Trophic relationships of hydrothermal vent and non-vent communities in the upper sublittoral and upper bathyal zones off Kueishan Island,Taiwan: a combined morphological,gut content analysis and stable isotope approach

This study used morphological, gut content analysis and carbon- and nitrogen-stable isotope analysis to investigate the trophic structure of upper sublittoral (15–30 m deep) and upper bathyal (200–300 m deep) hydrothermal vents and the adjacent non-vent upper bathyal environment off Kueishan Island. The sublittoral vents host no chemosynthetic fauna, but green and red algae, epibiotic biofilm on crustacean surfaces, and zooplankton form the base of the trophic system. Suspension-feeding sea anemones and the generalist omnivorous vent crab Xenograpsus testudinatus occupy higher trophic levels. The upper bathyal hydrothermal vent is a chemoautotrophic-based system. The vent mussel Bathymodiolus taiwanensis forms a chemosynthetic component of this trophic system. Bacterial biofilm, surface plankton, and algae form the other dietary fractions of the upper bathyal fauna. The vent hermit crab Paragiopagurus ventilatus and the vent crab X. testudinatus are generalist omnivores. The vent-endemic tonguefish Symphurus multimaculatus occupies the top level of the trophic system. The adjacent non-vent upper bathyal region contains decapod crustaceans, which function as either predators or scavengers. The assemblages of X. testudinatus from sublittoral and upper bathyal vents exhibited distinct stable isotope values, suggesting that they feed on different food sources. The upper bathyal Xenograpsus assemblages displayed large variations in their stable isotope values and exhibited an ontogenetic shift in their δ¹³C and δ¹⁵N stable isotope signatures. Some individuals of Xenograpsus exhibited δ¹⁵N values close to those of non-vent species, suggesting that the highly mobile Xenograpsus may transfer energy between the upper bathyal hydrothermal vents and the adjacent non-vent upper bathyal environment.     

Hydrothermal vent communities at the shallow subpolar Mid-Atlantic ridge

A new type of animal community has been found near hot vents in the subpolar Atlantic at 100 to 106 m depth off Kolbeinsey on the Jan-Mayen ridge. Incubation of high temperature fluids yielded cultures of undescribed hyperthermophilic eu- and archaebacteria, growing in a temperature range between 70° and 110°C depending on the isolates. Bacteria are closely related to species occurring within deep sea hydrothermal areas. In contrast to deep-sea vent sites of the Mid-Atlantic and other oceans, the Kolbeinsey macro- and meiofauna consists of species reported from non-vent areas in the boreal Atlantic and adjacent polar seas. The most abundant forms are a solitary hydroid polyp and two sponges. Kolbeinsey is an isolated and young area of hydrothermal activity at relatively low depth and in highly productive waters; these findings could indicate a model for an early evolutionary step towards the formation of a genuine specialized vent community.     

Identification of archaeogastropod larvae from a hydrothermal vent community

Dispersal is essential in order that endemic species living in ephemeral, patchy vent environments may persist over evolutionary time. Quantitative field studies of larval dispersal, however, require specieslevel identification of the larval forms because each individual must be distinguished from related vent species, and from non-vent species living in the surrounding deep-sea environment. Methods for culturing these larvae to an identifiable stage have not yet been developed. To solve the larval identification problem for the archaeogastropod molluscs (a prominent component of vent communities), we used a scanning electron microscope (SEM) to image shells of larvae collected in the water column near vents along the East Pacific Rise (9°40′ to 9°50′N; 104°W). Larval shell size, shape and ornamentation were compared to protoconchs retained in juvenile or adult shells of identified species, and used to assign five larval groups unequivocally to species (Cyathermia naticoides Warén and Bouchet, 1989; Neomphalus fretterae McLean, 1981; Clypeosectus delectus McLean, 1989; Rhynchopelta concentrica McLean, 1989; and Lirapex granularis Warén and Bouchet, 1989) and seven groups tentatively to species or genus [Lepetodrilus spp. (three groups); Gorgoleptis sp; Peltospira ?operculata McLean, 1989; and ?Melanodrymia sp. (two groups)].     

