Spatial scale and habitat-dependent diversity patterns in nematode communities in three seepage related sites along the Norwegian Sea margin

Assessing the relative contribution of local diversity to regional biodiversity may be the key to understanding large-scale and even global patterns in species diversity. Here, the contribution of habitat heterogeneity of cold seeps at three spatial scales [micro-scale (ms), macro-scale (10 to 100s of ms), and mega-scale (10 to 100s of km)] to the total nematode biodiversity (genus level) along the Norwegian continental margin is evaluated. Due to the development of higher resolution bathymetry and increased bottom sampling in recent years, continental margins, once regarded as monotonous landscapes, are now acknowledged to have a high degree of habitat complexity and diversity. By calculating the additive partitioning of gamma diversity in alpha and beta fractions, we examined to what extent habitat diversity of seep sites significantly increases the nematode genus composition and diversity at different spatial scales. Siboglinidae patches and control sediments yielded comparably high levels of nematode genus richness. They exhibited low turnover rates within and across the different seep sites. In contrast, the bacterial mats at Håkon Mosby Mud Volcano (HMMV) and the reduced sediments at the Nyegga pockmarks harboured genus-poor nematode communities with an equally high dominance of one or two species, which were different for each seep. Different habitats, in particular at the HMMV, contributed significantly to the seep nematode richness. This study demonstrates that the presence of distinct habitat types within multiple seep sites contributes to the high diversity of nematode communities inhabiting the seeps in the Norwegian deep sea.     

Macrobenthic infaunal communities associated with deep‐sea hydrocarbon seeps in the northern Gulf of Mexico

There are thousands of seeps in the deep ocean worldwide; however, many questions remain about their contributions to global biodiversity and the surrounding deep‐sea environment. In addition to being globally distributed, seeps provide several benefits to humans such as unique habitats, organisms with novel genes, and carbon regulation. The purpose of this study is to determine whether there are unique seep macrobenthic assemblages, by comparing seep and nonseep environments, different seep habitats, and seeps at different depths and locations. Infaunal community composition, diversity, and abundance were examined between seep and nonseep background environments and among three seep habitats (i.e., microbial mats, tubeworms, and soft‐bottom seeps). Abundances were higher at seep sites compared to background areas. Abundance and diversity also differed among microbial mat, tubeworm, and soft‐bottom seep habitats. Although seeps contained different macrobenthic assemblages than nonseep areas, infaunal communities were also generally unique for each seep. Variability was 75% greater within communities near seeps compared to communities in background areas. Thus, high variability in community structure characterized seep communities rather than specific taxa. The lack of similarity among seep sites supports the idea that there are no specific infauna that can be used as indicators of seepage throughout the northern Gulf of Mexico, at least at higher taxonomic levels.     

Community structure and nutrition of deep methane-seep macrobenthos from the North Pacific (Aleutian) Margin and the Gulf of Mexico (Florida Escarpment)

