南海北部冷泉平端深海偏顶蛤的主要生化成分及其与热液和近岸种的对比研究

Bathymodiolus属贻贝是广泛分布于全球深海冷泉和热液生态系统的优势种,在深海化能生态系统的物质循环和能量流动中起着重要作用。本文以我国南海北部冷泉的主要优势种平端深海偏顶蛤(Bathymodiolus platifrons)为研究对象,采用常规生化测定方法,研究了其主要生化成分和氨基酸组成的特征,并就特定组织中主要生化成分的含量与生活在热液区的平端深海偏顶蛤和近岸的远东偏顶蛤(Modiolus kurilensis)进行了对比。结果表明,南海冷泉平端深海偏顶蛤软体部含水量84.28%,粗蛋白含量7.18%,粗脂肪含量1.23%,糖类含量2.75%,与已报道的深海贝类组成相近。虽然主要生化成分含量在3种贻贝的鳃、外套膜、闭壳肌和消化腺4个组织中总体差异不大,但是冷泉平端深海偏顶蛤和热液平端深海偏顶蛤的外套膜糖类含量(25.20%、30.66%)显著高于远东偏顶蛤(6.97%,P0.05),这表明平端深海偏顶蛤的主要储存物质为外套膜中的糖类。在氨基酸组成上,冷泉平端深海偏顶蛤鳃中氨基酸总量为44.55%(干质量),外套膜中为34.83%(干质量),其中必需氨基酸分别占比41.73%和40.52%,总体与其他贝类相似。然而,在平端深海偏顶蛤中,与渗透压调节相关的甘氨酸和与硫代谢相关的牛磺酸含量较高,这与其适应深海高盐度高硫化氢浓度的环境相关。综上所述,南海冷泉平端深海偏顶蛤在常规生化组分和氨基酸组成上与近岸常见双壳类具有一定差异,这些差异与其特殊生境的关系还需要更加深入的研究。     

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.     

Growth, production, and mortality of the chemosynthetic vesicomyid bivalve, Calyptogena kilmeri from cold seeps off central California

The vesicomyid Calyptogena kilmeri is one of the most abundant bivalves inhabiting chemosynthetic environments shallower than c. 1500 m along central California. We estimated the population size structure, biomass, rates of individual growth, somatic production, and mortality for C. kilmeri, based on sampling of seep habitats and tag–recapture studies at chemosynthetic communities in Monterey Bay, California. The composite growth rate of C. kilmeri over all sites was relatively high (K = 0.25), reaching c. 80% of asymptotic length (104.7 mm) in 6.6 years. The density of C. kilmeri was estimated as 938.5 ind. m⁻² and biomass density varied from 704 to 2059 g ash free dry mass (AFDM) m⁻². Somatic production was also high (294–297 g·AFDM·m⁻² year⁻¹), and production/biomass ratios for C. kilmeri varied from 0.14 to 0.42 among sites, related to variation in size–frequency distributions among sites. Instantaneous mortality rates estimated from size distributions ranged from 0.17 to 0.24 year⁻¹. Growth and somatic production by C. kilmeri are in the range reported for chemosynthetic bivalves from hydrothermal vent and seep habitats, as well as photosynthetic‐based assemblages of inshore or intertidal bivalves, and greatly exceed rates reported for heterotrophic deep‐sea benthos.     

Depth-related structure and ecological significance of cold-seep communities—a case study from the Sea of Okhotsk