Stable isotopic compositions of hydrothermal vent organisms

Stable isotopic analyses were used to study trophic relationships in two communities of deep-sea hydrothermal vent organism in the Pacific Ocean. The community at Hanging Gardens on the East Pacific Rise (21°N), sampled in 1985, is dominated by two species of vestimentiferan tubeworms; communities at Alice Springs and Snail Pits on the Marianas Back Arc Spreading Center (western Pacific), sampled in 1987, are dominated by gastropod mollusks, barnacles, and anemones. In both locations, carbon and nitrogen isotopic values of vent invertebrates are significantly different from those of non-vent invertebrates collected at 11°N on the East Pacific Rise and elsewhere in the deep-sea. These distinct isotopic compositions reflect local sources of organic carbon and nitrogen used by vent consumers. Many vent invertebrates lacking chemoautotrophic endosymbionts have ¹³C-enriched values of-11 to-16%. compared to values of-17 to-22%. normally observed in deep-sea fauna. This suggests that a ¹³C-enriched food source is trophically important in both vent communities. Free-living bacteria colonizing surfaces and suspended in the water column may constitute this food resource. Nitrogen isotopic analyses show that the food web of the East Pacific Rise community has more trophic levels than the Marianas vent community.     

POM in macro-/meiofaunal food webs associated with three flow regimes at deep-sea hydrothermal vents on Axial Volcano,Juan de Fuca Ridge

Deep-sea hydrothermal vent ecosystems host both symbiotic and non-symbiotic invertebrates. The non-symbiotic vent fauna is generally assumed to rely on free-living chemoautotrophic bacteria as their main food source but other sources such as detritus have recently been suggested to be a part of the invertebrate diets. Little is known about how food availability influences the distribution of vent organisms on a small scale. In addition, the feeding ecology and role of small, often numerically dominant invertebrates, the meiofauna is poorly understood at vents. In this study, we used stable carbon and nitrogen isotopic analysis to investigate the role of particulate detritus in the diets of macro- and meiobenthic invertebrates within three vent assemblages at Axial Volcano, Juan de Fuca Ridge, and Northeast Pacific. Particulate organic matter of a detrital origin became more important in the diet of invertebrates in assemblages typically associated with low-hydrothermal flow intensities. Meiobenthic species occupied several different feeding guilds and trophic levels in the assemblages investigated. We conclude that small-scale spatial variability in food sources is an important feature of vent food webs and that spatial patterns observed here and elsewhere are shaped by variations in hydrothermal discharge.     

Molecular taxonomy of vestimentiferans of the western Pacific,and their phylogenetic relationship to species of the eastern Pacific III. Alaysia-like vestimentiferans and relationships among families

The phylogenetic status of Alaysia-like vestimentiferans, which were collected at eight sites in the western Pacific, was analyzed on the basis of the nucleotide sequence of part of a mitochondrial gene for cytochrome oxidase I. The 123 individuals analyzed were tentatively classified into four species, which inhabit, respectively, seep areas off the central part of the Japanese mainland, hydrothermal vent fields in the Mid-Okinawa Trough, hydrothermal vent fields in the southern Okinawa Trough, and both seep and vent sites in the Bismarck Sea. From six additional lamellibrachiid specimens from two sites in the Bismarck Sea, two new tentative species were recognized, namely, one from a seep area off Papua New Guinea and another from the hydrothermal vent field at the DESMOS site in the Manus Basin. Phylogenetic analysis using all available sequences of six vestimentiferan families and Alaysia-like vestimentiferans revealed that vestimentiferans could be divided into three groups, namely, lamellibrachiids, escarpiids, and others, even though the monophyly of the third group was supported by only low bootstrap probabilities. Within the first group, the earliest divergence of a tentative lamellibrachiid species from the DESMOS site was apparent. All Alaysia-like vestimentiferans formed a monophyletic group with Arcovestia ivanovi from the Manus Basin, and it is suggested that this group might be derived from vestimentiferans that are endemic to the hydrothermal sites in the Eastern Pacific Rise.     

