Vertical changes in abundance,biomass and community structure of copepods down to 3000 m in the southern Bering Sea

Vertical changes in abundance, biomass and community structure of copepods down to 3000 m depth were studied at a single station of the Aleutian Basin of the Bering Sea (53°28′N, 177°00′W, depth 3779 m) on the 14th June 2006. Both abundance and biomass of copepods were greatest near the surface layer and decreased with increase in depth. Abundance and biomass of copepods integrated over 0–3000 m were 1,390,000 inds. m^?2 and 5056 mg C m^?2, respectively. Copepod carcasses occurred throughout the layer, and the carcass:living specimen ratio was the greatest in the oxygen minimum layer (750–100 m, the ratio was 2.3). A total of 72 calanoid copepod species belonging to 34 genera and 15 families occurred in the 0–3000 m water column (Cyclopoida, Harpacticoida and Poecilostomatoida were not identified to species level). Cluster analysis separated calanoid copepod communities into 5 groups (A–E). Each group was separated by depth, and the depth range of each group was at 0–75 m (A), 75–500 m (B), 500–750 m (C), 750–1500 m (D) and 1500–3000 m (E). Copepods were divided into four types based on the feeding pattern: suspension feeders, suspension feeders in diapause, detritivores and carnivores. In terms of abundance the most dominant group was suspension feeders (mainly Cyclopoida) in the epipelagic zone, and detritivores (mainly Poecilostomatoida) were dominant in the meso- and bathypelagic zones. In terms of biomass, suspension feeders in diapause (calanoid copepods Neocalanus spp. and Eucalanus bungii) were the major component (ca. 10–45%), especially in the 250–3000 m depth. These results are compared with the previous studies in the same region and that down to greater depths in the worldwide oceans.     

Seasonal changes in zooplankton abundance,biomass, size structure and dominant copepods in the Oyashio region analysed by an optical plankton counter

To identify seasonal patterns of change in zooplankton communities, an optical plankton counter (OPC) and microscopic analysis were utilised to characterise zooplankton samples collected from 0 to 150 m and 0 to 500 m in the Oyashio region every one to three months from 2002 to 2007. Based on the OPC measurements, the abundance and biomass of zooplankton peaked in June (0–150 m) or August (150–500 m), depending on the depth stratum. The peak periods of the copepod species that were dominant in terms of abundance and biomass indicated species-specific patterns. Three Neocalanus species (Neocalanus cristatus, Neocalanus flemingeri and Neocalanus plumchrus) exhibited abundance peaks that occurred before their biomass peaks, whereas Eucalanus bungii and Metridia pacifica experienced biomass peaks before their abundance peaks. The abundance peaks corresponded to the recruitment periods of early copepodid stages, whereas the biomass peaks corresponded to the periods when the dominant populations reached the late copepodid stages (C5 or C6). Because the reproduction of Neocalanus spp. occurred in the deep layer (>500 m), their biomass peaks were observed when the major populations reached stage C5 after the abundance peaks of the early copepodid stages. The reproduction of E. bungii and M. pacifica occurred near the surface layer. These species first formed biomass peaks of C6 and later developed abundance peaks of newly recruited early copepodid stages. From the comparison between OPC measurements and microscopic analyses, seasonal changes in zooplankton biomass at depths of 0–150 m were governed primarily by E. bungii and M. pacifica, whereas those at depths of 150–500 m were primarily caused by the three Neocalanus species.     

