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.     

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.     

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.     

The Norwegian–Barents–Svalbard (NBS) continental margin: Introducing a natural laboratory of mass wasting, hydrates, and ascent of sediment, pore water, and methane

Side-scan sonar mapping and ground-truthing of the Norwegian–Barents–Svalbard continental margin shed new light on shelf glaciation, mass wasting, hydrates, and features like the Håkon Mosby mud volcano (HMMV), reflecting upward mobility of gas, pore fluids, and sediments. Detailed HMMV examination revealed thermal gradients to 10°/m, bottom-water CH₄ and temperature anomalies, H₂S- and CH₄-based chemosynthetic ecosystems, and subbottom methane hydrate (to 25%). Seismic and chemical data suggest HMMV origins at 2–3?km depth within the 6-km-thick depocenter. The HMMV and mound fields bordering the Bjørnøyrenna slide valley and pockmarks bordering the Storegga slide may all have formed in response to sediment failure.     

The ecology and distribution of benthic foraminifera at the Håkon Mosby mud volcano (SW Barents Sea slope)

To investigate a possible influence of submarine methane seepage on benthic foraminiferal communities, Rose Bengal stained (“live”) and empty tests of benthic foraminifera were studied from the sediment surface down to 15 cm sub-bottom depth of 12 sites at the Håkon Mosby mud volcano (HMMV). In addition, one reference site well away from the seep sites, but from similar water depths and the same general hydrographic setting was occupied for comparison. The HMMV is located at 1265 m water depth on the SW Barents Sea continental slope. Distinct living foraminiferal associations at the HMMV are linked to specific sedimentary, microbial, and macrofaunal habitats. In the center of the crater, and in crater areas completely covered by bacterial mats, Cassidulina reniforme is the only, albeit rare, living species. Below the top few millimeters, sediments are anoxic and devoid of living specimens. At the rim of the mud volcano, at sites densely populated by pogonophoran tube worms, three benthic foraminiferal associations are found; (i) a Fontbotia wuellerstorfi–Lobatula lobatula association living attached to the upper parts of pogonophoran tubes, which protrude into oxic water, (ii) a diverse Cassidulina neoteretis association populating dysoxic sediments of the surface centimeter, and (iii) a species-poor Bolivina pseudopunctata association colonizing the subsurface sediments down to four centimeters. Generally, we did not find endemic or seep indicative species or associations at the HMMV. However, the HMMV live faunas dominated by B. pseudopunctata are not found at the reference site nor are they described from comparable water depths and environments without gas seepages from the Norwegian-Greenland Seas.In the center and outer rim of the mud volcano, a C. neoteretis–Reophax guttifer dead association, similar to the one at the reference site, characterizes an assemblage of strongly corroded and partly displaced tests. At bacterial mat sites, a C. reniforme dead association corresponds to the living one. Thus both the living and the dead associations are indicative of a specific bacterial mat environment at the HMMV.     

Sea-floor terrains of Håkon Mosby Mud Volcano as surveyed by deep-tow video and still photography

A survey of the submarine Håkon Mosby mud volcano (HMMV) area by photo and video cameras permits the classification and mapping of sea-floor terrains. Approximate concentric zoning is seen in the distribution of the terrains, which correlates well with morphostructural elements of the mud volcano. A relatively limited biological community, dominated by tubeworms (Pogonophora and Polychaeta) and demersal fish, exists on the HMMV. Photo and video images show no evidence for gas bubbles in the water column, although methane is present in the mud volcano sediments. White patches, which comprise over 75% of the sea floor in some areas, are interpreted to be bacterial mats and/or gas hydrates.     

Microbial processes of carbon cycle as the base of food chain of Håkon Mosby Mud Volcano benthic community

Box and gravity cores recovered from the Håkon Mosby mud volcano (HMMV) during cruise 15 of the R/V Professor Logachev were analyzed for bacterial activity and benthic fauna distribution. The high bacterium number (up to 9.6×10⁹ cells cm^-3 of the sediment) and marked rates of sulfate reduction (up to 0.155?mg?S?dm^-3?day^-1) and methane oxidation (up to 9.9?μg?C?dm^-3?day^-1) were shown for the upper horizons of the sediments of the HMMV peripheral zone. The benthic community is characterized by the presence of two pogonophoran species, Oligobrachia sp. and Sclerolinum sp., harboring symbiotic methanotrophic bacteria.     