Influence of habitat on the reproductive biology of the deep-sea hydrothermal vent limpet <Emphasis Type="Italic">Lepetodrilus fucensis</Emphasis> (Vetigastropoda: Mollusca) from the Northeast Pacific

Habitat selection by the hydrothermal vent limpet, Lepetodrilus fucensis, in Northeast Pacific hydrothermal vent ecosystems, may influence its reproductive output, as it occupies habitats with varying physico-chemical conditions that reflect the availability of nutritional resources. Histological techniques were used to determine size at first reproduction, gametogenesis, reproductive output, and fecundity in relation to shell length (SL), through examination of the gonads of male and female L. fucensis, collected from five different hydrothermal vent habitat types with different temperature anomalies and hydrothermal fluid flow vigour: vigorous (VIG), diffuse (DIF), tubeworm bushes (TWB), peripheral (PER), and senescent areas (SEN). Both male and female L. fucensis exhibited early maturity, with the first reproductive event occurring at 3.8 and 3.9 mm shell length, respectively. All stages of gamete development were present in the gonads of males and females, suggesting continuous gametogenesis and asynchronous reproduction in this species. Gametogenic maturity of limpets did not vary among actively venting habitats (VIG, DIF, TWB, and PER), but was significantly lower in males and females from SEN habitats. Mean oocyte diameter was largest in females from VIG habitats, and smallest in females from SEN habitats, than in those from the other habitats (DIF, TWB, and PER). Females from actively venting habitats also had greater actual fecundity than those from senescent habitats. While the gametogenic pattern of L. fucensis appears phylogenetically constrained, selection of actively venting habitats by L. fucensis maximizes its reproductive output. The multiple feeding strategies of L. fucensis may allow for a constant supply of energy to be allocated to reproduction in any habitat except senescent vents. Early maturity, high fecundity, and continuous production of gametes suggest a reproductive strategy characteristic of an opportunistic species, and may be contributing to the extremely abundant populations of L. fucensis observed in the Northeast Pacific vent ecosystem.     

Evolution of habitat use by deep-sea mussels

Previous phylogenetic studies proposed that symbiont-bearing mussels of the subfamily Bathymodiolinae (Bivalvia: Mytilidae) invaded progressively deeper marine environments and evolved from lineages that decomposed wood and bone to specialized lineages that invaded cold-water hydrocarbon seeps and finally deep-sea hydrothermal vents. To assess the validity of the hypotheses, we examined two nuclear (18S and 28S rRNA) and two mitochondrial genes (COI and ND4) from a broad array of bathymodiolin species that included several recently discovered species from shallow hydrothermal seamounts. Bayesian phylogenetic analysis and maximum-likelihood estimates of ancestral character states revealed that vent species evolved multiple times, and that reversals in vent and seep habitat use occurred within the sampled taxa. Previous hypotheses regarding evolution from wood/bone-to-seeps/vents are supported in that mid-ocean hydrothermal vent species may represent a monophyletic group with one noticeable reversal. Earlier hypotheses about progressive evolution from shallow-to-deep habitats appear to hold with a few instances of habitat reversals.     

Chromosomal and nuclear characteristics of deep-sea hydrothermal-vent organisms: correlates of increased growth rate

A range of tissue and cell types from adult and juvenile stages of vent- and non-vent-dwelling deep-sea and shallow-water organisms were compared for signs of cell division, in preparation for a cytogenetic study of the different groups. Virtually all the Mid-Atlantic Ridge (MAR) vent species (bresiliid shrimp, bathymodiolid mussel, branchipolynoid polychaete, and a range of small gastropods) showed an abundance of metaphase chromosome spreads, indicating a generally high intrinsic rate of cell division, irrespective of their phylogenetic group. In comparison, similar tissues and life-history stages (i.e. gonad, developing eggs and gill) from two deep-sea bathypelagic crustaceans, Hymenodora and Gnathophausia, failed to yield any convincing evidence of cell division. This indicated that rates of growth and reproduction in the rest of the deep sea are extremely low. While much has been written about the contrasting growth rates of vent versus non-vent organisms, this is the first direct evidence of differences in cell-division rate between vent and non-vent species, which undoubtedly reflects contrasting conditions relating to food availability and temporal stability in these two deep-sea environments. The cells of shallow-water marine invertebrates (Mytilus, Nucella, Littorina, Pomatoceros and Crangon), based on the results of previous cytogenetic investigations, occupy an intermediate position with respect to their cell-division kinetics. When chromosome numbers, DNA content and nuclear diameters were compared between vent and non-vent species, this showed that no change had occurred in these characters since the time of vent colonisation. Representative chromosome spreads from hydrothermal-vent organisms are shown here for the first time.     