Depth zonation and bathymetric trends of deep-sea megafaunal scavengers of the Hawaiian Islands

The deep sea has been shown to exhibit strong depth zonation in species composition and abundance. Examination of these patterns can offer ecological insight into how organisms adapt and respond to changing environmental parameters that co-occur with depth. Here we provide the first tropical study on bathymetric zonation and other depth-related trends (size, abundance, and species richness) spanning shelf to abyssal depths of scavenging megafauna. Baited time-lapse free-vehicle cameras were used to examine the deep-sea benthic and demersal scavenging communities of the Hawaiian Islands, an area for which the biology and ecology have remained poorly studied below 2000 m. Twenty-two deployments ranging in depth from 250 to 4783 m yielded 37 taxa attracted to bait, including the first known occurrence of the family Zoarcidae in the Hawaiian Islands. Cluster analysis of Bray–Curtis similarity of species peak abundance (n_max) revealed four main faunal zones (250–500, 1000, 1500–3000, and ?4000 m) with significant separation (ANOSIM, global R=0.907, p=0.001) between designated depth groups. A major faunal break was identified at the 500–1000 m transition where species turnover was greatest, coinciding with the location of the local oxygen minimum zone. Dominance in species assemblage shifted from decapod crustaceans to teleosts moving from shallow to deeper faunal zones. Significant size differences in total length with depth were found for two of the four fish species examined. A logarithmic decline was observed in scavenger relative abundance with depth. Evidence of interaction between scavenging species was also noted between Synaphobranchus affinis and Neolithodes sp. (competition) and Histiobranchus sp. and aristeid shrimp (predation), suggesting that interactions between scavengers could influence indices of abundance generated from baited camera data.     

Serpulids living deep: calcareous tubeworms beyond the abyss

Although the carbonate compensation depth (CCD) for calcite, generally located in the depth range 4000–5000 m, is often proposed as a physiological barrier to deep-ocean colonization, many organisms with calcareous exoskeletons are found in the deepest oceanic trenches. Serpulid polychaetes inhabiting unprotected calcareous tubes are unlikely deep-sea inhabitants, yet, they are found at all oceanic depths from intertidal to hadal. Here we review and revise the published and unpublished records of Serpulidae from below 5000 m depth. We also describe tube ultrastructure and mineralogical content of available deep-sea serpulid tubes to obtain insights into their biomineralisation. Species belonging to the genera Bathyditrupa, Bathyvermilia, Hyalopomatus, Pileolaria (spirorbin) and Protis were found at depths from 5020 to 9735 m. However, only specimens of Protis sp. were truly hadal (>6000 m) being found at 6200–9700 m. Hadal specimens of Protis have irregularly oriented prismatic tube microstructure similar to that found in more shallow-water representatives of the genus. Initial EDX analysis suggested a mostly calcitic composition (i.e., the most stable CaCO₃ polymorph) on the basis of high Mg levels. Surprisingly, however, tubes of Bathyditrupa hovei and a species of Protis analysed using the more reliable method of laser Raman spectroscopy were found to be composed of aragonite. The compensation depth for this less stable CaCO₃ polymorph in the oceans is usually 2000–3000 m. We found no obvious structural adaptations to life at extreme depths in the studied serpulid tubes and how serpulids are able to biomineralise and maintain their tubes below the CCD remains to be explained.     

Growth rates,densities, and distribution of Lophelia pertusa on artificial structures in the Gulf of Mexico

Using industry inspection video and ROV imaging, we examined Lophelia pertusa (Linnaeus 1758) on 10 artificial structures of known ages (9 to 100 years) in the northern Gulf of Mexico (GoM). Five different types deep-water energy installations with depths ranging from 320 to 995 m, and three shipwrecks with depths ranging from approximately 530 to 615 m, were examined. Density, depth ranges, and growth rates of L. pertusa colonies were calculated from video and image analysis. L. pertusa colonies were present on all structures examined. Minimum calculated growth rates for the largest colonies ranged from 0.32 to 3.23 cm/yr on the different structures. The shallowest depth at which L. pertusa was observed was 201 m and the deepest was 801 m, considerably expanding the known depth range of this species in the northern GoM. Colony density varied with structure type, age, and depth, with the highest density between 503 and 518 m on the single structure that spanned the entire depth range of occurrence of L. pertusa observed in this study. L. pertusa colonies growing on thinner and deeper installations appear to have higher colonization rates, i.e. to develop higher densities over a shorter time period, compared to those on shallower and more massive types of installations. However, on average, colonies have slower growth rates on these installations than colonies on more massive, shallower installations (compliant and solid installations). In general, the calculated minimum growth rates were higher on the installations than on the shipwrecks, which were substantially older. A continuum of colony sizes was documented on all installations, suggesting multiple settlement events. L. pertusa thickets were observed on the oldest anthropogenic structures, with most of the components of these structures covered by colonies of L. pertusa. Brown, orange, and mottled color-variants were documented for the first time in the GoM. All installations examined for this study were colonized by L. pertusa and it is likely that most artificial surfaces in appropriate depths in the GoM will be as well.     