Methane seeps occur at depths extending to over 7000 m along the world's continental margins, but there is little information about the infaunal communities inhabiting sediments of seeps deeper than 3000 m. Biological sampling was carried out off Unimak Island (3200–3300 m) and Kodiak Island (4500 m) on the Aleutian margin, Pacific Ocean and along the Florida Escarpment (3300 m) in the Gulf of Mexico to investigate the community structure and nutrition of macrofauna at these sites. We addressed whether there are characteristic infaunal communities common to the deep‐water seeps or to the specific habitats (clam beds, pogonophoran fields, and microbial mats) studied here, and ask how these differ from background communities or from shallow‐seep settings sampled previously. We also investigated, using stable isotopic signatures, the utilization of chemosynthetically fixed and methane‐derived organic matter by macrofauna from different regions and habitats. Within seep sites, macrofaunal densities were the greatest in the Florida microbial mats (20,961 ± 11,618 ind·m⁻²), the lowest in the Florida pogonophoran fields (926 ± 132 ind·m⁻²), and intermediate in the Unimak and Kodiak seep habitats. Seep macrofaunal densities differed from those in nearby non‐seep sediments only in Florida mat habitats, where a single, abundant species of hesionid polychaete comprised 70% of the macrofauna. Annelids were the dominant taxon (>60%) at all sites and habitats except in Florida background sediments (33%) and Unimak pogonophoran fields (27%). Macrofaunal diversity (H′) was lower at the Florida than the Alaska seeps, with a trend toward reduced richness in clam bed relative to pogonophoran field or non‐seep sediments. Community composition differences between seep and non‐seep sediments were evident in each region except for the Unimak margin, but pogonophoran and clam bed macrofaunal communities did not differ from one another in Alaska. Seep δ¹³C and δ¹⁵N signatures were lighter for seep than non‐seep macrofauna in all regions, indicating use of chemosynthetically derived carbon. The lightest δ¹³C values (average of species’ means) were observed at the Florida escarpment (−42.8‰). We estimated that on average animal tissues had up to 55% methane‐derived carbon in Florida mats, 31–44% in Florida clam beds and Kodiak clam beds and pogonophoran fields, and 9–23% in Unimak seep habitats. However, some taxa such as hesionid and capitellid polychaetes exhibited tremendous intraspecific δ¹³C variation (>30‰) between patch types. Overall we found few characteristic communities or features common to the three deep‐water seeps (>3000 m), but common properties across habitats (mat, clam bed, pogonophorans), independent of location or water depth. In general, macrofaunal densities were lower (except at Florida microbial mats), community structure was similar, and reliance on chemosynthesis was greater than observed in shallower seeps off California and Oregon.     

Limited differences among habitats in deep‐sea macro‐infaunal communities off New Zealand: implications for their vulnerability to anthropogenic disturbance

The spread of human activities into the deep sea may pose a high risk to benthic communities and affect ecosystem integrity. The deep sea is characterized by physical and biological heterogeneity and different habitat types are likely to differ in their vulnerability to anthropogenic impacts. However, across‐habitat comparisons are rare, and no comprehensive ecological risk assessment has yet been developed. To address this gap in our knowledge, we compared macro‐infaunal community structure in four habitats (slope, canyons, seamounts and methane seeps) at depths between 700 and 1500 m in the Hikurangi Margin and Bay of Plenty regions off New Zealand. The most striking contrast in community structure was between the two study regions, due to an order of magnitude difference in macro‐infaunal abundance that we believe was caused by differences in surface productivity and food availability at the sea bed. We found differences in structural and functional attributes of macro‐infaunal communities among some habitats in the Hikurangi Margin (slope, canyon and seep), but not in the Bay of Plenty. We posit that differences between canyon and slope communities on the Hikurangi Margin are due to enhanced food availability inside canyons compared with adjacent slope habitats. Seep communities were characterized by elevated abundance of both symbiont‐bearing and heterotrophic taxa, and were the most distinct, and variable, among the habitats that we considered on the Hikurangi Margin. Communities of seamounts were not distinct from slope or canyon communities on the Hikurangi Margin, probably reflecting similar environmental conditions in these habitats. The communities of deep‐sea canyon and seep habitats on the Hikurangi Margin were sufficiently dissimilar from each other and from slope habitats to warrant separate management consideration. By contrast, the low dissimilarity between communities of canyon and slope habitats in the Bay of Plenty suggests that habitat‐based management is not required in this region, for macro‐infauna at least. Although the two study regions share similar species pools, populations of the Hikurangi Margin region may be less vulnerable than the sparser populations of the Bay of Plenty due to the higher availability of potential colonizers and faster population growth. Thus regions, and habitats in some regions, should be subject to separate ecological risk assessment to help identify the key risks and consequences of human activities, and to inform options for reducing or mitigating impacts.     