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.     

南海北部自由生活海洋线虫的群落结构和生物多样性

A quantitative study on the community structure and biodiversity of free-living marine nematodes and their relationship with environmental factors in the northern South China Sea were carried out based...     

Molluscan assemblages of seagrass-covered and bare intertidal flats on the Banc d'Arguin, Mauritania, in relation to characteristics of sediment and organic matter

The Banc d'Arguin, a non-estuarine area of shallows and intertidal flats off the tropical Saharan coast of Mauritania, is characterised by extensive intertidal and subtidal seagrass beds. We examined the characteristics of intertidal seagrass (Zostera noltii) meadows and bare areas in terms of the presence and abundance of molluscs (gastropods and bivalves). To explain observed differences between molluscan assemblages in seagrass and bare patches, some aspects of the feeding habitat (top-5 mm of the sediment) and of food (organic materials) of molluscs were examined. The novelty of this study is that phytopigments were measured and identified to assess source and level of decay (freshness) of organic material in the sediment and to study their importance as an explanatory variable for the distribution of molluscs. Over an area of 36 km² of intertidal flats, at 12 sites, paired comparisons were made between seagrass-covered and nearby bare patches. Within seagrass meadows, dry mass of living seagrass was large and amounted to 180 ±10 g AFDM m^− 2 (range 75–240). Containing twice the amount of silt per unit dry sediment mass, seagrass sediments were muddier than bare areas; the relative amount of organic material was also larger. The total number of species of bivalves and gastropods amounted to 27, 14 of which were found only in seagrass areas, 4 only in bare and 9 in both types of habitat. Among the three numerically most abundant species, the bivalves Anadara senilis, Dosinia hepatica and Loripes lacteus, the first was numerically most abundant in bare and the other two in seagrass-covered areas. Bare intertidal areas had greater mean total biomass of molluscs (80.5 g AFDM m^− 2) than seagrass meadows (30.0 g AFDM m^− 2). In both habitats, the bulk of the biomass was made up by A. senilis. Excluding this species, bare mudflats contained on average only 3.1 g AFDM m^− 2 and seagrass meadows 6.9 g AFDM m^− 2. As compared to previous surveys in 1980–1986, the biomass of A. senilis had increased almost 10-fold and D. hepatica, previously found in very small numbers, had become the most numerous species. However, the total biomass excluding that of A. senilis was similar. Concentrations of phytopigments were similar to those observed at temperate mudflats, indicating that the Banc d'Arguin might not be as oligotrophic as previously thought. Per unit of dry sediment mass, smaller amounts of phytopigments were found in bare than in seagrass areas. Per unit of dry organic material, bare sediments contained most (fresh) phytopigments. This suggests that in seagrass-covered meadows the organic material is more degraded than in bare sediments. Overall, the composition of phytopigments, quite surprisingly, indicated a benthic-diatom-dominated trophic system. Multivariate statistics revealed that patterns of zoobenthic assemblages were correlated with patterns of a combination of four environmental parameters: grain size of the sediment, amount of fresh phytopigments and amounts of leaves and roots of seagrass.     

Cold seep communities in the deep eastern Mediterranean Sea: composition, symbiosis and spatial distribution on mud volcanoes

