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A 2-Dimensional mathematical reaction-transport model was developed to study the impact of the mud-dwelling frenulate tubeworm Siboglinum sp. on the biogeochemistry of a~sediment (MUC15) at the Captain Arutyunov mud volcano (CAMV). By explicitly describing the worm in its surrounding sediment, we are able to make budgets of processes occurring in- or outside of the worm, and to quantify how different worm densities and biomasses affect the anaerobic oxidation of methane (AOM) and sulfide reoxidation (HSox). The model shows that, at the observed densities, the presence of a thin worm body is sufficient to keep the upper 10 cm of sediment well homogenised with respect to dissolved substances, in agreement with observations. By this "bio-ventilation" activity, the worm pushes the sulfate–methane transition (SMT) zone downward to the posterior end of its body, and simultaneously physically separates the sulfide produced during the anaerobic oxidation of methane from oxygen. While there is little scope for AOM to take place in the tubeworm's body, 70% of the sulfide that is produced by sulfate reduction processes or that is advected in the sediment is preferentially shunted via the organism where it is oxidised by endosymbionts providing the energy for the worm's growth. The process of sulfide reoxidation, occurring predominantly in the worm's body is thus very distinct from the anaerobic oxidation of methane, which is a diffuse process that takes place in the sediments in the methane-sulfate transition zone. We show how the sulfide oxidation process is affected by increasing densities and length of the frenulates, and by upward advection velocity. Our biogeochemical model is one of the first to describe tubeworms explicitly. It can be used to directly link biological and biogeochemical observations at seep sites, and to study the impacts of mud-dwelling frenulates on the sediment biogeochemistry under varying environmental conditions. Also, it provides a tool to explore the competition between bacteria and fauna for available energy resources.

Submarine canyons have often been identified as hotspots of secondary production with the potential to house distinct faunal assemblages and idiosyncratic ecosystems. Within these deep-sea habitats, assemblages of scavenging fauna play a vital role in reintroducing organic matter from large food falls into the wider deep-sea food chain. Free-fall baited traps were set at different depths within three submarine canyons on the Iberian Margin. Amphipods from the traps were identified to species level and counted. Scavenging amphipod assemblages were compared at different depths within each canyon and between individual canyon systems. Using data from literature, abyssal plain assemblages were compared to submarine canyon assemblages. Samples from canyons were found to contain common abyssal plain species but in greater than expected abundances. It is proposed that this is a result of the high organic carbon input into canyon systems owing to their interception of sediment from the continental shelf and input from associated estuarine systems. Community composition differed significantly between the submarine canyons and abyssal plains. The cause of this difference cannot be attributed to one environmental variable due to the numerous inherent differences between canyons and abyssal plains.

During the past two decades, European cold seep ecosystems have attracted the scientific interest and to date there are several studies which have investigated the community structure and biodiversity of individual sites. In order to gain a better insight into the biology, biodiversity, and biogeography of seep-associated microbial communities along Europe's continental margins, a comparative approach was applied in the present work. By exploiting the publicly available data on 16S rRNA gene sequences retrieved from sediments of the Håkon Mosby mud volcano, Gulf of Cádiz and the eastern Mediterranean mud volcanoes/pockmarks (Anaximander area and Nile Fan), we investigated the prokaryotic biological components connecting these geographically isolated systems. The construction of interaction networks for both archaeal and bacterial shared operational taxonomic units (OTUs) among the different sites, revealed the presence of persistent OTUs, which can be considered as "key-players". One archaeal OTU (HQ588641) belonging to the ANME-3 group and one δ-Proteobacteria (HQ588562) were found in all five investigated areas. Other Archaea OTUs shared between four sites or less, belonged to the ANME-2c, -2a, MBG-D, -B and Thaumarchaeota. All other shared Bacteria belonged to the δ- and γ-Proteobacteria, with the exception of one JS1 affiliate OTU. The distribution of the majority of the shared OTUs seems to be restricted in cold seeps, mud volcanoes and other marine methane-rich environments. Although the investigated sites were connected through a small number of OTUs, these microorganisms hold central ecophysiological roles in these sediments, namely methane- and sulfur-mediated mineralization.