Larvae of benthic invertebrates in hydrothermal vent plumes over Juan de Fuca Ridge

Larvae of benthic invertebrates collected in the water column above Juan de Fuca Ridge show distinct variations in abundance and composition in, and away from, the neutrally-buoyant hydrothermal plume emanating from underlying vents. Larvae of vent gastropods (Lepetodrilus sp. and two peltospirid species) occur in significantly higher abundances in the plume than away from it (mean abundance=21.0 individuals 1000 m^?3 vs 1.4 individuals 1000 m^?3), and larvae of vent bivalves (Calyptogena? sp.) occur exclusively in the plume (mean abundance=0.5 individuals 1000 m^?3). Larvae from other benthic taxa known not to be endemic to Juan de Fuca vent communities, such as anthozoans, pholad clams, bryozoans and echinoderms, are less abundant in the plume than away (mean abundance=47.5 vs 16.9 individuals 1000 m^?3) at comparable depths and heights above the bottom. These results support the hypothesis that larvae of vent species are entrained into buoyant hydrothermal plumes and transported at the level of lateral spreading several hundred meters above the seafloor. The discovery of vent-associated larvae in the plume suggests that models used to predict hydrodynamic processes in the plume will also be useful for modeling larval dispersal. Advanced imaging and new molecular-based approaches will be required to resolve taxonomic uncertainties in some larval groups (e.g. certain polychaete families) in order to distinguish vent species and make comprehensive flux estimates of all vent larvae in the neutrally-buoyant plume.     

Carotenoids indicate differences in diet of the hydrothermal vent crabBythograea thermydron (Brachyura)

Carotenoid pigments were used as markers to investigate the sources of energy to two deep-sea hydrothermal vent communities. Specimens of the hydrothermal-vent brachyuran crabBythograea thermydron were collected at 2 500 m depth from the Rose Garden vent site in the Galápagos Rift Valley in February, November and December 1979, and 2 600 m depth on the East Pacific Rise at 21°N in May 1982. Four carotenoids (astaxanthin, canthaxanthin, echinenone and beta-carotene) have been identified as the pigments responsible for the red color of the eggs of the crabs from the Galápagos Rift site. Consistent with the fact that animals are unable to synthesize carotenoidsde novo, precursors were not present in the crabs' tissues, affirming that these pigments are of dietary origin. The number of ovigerous female crabs and the concentrations of carotenoids in the eggs suggest a readily available source of these pigments in the Galápagos vent environment. In contrast, the developing eggs ofB. thermydron from the 21°N vent site were cream-colored, with only trace quantites of carotenoids and fewer types of carotenoids. Analysis of carotenoid distribution in both females and males in these two populations indicates a very low level of carotenoids in the diet of the 21°N vent crabs, and reflects differences in trophic interactions and primary production at the two vent sites. The few types and low concentration of carotenoids inB. thermydron indicate a diet that is different from non-vent, deep-sea crustaceans. We hypothesize that the source of carotenoids are bacteria within the vent community, and not ultimately from photosynthetic production.     

Scientists as Stakeholders in Conservation of Hydrothermal Vents

Abstract: Hydrothermal vents are deep‐sea ecosystems that are almost exclusively known and explored by scientists rather than the general public. Continuing scientific discoveries arising from study of hydrothermal vents are concommitant with the increased number of scientific cruises visiting and sampling vent ecosystems. Through a bibliometric analysis, we assessed the scientific value of hydrothermal vents relative to two of the most well‐studied marine ecosystems, coral reefs and seagrass beds. Scientific literature on hydrothermal vents is abundant, of high impact, international, and interdisciplinary and is comparable in these regards with literature on coral reefs and seagrass beds. Scientists may affect hydrothermal vents because their activities are intense and spatially and temporally concentrated in these small systems. The potential for undesirable effects from scientific enterprise motivated the creation of a code of conduct for environmentally and scientifically benign use of hydrothermal vents for research. We surveyed scientists worldwide engaged in deep‐sea research and found that scientists were aware of the code of conduct and thought it was relevant to conservation, but they did not feel informed or confident about the respect other researchers have for the code. Although this code may serve as a reminder of scientists’ environmental responsibilities, conservation of particular vents (e.g., closures to human activity, specific human management) may effectively ensure sustainable use of vent ecosystems for all stakeholders.     

