Demersal Continental Shelf and Upper Slope Cephalopod Assemblages from the Balearic Sea (North-Western Mediterranean). Biological Aspects of Some Deep-Sea Species

The analysis of 79 hauls performed by commercial bottom trawlers from 50 to 800 m depth in the Balearic Sea (north-western Mediterranean) from June 1995 to September 1996 yielded a total of 30 cephalopod species belonging to 12 families. Cluster analysis of these data gave as a result two main groups 50–200 m and 200–800 m each subdivided into two other groups (50–100 vs. 100–200 m and 200–600 vs. 600–800 m). These results suggested the existence of two assemblages that could be associated to the continental shelf (50–100 m) and the upper slope (600–800 m) separated by a wide transitional zone (100–600 m) representing a region of overlapping shelf and slope faunas (ecotone). The faunistic bathymetric gradient showed a continuous substitution of species with depth rather than discrete assemblages separated by distinct boundaries. The more coastal species such as Eledone moschata, Loligo vulgarisSepia officinalis and Octopus vulgaris were found on the continental shelf; on the transitional zone, apart from species characteristic of this zone (Illex coindetii, Sepietta oweniana, Rossia macrosoma, Scaeurgus unicirrhus and Pteroctopus tetracirrhus), we also observed species from both the continental shelf and slope. The upper slope was characterized by typical species of deeper waters, such asBathypolypus sponsalis , Histioteuthis reversa, H. bonnellii, Ancistroteuthis lichtensteinii andOnychoteuthis banksii . The octopod O. vulgaris was the dominant species on the continental shelf and upper transitional zone, being substituted by T. sagittatus on the lower transitional zone and upper slope. Mean biomass decreased abruptly from the continental shelf to the transitional zone and from there to the upper slope. Mean species richness and species diversity were higher in the transitional zone than in the continental shelf and upper slope. Finally, some biological aspects of the more abundant deep-sea cephalopod species are studied: Bathypolypus sponsalis, Octopus salutii,Pteroctopus tetracirrhus , Histioteuthis reversa and H. bonnellii.     

Unexpected records of ‘deep‐sea’ carnivorous sponges Asbestopluma hypogea in the shallow NE Atlantic shed light on new conservation issues

Marine cave communities have been a continued source of ecological surprises, among other things because of their close ecological and evolutionary ties with the deep sea. The discovery of cladorhizid sponges, the deepest occurring poriferan family, in shallow Mediterranean caves in the 1990s was one such surprise, leading to the generally accepted hypothesis that the whole family was carnivorous, an unprecedented feeding mode for sponges. The recent observation of the cave species Asbestopluma hypogea in the Mediterranean bathyal, confirmed the view that some shallow caves can occasionally shelter otherwise deep‐dwelling species. Here we present new distribution data of A. hypogea, from deep Mediterranean locations, and for the first time from Atlantic locations. Among the new Atlantic records, the most surprising ones are located in three different geographic areas (Ria de Arousa, Groix Island and Cherbourg) of the NW European coasts, from the Iberian Peninsula to the English Channel, where A. hypogea reaches SCUBA depths (5–50 m), while not sheltered in marine caves. The carnivorous sponge however reaches its shallowest occurrence (5 m), in a small cave at Groix Island. The ecological significance of these discoveries, particularly the very patchy distribution and peculiar dynamics, are noteworthy, and the shallow occurrence of A. hypogea, together with other deep‐water or uncommon species, constitute unique assemblages that must be considered in conservation plans.     

Benthic megafaunal and demersal fish assemblages on the Chilean continental margin: The influence of the oxygen minimum zone on bathymetric distribution