Community structure of infaunal macrobenthos around vestimentiferan thickets at the San Clemente cold seep,NE Pacific

The San Clemente cold seep lies within 100–200 km of other reducing habitats in the NE Pacific, offering an opportunity to compare diversity and species overlap among reducing habitats (i.e. whale‐, kelp‐, and wood‐falls) at similar depths within a single region. Video observations from the research submersible Alvin at the San Clemente seep (1800 m depth) indicated clumps (‘thickets’) of vestimentiferans distributed as meter‐scale patches interspersed with vesicomyid clam beds and black sediments. Sediment‐core samples were collected at distances of 0 to 80–200 m along randomly oriented transects radiating outward from vestimentiferan thickets to evaluate changes in macrofaunal community structure from thickets into the background community. Macrofaunal abundance was elevated at distances of 0–1 m compared to 80–200 m (i.e. the ‘background’ community). The tube‐building frenulate worms Siboglinum spp., along with peracarid crustaceans, dominated sediments within 1 m of vestimentiferan thickets. Species diversity was depressed within 1 m of thickets but with high rates of species accumulation, suggesting that seep sites greatly increase sediment heterogeneity and facilitate colonization by non‐background macrofaunal species. Stable isotope data indicate chemosynthetic nutrition for some dominant macrofaunal species within 1 m of tubeworm thickets. The macrofaunal community near vestimentiferan thickets in San Clemente seep contains intermediate levels of species richness and diversity compared to other deep‐sea seep areas in the northeast Pacific. There was low species overlap between the San Clemente seep macrofauna and communities in reducing habitats near wood‐, whale‐, and kelp‐falls at similar depths within the region, suggesting that seeps harbor a distinct infaunal community.     

Diversity patterns along and across the Chilean margin: a continental slope encompassing oxygen gradients and methane seep benthic habitats

In the present study we review datasets available for the Chilean margin to assess the relationship between environmental (or habitat) heterogeneity and benthic diversity. Several factors, such as the presence of different water masses, including the oxygen-deficient Equatorial Sub-surface Waters (ESSW) at the continental shelf and upper slope, and the Antarctic Intermediate Waters (AIW) at mid slope depths appear to control the bathymetric distribution of benthic communities. The presence of methane seeps and an extended oxygen minimum zone (OMZ) add complexity to the benthic distribution patterns observed. All these factors generate environmental heterogeneity, which is predicted to affect the diversity patterns both along and across the Chilean continental margin. The response to these factors differs among different faunal size groups: meio-, macro-, and megafauna. Physiological adaptations to oxygen deficiency and constraints related to body size of each group seem to explain the larger-scale patterns observed, while sediment/habitat heterogeneity ( e.g. at water mass boundaries, hardgrounds, biogeochemical patchiness, sediment organic content, grain size) may influence the local fauna diversity patterns.     

Surficial Hydrocarbon Seep Infauna from the Blake Ridge (Atlantic Ocean, 2150 m) and the Gulf of Mexico (690–2240 m)

Abstract.  Infauna, including foraminifera and metazoans, were enumerated and identified from five types of seep habitats and two adjacent non-seep habitats. Collections were made with the deep submergence research vessel 'Alvin' from three areas of active seepage in the Gulf of Mexico (Alaminos Canyon [2220 m], Atwater Canyon [1930 m], and Green Canyon lease block 272 [700 m]) and on the Blake Ridge Diapir [2250 m], which is located off the southeastern coast of the United States. The seep habitats sampled included four types of microbial mats ( Beggiatoa , Thioploca , thin and thick Arcobacter ) and the periphery of a large mussel bed. Sediments under large rhizopod protists, xenophyophores, were sampled adjacent to the mussel bed periphery. A non-seep site, which was >1 km away from active seeps, was also sampled for comparison. Densities of most taxa were higher in the Gulf of Mexico seeps than in Blake Ridge samples, largely because densities in the thick microbial mats of Blake Ridge were significantly lower. Diversity was higher in the Thioploca mats compared to other microbial-mat types. Within an ocean basin ( i.e. , Atlantic, Gulf of Mexico) we did not observe significant differences in meiofaunal or macrofaunal composition in Beggiatoa versus Thioploca mats or thin versus thick Arcobacter mats. Foraminifera represented up to 16% of the seep community, a proportion that is comparable to their contribution at adjacent non-seep communities. In general, the observed densities and taxonomic composition of seep sites at the genus level was consistent with previous observations from seeps ( e.g. , the foraminifers Bolivina and Fursenkoina , the dorvilleid polychaete Ophryotrocha ).     