We discovered and investigated several cold-seep sites in four depth zones of the Sea of Okhotsk off Northeast Sakhalin: outer shelf (160–250 m), upper slope (250–450 m), intermediate slope (450–800 m), and Derugin Basin (1450–1600 m). Active seepage of free methane or methane-rich fluids was detected in each zone. However, seabed photography and sampling revealed that the number of chemoautotrophic species decreases dramatically with decreasing water depth. At greatest depths in the Derugin Basin, the seeps were inhabited by bacterial mats and bivalves of the families Vesicomyidae (Calyptogena aff. pacifica, C. rectimargo, Archivesica sp.), Solemyidae (Acharax sp.) and Thyasiridae (Conchocele bisecta). In addition, pogonophoran tubeworms of the family Sclerolinidae were found in barite edifices. At the shallowest sites, on the shelf at 160 m, the seeps lack chemoautotrophic macrofauna; their locations were indicated only by the patchy occurrence of bacterial mats.Typical seep-endemic metazoans with chemosynthetic symbionts were confined to seep sites at depths below 370 m. A comparative analysis of the structure of seep and background communities suggests that differences in predation pressure may be an important determinant of this pattern. The abundance of predators such as carnivorous brachyurans and asteroids, which can invade seeps from adjacent habitats and efficiently prey on sessile seep bivalves, decreased very pronouncedly with depth. We conclude from the obvious correlation with the conspicuous pattern in the distribution of seep assemblages that, on the shelf and at the upper slope, predator pressure may be high enough to effectively impede any successful settlement of viable populations of seep-endemic metazoans. However, there was also evidence that other depth-related factors, such as bottom-water current, sedimentary regimes, oxygen concentrations and the supply of suitable settling substrates, may additionally regulate the distribution of seep fauna in the area.As a consequence of the pronounced pattern in the distribution of seep communities, their ecological significance as food sources of surrounding background fauna increased with water depth. Isotopic analyses suggest that in the Derugin Basin seep colonists feed on chemoautotrophic seep organisms, either directly or by preying on metazoans with chemosynthetic symbionts. In contrast, seep organisms apparently do not contribute to the nutrition of the adjacent background fauna on the shelf and at the slope. In this area, elevated epifaunal abundances at seep sites were caused primarily by the availability of suitable settling substrates rather than by an enrichment of food supply.     

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.     

基于贝壳生长轮方法的平端深海偏顶蛤的生长和年龄结构研究

平端深海偏顶蛤(Gigantidasplatifrons)是广泛分布于西太平洋深海热液和冷泉生态系统中的优势种和共有种,也是深海化能生态系统中重要的生境营造种。以南海F冷泉平端深海偏顶蛤为研究对象,基于贝壳日生长轮方法,分析了平端深海偏顶蛤的年龄与生长速率,建立了其年龄与贝壳壳长关系的生长方程。同时,结合获得的生长方程,分析了南海F冷泉采样点(119.285 6°E,22.115 4°N)平端深海偏顶蛤种群的年龄结构。研究结果表明,采样获得的平端深海偏顶蛤种群的最大壳长为11.4 cm,最大寿命为13.5龄。研究区域的贝壳长度分布集中在4～7 cm,占比60%;年龄分布集中在2～4龄,占比49.7%。相关研究结果为进一步开展平端深海偏顶蛤的生长研究提供了基础数据,有助于深入了解冷泉区域的种群动态变化规律。     

Distribution and temporal variation of mega‐fauna at the Regab pockmark (Northern Congo Fan), based on a comparison of videomosaics and geographic information systems analyses

The Regab pockmark is a large cold seep area located 10 km north of the Congo deep sea channel at about 3160 m water depth. The associated ecosystem hosts abundant fauna, dominated by chemosynthetic species such as the mussel Bathymodiolus aff. boomerang, vestimentiferan tubeworm Escarpia southwardae, and vesicomyid clams Laubiericoncha chuni and Christineconcha regab. The pockmark was visited during the West African Cold Seeps (WACS) cruise with RV Pourquoi Pas? in February 2011, and a 14,000‐m² high‐resolution videomosaic was constructed to map the most populated area and to describe the distribution of the dominant megafauna (mussels, tubeworms and clams). The results are compared with previous published works, which also included a videomosaic in the same area of the pockmark, based on images of the BIOZAIRE cruise in 2001. The 10‐year variation of the faunal distribution is described and reveals that the visible abundance and distribution of the dominant megafaunal populations at Regab have not changed significantly, suggesting that the overall methane and sulfide fluxes that reach the faunal communities have been stable. Nevertheless, small and localized distribution changes in the clam community indicate that it is exposed to more transient fluxes than the other communities. Observations suggest that the main megafaunal aggregations at Regab are distributed around focused zones of high flux of methane‐enriched fluids likely related to distinct smaller pockmark structures that compose the larger Regab pockmark. Although most results are consistent with the existing successional models for seep communities, some observations in the distribution of the Regab mussel population do not entirely fit into these models. This is likely due to the high heterogeneity of this site formed by the coalescence of several pockmarks. We hypothesize that the mussel distribution at Regab could also be controlled by the occurrence of zones of both intense methane fluxes and reduced efficiency of the anaerobic oxidation of methane possibly limiting tubeworm colonization.     

