraminiferal assemblages. The presence of hummocky and swaley cross-stratification in these latter deposits suggests that they were affected by storms. Pervasive fluid-escape structures are also observed throughout the carbonates. The three subunits consist of bioclastic packstones to rudstones made up of abundant fragments of small mytilids. Isotopic data from serpulid polychaete Ditrupa tubes show 13C-depleted values (up to − 16.1‰), whereas δ18O yields normal marine values. Additional isotopic data on shells of scallops, oysters, and small mussels, as well as bulk sediment, show diagenetic alterations. Based on actualistic examples of massive concentrations of mussels, the nearly monospecific composition of the El Alcor deposits, together with negative δ13C values of Ditrupa tubes, indicates that cold seeps presumably promoted carbonate formation. However, the absence of typical features of cold-seep deposits, such as authigenic carbonates mediated by anaerobic bacterial activity and the typical chemosynthetic shelly organisms, makes the large carbonate body of El Alcor an unusual cold-seep deposit.

The Gulf of Cadiz is an extensive seepage area in the south Iberian margin (NE Atlantic) encompassing over 40 mud volcanoes (MVs) at depths ranging from 200 to 4000 m. The area has a long geologic history and a central biogeographic location with a complex circulation ensuring oceanographic connectivity with the Mediterranean Sea, equatorial and North Atlantic regions. The geodynamics of the region promotes a notorious diversity in the seep regime despite the relatively low fluxes of hydrocarbon-rich gases. We analyse quantitative samples taken during the cruises TTR14, TTR15 and MSM01-03 in seven mud volcanoes grouped into Shallow MVs (Mercator: 350 m, Kidd: 500 m, Meknès: 700 m) and Deep MVs (Captain Arutyunov: 1300 m, Carlos Ribeiro: 2200 m, Bonjardim: 3000 m, Porto: 3900 m) and two additional Reference sites (ca. 550 m). Macrofauna (retained by a 500 μm sieve) was identified to species level whenever possible. The samples yielded modest abundances (70–1567 individuals per 0.25 m2), but the local and regional number of species is among the highest ever reported for cold seeps. Among the 366 recorded species, 22 were symbiont-hosting bivalves (Thyasiridae, Vesicomyidae, Solemyidae) and tubeworms (Siboglinidae). The multivariate analyses supported the significant differences between Shallow and Deep MVs: The environmental conditions at the Shallow MVs make them highly permeable to the penetration of background fauna leading to high diversity of the attendant assemblages (H′: 2.92–3.94; ES(100): 28.3–45.0; J′: 0.685–0.881). The Deep MV assemblages showed, in general, contrasting features but were more heterogeneous (H′: 1.41–3.06; ES(100): 10.5–30.5; J′: 0.340–0.852) and often dominated by one or more siboglinid species. The rarefaction curves confirmed the differences in biodiversity of Deep and Shallow MVs as well as the convergence of the latter to the Reference sites. The Bray–Curtis dissimilarity demonstrated the high β-diversity of the assemblages, especially in pairwise comparisons involving samples from the Deep MVs. Diversity partitioning assessed for species richness, Hurlbert's expected number of species and Shannon–Wiener index confirmed the high β-diversity across different spatial scales (within MVs, between MVs, between Deep and Shallow MVs). We suggest that historical and contemporary factors with differential synergies at different depths contribute to the high α-, β- and γ-diversity of the mud volcano faunal assemblages in the Gulf of Cadiz.

The industrialization of the deep sea is expanding world wide. Increasing oil and gas exploration activities in the absence of sufficient baseline data in deep-sea ecosystems has made environmental management challenging. Here, we review the types of activities that are associated with global offshore oil and gas development in water depths over 200 m, the typical impacts of these activities, some of the more extreme impacts of accidental oil and gas releases, and the current state of management in the major regions of offshore industrial activity including 18 exclusive economic zones. Direct impacts of infrastructure installation, including sediment resuspension and burial by seafloor anchors and pipelines, are typically restricted to a radius of ∼100 m on from the installation on the seafloor. Discharges of water-based and low toxicity oil based drilling muds and produced water can extend over 2 km, while the ecological impacts at the population and community levels on the sea floor are most commonly on the order of 200–300 m from their source. These impacts may persist in the deep sea for many years and likely longer for its more fragile ecosystems, such as cold-water corals. This synthesis of information provides the basis for a series of recommendations for the management of offshore oil and gas development. An effective management strategy, aimed at minimizing risk of significant environmental harm, will typically encompass regulations of the activity itself (e.g., discharge practices, materials used), combined with spatial (e.g., avoidance rules and marine protected areas), and temporal measures (e.g., restricted activities during peak reproductive periods). Spatial management measures that encompass representatives of all of the regional deep-sea community types is important in this context. Implementation of these management strategies should consider minimum buffer zones to displace industrial activity beyond the range of typical impacts: at least 2 km from any discharge points and surface infrastructure and 200 m from seafloor infrastructure with no expected discharges. Although managing natural resources is, arguably, more challenging in deep-water environments, inclusion of these proven conservation tools contributes to robust environmental management strategies for oil and gas extraction in the deep sea.