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.     

Geographical variation in metazoan parasites of the deep-sea fish Bathypterois mediterraneus Bauchot, 1962 (Osteichthyes: Ipnopidae) from the Western Mediterranean

This study examines the parasite fauna of Bathypterois mediterraneus, the most common fish below 1500 m in Western Mediterranean waters. Samples were obtained during July 2010 from the continental slope of two different areas (off Catalonia and Balearic Islands) in three different bathymetric strata at depths between 1000 and 2200 m. The parasite fauna of B. mediterraneus included a narrow range of species: Steringophorus cf. dorsolineatum, Scolex pleuronectis, Hysterothylacium aduncum, Anisakis sp. larva 3 type II and Sarcotretes sp. Steringophorus cf. dorsolineatum and H. aduncum were the most predominant parasites. H. aduncum showed significant differences in abundance between depths of 2000–2200 m with 1000–1400 m and 1400–2000 m, irrespective of locality, whereas S. cf. dorsolineatum showed significant differences between the two localities at all depths except for 2000–2200 m. We suggest the possible usefulness of these two parasites as geographical indicators for discriminating discrete stocks of B. mediterraneus in Western Mediterranean waters.     

Seasonal oxygen and carbon isotope variability in euthecosomatous pteropods from the Sargasso Sea

We examine seasonal variations in the stable carbon and oxygen isotopic composition of individual shells of the pteropods Limacina inflata and Styliola subula, collected from Oceanic Flux Program sediment traps (at 500 m depth) near Bermuda in the western Sargasso Sea. Calcification depths estimated from L. inflata δ¹⁸O vary between 200 and 650 m in late winter and spring, and between 50 and 250 m in late summer and fall. S. subula shows similar seasonal variability with calcification depths between 250 and 600 m in late winter and spring and 50–400 m in late summer and fall. These results suggest that both species calcify across a greater range of depths than indicated by previous geochemical studies. Furthermore, the data indicate that these species change their calcification depth in conjunction with changes in thermal stratification of the water column. Pteropod shell δ¹³C values vary inversely with δ¹³C_DIC but show a positive correlation with seawater [CO₃²⁻] and temperature after depth differences in δ¹³C_DIC are accounted for. We hypothesize that either the influence of temperature on metabolic CO₂ incorporation during shell growth and/or the influence of ambient [CO₃²⁻] on shell geochemistry can explain these relationships. Taken together, the individual shell δ¹⁸O and δ¹³C data suggest that shell calcification, and by inference the life cycle, of these pteropods is several months or less. Individual pteropod shell analyses have potential for contributing to our understanding of the environmental parameters that play a role in seasonal calcification depth shifts, as well as to our knowledge of past upper ocean thermal structure.     