Benthic megafaunal and demersal fish assemblages were sampled in three areas off Chile during the German–Chilean Expedition PUCK (SO-156) onboard the R/V Sonne from March to May 2001, at depths ranging from 120 to 2201 m. These samples, taken with an Agassiz trawl, are among the deepest ever taken in Chilean waters. A total of 147 species were recorded, mainly decapod crustaceans (Galatheidae, Pandalidae, Crangonidae), gastropods (Trochidae, Muricidae, Volutidae), ophiuroids (Asteronychidae, Gorgonocephalidae, Ophiolepididae, Ophiurinae), asteroids (Pterasteridae, Solasteridae, Goniopectinidae), polychaetes (Onuphidae, Aphroditidae, Maldanidae), and demersal fish (Macrouridae, Ipnopidae, Squalidae). Species richness and rarefaction analyses suggest that the fauna was undersampled. From the 147 species identified in this study, 36 species (24.5%) occurred only once and another 24 species occurred only twice (16.3%). Depth and dissolved oxygen levels were found to be the main factors influencing megafaunal changes along the continental shelf and in bathyal areas, as indicated by principal component and Pearson's correlation analyses. Some species appear to be limited to distinct areas in the upper and lower bathyal zones, whereas other species have a wider range, extending from the continental shelf to lower bathyal zones. Biogeographic relations exist with the Pacific, South Atlantic, and Southern Oceans, but the latter seem to be weaker than would be expected considering the connection by Antarctic intermediate water.     

Depth-related distribution and abundance of seastars (Echinodermata: Asteroidea) in the Porcupine Seabight and Porcupine Abyssal Plain,N.E. Atlantic

The depth-related distribution of seastar (Echinodermata: Asteroidea) species between 150 and 4950 m in the Porcupine Seabight and Porcupine Abyssal Plain is described. 47 species of asteroid were identified from ∼14,000 individuals collected. The bathymetric range of each species is recorded. What are considered quantitative data, from an acoustically monitored epibenthic sledge and supplementary data from otter trawls, are used to display the relative abundance of individuals within their bathymetric range. Asteroid species are found to have very narrow centres of distribution in which they are abundant, despite much wider total adult depth ranges. Centres of distribution may be skewed. This might result from competition for resources or be related to the occurrence of favourable habitats at particular depths. The bathymetric distributions of the juveniles of some species extend outside the adult depth ranges. There is a distinct pattern of zonation with two major regions of faunal change and six distinct zones. An upper slope zone ranges from 150 to ∼700 m depth, an upper bathyal zone between 700 and 1100 m, a mid-bathyal zone from 1100 to1700 m and a lower bathyal zone between 1700 and 2500 m. Below 2500 m the lower continental slope and continental rise have a characteristic asteroid fauna. The abyssal zone starts at about 2800 m. Regions of major faunal change are identified at the boundaries of both upper and mid-bathyal zones and at the transition of bathyal to abyssal fauna. Diversity is greatest at ∼1800 m, decreasing with depth to ∼2600 m before increasing again to high levels at ∼4700 m.     

Macrofaunal nematodes of the deep Whittard Canyon (NE Atlantic): assemblage characteristics and comparison with polychaetes

Large nematodes form an important component of deep‐sea macrofaunal assemblages, but are often considered to be part of the meiobenthos and are rarely studied. We analysed the standing stocks, diversity, and functional group and genus‐level composition of macrofaunal nematodes at lower bathyal depths (3,500 m water depth) in the Whittard Canyon system (NE Atlantic) and on the adjacent continental slope. Five replicate sample sets were obtained using a Megacorer, at four locations (three canyon branches, one slope). Sediment samples were processed on a 500‐μm mesh to provide both nematode and polychaete data from the same samples. The dominant nematode genera included Paramesacanthion, Metacylicolaimus, Cylicolaimus and Phanodermopsis. Nematode standing stocks (density and biomass) increased significantly from slope to canyon locations. Similarly, nematode dominance increased substantially (and diversity decreased) from slope to canyon locations. Nematode feeding groups and tail shape groups also appeared to exhibit common trends across study locations. Nematode genus‐level faunal composition varied significantly between slope and canyon locations. We describe and discuss the broadly similar trends detected in the matching polychaete data, noting some differences in the polychaete density and diversity responses. We suggest that the similar trends in macrofaunal nematode and polychaete ecology across our four study locations reflect responses to both changing sedimentology and food availability.     