台湾岛西南海域福尔摩沙海脊冷泉区地形地貌特征分析

利用水深数据和ROV近海底影像资料,对福尔摩沙海脊冷泉区的海底地形地貌和冷泉系统的海底表征进行了描述和分析,并讨论了二者之间的响应关系。结果表明,相对于船载多波束数据而言,近海底多波束测深系统所获得的数据能更高精度地反映冷泉区海底地形地貌特征,是研究冷泉系统不可或缺的基础资料。基于ROV近海底观测影像资料,福尔摩沙海脊冷泉系统整体表现为局部被化能自养生物群落覆盖并有流体喷口零星分布的巨大自生碳酸盐岩岩丘,海底表征主要包括形态各异的自生碳酸盐岩结壳或岩体、化能自养生物群落、流体喷口、还原性沉积物等几种形式。研究表明,福尔摩沙海脊冷泉区的地形地貌特征与冷泉系统海底表征具有良好的响应关系,并且该区的地形地貌特征主要受控于出露于海底的自生碳酸盐岩的形态特征及规模。首次揭示了福尔摩沙海脊冷泉区地形地貌特征与其海底表征之间的响应关系,以期为后续的冷泉研究提供必要的背景资料支持。     

Patterns of growth in cold-seep vestimenferans including Seepiophila jonesi: a second species of long-lived tubeworm

Seepiophila jonesi is a vestimentiferan tubeworm (Siboglinidae: Polychaeta) inhabiting the cold seeps of the upper slope of the Gulf of Mexico. It commonly co‐occurs with Lamellibrachia luymesi, which is among the most long‐lived non‐clonal animals known. The growth pattern of S. jonesi is best described by a model including a size‐specific probability of growth and an average growth rate that does not vary with individual size. This model, based on growth data from in situ staining and collection approximately 1 year later, predicts that S. jonesi is very slow growing and may attain ages comparable with L. luymesi. The efficacy of this model in describing L. luymesi growth rate was assessed, but the previously employed model of declining growth rate with individual size provided the better fit to the empirical data. Comparisons of both S. jonesi and L. luymesi growth rates among sites and among aggregations within a site indicate that there is some degree of habitat variability contributing to differences in growth rates. However, position of the anterior end of the worm within an aggregation did not have a significant effect on growth rate in comparisons among groups of L. luymesi from different distances from the center of an aggregation. The evolution of longevity in these species of vestimentiferans was favored by the relative stability of the seep habitat and sulfide sources, in contrast to the hydrothermal vent environment inhabited by relatively short‐lived and fast‐growing vestimentiferan species.     

Physiological and behavioral responses of Bathynerita naticoidea (Gastropoda: Neritidae) and Methanoaricia dendrobranchiata (Polychaeta: Orbiniidae) to hypersaline conditions at a brine pool cold seep