Observations on depth distribution,diversity and abundance of pelagic nemerteans from the Pacific Ocean off California and Hawaii

Meso- and bathypelagic nemerteans were collected during five cruises west of Pt. Conception, California, and one west of Oahu, Hawaii, from depths of 0–3800 m, using a 10 m² Tucker Trawl modified with a thermally protected closing cod end, as well as MOCNESS-10 and MOCNESS-1D trawls. In this study, about 53 putative species were collected off California and about 23 off Hawaii, with little taxonomic overlap between the two groups, as compared to 16 and 0 species previously known from these regions. Numbers of nemertean individuals and species were greatest between 625 and 1750 m off California, and 1300–2500 m off Hawaii. Abundance of nemerteans off California was about 10 times that off Hawaii. The Shannon– Wiener diversity index (H′) was higher for Hawaii than for California due to numerical dominance by two species, Phallonemertes cf murrayi and Nectonemertes cf mirabilis, off California and a more even species distribution off Hawaii. Neither of the two dominant California species showed evidence of nocturnal vertical migration to shallow water. This is the first report of a Phallonemertes species from the Pacific and the first report of pelagic nemerteans from waters off Hawaii.     

Life history of the deep-sea cephalopod family Histioteuthidae in the western Mediterranean

The life cycle of the two species of the deep-sea family Histioteuthidae inhabiting the Mediterranean Sea (Histioteuthis reversa and Histioteuthis bonnellii) was studied from monthly samples taken throughout the year during daytime hours by bottom trawl gears. A small sample of individuals found floating dead on the sea surface was also analyzed. Both species were caught exclusively on the upper slope at depths greater than 300 m. Their frequency of occurrence increased with depth and showed two different peaks, at 500–600 m and 600–700 m depth in H. bonnellii and H. reversa, respectively, which might indicate spatial segregation. Maturity stages were assigned using macroscopic determination and confirmed with histological analyses. Although mature males were caught all year round, no mature females were found, which suggests that their sexual maturation in the western Mediterranean takes place deeper than the maximum depth sampled (800 m). In fact, the increase in mean squid size with increasing depth in H. reversa indicates an ontogenetic migration to deeper waters. The individuals of both species found floating dead on the sea surface were spent females which had a relatively large cluster of small atresic eggs and a small number of remaining mature eggs scattered in the ovary and mantle cavity. The sizes of these females were clearly larger than the largest individuals caught with bottom trawls. A total of 12 and 7 different types of prey, belonging to three major taxonomic groups (crustaceans, osteichthyes and cephalopods), were identified in the stomach contents of H. reversa and H. bonnellii, respectively. In both species fishes were by far the main prey followed by crustaceans, whereas cephalopods were found only occasionally. The preys identified, mainly myctophids and natantian crustaceans, indicate that both histioteuthids base their diet on pelagic nictemeral migrators.     

Depth distribution of algal species on the deep insular fore reef at Lee Stocking Island,Bahamas

Deep-water benthic algal composition and cover were studied with a submersible on the deep fore reef of Lee Stocking Island, Bahamas, from 45 to 150 m. Algal cover decreased from 57% to 16% over this depth range. Although there was substantial overlap in depth distributions, different species or groups of species dominated benthic cover at different depths. Lobophora and Halimeda copiosa co-dominated the fore reef from 45 to 60 m. A Corallinales/Peyssonnelia group was abundant from 60 to 120 m. The Corallinales/Peyssonnelia group shared dominance with Ostreobium between 90 and 120 m. Ostreobium was the only alga observed below 150 m and remained abundant below 200 m. Movement of sand down the fore reef is recognized as having substantial influence on algal cover.     

Population structure and species dynamics of Spisula solida,Diogenes pugilator and Branchiostoma lanceolatum along a temporal–spatial gradient in the south coast of Portugal