Microbial activities in the emitted hydrothermal waters of the Galápagos rift vents

In-situ measurement of chemolithotrophic and some heterotrophic microbial activities were made in the immediate vicinity of actively discharging hydrothermal vents of the Galápagos Rift region at depths of 2 500 to 2 600 m. The CO₂-assimilation or chemosynthesis productivity in the emitted vent waters, freshly mixed with oxygenated ambient seawater of 2°C, was minor compared to the bacterial biomass produced within the subsurface vent system prior to emission. Uptake of acetate and glucose indicated the presence of mixotrophic or facultatively chemolithotrophic bacteria in the emitted vent waters in agreement with isolations. Demonstration of ribulose bisphosphate carboxylase and phosphoenol pyruvate carboxylase in cultures of thiobacilli isolated from these vent water supports the notion that chemoautotrophic sulfur-oxidizing bacteria are one of the sources of primary production in the form of particulate organic carbon for filtering organisms in the deep sea hydrothermal environment. The rates of bacterial metabolic activities in emitted vent water are too low for the amount of invertebrate biomass and the rate of its growth and maintenance. Therefore, the larger portion of chemosynthetic sustenance of deep sea vent ecosystems appears to be based on symbiotic associations between bacteria and invertebrates and on surface attached bacteria.     

Benthic fauna of a shallow-water gaseohydrothermal vent area in the Aegean Sea (Milos, Greece)

The benthic community of shallow-water hydrothermal vents (10 m water depth) in a sandy bay on the south coast of Milos, Greece was studied. The macro- and meiofauna, as well as environmental factors such as temperature, salinity, sulfide concentration, grain size composition, ATP concentration, organic carbon, chlorophyll a and phaeopigments were evaluated. Samples were taken along transects from a seagrass meadow into a hydrothermally active area. Hydrothermally active spots were distinguished from the surrounding sediment by their black sediment coloration and an overlying white bacterial mat. For comparison, a control transect from a seagrass meadow into bare sand in a non-vent area in the same bay was studied. Overall, we found decreasing faunal diversity from the seagrass bed towards the hydrothermally active area. Along this gradient, four different zones could be distinguished. (a) The seagrass area had a relatively low diversity compared to the control stations, but much higher diversity than in the areas directly influenced by hydrothermal activity. (b) The border zone between the seagrass and the hydrothermal area, with moderate sulfide concentrations, pH values, seawater salinity and temperatures, was inhabited by an impoverished community dominated by the polychaete Capitella capitata, the nematodes Oncholaimus campylocercoides and Sabatieria sp., and an as yet undescribed chromadorid nematode (Chromadorina sp.). (c) The transition zone at the border of the hydrothermal area proper, with temperatures and sulfide concentrations higher than in the border zone, was characterized by high salinity (60‰) and low pH values. Along with the gastropod Cyclope neritea, single specimens of C. capitata and O. campylocercoides occurred. (d) The hydrothermal area proper, characterized by high sulfide concentrations, had high sediment temperatures and high salinities. The pH was as low as 6. Here, the gastropod Cyclope neritea was the only species occurring in considerable numbers. The gaseohydrothermal activity seems to largely exclude the surrounding fauna in a graded zonal pattern. The increasing hydrothermal stress entrains a change of the ambient biota towards a depauperate consortium of tolerant, opportunistic species, e.g. Capitella capitata (Polychaeta) and Oncholaimus campylocercoides (Nematoda). Although the hydrothermal area proper was characterized by a thick bacterial mat, the benthic community was characterized not by bacteria feeders but by scavenging species. Received: 13 September 1996 / Accepted: 24 October 1996     