Two mud volcano fields were explored during the French–Dutch MEDINAUT cruise (1998) with the submersible NAUTILE, one south of Crete along the Mediteranean Ridge at about 2000 m depth (Olimpi mud field) and the other south of Turkey between 1700 and 2000 m depth (Anaximander mud field) where high methane concentrations were measured. Chemosynthetic communities were observed and sampled on six mud volcanoes and along a fault scarp. The communities were dominated by bivalves of particularly small size, belonging to families commonly found at seeps (Mytilidae, Vesicomyidae, Thyasiridae) and to Lucinidae mostly encountered in littoral sulfide-rich sediments and at the shallowest seeps. Siboglinid polychaetes including a large vestimentiferan Lamellibrachia sp. were also associated. At least four bivalve species and one siboglinid are associated with symbiotic chemoautotrophic bacteria, as evidenced by Transmission Electronic Microscopy and isotopic ratio measurements. Among the bivalves, a mytilid harbors both methanotrophic and sulfide-oxidizing bacteria. Video spatial analysis of the community distribution on three volcanoes shows that dense bivalve shell accumulations (mainly lucinids) spread over large areas, from 10% to 38% of the explored areas (2500–15000 m²) on the different volcanoes. Lamellibrachia sp. had different spatial distribution and variable density in the two mud volcano fields, apparently related with higher methane fluxes in the Anaximander volcanoes and maybe with the instability due to brines in the Olimpi area. The abundance and richness of the observed chemosynthetic fauna and the size of some of the species contrast with the poverty of the deep eastern Mediterranean. The presence of a specialized fauna, with some mollusk genera and species shared with other reduced environments of the Mediterranean, but not dominated by the large bivalves usually found at seeps, is discussed.     

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.     

The impact of Melinna palmata grube, a tube-building polychaete, on meiofaunal community structure in a soft-bottom subtidal habitat

Field investigations were performed in a low-energy subtidal muddy sediment in Loch Creran, a sea-loch on the west coast of Scotland where the impact of Melinna palmata, a tube-building polychaete, on the meiofaunal community structure was assessed. This polychaete was found in high densities and forms prominent faecal casts on the sediment surface. Samples were retrieved from the casts, the feeding area of the polychaete and from an unaffected control area. The number of nematodes, the dominant taxon, did not vary significantly between areas. Of the five most abundant nematode species one was significantly more abundant in the cast area. Diversity and trophic structure of the nematode component of the meiofauna was very similar in all three areas. Copepods were more than twice as abundant in background sediments as in faecal mounds and feeding areas; however, Longipedia spp. were more abundant in the cast areas. For the copepods these findings support models which state that natural disturbances create a mosaic of patches that have different species compositions depending on their states of recovery.     

Microhabitat type determines the composition of nematode communities associated with sediment-clogged cold-water coral framework in the Porcupine Seabight (NE Atlantic)

The nematofauna associated with a cold-water coral degradation zone in the Porcupine Seabight (NE Atlantic) was investigated. This is the first comprehensive study of nematodes associated with cold-water corals. This research mainly aimed to investigate the influence of microhabitat type on nematode community structure. Three distinct microhabitats for nematodes were distinguished: dead coral fragments, glass sponge skeletons and the underlying sediment. The nematode assemblages associated with these three microhabitats were significantly different from each other. Coral and sponge substrata lie relatively unprotected on the seafloor and are consequently more subjected to strong currents than the underlying sediment. As a result, both large biogenic substrata were characterized by higher abundances of taxa that are less vulnerable and more adapted to physical disturbance, whereas the underlying sediment yielded more slender, sediment-dwelling taxa. Typically epifaunal taxa, such as Epsilonematidae and Draconematidae, were especially abundant on dead coral fragments, where they are thought to feed on the microbial biofilm which covers the coral surface. Several epifaunal genera showed significant preferences for this microhabitat, and Epsilonema (Epsilonematidae) was dominant here. Sponge skeletons are thought to act as efficient sediment traps, resulting in a lower abundance of epifaunal taxa compared to coral fragments. The underlying sediment was dominated by taxa typical for slope sediments. The considerable degree of overlap between the communities of each microhabitat is attributed to sediment infill between the coral branches and sponge spicules. It is assumed that the nematofauna associated with large biogenic substrata is composed of a typical sediment-dwelling background community, supplemented with taxa adapted to an epifaunal life strategy. The extent to which these taxa contribute to the community depends on the type of the substratum. Selective deposit feeders were dominant on sponge skeletons and in the underlying sediment, whereas coral fragments were dominated by epistratum feeders. The presence of a microbial biofilm on the coral fragments is proposed as an explanation for the significant preference of epistratum feeders for this microhabitat. Densities in the underlying sediment were low in comparison with other studies, but biodiversity was higher here than on the coral and sponge fragments, a difference which is attributed to lower disturbance. Nevertheless, the large biogenic substrata provide a microhabitat for rare, epifaunal taxa, and fragments of both substrata within the sediment increase habitat complexity and hence biodiversity.     