The population structure, dynamics and distribution of Spisula solida, Diogenes pugilator and Branchiostoma lanceolatum, common species in the south coast of Portugal, were studied in a spatial–temporal manner in order to understand the influence of cross-shore sediment transport and anthropogenic activities. Spisula solida is harvested commercially, whereas D. pugilator and B. lanceolatum are non-target species, with little information available on the population dynamics of these species. The study was performed in 2001–2002, and along a gradient of 100–5000 m from the coastline, corresponding to a depth gradient of between 1 and 32 m deep. Spisula solida was distributed preferentially at 3–12 m deep, and its distribution appeared to be influenced seasonally by the cross-shore sediment dynamics. Results suggested benthic recruitment in June. Some recruitments had no expression (year 2001) since adults were not present, which seems to be a direct impact of clams' fisheries. Diogenes pugilator showed preferential distribution at shallow depths, from 1.3 to 8 m. Results suggested recruitments every 4 months, in June, February and October. Branchiostoma lanceolatum showed the widest distribution, from 7 to 26 m deep. Recruitment seemed to start in June until October, when it attained an abundance peak of juveniles. For both non-target species no clear effects of the cross-shore dynamics or the clams' fisheries impact were visible.     

Deep-sea suprabenthos assemblages (Crustacea) off the Balearic Islands (western Mediterranean): Mesoscale variability in diversity and production

The composition of suprabenthic crustacean assemblages, their diversity, production (P) and production/biomass (P/B) ratios, were analyzed at species level along two transects situated to the north (N) and south (S) of Mallorca (Balearic Islands, western Mediterranean) at depths between 134 m and 760 m, based on a ca. bi-monthly sampling performed between August 2003 and June 2004. Differences with depth and season in assemblage composition and diversity were analyzed as a function of the contrasting environmental features (e.g. water mass dynamics) of the two areas. We identified 187 species (18 decapods, 5 euphausiids, 16 mysids, 76 gammaridean amphipods, 13 hyperiids, 1 caprellid, 21 isopods and 37 cumaceans). Substantial mesoscale variability in the deep-sea suprabenthic assemblages coupled with diversity trends between the N and S transects were found. Seasonality was the most important gradient influencing the dynamics of suprabenthos over the upper (350 m) and middle (650–750 m) slope in the N area. Conversely, the S area appeared to be more stable temporally with depth as the main gradient inducing assemblage differences. Different depth-related patterns were observed both for diversity and P/B. To the north diversity was very low at the shelf-break, increasing on the upper-slope (H′ > 3.00) and then decreasing again on the middle-slope. To the south diversity increased smoothly downward, reaching the highest values on the middle-slope. Regarding productivity, P/B was highest at intermediate depths to the north (over ca. 450–500 m), while to the south highest P/Bs were found deeper (over ca. 600–650 m). The higher P/B at intermediate depths found along N are likely due to higher % of organic matter (OM) in sediments, a product of oceanographic frontal systems. In particular, P/B was higher along N among omnivores and detritus feeders (e.g. Andaniexis mimonectes, Lepechinella manco and combined cumaceans), coupled to enriched OM in sediments, while along S mesoplanktonic carnivores (Rhachotropis spp.) had higher P/Bs. We conclude that on the north slope the influence of frontal systems and more active flow dynamics of different water masses (WIW and LIW) increases natural disturbance in the area, increasing productivity and diversity of suprabenthic peracarids in the Benthic Boundary Layer. Also, species showed a displacement of their average distributions (their Centres of Gravity, CoG) to shallower depths along N, which is another indicator of more favorable habitat conditions for suprabenthos in the 400–500 m range at N.     

Living benthic foraminifera of the Hess Rise and Suiko Seamount,central North Pacific