Macrobenthic polychaete assemblages of the continental shelf and upper slope off the west coast of the South Island,New Zealand

Polychaete assemblages are described from replicate box‐core samples collected in summer 1983 at 18 stations on the continental shelf and upper slope (28–943 m) off the west coast of the South Island, New Zealand, south‐eastern Tasman Sea (c. 41–43°S, 169–172°E). Three main station groupings were identified by multivariate analysis: (1) inner shelf sandy stations characterised by Prionospio australiensis, Aricidea (Acesta) sp., Magelona cf. dakini, Paraprionospio aff. pinnata, Aglaophamus sp., Heteromastus filiformis, and Magelona sp.; (2) middle to outer shelf muddy stations characterised by Levinsenia cf. gracilis, Prionospio australiensis, Paraprionospio coora, Aglaophamus verrilli, and Auchenoplax mesos; and (3) upper slope sandy mud or mud stations characterised by Prionospio ehlersi. A combination of water depth and sediment clay content provided the best correlation with the biotic pattern. Spionidae was the most abundant family (49% of polychaete individuals), which may reflect the scope for opportunistic species in a shelf environment characterised by a high input of terrigenous sediment and episodic upwelling.     

Diversity patterns of free‐living marine nematodes in the southwest continental shelf off Bay of Bengal and their link to abiotic variables

Marine sediments in continental shelf ecosystems harbor a rich biodiversity of benthic communities. In this study, the spatial and temporal diversity and community assemblages of free‐living marine nematodes were studied by sampling at six depths and over 3 years from the southwest continental shelf off Bay of Bengal, one of the least explored tropical shelf ecosystems. The dominant marine nematode species were related with abiotic variables as part of this study. The effects of sediment granulometry generally decreased with increasing depth and the highest nematode density and species diversity were recorded on coarse sand (shallower depths). Multivariate analysis of the nematode community data showed that community structure differed significantly among depths as well as among years. Statistical analyses showed significant correlations between the nematode community and abiotic variables. Sediment texture, organic matter, water pressure and depth profile were crucial factors for determining diversity, vertical profile and feeding types of the nematode community. Other environmental factors, including anthropogenic pressure, did not have an effect on nematode diversity except for the presence of some tolerant species (Metachromadora spp., Sabatieria spp. and Siplophorella sp.). This study represents a baseline of knowledge of free‐living marine nematode communities that can be used in the future to compare nematode assemblages from temperate shelf ecosystems.     

Polychaete community structure in the South Eastern Arabian Sea continental margin (200–1000 m)

Macrofaunal polychaete communities (>500 µm) in the South Eastern Arabian Sea (SEAS) continental margin (200–1000 m) are described, based on three systematic surveys carried out in 9 transects (at ~200 m, 500 m and 1000 m) between 7°00′and 14°30′N latitudes. A total of 7938 polychaetes belonging to 195 species were obtained in 136 grab samples collected at 27 sites. Three distinct assemblages were identified in the northern part of the SEAS margin (10–14°30′N), occupying the three sampled depth strata (shelf edge, upper and mid-slope) and two assemblages (shelf edge and slope) in the south (7–10°N). Highest density of polychaetes and dominance of a few species were observed in the shelf edge, where the Arabian Sea oxygen minimum zone (OMZ) impinged on the seafloor, particularly in the northern transects. The resident fauna in this region (Cossura coasta, Paraonis gracilis, Prionospio spp. and Tharyx spp.) were characteristically of smaller size, and well suited to thrive in the sandy sediments in OMZ settings. Densities were lowest along the most northerly transect (T9), where dissolved oxygen (DO) concentrations were extremely low (<0.15 ml l⁻¹, i.e.<6.7 μmol l⁻¹). Beyond the realm of influence of the OMZ (i.e. mid-slope, ~1000 m), the faunal density decreased while species diversity increased. The relative proportion of silt increased with depth, and the dominance of the aforementioned species decreased, giving way to forms such as Paraprionospio pinnata, Notomastus sp., Eunoe sp. and lumbrinerids. Relatively high species richness and diversity were observed in the sandy sediments of the southern sector (7–9°N), where influence of the OMZ was less intense. The area was also characterized by certain species (e.g. Aionidella cirrobranchiata, Isolda pulchella) that were nearly absent in the northern region. The gradients in DO concentration across the core and lower boundary of the OMZ, along with bathymetric and latitudinal variation in sediment texture, were responsible for differences in polychaete size and community structure on the SEAS margin. Spatial and temporal variations were observed in organic matter (OM) content of the sediment, but these were not reflected in the density, diversity or distribution pattern of the polychaetes.     