We investigated how interactions between hydrography, topography and species ecology influence the assembly of species and functional traits across multiple spatial scales of a cold-water coral reef seascape. In a novel approach for these ecosystems, we used a spatially resolved complex three-dimensional flow model of hydrography to help explain assembly patterns. Forward-selection of distance-based Moran's eigenvector mapping (dbMEM) variables identified two submodels of spatial scales at which communities change: broad-scale (across reef) and fine-scale (within reef). Variance partitioning identified bathymetric and hydrographic gradients important in creating broad-scale assembly of species and traits. In contrast, fine-scale assembly was related more to processes that created spatially autocorrelated patches of fauna, such as philopatric recruitment in sessile fauna, and social interactions and food supply in scavenging detritivores and mobile predators. Our study shows how habitat modification of reef connectivity and hydrography by bottom fishing and renewable energy installations could alter the structure and function of an entire cold-water coral reef seascape.

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.

Benthic foraminifera are an important component of the marine biota, but protocols for investigating their viability and metabolism are still extremely limited. Classical studies on benthic foraminifera have been based on direct counting under light microscopy. Typically, these organisms are stained with Rose Bengal, which binds proteins and other macromolecules, but does not allow discrimination between viable and recently dead organisms. The fluorescent in situ hybridization technique (FISH) represents a new and useful approach to identify living cells possessing an active metabolism. Our work is the first test of the suitability of the FISH technique, based on fluorescent probes targeting the 18S rRNA, to detect live benthic foraminifera. The protocol was applied on Ammonia group and Miliolids, as well as on agglutinated polythalamous (i.e., Leptohalysis scottii and Eggerella scabra) and soft-shelled monothalamous (i.e., Psammophaga sp. and saccamminid morphotypes) taxa. The results from FISH analyses were compared with those obtained, on the same specimens assayed with FISH, from microscopic analysis of the cytoplasm colour, presence of pigments and pseudopodial activity. Our results indicate that FISH targets only metabolically active foraminifera, and allows discerning from low to high cellular activity, validating the hypothesis that the intensity of the fluorescent signal emitted by the probe is dependent upon the physiological status of cells. These findings support the usefulness of this molecular approach as a key tool for obtaining information on the physiology of living foraminifera, both in field and experimental settings.

Interest in seamount research has gathered momentum over the past five years in an effort to understand the physical, geochemical and biological characteristics as well as the interconnectedness of seamount ecosystems. The majority of biological seamount research has concentrated upon the rich and diverse suspension feeding organisms that dominate the megafauna, such as gorgonians and antipatharian corals; by comparison there have been few studies that have investigated the no less enigmatic, but possibly just as important infauna. To help fill this knowledge gap, the macrofaunal community was sampled from a total of five stations along a northerly transect (capturing water depths from ∼130 m to ∼3300 m), on Senghor Seamount (NE Atlantic). The focus of this study is on the polychaete communities. Polychaete abundance peaked at the summit and a mid-slope station (∼1500 m), a pattern mirrored by the biomass values. The polychaete community along the transect appeared to be particularly diverse, with 135 species nominally identified to putative species from a total of 954 individuals. A diversity maximum was identified on the upper slope at ∼800 m depth, with species diversity, richness and evenness also all peaking at this station. Depth is likely to be a significant factor in determining levels of similarity between stations.

Submarine canyons are sites of intense energy and material exchange between the shelf and the deep adjacent basins. To test the hypothesis that active submarine canyons represent preferential conduits of available food for the deep-sea benthos, two mooring lines were deployed at 1200 m depth from November 2008 to November 2009 inside the Blanes canyon and on the adjacent open slope (Catalan Margin, NW Mediterranean Sea). We investigated the fluxes, biochemical composition and food quality of sinking organic carbon (OC). OC fluxes in the canyon and the open slope varied among sampling periods, though not consistently in the two sites. In particular, while in the open slope the highest OC fluxes were observed in August 2009, in the canyon the highest OC fluxes occurred in April–May 2009. For almost the entire study period, the OC fluxes in the canyon were significantly higher than those in the open slope, whereas OC contents of sinking particles collected in the open slope were consistently higher than those in the canyon. This result confirms that submarine canyons are effective conveyors of OC to the deep sea. Particles transferred to the deep sea floor through the canyons are predominantly of inorganic origin, significantly higher than that reaching the open slope at a similar water depth. Using multivariate statistical tests, two major clusters of sampling periods were identified: one in the canyon that grouped trap samples collected in December 2008, concurrently with the occurrence of a major storm at the sea surface, and associated with increased fluxes of nutritionally available particles from the upper shelf. Another cluster grouped samples from both the canyon and the open slope collected in March 2009, concurrently with the occurrence of the seasonal phytoplankton bloom at the sea surface, and associated with increased fluxes of total phytopigments. Our results confirm the key ecological role of submarine canyons for the functioning of deep-sea ecosystems, and highlight the importance of canyons in linking episodic storms and primary production occurring at the sea surface to the deep sea floor.