Bathynerita naticoidea (Gastropoda: Neritidae) and Methanoaricia dendrobranchiata (Polychaeta: Orbiniidae) are two of the most abundant invertebrates associated with cold‐seep mussel beds in the Gulf of Mexico. At the methane seep known as Brine Pool NR‐1 (27 °43.415 N, 91 °16.756 W; 650 m depth), which is surrounded by a broad band of mussels (Bathymodiolus childressi), these species have distinctly different patterns of abundance, with the gastropod being found mostly at the outer edge of the mussel bed (average density in November 2003: 817 individuals·m⁻² in outer zone, 20·m⁻² in inner zone) and the polychaete being found almost exclusively near the inner edge (average density in November 2003: 3155 individuals·m⁻² in inner zone, 0·m⁻² in outer zone), adjacent to the brine pool itself. The salinity of the brine pool exceeds 120, so we hypothesized that M. dendrobranchiata should be more tolerant of high salinities than B. naticoidea. The opposite proved to be true. The gastropods were capable of withstanding salinities at least as high as 85, whereas the polychaetes died at salinities higher than 75. Both species were osmoconformers over the range of salinities (35–75) tested. Behavioral responses of B. naticoidea to salinities of 50, 60, and 70 were investigated in inverted vertical haloclines. Gastropods generally did not enter water of salinity greater than 60, but tolerated short periods at 60. Behavioral avoidance of brine should limit the vertical distribution of B. naticoidea in the inner zone to the top 2.5–5 cm of the mussel bed. Behavior is also a likely (though unproven) mechanism for controlling horizontal distribution of this species across the mussel bed. Methanoaricia dendrobranchiata can tolerate short excursions into the brine, but probably avoids hypersaline conditions by aggregating on the tops of the mussels.     

Biologic and geologic characteristics of cold seeps in Monterey Bay, California

Cold seep communities discovered at three previously unknown sites between 600 and 1000 m in Monterey Bay, California, are dominated by chemoautotrophic bacteria (Beggiatoa sp.) and vesicomyid clams (5 sp.). Other seep-associated fauna included galatheid crabs (Munidopsis sp.), vestimentiferan worms (Lamellibrachia barhami?), solemyid clams (Solemya sp.), columbellid snails (Mitrella permodesta, Amphissa sp.), and pyropeltid limpets (Pyropelta sp.). More than 50 species of regional (i.e. non-seep) benthic fauna were also observed at seeps. Ratios of stable carbon isotopes (δ¹³C) in clam tissues near 36‰ indicate sulfur-oxidizing chemosynthetic production, rather than non-seep food sources, as their principal trophic pathway. The “Mt Crushmore” cold seep site is located in a vertically faulted and fractured region of the Pliocene Purisima Formation along the walls of Monterey Canyon ( 635 m), where seepage appears to derive from sulfide-rich fluids within the Purisima Formation. The “Clam Field” cold seep site, also in Monterey Canyon ( 900 m) is located near outcrops in the hydrocarbon-bearing Monterey Formation. Chemosynthetic communities were also found at an accretionary-like prism on the continental slope near 1000 m depth (Clam Flat site). Fluid flow at the “Clam Flat” site is thought to represent dewatering of accretionary sediments by tectonic compression, or hydrocarbon formation at depth, or both. Sulfide levels in pore waters were low at Mt Crushmore (ca 0.2 mM), and high at the two deeper sites (ca 7.011.0 mM). Methane was not detected at the Mt Crushmore site, but ranged from 0.06 to 2.0 mM at the other sites.     

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.     

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.     

基于单波束测深资料的海底冷泉动力学特征反演方法研究

为了准确理解冷泉水体中甲烷气体的分布规律,综合运用单波束测深数据和冷泉水体流场数据,建立了冷泉气泡上升、溶解速率的定量反演方法,利用现场海试资料对反演方法进行了验证,并对海试区域的冷泉气体溢出、溶解通量以及冷泉水体的甲烷浓度进行了估算。计算结果表明,考察船航向与冷泉水体流向的差异会对声学探测结果产生影响,当航向与流向的夹角大于±60°时,声学成像中冷泉倾角的误差将超过50%。同时,冷泉气泡上升速率的衰减与溢出口水深显著相关,相关系数可达0.9,并且冷泉上升流对上升速率的提升效果明显。水合物稳定带的分布对冷泉气泡的收缩速率影响显著,稳定带内、外收缩速率的差异可达3～4倍。冷泉气体通量的计算结果表明,调查区域内冷泉的溢出强度整体较大,同时溶解通量与水深之间呈现明显的分段效应。根据冷泉气体溶解通量估算的冷泉水体甲烷浓度与色谱分析结果具有较好的一致性。相关研究有助于实现对冷泉水体中甲烷气体分布的定量评估,为潜在海底冷泉区的圈划和海域天然气水合物的调查提供技术支撑。     

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.     