The deep-sea macrobenthos on the continental slope of the northwestern Mediterranean Sea: a quantitative approach

As part of the ECOMARGE operation (J.G.O.F.S. France), macrobenthic assemblages in the Toulon Canyon were described and quantified on the basis of sampling carried out between 250 and 2000 m depth on the Mediterranean continental slope. Results show that Mediterranean bathyal assemblages are made up mainly of continental shelf eurybathic species. The qualitative and quantitative composition of populations varies with depth on the slope and also varies with station position at equivalent depth, whether on the flanks or in the canyon channel. Various analyses have provided evidence on the factors responsible for this population distribution pattern. No single factor emerges as predominant, but rather a group of factors, which are related to the nature and origin of sediments and more particularly their grain size distribution, geochemical composition and mode of transportation and sedimentation (benthic nepheloid or originating from the water column), act in conjunction to determine the pattern. Comparison with ocean continental slopes shows that in the Mediterranean Sea the absence of tidal current modifies the trophic structure of the macrobenthic assemblages, which are characterized by a dominance of surface and subsurface deposit feeders as compared to a dominance of suspension feeders and carnivores in the upper and median part of the slope in the ocean. Surface dumping of dredge spoil at the canyon head and channelling of waste induces an increase of organic matter and pollutant concentrations in sediment from the upper part of the canyon channel but does not give rise to any marked population degradation.     

Patterns of bathymetric zonation of bivalves in the Porcupine Seabight and adjacent Abyssal plain,NE Atlantic

Although the organization patterns of fauna in the deep sea have been broadly documented, most studies have focused on the megafauna. Bivalves represent about 10% of the deep-sea macrobenthic fauna, being the third taxon in abundance after polychaetes and peracarid crustaceans. This study, based on a large data set, examined the bathymetric distribution, patterns of zonation and diversity–depth trends of bivalves from the Porcupine Seabight and adjacent Abyssal Plain (NE Atlantic). A total of 131,334 individuals belonging to 76 species were collected between 500 and 4866 m. Most of the species showed broad depth ranges with some ranges extending over more than 3000 m. Furthermore, many species overlapped in their depth distributions. Patterns of zonation were not very strong and faunal change was gradual. Nevertheless, four bathymetric discontinuities, more or less clearly delimited, occurred at about 750, 1900, 2900 and 4100 m. These boundaries indicated five faunistic zones: (1) a zone above ∼750 m marking the change from shelf species to bathyal species; (2) a zone from ∼750 to 1900 m that corresponds to the upper and mid-bathyal zones taken together; (3) a lower bathyal zone from ∼1900 to 2900 m; (4) a transition zone from ∼2900 to 4100 m where the bathyal fauna meets and overlaps with the abyssal fauna and (5) a truly abyssal zone from approximately 4100–4900 m (the lower depth limit of this study), characterized by the presence of abyssal species with restricted depth ranges and a few specimens of some bathyal species with very broad distributions. The ∼4100 m boundary marked the lower limit of distribution of many bathyal species. There was a pattern of increasing diversity downslope from ∼500 to 1600 m, followed by a decrease to minimum values at about 2700 m. This drop in diversity was followed by an increase up to maximum values at ∼4100 m and then again, a fall to ∼4900 m (the lower depth limit in this study).     

Dense infaunal assemblages on the continental slope off Cape Hatteras, North Carolina