Benthic microalgal biomass in sediments of Onslow Bay, North Carolina

Sediment samples were collected at stations along cross-shelf transects in Onslow Bay, North Carolina, during two cruises in 1984 and 1985. Station depths ranged from 11 to 285 m. Sediment chlorophyll a concentrations ranged from 0·06 to 1·87 μg g⁻¹ sediment (mean, 0·55), or 2·6–62·0 mg m². Areal sediment chlorophyll a exceeded water column chlorophyll a a at 16 of 17 stations, especially at inshore and mid-shelf stations. Sediment ATP concentrations ranged from 0 to 0·67 μg g⁻¹ sediment (mean, 0·28). Values for both biomass indicators were lowest in the depth range including the shelf break (50–99 m). Organic carbon contents of the sediments were uniformly low across the shelf, averaging 0·159% by weight. Photography of the sediments revealed extensive patches of microalgae on the sediment surface.Our data suggest that viable benthic microalgae occur across the North Carolina continental shelf. The distribution of benthic macroflora on the North Carolina shelf indicates that sufficient light and nutrients are available to support primary production out to the shelf break. Frequent storm-induced perturbations do not favour settling of phytoplankton, an alternative explanation for the presence of microalgal pigments in the sediments. Therefore, we propose that a distinct, productive benthic microflora exists across the North Carolina continental shelf.     

Meiofauna in a cold-seep community off Hatsushima,central Japan

The community structure of the bathyal meiofauna of a cold-seep community found off Hatsushima in Sagami Bay, central Japan, was compared with the community composition outside the influence of the seep, using sediments collected during dives 226 and 227 of the deep-sea submersibleShinkai 2000. The sediment from the Hatsushima seep site (HSS) was very coarse, black in color, and with an odor of hydrogen sulfide, suggesting reduced thiobiotic conditions. The sediment from the control area was well-oxygenated, fine silt. Despite the differences in the characteristics of the sediments, the abundance of meiofauna in the HSS was not very different from that in the control area. However, its composition even at the major taxonomic group level was distinct; for example, a high nematode/copepod ratio occurred in one of the samples collected at the HSS. At the species level, nematodes were less diverse at the HSS than at the control area. The composition of the nematode fauna at the HSS showed stronger affinity with that collected at the adjacent control area than with a community sampled from other deep-sea environments or another seep community in shallow water. This emphasizes that the adaptation of nematodes to the thiobiotic condition is controlled by local conditions.     

Biodiversity patterns of free-living marine nematodes in a tropical bay: Cienfuegos, Caribbean Sea

Spatial and temporal biodiversity patterns of free-living marine nematodes were studied in Cienfuegos Bay, a tropical semi-enclosed basin in the Caribbean Sea. Taxonomic (to species level) and functional (biological trait) approaches were applied for describing the assemblage structure and relating it to abiotic environment based on a sampling scheme in six subtidal stations and three months. Biological trait approach added relevant information to species pattern regarding relationships between diversity patterns and the abiotic environment. The most common morphotypes were deposit feeding nematodes, with colonising abilities of 2–3 (in a scale from 1 to 5), tail conical cylindrical or filiforme and body slender; and their abundance were correlated with depth, organic matter and silt/clay fraction. In spite of a high turnover of species, functional diversity of assemblages did not change notably in space and time. A result probably due to sampling of the habitat pool of species and to low heterogeneity of the studied muddy bottoms. Chemical pollution (organic enrichment and heavy metals) and hydrodynamic regime possibly drove the biodiversity patterns. Spatial distribution of assemblages support the existence of two well differentiated basins inside the bay, the northern basin more polluted than the southern one. The low hydrodynamic regime would determine a poor dispersion of nematodes resulting in high spatial variance in the assemblage structure; and also the associated hypoxic conditions and pollutants in sediments can explain the dominance of tolerant nematode species such as Daptonema oxycerca, Sabatieria pulchra, Terschellingia gourbaultae, and Terschellingia longicaudata. A comparison of spatial–temporal patterns of biodiversity between Cienfuegos Bay and other semi-enclosed bays in temperate regions suggests several similarities: nematode assemblages are strongly influenced by anthropogenic disturbance, temporal trends are weak or overridden by spatial ones, and few cosmopolitan genera/species tolerant to pollution and hypoxic conditions are dominant.     