Ecophysiological differences between vesicomyid species and metabolic capabilities of their symbionts influence distribution patterns of the deep‐sea clams

This study provides an analysis of vesicomyid bivalve–symbiont community distribution across cold seep and hydrothermal vent areas in the Guaymas Basin (Gulf of California, Mexico). Using a combination of morphological and molecular approaches including fluorescent in situ hybridization (FISH), and electronic microscopy observations, vesicomyid clam species and their associated symbionts were characterized and results were analyzed in light of geochemical conditions and other on‐site observations. A greater diversity of vesicomyids was found at cold seep areas, where three different species were present (Phreagena soyoae [syn. kilmeri], Archivesica gigas, and Calyptogena pacifica). In contrast, A. gigas was the only species sampled across the hydrothermal vent area. The same haplotype of A. gigas was found in both hydrothermal vent and cold seep areas, highlighting possible contemporary exchanges among neighboring vents and seeps. In either ecosystem, molecular characterization of the symbionts confirmed the specificity between symbionts and hosts and supported the hypothesis of a predominantly vertical transmission. In addition, patterns of clams could reflect potential niche preferences for each species. The occurrence of numerous traces of vesicomyid movements on sediments in the sites colonized by A. gigas seemed to indicate that this species might have a better ability to move. Furthermore, variation in gill sulfur content could reveal a higher plasticity and sulfur storage capacity in A. gigas. Thus, the distribution of vesicomyid species across the chemosynthetic areas of the Guaymas Basin could be explained by differences in biological traits of the vesicomyid species that would allow A. gigas to more easily exploit transient and punctual sources of available sulfide than P. soyoae.     

Nematode species distribution patterns at the Håkon Mosby Mud Volcano (Norwegian Sea)

The Håkon Mosby Mud Volcano (HMMV) is a methane seep that is densely populated by the bacteria Beggiatoa spp. as well as by tubeworms of the Family Siboglinidae. Analyses of samples from seven stations located in five different habitats (bare sediments, bacterial mats, siboglinid fields, sediments with small patches of siboglinids and areas of ‘normal’ sea floor) revealed differences in the density and species‐level diversity of nematode communities. Total densities increased from stations inside the mud volcano and on its rim towards the ‘normal’ areas outside the volcano. Nematode species diversity was similar in samples from the siboglinid fields and the bare HMMV sediments. However, the sediment with patches of siboglinids collected outside the volcano was characterised by a diverse nematode community comprising 47 species, whereas the control sediments without siboglinids yielded only 27 species. The nematode assemblage at the bacterial mat site included only two identified species, Halomonhystera disjuncta and Sabatieria ornata, with H. disjuncta being strongly dominant. Terschellingia distlamphida, S. ornata and Aponema ninae dominated nematode communities at the siboglinid fields and in bare HMMV sediments. Species dominating at stations inside the caldera were found in comparable low densities in the sediment with siboglinid patches collected outside the volcano, but were missing in the background samples, where species of Endeolophos, Acantholaimus and Desmoscolex were dominant. Species diversity generally decreased with sediment depth. A subsurface‐dwelling nematode community was observed in the siboglinid fields and the bare sediments. Background sediments showed a sharp decline with 18–20 species at 0–2 cm versus three species at 3–5 cm sediment depth. The trophic composition of the nematode fauna at the HMMV showed a prevalence of deposit feeders at almost all stations, whereas background sediments exhibited a uniform distribution of non‐selective and selective deposit feeders and epistrate feeders. The high percentage of deposit feeders inside the mud volcano could be related to the grain‐size distribution and the significantly higher bacterial biomass compared to the control sediments.     