Rose-Bengal-stained benthic foraminifera in six pilot-core samples and one multicore sample collected from the Hess Rise and Suiko Seamount in August 1994 were studied in order to understand foraminiferal distributions between two areas divided by an oceanic front in the central North Pacific. Samples from the Hess Rise were collected in depths of 2167–3354 m under the warm, saline Kuroshio Extension, while samples from Suiko Seamount came from depths of 1811–1955 m under the cold, less-saline subarctic current. Sediment-trap results for the year prior to our sediment sampling show that organic matter fluxes were about 2.5 times greater at Suiko Seamount than at the Hess Rise. However, the hydrographic structure between 1800 and 3400 m, based on CTD observations, is almost the same at both sites. Temperature decreases from 2.2 to 1.7°C over the depth range of 1800–3400 m, salinity increases from 34.5 to 34.7, and the dissolved oxygen content gradually increases from 1.5 to 3.0 ml l⁻¹. The faunal populations at the Hess Rise are quite different from those at Suiko Seamount. The abundant species at the Hess Rise are Epistominella exigua, Brizalina pacifica, Fursenkoina cedrosensis and Alabaminella weddellensis. These species characteristically inhabit phytodetrital aggregates deposited on an oligotrophic seafloor. The populations at Suiko Seamount are dominated by Triloculina frigida, Lagenammina cf. arenulata, Reophax subfusiformis, and Reophax scorpiurus. The reason for differences between these populations is unclear. However, the typical phytodetritus-dwelling species E. exigua is dominant at the Hess Rise, which is located in a subtropical area that has a pulsed supply of settling organic matter in the spring. On the other hand, E. exigua is rare at Suiko Seamount, a subarctic site where there are more stable and greater fluxes of organic matter in summer and autumn. Occurrences of this species may be related to the seasonally short supply of organic matter that reaches the seafloor in the oceanic North Pacific.     

Contrasting life-histories,secondary production,and trophic structure of Peracarid assemblages of the bathyal suprabenthos from the Bay of Biscay (NE Atlantic) and the Catalan Sea (NW Mediterranean)

The life-histories and the secondary production of four dominant peracarid crustaceans (the mysids Boreomysis arctica and Parapseudomma calloplura, the amphipod Rhachotropis caeca, and the isopod Ilyarachna longicornis) in bathyal depths of the Bay of Biscay (NE Atlantic; between 383 and 420 m) and the Catalan Sea (Northwestern Mediterranean; between 389 and 1355 m) were established. Both the Atlantic and the Mediterranean populations of the major part of the target-species had two generations/year with mean cohort-production intervals (CPI) ranging from 5.5 mo for Ilyarachna longicornis to 6.3 mo for Parapseudomma calloplura. The Hynes method showed secondary production to vary in the Bay of Biscay between 0.113 mg DW m⁻² yr⁻¹ for I. longirostris and 3.069 mg DW m⁻² yr⁻¹ for P. calloplura, with P/B ratios between 4.57 (I. longirostris) and 7.93 (Boreomysis arctica). In the Catalan Sea, production varied between 0.286 mg DW m⁻² yr⁻¹ for I. longirostris and 1.096 mg DW m⁻² yr⁻¹ for P. calloplura, with P/B between 5.72 (I. longirostris) and 6.66 (P. calloplura). Application of two different empiric models to the whole peracarid assemblage gave similar levels of secondary production in both study areas (between 29.26 and 32.14 mgDWm⁻² yr⁻¹ in the Bay of Biscay; between 26.23 and 26.54 mg DW m⁻² yr⁻¹ in the Catalan Sea). From the analysis of gut contents of 22 species the dominant species in each study area were assigned to two basic trophic levels, detritus feeders and predators. Also, cumulative curves of dominance showed high diversity (low dominance) for peracarid assemblages distributed at mid-bathyal depths (524–693 m) both in the Bay of Biscay off Arcachon and in the Catalan Sea off Barcelona. We also discuss and compare, both within and between areas, how environmental features may explain the observed diversity patterns, the trophic structure, and the production results obtained for the suprabenthos assemblages.     