Small is beautiful: An inverted view of habitat fragmentation in seagrass beds

An intertidal Zostera marina landscape in Torbay, Devon, UK, was sampled to investigate the relationship between patch size, diversity and infaunal assemblage composition with the intention of defining a minimum Zostera patch size where the infaunal seagrass assemblage becomes distinct from the bare sand assemblage. All Zostera patches were found to support a higher level of biodiversity than the surrounding bare sand. However, the size of the Zostera patch had no impact on the level of diversity; it was just the presence or absence of seagrass that made a difference. The sediment and seagrass variables were not significantly different across the range of Zostera patch sizes, indicating that the environment characteristics were homogeneous within the Zostera patches at the patch scale. Multivariate analysis revealed that assemblage composition did vary between the patch types, although the opportunistic polychaete Capitella capitata was present in all patch types and was the most abundant species overall. The presence of opportunistic species and the homogeneity of the Zostera patch variables may be due to the location of this intertidal seagrass bed, which is relatively exposed compared to the locations of other seagrass beds along the south coast of Devon, resulting in a more dynamic and disturbed environment. Nevertheless, our results demonstrate that even small patches of seagrass comprising a few plants support a higher abundance and diversity of infaunal invertebrates than bare sand, indicating that Zostera patches have conservation value whatever their size.     

The impact of implanted whale carcass on nematode communities in shallow water area of Peter the Great Bay (East Sea)

In May, 2007 we sank the remains of a Minke whale (Balaenoptera acutorostrata) in the East Sea, Peter the Great Bay, at 30 m of water near the coast of Big Pelis Island. In the present study we describe the nematode communities in sediments under the implanted whale carcass. Abundance of nematodes increased with the distance from the carcass. Dominant trophic group was non-selective deposit feeders. The highest values of indexes of a specific diversity and evenness were noted in sediments under the whale, while domination index occurred at the highest distance from the whale. The suggestion is made that the cause of low density of nematodes in sediments under the whale is an extreme increase in number of macrofaunal animals, and predation and food competition between macro- and meiofauna. The changes noted in nematode assemblages living in an implanted whale in shallow waters are similar to those in deep-sea assemblages.     

Trophic resource use by macrozoobenthic primary consumers within a semi-enclosed coastal ecosystem: Stable isotope and fatty acid assessment

The diet of different macrozoobenthic trophic groups was investigated in the Arcachon Bay—a semi-enclosed macrotidal ecosystem that shelters the largest Zostera noltei seagrass meadow in Europe—in early spring and late summer 2009, using stable isotopes and fatty acids. Fatty acid profiles and literature information about the biology and physiology of benthic consumers were combined to identify the main organic matter sources for the benthic primary consumers. An isotope mixing model was then run to evaluate the contribution of each organic matter source to each identified trophic group (suspension feeders, sub-surface deposit feeders, micro-and macrograzers, suspension-oriented interface feeders and deposit-oriented interface feeders). Variations in organism' diets with respect to both habitats (intertidal seagrass meadows, intertidal bare sediments and subtidal bare sediments) and study periods were also investigated. At the scale of this study, it appeared that the diet of macrozoobenthos primary consumers was based exclusively on autochthonous material (no use of terrestrial organic matter): mainly microphytobenthos, seagrasses and their epiphytes, and phytoplankton. In addition, the different trophic groups relied on different organic matter pools: for instance, suspension feeders mainly fed on microphytobenthos and phytoplankton, whereas subsurface deposit feeders fed on microphytobenthos, decayed seagrasses and bacteria, and grazers mainly fed on microphytobenthos, and seagrasses and their epiphytes. The same pattern was observed in both early spring and late summer, indicating a stability of the benthic system at a six-month time scale. Finally our results showed that, in Arcachon Bay, the seagrass meadow directly or indirectly (through detritus) plays a significant role in the diet of most benthic consumers.