Unusually dense assemblages of benthic infaunal invertebrates have been discovered in continental slope sediments off Cape Hatteras, North Carolina. Densities were highest on the upper slope, ranging from 24,055 to 61,244 (X¯=46,255) individuals m⁻² in nine samples taken at a 600-m site in 1984 and 1985, and from 15,522 to 89,566 (X¯=37,282) individuals m⁻² in single samples at 15 stations over a wider depth range of 530 to 1535 m in 1992. A lower slope station at 2000 m sampled six times in 1984–1985 and again in 1992, had densities consistently higher than 8500 individuals m⁻². Species richness and diversity are consistently lower on the Cape Hatteras slope than at other locations off North Carolina and elsewhere in the western North Atlantic. The 1992 studies indicated that the upper slope infaunal assemblages (600m) were dominated by oligochaetes, while the middle slope assemblages (800–1400 m) were dominated by the polychaeteScalibregma inflatum. This latter depth range could be defined into two assemblages based upon suites of less abundant species. At depths of 1500–2000 m, a lower slope assemblage dominated by various deposit feeding polychaetes and oligochaetes was found. Results from the 1984–1985 studies suggest seasonal or year-to-year patterns in the dominance ofS. inflatum andCossura longocirrata. Unusually high sedimentation rates and organic carbon flux have been recorded from the slope off Cape Hatteras and may account for the high infaunal productivity in the area. Most of the dominant infaunal organisms are species more typical of shallow, coastal habitats rather than deep-sea species that dominate other areas of the U.S. Atlantic continental slope. Parallel investigations regarding the nature of organic matter in the Cape Hatteras sediments have revealed a mixture of both marine and terrestrially derived carbon, only a small percentage of which is composed of the smaller molecular weight polyunsaturated fatty acids more typical of continental slope sediments. It is likely that the high percentage of refractory organic matter would favor the survival of preadapted shelf species over those from adjacent slope environments.     

Protracted oogenesis and annual reproductive periodicity in the deep‐sea pennatulacean Halipteris finmarchica (Anthozoa,Octocorallia)

Halipteris finmarchica is one of the most common species of deep‐sea pennatulacean corals in the Northwest Atlantic; it was recently determined to act as a biogenic substrate for other species and as a nursery for fish larvae. Its reproductive cycle was investigated in colonies sampled in 2006 and 2007 along the continental slope of Newfoundland and Labrador (Canada). Halipteris finmarchica exhibits large oocytes (maximum diameter of 1000 μm), which are consistent with lecithotrophic larval development. Female potential fecundity based on mature oocytes just before spawning was ~6 oocytes · polyp^?1 (500–6300 oocytes · colony^?1); male potential fecundity was 16 spermatocysts · polyp^?1 (5500–57,400 spermatocysts · colony^?1). Based on statistical analysis of size‐probability frequency distributions, males harboured one cohort of spermatocysts that matured inside 8–11 months, whereas females harboured two distinct cohorts of oocytes; a persistent pool of small ones (≤400 μm) and a small number (~20%) of larger ones that grew from ~400 to >800 μm over a year. Despite this difference in the tempo of oogenesis and spermatogenesis, a synchronic annual spawning was detected. A latitudinal shift in the spawning period occurred from south (April in the Laurentian Channel) to north (May in Grand Banks and July–August in Labrador/Lower Arctic), following the development of the phytoplankton bloom (i.e. sinking of phytodetritus). Prolonged oogenesis with the simultaneous presence of different oocyte classes in a given polyp is likely not uncommon in deep‐sea octocorals and could hamper the detection of annual/seasonal reproduction when sample sizes are low and/or time series discontinued or brief.     

Demersal fish and habitat associations from visual surveys on the central California shelf

In 2004, we surveyed demersal fishes and habitats on the continental shelf off central California (65–110 m depth) using the occupied submersible Delta. Our objectives were to estimate the relative abundance of habitats and to examine demersal fish species composition, diversity, density, and sizes relative to these habitats. A total of 112 transects were completed covering 32 km of seafloor. A higher density of fishes was estimated in boulder and cobble habitats than in mud and brachiopod beds. More than 80% of the fishes were small, measuring 20 cm or less in total length. Species with small maximum size (primarily pygmy rockfish, Sebastes wilsoni, and blackeye gobies, Rhinogobiops nicholsii) accounted for nearly half (49%) of the total number of 12,441 fishes. Most fishes were immature, with only 4 of 20 harvested species having more than 50% of the individuals larger than the size at first maturity. Our study area on the continental shelf may be an ontogenetic transition zone for immature fishes before they move to their adult habitat on the slope. Alternatively, historical fishing pressure may have contributed to the lack of large, mature fishes in the survey area. Understanding the importance of these habitats to fishes at various life stages will improve our ability to assess these deepwater fish stocks and effectively manage these living resources on an ecosystem basis.     