Habitat heterogeneity influences cold‐seep macrofaunal communities within and among seeps along the Norwegian margin – Part 2: contribution of chemosynthesis and nutritional patterns

The relative contribution of chemosynthesis in heterotrophic fauna at seeps is known to be influenced by depth and by habitat. Using stable isotopes of carbon and nitrogen, we investigated macro‐ and megafaunal nutritional patterns in Norwegian margin cold seeps by comparing food webs both among habitats within a seep site and between different sites. The very active Håkon Mosby mud volcano (HMMV) is characterized by geochemical gradients, microbial activity and faunal zonation from the centre to the periphery. The Storegga Slide (600–900 m depth) has pockmarks with patchy less active seeps, and also shows concentric zonation of habitats but at much smaller spatial scale. The dominant carbon source for macrofaunal nutrition in both areas was chemosynthetically fixed and the bulk of organic carbon was derived from sulphur‐oxidizing bacteria. In HMMV, food chains were clearly separated according to habitats, with significantly lighter δ¹³C signatures on microbial mats and adjacent sediment (?33.06 to ?50.62‰) than in siboglinid fields (?19.83 to ?35.03‰). Mixing model outputs revealed that the contribution of methane‐derived carbon was small in siboglinid fields (0–17%) but significant (39–61%) in the microbial mats. Moreover, the variability of macrofauna signatures within this later habitat suggests the co‐occurrence of two food chains, one based on primary production via methanotrophy and the other via sulphide oxidation. The length of the food chains also varied among habitats, with at least one more trophic level in the siboglinid fields located at the periphery of the volcano. Conversely, in Storrega pockmarks, faunal δ¹³C signatures did not vary among habitats but among species, although separate food chains seem to co‐occur. The small size of the seepage areas and their lower fluxes compared to HMMV allow more background species to penetrate the seep area, increasing the range of δ¹⁵N and the trophic level number. Probably due to the higher flux of photosynthetic particulate organic carbon, the overall chemosynthesis‐based carbon contribution in invertebrate nutrition was lower than that in HMMV.     

Sidescan backscatter variations of cold seeps on the Hikurangi Margin (New Zealand): indications for different stages in seep development

Cold seeps on the Hikurangi Margin off New Zealand exhibit various seabed morphologies producing different intensity patterns in sidescan backscatter images. Acoustic backscatter characteristics of 25 investigated seep sites fall into four distinct types characterised by variations in backscatter intensity, distribution and inferred structural heights. The types reflect different carbonate morphologies including up to 20-m-high structures (type 1), low-relief crusts (type 2), scattered blocks (type 3) and carbonate-free sites (type 4). Each seep corresponds to a single type; intermediates were not observed. This correlates well with published data on seep fauna at each site, with the four types representing three different faunal habitats of successive stages of seep development. Backscatter signatures in sidescan sonar images of cold seeps may therefore serve as a convenient proxy for variations in faunal habitats.     

Metazoan meiofaunal communities at cold seeps along the Norwegian margin: Influence of habitat heterogeneity and evidence for connection with shallow-water habitats

