Bathymodiolus Thermophilus Research Articles

Faunal colonists, including mussel settlers, respond to microbial biofilms at deep-sea hydrothermal vents

Colonization processes at dynamic deep-sea hydrothermal vent ecosystems ultimately determine ecosystem structure, function, resilience, and recovery. Microbial biofilms form rapidly on surfaces near hydrothermal vents and are continuously exposed to the highly variable abiotic environment. Thus, biofilm microbes may provide a temporally integrated signal that can indicate whether the habitat is suitable for faunal colonists. This study explored the role of microbial biofilms in controlling faunal colonization through in-situ colonization experiments at Tica Vent in the 9°50’ N region of the East Pacific Rise (EPR). Short-term experiments (∼2 weeks) were conducted by deploying colonization surfaces (“sandwiches”) either with an established biofilm (developed for >1 year) or a fresh biofilm (developed throughout experiment) in zones characterized by different faunal assemblages. Differences in associated larval settlers, faunal immigrants, and microbial communities according to biofilm age across multiple biogenic zones were investigated. Faunal and microbial community compositions significantly differed according to whether the sandwiches had established or fresh biofilms as well as the biogenic zone they were deployed in. Several faunal colonists, including settlers such as the foundational chemosymbiotic mussel Bathymodiolus thermophilus and the nectochaete Archinome sp., were found associated more with established biofilms than fresh biofilms. Microbial biofilm communities were dominated by putative chemoautotrophic members of the Campylobacterota phylum and Gammaproteobacteria class and several microbial taxa were found to covary with faunal colonists. Overall, these findings show that microbial community composition plays a role in larval settlement and animal migration in hydrothermal vent systems and the detection of microbial and faunal interactions provides a starting point for identifying key microbial characteristics influencing colonization processes at hydrothermal vents.

A regulatory hydrogenase gene cluster observed in the thioautotrophic symbiont of Bathymodiolus mussel in the East Pacific Rise

The mytilid mussel Bathymodiolus thermophilus lives in the deep-sea hydrothermal vent regions due to its relationship with chemosynthetic symbiotic bacteria. It is well established that symbionts reside in the gill bacteriocytes of the mussel and can utilize hydrogen sulfide, methane, and hydrogen from the surrounding environment. However, it is observed that some mussel symbionts either possess or lack genes for hydrogen metabolism within the single-ribotype population and host mussel species level. Here, we found a hydrogenase cluster consisting of additional H2-sensing hydrogenase subunits in a complete genome of B. thermophilus symbiont sampled from an individual mussel from the East Pacific Rise (EPR9N). Also, we found methylated regions sparsely distributed throughout the EPR9N genome, mainly in the transposase regions and densely present in the rRNA gene regions. CRISPR diversity analysis confirmed that this genome originated from a single symbiont strain. Furthermore, from the comparative analysis, we observed variation in genome size, gene content, and genome re-arrangements across individual hosts suggesting multiple symbiont strains can associate with B. thermophilus. The ability to acquire locally adaptive various symbiotic strains may serve as an effective mechanism for successfully colonizing different chemosynthetic environments across the global oceans by host mussels.

Synthesis of the Proposed Isomers of the Deep-Sea Mussel Metabolites Bathymodiolamides A and B.

The first total synthesis of bathymodiolamides A and B, ceramide-like metabolites of the deep-sea hydrothermal vent mussel Bathymodiolus thermophilus, was accomplished in eight linear steps starting from Garner's aldehyde and three carboxylic acids. A sequence of vinylation of Garner's aldehyde, N-acylation with lauric acid, dihydroxylation of the terminal alkene, and stepwise Steglich-Hassner esterifications of the resulting vicinal diol with the respective saturated and unsaturated carboxylic acids, which had to be prepared separately, afforded the target products in 38 and 39% yield. We found distinct discrepancies between their NMR data and antiproliferative activities and those reported for the natural isolates.

Molecular analyses of the gill symbiosis of the bathymodiolin mussel Gigantidas platifrons.

SummaryAlthough the deep-sea bathymodiolin mussels have been intensively studied as a model of animal-bacteria symbiosis, it remains challenging to assess the host-symbiont interactions due to the complexity of the symbiotic tissue—the gill. Using cold-seep mussel Gigantidas platifrons as a model, we isolated the symbiont harboring bacteriocytes and profiled the transcriptomes of the three major parts of the symbiosis—the gill, the bacteriocyte, and the symbiont. This breakdown of the complex symbiotic tissue allowed us to characterize the host-symbiont interactions further. Our data showed that the gill's non-symbiotic parts play crucial roles in maintaining and protecting the symbiosis; the bacteriocytes supply the symbiont with metabolites, control symbiont population, and shelter the symbiont from phage infection; the symbiont dedicates to the methane oxidation and energy production. This study demonstrates that the bathymodiolin symbiosis interacts at the tissue, cellular, and molecular level, maintaining high efficiency and harmonic chemosynthetic micro niche.