Determining seafood geographic origin is critical for controlling its quality and safeguarding the interest of consumers. Here, we use trace element fingerprinting (TEF) of bivalve shells to discriminate the geographic origin of specimens. Barium (Ba), manganese (Mn), magnesium (Mg), strontium (Sr) and lead (Pb) were quantified in cockle shells (Cerastoderma edule) captured with two fishing methods (by hand and by hand-raking) and from five adjacent fishing locations within an estuarine system (Ria de Aveiro, Portugal). Results suggest no differences in TEF of cockle shells captured by hand or by hand-raking, thus confirming that metal rakes do not act as a potential source of metal contamination that could somehow bias TEF results. In contrast, significant differences were recorded among locations for all trace elements analysed. A Canonical Analysis of Principal Coordinates (CAP) revealed that 92% of the samples could be successfully classified according to their fishing location using TEF. We show that TEF can be an accurate, fast and reliable method to determine the geographic origin of bivalves, even among locations separated less than 1 km apart within the same estuarine system. Nonetheless, follow up studies are needed to determine if TEF can reliably discriminate between bivalves originating from different ecosystems.

Five mammal carcasses were experimentally deployed at 1000 m depth in the Setubal Canyon (NE Atlantic) in March 2011 and the remaining bones were collected after 18 and 28 months. High numbers (1.92–6.27 individuals·cm2) of small mussels were found among a diverse invertebrate assemblage colonizing surfaces, crevices and cavities in the trabecular matrix of bones. In this study we characterized the mussel population and their associated bacteria and investigated population structure and settlement patterns. The results of molecular analysis indicated that the mussels are conspecific with the widely distributed bathymodiolin species ‘Idas’ simpsoni and harbor sulfur-oxidizing bacterial symbionts. Modal decomposition of length-frequency distributions and estimated shell growth rate suggested that settlement of ‘I.’ simpsoni is continuous and that post-settlement mortality affects the population structure. This study reports the highest densities and fastest growth rates within bathymodiolin mussels occurring on organic substrates yet recorded and provides evidence for effective mussel recruitment in organic falls in the NE Atlantic Ocean. These results highlight the importance of ephemeral habitats on the biogeographic distribution and dispersal of chemosymbiotic species in deep-sea ecosystems.

Sunken whale carcasses, known as ‘whale falls’, deliver large pulses of organic material to the seafloor and serve as habitat islands for unique assemblages of deep-sea fauna that include generalist scavenging species, chemosynthetic fauna related to those from hydrothermal vents and cold seeps, and bone-specialist species. Despite the growing interest in this peculiar habitat in the last decade, all long-term studies of deep-sea whale-falls have been restricted to the Pacific and the ecology of these habitats in the Atlantic Ocean has been overlooked. Here we report the first study on large mammal carcasses in the deep Atlantic Ocean. Cow carcasses amounting to 570 kg were deployed at 1000 m in the Setubal canyon, where cetaceans naturally occur. Eighteen months after deployment, all soft tissues had been consumed and the visible remains of the carcasses consisted solely of skeletal material that supported a diverse macrofaunal assemblage distinct from that of the background, including organic-enrichment respondents, sulphophiles and bone-specialists. The trophic structure of this assemblage denotes an overlap of successional stages previously described in whale falls. Our results provide important insights into the contribution of mammal falls to deep-sea biodiversity in the NE Atlantic, species distribution and biogeography of bone specialists, and the potential role of these habitats as dispersal stepping-stones for vent and seep species. Additionally we show that cow carcasses may serve as an accessible model system for the study of ecological processes at deep-sea whale-falls. published

Among chemosymbiotic metazoans found at deep-sea hydrothermal vents, cold seeps and organic falls, members of the mussel clade Bathymodiolinae (Bivalvia: Mytilidae) have evolved interactions with a higher diversity of bacterial lineages than other bivalve groups. Here, we characterized the bacteria associated with "Bathymodiolus" mauritanicus and Idas-like specimens from three sites in the Northeast Atlantic (two mud volcanoes in the Gulf of Cadiz and one seamount of the Gorringe Bank). Phylogenetic analysis of bacterial 16S rRNA-encoding gene sequences demonstrated that "B". mauritanicus has a dual symbiosis dominated by two phylotypes of methane-oxidising bacteria and a less abundant phylotype of a sulphur-oxidising bacterium. The latter was the dominant phylotype in a sympatric population of Idas-like mussels at the Darwin mud volcano. These results are the first report of a bacterial phylotype shared between two deep-sea mussels from divergent clades. This sulphur-oxidising bacterium was absent from Idas-like specimens from the other two sites (Gorringe Bank and Meknès mud volcano), in which bacterial clone libraries were dominated by other Gammaproteobacteria related to symbionts previously identified in Idas modiolaeformis from the Eastern Mediterranean. All Idas-like specimens studied herein are closely related and also related to I. modiolaeformis. However, they probably display different associations with bacteria, with the possible absence of both methane- and sulphur-oxidising symbionts at the Gorringe Bank. These results draw a very complex picture of associations between mussels and bacteria in the Northeast Atlantic, which could be highly variable depending on locale characteristics of the habitats.