Cold-seep environments and their associated symbiont-bearing megafaunal communities create islands of primary production for macro- and meiofauna in the otherwise monotonous and nutrient-poor deep-sea environment. To examine the spatial variation and distribution patterns of metazoan meiobenthos in different seepage-related habitats, samples were collected in two regions off Norway: several pockmarks associated with the Storegga Slide including the Nyegga pockmark area (730 m; 64°N), and the active, methane-venting Håkon Mosby Mud Volcano (HMMV) west of the Barents Sea (1280 m; 72°N). Based on sediment geochemistry and associated epifauna, three different habitat types were distinguished across the two regions: (1) reduced sediment with suboxic conditions, sometimes covered by bacterial mats, (2) sediment colonised by chemosynthetic, siboglinid tubeworms, and (3) sediment outside the influence of seepage and without a large chemosynthetic fauna. Meiofaunal communities varied strongly in terms of generic diversity and dominance among the different habitat types. Control sites and Siboglinidae polychaete fields both supported high nematode genus richness similar to normal deep-sea sediments, whereas the reduced sediments yielded a genus-poor nematode community dominated by one or two successful species. Meiofaunal densities in the different habitats were negatively correlated with macrobenthic densities. An extremely dense (>11,000 ind. 10 cm^–2), mono-specific nematode population appeared to be restricted to the bacterial mats at HMMV. It consisted of a new cryptic species of the Halomonhystera disjuncta complex, which has been described from intertidal habitats in the North Sea. The reduced seep sediments at Nyegga did not yield H. disjuncta but were dominated by Terschellingia longicaudata, another cosmopolitan nematode species known to be abundant in organic-rich, oxygen-poor, shallow-water environments. These observations point to a past or recent connection between margins and shallow-water habitats.     

Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins

Biological structures exert a major influence on species diversity at both local and regional scales on deep continental margins. Some organisms use other species as substrates for attachment, shelter, feeding or parasitism, but there may also be mutual benefits from the association. Here, we highlight the structural attributes and biotic effects of the habitats that corals, sea pens, sponges and xenophyophores offer other organisms. The environmental setting of the biological structures influences their species composition. The importance of benthic species as substrates seems to increase with depth as the complexity of the surrounding geological substrate and food supply decline. There are marked differences in the degree of mutualistic relationships between habitat-forming taxa. This is especially evident for scleractinian corals, which have high numbers of facultative associates (commensals) and few obligate associates (mutualists), and gorgonians, with their few commensals and many obligate associates. Size, flexibility and architectural complexity of the habitat-forming organism are positively related to species diversity for both sessile and mobile species. This is mainly evident for commensal species sharing a facultative relationship with their host. Habitat complexity is enhanced by the architecture of biological structures, as well as by biological interactions. Colony morphology has a great influence on feeding efficiency for suspension feeders. Suspension feeding, habitat-forming organisms modify the environment to optimize their food uptake. This environmental advantage is also passed on to associated filter-feeding species. These effects are poorly understood but represent key points for understanding ecosystems and biodiversity on continental margins. In this paper we explore the contributions of organisms and the biotic structures they create (rather than physical modifications) to habitat heterogeneity and diversity on the deep continental margins.     

An ancient linked fluid migration system: cold-seep deposits and sandstone intrusions in the Panoche Hills, California, USA

In central California, Maastrichtian–Danian shales of the Moreno Formation preserve a fluid migration system that developed along the western margin of the former Great Valley forearc basin. The system consists of a network of interconnected sandstone intrusions linked to overlying fossiliferous carbonates whose geochemistry, fauna, and petrology are characteristic of active cold seeps. The system is approximately 800 m thick and represents episodic migration and seafloor expulsion of fluids over at least 0.5×10⁶ years. This locality has the most extensive exposure yet discovered of a complete seep system, from underlying fluid pathways to seep deposits and associated communities.     