Factors influencing the abundance of deep pelagic bioluminescent zooplankton in the Mediterranean Sea

Measurements of the density of deep pelagic bioluminescent zooplankton (BL) were made with the Intensified Silicon Intensifier Target (ISIT) profiler in the Ligurian, Tyrrhenian, Adriatic, Ionian Seas and the Strait of Sicily from ～300 m to near seafloor. Mean BL densities ranged from 2.61 m^?3 at 500–1000 m depth in the Adriatic Sea to 0.01 m^?3 at 4000–5000 m depth in the E Ionian Sea. We investigated drivers of spatial variation of deep pelagic BL density. Linear regression was applied between surface chlorophyll a (Chl a) concentration and underlying BL density. Chl a values were determined from satellite derived 100 km radius composites (six 10-day means per ISIT deployment, over preceding 60 days). At 500–1000 m depth we found a significant positive relationship between mean BL density and mean Chl a in the period prior to 0–10 days (at 1% level) and prior to 10–40 days (at 5% level). Beyond 40 days no relationship between BL density and Chl a was found at this depth. At depths 1000–1500 m BL density values were low and no significant relationship with Chl a was detected. Generalised additive modelling (GAM) was used to assess the influence of benthic hotspots (seamount; cold water coral mound; mud volcano) on overlying BL density. A reduction in BL density was found downstream of the Palinuro seamount from 300 to 600 m. No effect on BL density in the overlying water column was detected from the presence of cold water corals. Higher BL densities were detected over the W Madonna dello Ionio mud volcano than at other sites sampled in the NW Ionian Sea. We find surface Chl a to be a good predictor of BL density in the mesopelagic zone; below this depth we hypothesise that processes affecting the efficiency of particle export to deep water may exert greater influence on BL density.     

Reproductive pattern and egg development of the deep-sea crab Paromola cuvieri (Brachyura,Homolidae) around the Canary Islands (NE Atlantic)

This work investigated the biology of Paromola cuvieri in the Atlantic Ocean, including reproductive patterns, sex structure, depth distribution and egg development. Females were more abundant than males, mainly at depths greater than 600 m. Males were dominant in size classes larger than 110 mm carapace width (CW), especially over 120 mm, for which only males were observed. The mean length of both sexes decreased with depth. P. cuvieri appears to have continuous reproduction throughout the year. Large females tend to spawn mainly between summer and winter, but smaller females spawn during the summer. Four maturity categories were identified based on the ovary colour, and these categories were histologically validated. Three categories of testes were identified according to their colour and morphology. Sexual maturity was estimated at sizes between 71.6 and 74.0 mm CW for females and at 91.0 mm CW for males. The relative growth showed changes along the ontogeny, at 73.6 mm CW (females) and 91.3–92.2 mm CW (males). Egg development appears to not be completely synchronous, and two colour patterns can be observed simultaneously. Four stages of egg development were found: eggs undivided and fully filled with yolk in orange egg masses; eggs with a free region of yolk visible in orange or red egg masses; embryos with slightly pigmented, crescent-shaped eyes in brown egg masses; and embryos with visible pigmented structures, enlarged eyes, segmented appendages and abdomen in brown egg masses. The mean number of external eggs carried by females in stage I was estimated to be 315,753±19,267. Three species of barnacle were observed on the exoskeleton: Poecilasma aurantia, Poecilasma crassa and Heteralepas microstoma (Cirripedia, Thoracica).     

Vertical zonation patterns of scavenging amphipods from the Hadal zone of the Tonga and Kermadec Trenches

Deep-sea trenches present an ideal system in which to study vertical zonation, though the difficulties inherent in studying these great depths have thus far deterred such research. Here, we present the first account of the structure and vertical zonation of the scavenging guild residing in the hadal habitat of the Tonga and Kermadec Trenches (SW Pacific Ocean). Four species of scavenging amphipod (Eurythenes gryllus, Scopelocheirus schellenbergi, Hirondellea dubia, and Uristes sp. nov.) were captured with baited free-vehicle traps set 1 m above the seafloor and deployed to bottom depths between 5155 and 10,787 m. Each species occupied a distinct vertical zone of 3.5 km or less, and the scavenging amphipod assemblage vertically partitioned the hadal zone. Scavenging amphipod diversity and evenness sharply declined below 9 km. For S. schellenbergi and H. dubia, population structure was stratified ontogenetically. Early instars appeared to reside exclusively in the shallower depths of each species’ vertical zone, and were functionally missing from the median and deeper regions. We captured brooding Uristes sp. nov. females, documenting the first bait-attendance event of brooding amphipods in the hadal zone. Separation of habitat in conjunction with deviations in reproductive traits, behaviors, and nutritional strategies may facilitate the coexistence of these four related species in this harsh and confining habitat.     