Interranual variability in horizontal patterns of larval fish assemblages in the northeastern Aegean Sea (eastern Mediterranean) during early summer

Larval fish community structure was studied in the northeastern Aegean Sea (NEA) over an area influenced by the advection of Black Sea water (BSW). Sampling was carried out in early summer during a period of 4 years (2003–2006). Taxonomic composition and abundance presented high variability in space that remained relatively constant among years. Tow depth and indicators of trophic conditions in the upper water column (i.e., zooplankton displacement volume, fluorescence) explained significantly the structure of larval assemblages during all surveys. The northern continental shelf (Thracian and Strymonikos shelf), where a large amount of enriched, low salinity BSW is retained, was dominated by larvae of epipelagic species, mainly anchovy (Engraulis encrasicolus). Interannual changes in horizontal extension of the BSW seemed to match closely observed changes in the distribution of anchovy larvae. Mesopelagic fish larvae were particularly abundant beyond the continental shelf (over the North Aegean Trough) where a strong frontal structure is created between the low salinity waters of BSW origin and the high salinity waters of the Aegean Sea. Larvae of certain mesopelagic species (e.g., Ceratoscopelus maderensis) may occasionally be transported inshore when the prevailing current meanders towards the coast or feeds anticyclonic gyres over the continental shelf.     

白令海特征区域的表层沉积硅藻分布及其古海洋学意义

对白令海表层沉积物样品进行硅藻分析,共鉴定硅藻30属56种(含变种),并确定17种硅藻新记录,其中包括Kisseieviella carina等3种化石种。白令海表层沉积物中的硅藻优势种为Neodenticula seminae,Fragilariopsis cylindrus和Fragilariopsis oceanica,在白令海北部陆坡深水区附近以Neodenticula seminae为主,而在白令海北部陆架以Fragilariopsis cylindrus和Fragilariopsis oceanica为主。对硅藻结果进行聚类分析,可以划分出3个硅藻组合,硅藻组合Ⅰ代表海冰种硅藻组合,组合Ⅱ代表受阿拉斯加流影响的大洋浮游硅藻组合,组合Ⅲ代表上述两个硅藻组合之间的过渡组合。Fossula arctica是17种硅藻新记录之一,首次记录于白令海表层沉积物中,其百分含量分布趋势与Fragilariopsis cylindrus和Fragilariopsis oceanica的相近,在白令海北部陆架为11.7%~17.1%,而在陆坡深水区附近明显减少,是继Fragilariopsis cylindrus和Fragilariopsis oceanica之后白令海又一海冰指示种,并有望成为一种有效的海冰变化替代物运用于晚第四纪以来白令海海冰进退历史研究。     

Distribution of deep-water corals along the North American continental margins: Relationships with environmental factors

Despite the increasing attention to assemblages of deep-water corals in the past decade, much of this research has been focused on documenting and enumerating associated fauna. However, an understanding of the distribution of most species of coral and the ecological processes associated with these assemblages is still lacking. In this study, we qualitatively and quantitatively described the habitats of two families of deep-water corals in relation to six oceanographic factors (depth, slope, temperature, current, chlorophyll a concentration and substrate) on the Pacific and Atlantic Continental Margins of North America (PCM and ACM study areas, respectively). This study focused primarily on the distributions of Primnoidae and Paragorgiidae because of the large number of documented occurrences. For each environmental factor, deep-water coral locations were compared to the surrounding environment using χ² tests. On both continental margins, coral locations were found to be not randomly distributed within the study areas, but were within specific ranges for most environmental factors. In the PCM study area, Paragorgiidae and Primnoidae locations were found in areas with slopes ranging from 0° to 10.0°, temperature from −2.0 to 11.0 °C and currents from 0 to 143 cm s⁻¹. In the ACM study area, Paragorgiidae and Primnoidae locations were found in areas with slopes ranging from 0° to 1.4°, temperature ranging from 0 to 11.0 °C and currents ranging from 0 to 207 cm s⁻¹. Although the patterns in habitat characteristics were similar, differences existed between families with respect to particular environmental factors. In both study areas, most environmental parameters in locations where corals occurred were significantly different from the average values of these parameters as determined with χ² tests ($p<0.05$) except for substrate in Paragorgiidae locations and depth in Primnoidae locations on the PCM. This is the first study to show coral distributional patterns at the continental shelf/slope scale.     