Spatial heterogeneity of epibenthos on artificial reefs: fouling communities in the early stages of colonization on an East Australian shipwreck

Artificial reefs are spatially complex habitats and serve as good model systems to study patterns of community succession and the response of epibiota to environmental clines over small spatial scales. Here, we quantified spatial heterogeneity in community composition and diversity of fouling communities across a number of environmental gradients that included water depth, surface orientation of habitats, exposure to currents, and shelter. Assemblage structure was quantified by spatially replicated photo transects on a recently scuttled large navy ship off the East Australian coast, lying in 27 m of water. A rich assemblage of epifauna had colonized the wreck within a year, dominated by barnacles, sponges and bryozoans. Community structure varied significantly over small spatial scales of meters to tens of meters. Depth, surface orientation and exposure were the major environmental drivers. Assemblages were substantially less diverse and abundant on the deepest (23 m near the seafloor) part of the hull with residual antifouling paint, on sheltered surfaces inside the wreck, and on the sediment‐laden horizontal surfaces. Overall, the wrecks’ habitat complexity corresponds with small‐scale heterogeneity in the fouling communities. This study supports the notion that wrecks enhance local diversity and biomass within the habitat mosaic of their location, and habitat complexity may be an important mechanism for this, as demonstrated by the large spatial variability in the assemblages documented here.     

Monterey Bay cold-seep biota: Assemblages,abundance, and ultrastructure of living foraminifera

Although there is a growing body of evidence indicating benthic foraminifera inhabit hydrocarbon and cold seep environments, biochemical and ultrastructural data on seep foraminiferal communities are not available. Therefore, sediments collected from cold seeps in Monterey Bay, CA (900–1000 m), were examined for the presence of live benthic foraminifera. Results from three independent methods (ATP assay, ultrastructural analysis, rose Bengal staining) indicate that certain species inhabit the Clam Flat and Clam Field seeps. Abundances in our seep samples were lower than in comparable non-seep sites, although not atypical for these bathyal depths. Of 38 species represented at these two seep sites by cytoplasm-containing specimens, only Spiroplectammina biformis was restricted to the seep environment. However, because S. biformis is also known from non-seep sites in other areas, it should not be considered as endemic to seeps. Ultrastructural studies show abundant peroxisomes in seep specimens, which may allow inhabitation of such environments. One specimen of Uvigerina peregrina had prokaryotes nestled in test pores, suggesting that bacteria may play a role in the survival of foraminifera in this seep environment.     

Deep-sea habitat heterogeneity influence on meiofaunal communities in the Gulf of Guinea

To estimate the degree of spatial heterogeneity of benthic deep-sea communities, we carried out a multiple-scale (from m's to 200 km) investigation in the Congo-Angola margins (Equatorial West African margin, 3150–4800 m) in which we examined the metazoan meiofauna at a variety of habitats along the Congo Channel system and in the associated cold seep. We investigate the structure, density, vertical distribution patterns in the sediment and biomass of meiofaunal communities in the Gulf of Guinea and how they are controlled by hydrologic and biogeochemical processes. The meiofaunal communities in the Gulf of Guinea were shaped by heterogeneous conditions on the margin, and reflect the multiple-scale spatial variability that corresponds with the different identified habitats. The two control sites, located at >100 km away from the canyon, were inhabited by very dense and the most diverse meiobenthic communities. Similar meiobenthic communities inhabited the transition zone between the canyon and the cold seep. Sites located along the Congo Channel were obviously affected by the local high-velocity bottom currents and unstable sedimentary conditions in this active submarine system. Extremely low meiobenthic densities and very low proportions in the most surficial sediment layers provided evidence for recently highly disturbed sediments at these sites. The remote operated vehicle (ROV) Victor 6000 provided images of the cold seep, showing a patchy distribution of several types of patchy distributed megafaunal communities dominated by three key symbiotic taxa (Mytilidae, Vesicomyidae and Siboglinidae). These cold seep sediments were colonised by a unique meiobenthic community, characterised by a high small-scale (m's) patchiness, low species richness and the prominent dominance of two large-sized nematode species: Sabatieria mortenseni, which is a cosmopolitan nematode known from littoral habitats, and an undescribed Desmodora species. The high individual body weight of S. mortenseni and its dominance at the cold seep site resulted in a significantly higher nematode biomass at the seep compared to the surrounding sites. In addition, the vertical nematode profiles, with maximum proportions in subsurficial layers, points to a chemosynthesis-based meiobenthic community in this cold seep, in contrast to the phytodetritus-based communities at the control sites and at the transition zone.     