The Amon mud volcano (MV), located at 1250 m water depth on the Nile deep-sea fan, is known for its active emission of methane and non-methane hydrocarbons into the hydrosphere. Previous investigations showed a low efficiency of hydrocarbon-degrading anaerobic microbial communities inhabiting the Amon MV center in the presence of sulfate and hydrocarbons in the seeping subsurface fluids. By comparing spatial and temporal patterns of in situ biogeochemical fluxes, temperature gradients, pore water composition, and microbial activities over 3 yr, we investigated why the activity of anaerobic hydrocarbon degraders can be low despite high energy supplies. We found that the central dome of the Amon MV, as well as a lateral mud flow at its base, showed signs of recent exposure of hot subsurface muds lacking active hydrocarbon degrading communities. In these highly disturbed areas, anaerobic degradation of methane was less than 2% of the methane flux. Rather high oxygen consumption rates compared to low sulfide production suggest a faster development of more rapidly growing aerobic hydrocarbon degraders in highly disturbed areas. In contrast, the more stabilized muds surrounding the central gas and fluid conduits hosted active anaerobic hydrocarbon-degrading microbial communities. The low microbial activity in the hydrocarbon-vented areas of Amon MV is thus a consequence of kinetic limitations by heat and mud expulsion, whereas most of the outer MV area is limited by hydrocarbon transport.

The geo-biological exploration of a pockmark field located at ca. 800 m below sea level in the Gela basin (Strait of Sicily, Central Mediterranean) provided a relatively diverse chemosymbiotic community and methane-imprinted carbonates. To date, this is the first occurrence of such a type of specialised deep-water cold-seep communities recorded from this key region, before documented in the Mediterranean as rather disjunct findings in its eastern and westernmost basins. The thiotrophic chemosymbiotic organisms recovered from this area include empty tubes of the vestimentiferan Lamellibrachia sp., loose and articulated shells of lucinids (Lucinoma kazani, Myrtea amorpha), vesicomyids (Isorropodon perplexum), and gastropods (Taranis moerchii). A callianassid decapod (Calliax sp.) was consistently found alive in large numbers in the pockmark mud. Their post-mortem calcified parts mixed with molluscs and subordinately miliolid foraminifers form a distinct type of skeletal assemblage. Carbonate concretions display δ13C values as low as −40‰ PDB suggesting the occurrence of light hydrocarbons in the seeping fluids. Since none of the truly chemosymbiotic organisms was found alive, although their skeletal parts appear at times very fresh, some specimens have been AMS-14C dated to shed light on the historical evolution of this site. Lamellibrachiav and Lucinoma are two of the most significant chemosymbiotic taxa reported from various Mediterranean cold seep sites (Alboran Sea and Eastern basin). Specimens from station MEDCOR78 (pockmark #1, Lat. 36°46´10.18" N, Long. 14°01´31.59" E, 815 m below sea level) provided ages of 11736 ± 636 yr cal BP (Lamellibrachia sp.), and 9609.5 ± 153.5 yr cal BP (L. kazani). One shell of M. amorpha in core MEDCOR81 (pockmark #6, Lat 36°45´38.89" N, Long 14°00´07.58" E, 822 m below sea level) provided a sub-modern age of 484 ± 54 yr cal BP. These ages document that fluid seepage at this pockmark site has been episodically sustaining thiotrophic macrobenthic communities since the end of the Younger Dryas stadial up to sub-recent times.