A new bathymodioline mussel symbiosis at the Juan de Fuca hydrothermal vents

Until recently, the only major hydrothermal vent biogeographic province not known to include bathymodioline mussels was the spreading centers of the northeast Pacific, but deep-sea dives using DSV Alvin on the Endeavor segment of the Juan de Fuca Ridge (47°56N 129°06W; ∼2,200 m depth) in August 1999 yielded the only recorded bathymodioline mytilids from these northeastern Pacific vents. One specimen in good condition was evaluated for its relatedness to other deep-sea bathymodioline mussels and for the occurrence of chemoautotrophic and/or methanotrophic symbionts in the gills. Phylogenetic analyses of the host cytochrome oxidase I gene show this mussel shares evolutionary alliances with hydrothermal vent and cold seep mussels from the genus Bathymodiolus, and is distinct from other known species of deep-sea bathymodiolines, suggesting this mussel is a newly discovered species. Ultrastructural analyses of gill tissue revealed the presence of coccoid bacteria that lacked the intracellular membranes observed in methanotrophic symbionts. The bacteria may be extracellular but poor condition of the fixed tissue complicated conclusions regarding symbiont location. A single gamma-proteobacterial 16S rRNA sequence was amplified from gill tissue and directly sequenced from gill tissue. This sequence clusters with other mussel chemoautotrophic symbiont 16S rRNA sequences, which suggests a chemoautotrophic, rather than methanotrophic, symbiosis in this mussel. Stable carbon (δ¹³C = −26.6%) and nitrogen (δ¹⁵N = +5.19%) isotope ratios were also consistent with those reported for other chemoautotroph-mussel symbioses. Despite the apparent rarity of these mussels at the Juan de Fuca vent sites, this finding extends the range of the bathymodioline mussels to all hydrothermal vent biogeographic provinces studied to date.     

Morphological adaptations to chronic hypoxia in deep-sea decapod crustaceans from hydrothermal vents and cold seeps

Animals inhabiting hydrothermal vents and cold seeps face conditions that are challenging for survival. In particular, these two habitats are characterized by chronic hypoxia, sometimes reaching complete anoxia. The characteristics of the scaphognathite and gills were studied in four species of shrimp and three species of crabs from hydrothermal vents and cold seeps, in order to highlight potential adaptations that could enhance oxygen acquisition in comparison with shallow-water relatives. All the vent and seep species studied here exhibit significantly larger scaphognathites, likely allowing more water to flow over their gills per stroke of this appendage. This is probably more energetically efficient that prolonged hyperventilation. In contrast to annelids, vent and seep decapods usually do not possess enlarged gills, a phenomenon likely due to the physical limitations imposed by the size of the gill chamber. In the vent shrimp Rimicaris exoculata and the vent crab Bythograea thermydron, however, there is a significantly higher specific gill surface area linked to a higher number of lamellae per gram of gill. Again in contrast to annelids, the diffusion distance through the gills is not strikingly different between the vent shrimp Alvinocaris komaii and the shallow-water species Palaemon spp. This may indicate that the epithelium and cuticle of the decapod gills are already optimized for oxygen uptake and that reducing the thickness of these compartments is not physically possible without affecting the physical integrity of the gills.     

Reproductive biology and population structure of the deep-sea hydrothermal vent worm Paralvinella grasslei (Polychaeta: Alvinellidae) at 13°N on the East Pacific Rise

Paralvinella grasslei is a polychaetous annelid living in the harsh, unstable and heterogeneous environmental conditions found at deep-sea hydrothermal vent sites in the eastern Pacific. The aim of this work was to examine the possible influence of the reproductive biology of P. grasslei on the structure of its populations. Maximum observed oocyte size inside the oviduct is 275 m, and fecundity is relatively low. Examination of gametes and young specimens suggested a direct benthic development for this species. The population structure of P. grasslei at 13°N/EPR (EPR=East Pacific Rise) revealed a discontinuous recruitment which seems to be synchronized within vent sites and fields. The data also suggested the occurrence of discrete breeding periods. P. grasslei probably reproduces several times a years, with an apparent periodicity. Tidal signals could be a possible cue for the coordination of the reproductive cycle. The life-history of P. grasslei is discussed in light of the reproductive biology of other terebellomorph polychaetes, and seems to be well adapted for colonizing the unstable environment of hot vents. Two main hypotheses can explain the dissemination processes of this species along axial oceanic ridges. The influence of nearbottom currents occurring along the central graben of the East Pacific Rise can be cosidered to account for part of the transport of larvae and juveniles, but the observations of polychaete erpochaetes on the test of hydrothermal bythograeid crabs and evidence that crab migrations occur between vents also support the possibility of zoochory for the dissemination of alvinellid polychaetes.     