Interactions between sediment chemistry and frenulate pogonophores (Annelida) in the north-east Atlantic

The small frenulate pogonophores (Annelida: Pogonophora a.k.a. Siboglinidae) typically inhabit muddy sediments on the continental slope, although a few species occur near hydrothermal vents and cold seeps. We present data on the distribution and habitat characteristics of several species on the European continental shelf and slope from 48°N to 75°N and show how the animals interact with the chemistry of the sediments. The environments inhabited include: shallow (30 m), organic-rich, fjord sediments; slope sediments (1000–2200 m) and methane seeps at 330 m depth. All the species studied obtain nutrition from endosymbiotic bacteria. They take up reduced sulphur species, or in one case, methane, through the posterior parts of their tubes buried in the anoxic sediment. We conclude that most species undertake sulphide ‘mining’, a mechanism previously demonstrated in the bivalves Lucinoma borealis and Thyasira sarsi. These pogonophores participate in the sulphur cycle and effectively lower the sulphide content of the sediments. Our results show that the abundance of frenulate pogonophores increases with increasing sedimentation and with decreasing abundance of other benthos, particularly bioturbating organisms. The maximum sustainable carrying capacity of non-seep sediments for frenulate pogonophores is limited by the rate of sulphate reduction.     

New Zealand xenophyophores: Upper bathyal distribution,photographs of growth position,and a new species

Xenophyophores are a group of giant, but extremely fragile, rhizopod protozoans generally found at lower bathyal or abyssal depths. Recent dredge samples and photographs suggest that there is a largely endemic fauna of xenophyophores with anastomosing branches living at upper bathyal depths around New Zealand. Their distribution may be related to the layer of Antarctic Intermediate water that bathes the upper slope of the New Zealand Plateau.

Photographs of the seafloor at depths of 800–1300 m on the Lord Howe Rise show hemispherical masses, 5 cm across, of which the majority appear to be Syringammina tasmanensis Lewis, 1966 and at least one specimen may be Reticulammina lamellata Tendal, 1972. The photographs indicate that these species live on the seabed, not beneath it, that they are probably filter feeders and that they can reach a density of about one specimen per square metre.

A new species, Reticulammina maini, is described from the continental slope east of Auckland. A key to the New Zealand species of xenophyophore is presented.     

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.     

Assessment of the effectiveness of natural coral fragmentation as a dispersal mechanism for coral reef‐boring sponges

Eastern Pacific reefs are mostly made up of interlocking coral branches of Pocillopora, which are easily broken by physical forces associated with heavy swells and winds. In this study we investigated the potential of these coral fragments to enable propagation of boring sponges. For this, we quantified the frequency of occurrence and diversity of boring sponges in fragments of corals recently trapped among the branches of live colonies, and later tested the hypothesis that these sponges colonize new branches of corals. Nearly 80% of the coral colonies investigated had coral fragments among their branches, and 69% of these coral fragments contained boring sponges (11 species), some of these sponges in reproduction (23% of them carried oocytes). To test whether sponges inhabiting coral fragments could colonize new branching corals we transplanted them to healthy branches, and to branches whose living tissue was mechanically eliminated to simulate damage produced by grazing and death after bleaching and other causes of coral tissue mortality. All the transplanted coral fragments cemented to each new colony by means of calcification, and of the three sponge species tested (Cliona vermifera, Cliona tropicalis and Thoosa mismalolli) only C. vermifera was able to colonize both new living branches (26.9%) and cleaned branches (65.5%). The apparent capability of C. vermifera to colonize by direct contact may be another key ability of this species to maintain high frequency of occurrence in Pacific coral reefs. However, although C. tropicalis and T. mismalolli were not able to colonize new coral substrata by direct contact, coral fragments have the potential to contribute to local persistence of these sponges and to their dispersal, both by asexual (fragments) and sexual means (transport of sexual products). The present findings may partly explain the current increase of excavating sponges on deteriorating reefs with a large availability of dead branching corals.