Cold-seep assemblages on a giant pockmark off West Africa: spatial patterns and environmental control

A giant pockmark colonised by dense cold‐seep assemblages near 3160 m depth along the Congo‐Angola margin has been surveyed by the ROV Victor 6000. The quantitative distribution of chemosynthetic communities was mapped along the dive tracks from a video study using GIS and image mosaicking. Several types of faunal assemblages, either dominated by bivalves of the families Mytilidae (Bathymodiolus sp.) or Vesicomyidae (Calyptogena sp., ‘Vesicomya’ aff. chuni), or by Siboglinidae polychaetes (Escarpia southwardae) were mapped over the 800‐m diameter pockmark area and sampled for fauna, water and sediment. The isotopic analyses (δ¹³C) of tissues from symbiont‐bearing species were within the range typical of nutrition via symbiosis using methane for mussels and sulphide for vesicomyids and siboglinids. The living chemosynthetic communities were distributed on a SW‐NE axis, corresponding to the expression at the sediment surface of a main buried channel providing fluids to the pockmark. The site was characterised by a more active central part in a depression with abundant carbonate concretions where high‐density clusters of siboglinids and mytilids dominate. Large fields of dead and live vesicomyids with a lower mean density were observed in the external areas. The mean coverage of each of the three symbiotic taxa in these two contrasted areas was estimated from mosaic analysis and was up to 30% in the central area dominated by E. southwardae bushes (23%). Symbiont‐bearing species distribution was consistent with methane concentrations in seawater that were generally higher in mytilid beds than in the vicinity of siboglinids and vesicomyids. A Principal Component Analysis performed on environmental factors at the ten sampling sites revealed that 37% of the observed variance in the distribution of symbiont‐bearing species may be explained by variation in both methane and oxygen concentrations, while a Canonical Redundancy Analysis selected methane concentration as the only variable which explains symbiont‐bearing species densities. This spatial distribution of chemosynthetic species at the pockmark scale may reflect temporal patterns of succession of both substrate and fauna, and may be related to different individual pockmarks visible on the microbathymetry mapped using ROV data.     

Similar reproductive cycles and life-history traits in congeneric limid bivalves with different modes of nutrition

We have studied two congeneric limid bivalves, Acesta species novum and Acesta excavata (Bivalvia: Limidae), that live in similar physical conditions but use different food sources. Acesta sp. nov. live on the giant siboglinid tubeworm Lamellibrachia luymesi at cold seeps and feed on their eggs, a continuous food source that ultimately is derived from chemosynthesis, whereas A. excavata relies on planktonic food of photosynthetic origin, which varies seasonally. We hypothesized that these two species might show differences in the periodicity of reproduction; specifically, we predicted that A. sp. nov. would breed continuously and that A. excavata would demonstrate an annual reproductive cycle. Our hypothesis was refuted, as both species have a semi‐continuous reproductive cycle. Food quality, quantity or seasonality had no clear effect. Reproductive adults are functionally gonochoristic. Females broadcast buoyant eggs with an average size of 179 μm in A. sp. nov. and 160 μm in A. excavata. Both the size of the larval shell and the size of the spawned eggs are suggestive of pelagic lecithotrophic development. Both species also appear to be protandric hermaphrodites, changing from male to female at approximately 77 and 90 mm shell height, respectively, although not all individuals change sex. In the material investigated in this study there was a biased sex ratio favouring males.     

Deep-sea hydrocarbon seep gastropod Bathynerita naticoidea responds to cues from the habitat-providing mussel Bathymodiolus childressi

Bathynerita naticoidea (Gastropoda: Neritidae) is a numerically dominant heterotrophic gastropod found at hydrocarbon seep sites on the upper Louisiana slope of the Gulf of Mexico. Snails of this species are commonly associated with beds of the methanotrophic mussel Bathymodiolus childressi (Bivalvia: Mytilidae), and their population structure mirrors that of the mussels they are found among. Previous studies have shown that these snails feed on bacteria and decomposing periostracum on the B. childressi shell. We predicted that B. naticoidea might be attracted to cues specific to its preferred habitat, such as dissolved methane, mucus from conspecific snails, or metabolites produced by B. childressi mussels. To examine this, we used a flow‐through Y‐maze system to investigate the behavior of B. naticoidea exposed to these potential cues. We found that the nerite is not attracted to methane, but is strongly attracted to seawater conditioned with B. childressi. The attractant appears to be specific to this type of mussel, and is not a soluble cue produced by conspecific snails.