The effect of cadmium on antioxidant responses and the susceptibility to oxidative stress in the hydrothermal vent mussel Bathymodiolus azoricus

Hydrothermal vents are a unique environment of extreme physical–chemical characteristics and biological species composition. Cd is a toxic non-essential metal present in high concentrations in the hydrothermal vent environment, contrary to those found in marine coastal areas. Cd toxicity has been related, among other things, with reactive oxygen species production, even though this is a non-redox metal. Bathymodiolus azoricus is a deep-sea Mytilid bivalve very common in the Mid Atlantic Ridge (MAR) hydrothermal vent fields and very little is known about the antioxidant defence system in this specie. Because lethal Cd concentration in B. azoricus is unknown, the aim of this study was to assess the effects of a Cd concentration higher than that found in the hydrothermal vents on oxidative stress biomarkers, such as antioxidant enzymes. Mussels were exposed to 100 μg l⁻¹ Cd during 24, 48 and 144 h, respectively, in a pressurized aquarium (IPOCAMP). Superoxide dismutase (SOD), catalase (CAT), glutathione peroxidases (GPx), total oxyradical scavenging capacity (TOSC), metallothionein (MT) and lipid peroxidation (LPO) were measured in the gills and mantle of B. azoricus. The results indicate that gills are first affected by Cd toxicity. This may be due to different physiological functions of the tissues and by the presence of thio and methanotrophic symbiotic bacteria in the gills. The SOD and CAT are inhibited during the first day of exposure in the gills, although TOSC and MT concentrations were the same in control and exposed mussels. In the mantle, enzymatic activation only occurred after 6 days, and no significant differences in MT concentrations were found in the control and exposed mussels during the first day, as observed in the gills.     

Transparent exopolymer particles in a deep-sea hydrothermal system: Guaymas Basin,Gulf of California

This is the first report of transparent exopolymer particle (TEP) concentrations within deep-sea hydrothermal vent systems using colorimetric methods, measuring TEP in supernatants of sediments surrounding the vents, in fluids emanating directly from hydrothermal vents and in neutrally buoyant hydrothermal plumes. Samples were collected at Guaymas Basin (Gulf of California), a sedimented hydrothermal system. TEP concentrations within the hydrothermal fluids were significantly greater than the only other report of TEP in deep water. The range of values for TEP abundance were 8–6,451 μg/L of gum xanthan equivalents, the highest values being associated with supernatants of microbial mat-covered sediments. The potential sources and significance of the high concentrations of TEP observed in this deep-sea hydrothermal environment are discussed.     

Scarred limpets at hydrothermal vents: evidence of predation by deep-sea whelks

The chaotic physical and chemical environment at deep-sea hydrothermal vents has been associated with an ecosystem with few predators, arguably allowing the habitat to provide refuge for vulnerable species. The dominance of endemic limpets with thin, open-coiled shells at north Pacific vents may support this view. To test their vulnerability to predation, the incidence of healed repair scars, which are argued to reflect non-lethal encounters with predators, were examined on the shells of over 5,800 vent limpets of Lepetodrilus fucensis McLean (1988) that were collected from 13 to 18 August 1996. Three vent fields on the Juan de Fuca Ridge at ca. 2,200 m depth were sampled, two within 70 m of 47°56.87′N 129°05.91′W, and one at 47°57.85′N 129°05.15′W with the conspicuous potential limpet predators, the zoarcid fish Pachycara gymninium Anderson and Peden (1988), the galatheid crab Munidopsis alvisca Williams (1988), and the buccinid snail Buccinum thermophilum Harasewych and Kantor (2002). Limpets from the predator-rich vent were most often scarred, a significant difference created by the high incidence of scars on small (<4 mm long) limpets in this sample. Collected with the limpets were small (median shell diameter 4.4 mm) buccinids. They, rather than the larger, more conspicuous mobile fishes and crabs are argued to be the shell-damaging predator. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.