Rare species, habitat diversity and functional redundancy in marine benthos

Macro-ecological theories relating species richness, abundance, range size, biological traits and environmental tolerance have rarely been tested in marine soft-sediments, despite the spatial extent of these habitats and the inherent richness of resident communities. This study examines the contribution of rare species to marine soft-sediment communities from New Zealand, focussing on the relationships of range size with abundance, environment, habitat diversity and life history traits. 54% of the 351 species sampled exhibited restricted ranges (found at ≤ 2 sites). In contrast to many terrestrial systems, we observed only a weak positive relationship between abundance and frequency of occurrence. Restricted-range species were not randomly distributed, with their distribution related to habitat characteristics, suggesting an important link between habitat diversity and rarity. They exhibited a similar range of traits to the total observed species pool, suggesting that they are not only important to biodiversity but could play a role in stability. Restricted range species were generally not small and this, together with the number of different biological traits represented, suggests that rare species are important to the functioning of marine systems. Thus, our results highlight the importance of considering rare species in habitat-based approaches to conservation.     

Rare earth elements in cold seep carbonates from the southwestern Dongsha area,northern South China Sea

The circulation of methane-rich fluids at cold seeps often leads to the precipitation of seep carbonates close to the seafloor along continental margins, which can be used as records of past fluid seepage. Rare earth element (REE) concentrations in seep carbonates have been used to trace fluid sources and provide information on associated biogeochemical processes at cold seeps. The REE concentrations of a series of carbonates collected from cold seeps in the southwestern Dongsha area of the northern South China Sea are analyzed in this study. The total REE contents (ΣREE) of the seep carbonates analyzed show a wide variation from 17 ppm to 523 ppm with an average ΣREE value of 54 ppm, which are higher than the typical marine carbonate values of ∼28 ppm commonly reported and also higher than those of the carbonates from other cold seep areas. A positive correlation between Fe–Mn content and ΣREE was observed. These results suggest that the seep carbonates of this study were primarily controlled by the methane-derived fluid from which they precipitated. The Fe-rich dolomite and siderite, which are the main components of the carbonates, are responsible for the enrichment of the REE. A slight positive Ce anomaly observed in the shale-normalized REE patterns of the studied seep carbonates suggests that they formed in anoxic conditions, and the correlations between Ce/Ce* and La_N/Sm_N, Ce/Ce* and Dy_N/Sm_N, Ce/Ce* and ΣREE further reveal that the REE characteristics of most seep carbonate samples preserve the original redox conditions in which they precipitated and late diagenesis has had little effect on the REE. However, the REE characteristics of sub-samples DS2-2B, DS1-6A and DS1-7A are very different from those of the other sub-samples, indicating a greater impact of late diagenesis and post-oxidation favored REE enrichment.     

Epizooic metazoan meiobenthos associated with tubeworm and mussel aggregations from cold seeps of the northern Gulf of Mexico

The abundance and higher taxonomic composition of epizooic metazoan meiobenthic communities associated with mussel and tubeworm aggregations of hydrocarbon seeps at Green Canyon, Atwater Valley, and Alaminos Canyon in depths between 1400 and 2800?m were studied and compared to the infaunal community of non-seep sediments nearby. Epizooic meiofaunal abundances of associated meiobenthos living in tubeworm bushes and mussel beds at seeps were extremely low (usually <100?ind. 10?cm(-2)), similar to epizooic meiofauna at deep-sea hydrothermal vents, and the communities were composed primarily of nematodes, copepods, ostracods, and halacarids. In contrast, epizooic meiobenthic abundance is lower than previous studies have reported for infauna from seep sediments. Interestingly, non-seep sediments contained higher abundances and higher taxonomic diversity than epizooic seep communities, although in situ primary production is restricted to seeps.     