Large Vesicomyidae (Mollusca: Bivalvia) from cold seeps in the Gulf of Guinea off the coasts of Gabon,Congo and northern Angola

Two new genera and three new species of large Vesicomyidae are described from cold-seep sites on pockmarks and other sulfide-rich environments in the Gulf of Guinea (tropical east Atlantic) off Gabon, Congo (Brazzaville) and northern Angola, from 500 to 4000 m depth: “Calyptogena” (s.l.) regab n. sp., Wareniconcha (n.g.) guineensis (Thiele and Jaeckel 1931), Elenaconcha guiness n.g. n. sp., and Isorropodon atalantae n. sp. For two other species already taken by the R/V Valdivia in 1898, Calyptogena valdiviae (Thiele and Jaeckel 1931) and Isorropodon striatum (Thiele and Jaeckel 1931) new localities were discovered, and the species are rediscussed. E. guiness n.g. n.sp. is also recorded from off Banc d’Arguin, Mauritania, collected by commercial fishing vessels. The vesicomyid species here treated were encountered in different depth ranges along the Gabon–Congo–Angola margin, between 500 and 4000 m depth, and it was found that, in comparison with the dredge samples taken by the Valdivia expedition off southern Cameroon and off Rio de Oro (both at 2500 m), the same species occur in other depth ranges, in some cases with a vertical difference of more than 1000 m. .That means that the species are not confined to a given depth thought being typical for them and that the characteristics of the biotope are likely to play a major role in the distribution of the vesicomyids associated to cold seeps or other reduced environments along the West African margin.     

Spatial and temporal variability of POC in the northeast Subarctic Pacific

Particulate organic carbon (POC) concentrations from 0 to 1000 m were quantified in size-fractionated particulate matter samples obtained by the multiple unit large volume in situ filtration system (MULVFS) in 1996 and 1997 along the 1600 km long “line P” transect from continental slope waters near southern Vancouver Island to Ocean Station PAPA (OSP, 50°N, 145°W). Regression of in situ POC vs. beam attenuation coefficient, c, from a simultaneously deployed 1-m pathlength SeaTech transmissometer gave slope, intercept and r² values of 6.15±0.19×10⁻⁵ m⁻¹ (nmol C l⁻¹)⁻¹, 0.363±0.003 m⁻¹, and 0.951 (n=145), respectively. This result agreed within several percent of calibrations obtained from two 2600-km-long transects of the equatorial Pacific in 1992 (Bishop, 1999). Data from other, more frequently deployed transmissometers were standardized against the 1-m instrument, and the combined optical data set was used to document POC variability at finer spatial and temporal scales than could be sampled directly using either conventional water bottle casts or MULVFS. Published bottle POC vs. c relationships show much more variability and remain problematic. Along the line P transect in the salinity-stratified upper 100 m, POC isolines shoaled from winter to summer in concert with seasonal stratification. At the same time, POC was progressively enriched in subeuphotic zone waters to depths greater than 500 m. Near-surface POC fields sampled in the winter time showed strong temporal POC variability over time scales of days as well as between years. POC concentrations at OSP in February 1996 were higher than those found at any other time of year. Less variability was found along line P in other seasons. In May 1996, kilometer-scale spatial variability of POC at OSP was small; dawn vs. dusk variations of c were used to calculate 0–100 m POC turnover times shorter than 6 d. Calculations also suggest that 25–50% of primary productivity was expressed as dissolved organic carbon at OSP in May 1996.