Bridging the Atlantic and Mediterranean continental margins, the South Iberian region has recently been the focus for geological and biological investigations. In this region, the Gulf of Cadiz (GoC) encompasses a great variety of deep-sea habitats that harbour highly diverse biological communities. In this study, we describe the composition of gastropod assemblages obtained from in situ colonization experiments and benthic sampling of deep-sea habitats in the GoC. Gastropod distributional patterns, such as bathymetric ranges, bathymetric turnover, affinity to substrate types and abundance-occupancy relationships, are analysed and interpreted in relation to their inferred dispersal capabilities and substrate availability. Overall, the GoC comprises a high diversity of gastropods (65 species), and distinct assemblages were found in typical sedimentary environments at mud volcanoes and in association with carbonate and coral samples or organic substrata. The number of taxa peaked at the Carbonate Province in the middle slope (600–1200 m depth), a highly heterogeneous area with numerous mud volcanoes, carbonate mounds and corals. Darwin (1100 m) and Captain Arutyunov (1300 m) mud volcanoes harboured the most species-rich and abundant gastropod assemblages, respectively. Colonization experiments with organic substrata (wood and alfalfa grass) also yielded diverse and abundant gastropod assemblages. These organic inputs allowed the recruitment of local species but mainly of wood specialist taxa that were not previously known from the GoC. Our results suggest that the distribution of gastropod assemblages may be primarily determined by the occurrence of suitable habitats, probably due to the effect of the substrate type on the structural complexity of the habitat and availability and diversity of adequate food sources. The type of larval development is apparently not a limiting factor for colonization of deep-sea habitats. However, the predominance of non-planktotrophy, and especially lecithotrophy, suggests that a trade-off between more limited dispersal capability and higher potential for self-recruitment may be a recurrent pattern in gastropod species inhabiting reducing environments and other patchily distributed deep-sea habitats. A network of suitable habitats that ensures effective population connectivity would explain the predominance and relatively wide distribution of short-distance dispersing non-planktotrophic species in the GoC deep-sea habitats and other geographical regions.

The increasing number of bathymodiolin mussel species being described from deep-sea chemosynthetic environments worldwide has raised many questions about their evolutionary history, and their systematics is still being debated. Mussels are also abundant in fossil chemosynthetic assemblages, but their identification is problematic due to conservative shell morphology within the group and preservation issues. Potential resolution of bathymodiolin taxonomy requires new character sets, including morphological features that are likely to be preserved in fossil specimens. To investigate the phylogenetic significance of shell microstructural features, we studied the shell microstructure and mineralogy of 10 mussel species from hydrothermal vents and hydrocarbon seeps, and 15 taxa from sunken wood and bone habitats, and compared these observations with current molecular phylogenies of the sub-family Bathymodiolinae. In addition, we analyzed the shell microstructure in Adipicola chickubetsuensis from fossil whale carcasses, and in Bathymodiolus cf. willapaensis and "Modiola exbrocchii" from fossil cold seeps, and discussed the usefulness of these characters for identification of fossil chemosymbiotic mussels. Microstructural shell features are quite uniform among vent, seep, wood and bone mussel taxa, and therefore established bathymodiolin lineages cannot be discriminated, nor can the relations between fossil and modern species be determined with these characters. Nevertheless, the uniformity of shell microstructures observed among chemosymbiotic mussels and the similarity with its closest relative, Modiolus modiolus, does not challenge the monophyly of the group. Slight differences are found between the large vent and seep mussels and the small mytilids commonly found in habitats enriched in organic matter. Together with previous data, these results indicate that a repeated pattern of paedomorphism characterizes the evolutionary history of deep-sea mussels, and the occurrence of neotenous features should be considered in the taxonomic revision of this group. (C) 2012 Elsevier Ltd. All rights reserved.

The "Bathymodiolus" childressi group is the most geographically diverse assemblage of deep-sea mussel species. In this paper we consider several possible hypotheses to explain the present biogeographic distribution of the "B." childressi species complex. Mussels were collected for the first time from mud volcanoes in the Gulf of Cadiz (NE Atlantic Ocean) during the training through research (TTR) 16 research expedition in 2006. Preliminary observations of the shell features indicate that they belong to the "B." childressi species complex, which has been recognized as morphologically and genetically distinct from other Bathymodiolus species. Molecular analyses of two mitochondrial genes (COI-5 and ND4) were used to characterize the new mussel population from the Gulf of Cadiz (GOC) and to determine their phylogenetic relationships with other members of the "B." childressi group. The results indicate that the GOC mussels are conspecific with "Bathymodiolus" mauritanicus Cosel (2002), described from West Africa margin, and support a previous hypothesis that "B." mauritanicus is an amphi-Atlantic species. FCT-SFRH/BD/23231/2005 US National Science Foundation (OCE0241613) EC - GOCE-CT-2005-511234