Cold-seep habitat mapping: High-resolution spatial characterization of the Blake Ridge Diapir seep field

Relationships among seep community biomass, diversity, and physiographic controls such as underlying geology are not well understood. Previous efforts to constrain these relationships at the Blake Ridge Diapir were limited to observations from piloted deep-submergence vehicles. In August 2012, the autonomous underwater vehicle (AUV) Sentry collected geophysical and photographic data over a 0.131 km² area at the Blake Ridge Diapir seeps. A nested survey approach was used that began with a regional or reconnaissance-style survey using sub-bottom mapping systems to locate and identify seeps and underlying conduits. This survey was followed by AUV-mounted sidescan sonar and multibeam echosounder systems mapping on a mesoscale to characterize the seabed physiography. At the most detailed survey level, digital photographic imaging was used to resolve sub-meter characteristics of the biology. Four pockmarks (25–70 m diameter) were documented, each supporting chemosynthetic communities. Concentric zonation of mussels and clams suggests the influence of chemical gradients on megafaunal distribution. Data collection and analytical techniques used here yield high-resolution habitat maps that can serve as baselines to constrain temporal evolution of seafloor seeps, and to inform ecological niche modeling and resource management.     

Habitat heterogeneity influences cold‐seep macrofaunal communities within and among seeps along the Norwegian margin. Part 1: macrofaunal community structure

Håkon Mosby mud volcano (HMMV) is one of the most active and most studied seep sites in European waters. Many authors have described its thermal activity, dynamic of mud flows, and geochemical and microbial processes. It is characterised by a concentric zonation of successive biogenic habitats related to an activity and geochemical gradient from its centre to its periphery. Around the central area covered by mud flows, white and grey microbial mats occur among areas of bare sediment, whereas siboglinid tubeworm fields of Sclerolinum contortum and/or Oligobrachia haakonmosbiensis colonise the peripheral areas. The meiofaunal community is known to be structured among habitats, but the macrofauna has rarely been investigated and has never been sampled in situ. As part of the European project HERMES, using the ROVs Victor 6000 and Quest 4000, we sampled quantitatively the different habitats of the volcano for macrofauna sensus lato, retained on a 250‐ or 500‐μm sieve. We also sampled a newly discovered pockmark on Storegga slide (cne 5.6) and two pockmarks (G11, G12) in the Nyegga area. Macrofauna was identified and counted from phylum to family level. Our results on HMMV showed a gradient of increasing density and diversity from the volcano centre (1–3 taxa; 260 ind·m^?2) to the peripheral siboglinid fields (8–14 taxa, 93,000 ind·m^?2), with an intermediate situation for microbial mats. For macrofauna ≥500 μm, non‐siboglinid polychaetes dominated the communities of the central mud volcano area, white mats and S. contortum fields (83, 89 and 37% of the total, respectively), whereas gastropods dominated grey mats and O. haakonmosbiensis fields (89 and 44% of the total, respectively). Polychaete families followed the same pattern of diversity according to habitats within HMMV. Of 23 polychaete families identified, only one occurred in the centre, and three in the microbial mats. Capitellidae and Dorvilleidae (typical of organically and sulphide‐enriched areas) occurred at remarkably high densities in white microbial mats and in O. haakonmosbiensis fields. The S. contortum fields were the most diverse habitat with 12 polychaete families. The 250‐μm fraction showed similar taxa dominating the habitats, but taking meiofauna into account, nematodes became the major taxon in white mats and in S. contortum fields, where they were particularly large in size, whereas copepods dominated in other habitats. Meiofauna and macrofauna did not show the same patterns of density according to habitats. Using principal components analysis the habitats at HMMV were clearly distinct, and clustered according to dominant species of siboglinids and type of microbial mats. Pockmarks at Nyegga showed a similar concentric pattern of habitats around fluid sources as on the volcano, which seemed similarly to influence macrofauna composition, but at a much smaller scale. Total taxa and polychaete diversity are high in the S. contortum fields in these pockmarks as well. Regional‐scale comparisons including HMMV and Storegga suggested a higher influence of habitat‐type than seep‐site on the community structure.