Spanning the depths or depth-restricted: Three new species of Bathymodiolus (Bivalvia, Mytilidae) and a new record for the hydrothermal vent Bathymodiolus thermophilus at methane seeps along the Costa Rica margin

Bathymodiolus Kenk and Wilson, 1985 includes fourteen currently recognized species from deep-sea chemosynthetic environments in the Pacific, Atlantic, and Indian oceans. In this study, phylogenetic analyses of mytilid mussels sampled from seeps along the Costa Rica margin reveal the presence of three new Bathymodiolus species, sampled from depths across ~1000–1900 m. Bathymodiolus billschneideri n. sp., B. earlougheri n. sp., and B. nancyschneiderae n. sp. differ genetically from congeneric species of Bathymodiolus and show some stratification by depth. The depth ranges of Bathymodiolus billschneideri n. sp., B. nancyschneiderae n. sp., and B. earlougheri n. sp. were ~1400–1900 m, 1000–1100 m, and ~1000–1900m, respectively. Bathymodiolus billschneideri n. sp. and B. earlougheri n. sp. were found to be closely related to Bathymodiolus thermophilus Kenk and Wilson, 1985, while B. nancyschneiderae n. sp. exhibited closer relationships to non-East-Pacific taxa. Bathymodiolus thermophilus was the first Bathymodiolus mussel discovered, sampled from a vent along the Galápagos Rift Zone in 1985 and later recorded along much of the East Pacific Rise. This study confirms the presence of B. thermophilus at the Costa Rica margin, representing the first DNA and morphological samples of B. thermophilus at a seep environment. Analysis of habitat evolution suggests that B. thermophilus and its closest relative, B. antarcticus Johnson and Vrijenhoek, 2013, are of seep ancestry.

A mitochondrial genome phylogeny of Mytilidae (Bivalvia: Mytilida)

The family Mytilidae is a family of bivalve mussels that are distributed worldwide in diverse marine habitats. Within the family, classification systems and phylogenetic relationships among subfamilies remain not yet fully resolved. In this study, we newly determined 9 mitochondrial genome sequences from 7 subfamilies: Bathymodiolus thermophilus (Bathymodiolinae), Modiolus nipponicus (Modiolinae), Lithophaga curta (the first representative of Lithophaginae), Brachidontes mutabilis (Brachidontinae), Mytilisepta virgata (Brachidontinae), Mytilisepta keenae (Brachidontinae), Crenomytilus grayanus (Mytilinae), Gregariella coralliophaga (Crenellinae), and Septifer bilocularis (the first representative of Septiferinae). Phylogenetic trees using maximum likelihood and Bayesian inference methods for 28 mitochondrial genomes (including 19 previously published sequences) showed two major clades with high support values: Clade 1 ((Bathymodiolinae + Modiolinae) + (Lithophaginae + Limnoperninae)) and Clade 2 (((Mytilinae + Crenellinae) + Septiferinae) + Brachidontinae). The position of the genus Lithophaga (representing Lithophaginae) differed from a previously published molecular phylogeny. Divergence time analysis with a molecular clock indicated that lineage splitting among the major subfamilies of Mytilidae (including the habitat transition from marine to freshwater environments by ancestral Limnoperninae) occurred in the Mesozoic period, coinciding with high diversification rates of marine fauna during that time. This is the first mitochondrial genome-based phylogenetic study of the Mytilidae that covers nearly all subfamily members, excluding the subfamily Dacrydiinae.

Bathymodiolus growth dynamics in relation to environmental fluctuations in vent habitats

The deep-sea mussel Bathymodiolus thermophilus is a dominant species in the East Pacific Rise (EPR) hydrothermal vent fields. On the EPR volcanically unstable area, this late colonizer reaches high biomass within 4–5 years on new habitats created by lava flows. The environmental conditions and growth rates characterizing the reestablishment of B. thermophilus populations are however largely unknown, leaving unconstrained the role of this foundation species in the ecosystem dynamics. A typical example from the vent field at 9°50’N that was affected by the last massive eruption was the Bio-9 hydrothermal vent site. Here, six years later, a large mussel population had reestablished. The von Bertalanffy growth model estimates the oldest B. thermophilus specimens to be 1.3 year-old in March 2012, consistent with the observation of scarce juveniles among tubeworms in 2010. Younger cohorts were also observed in 2012 but the low number of individuals, relatively to older cohorts, suggests limited survival or growth of new recruits at this site, that could reflect unsuitable habitat conditions. To further explore this asumption, we investigated the relationships between mussel growth dynamics and habitat properties. The approach combined sclerochronology analyses of daily shell growth with continuous habitat monitoring for two mussel assemblages; one from the Bio-9 new settlement and a second from the V-vent site unreached by the lava flow. At both vent sites, semi-diurnal fluctuations of abiotic conditions were recorded using sensors deployed in the mussel bed over 5 to 10 days. These data depict steep transitions from well oxygenated to oxygen-depleted conditions and from alkaline to acidic pH, combined with intermittent sulfide exposure. These semi-diurnal fluctuations exhibited marked changes in amplitude over time, exposing mussels to distinct regimes of abiotic constraints. The V-vent samples allowed growth patterns to be examined at the scale of individual life and compared to long-term records of habitat temperature and oceanographic mooring data in the years following the eruption. Both shell growth and habitat temperature at V-vent varied over the spring-neap tidal cycle and over longer periods of c.a. 60 days. The correlation of growth rate with temperature and, for some individuals, with current velocities supports the idea that tidal forcing impacts growth. Its influence on habitat conditions includes the spring-neap cycle, which is not reflected in current velocities but influences the venting rate. Additionally, it is expected that mesoscale eddies periodically passing across the ridge imprint shell growth through the influence of bottom current on the decimeter-thick mixing interface where mussels thrive. We conclude that diurnal-semidiurnal tidal fluctuations exert major abiotic constraints on B. thermophilus mussels and that low-frequency fluctuations act as significant determinants on growth. Finally, we postulate that the modulation of tidal fluctuations by large-scale hydrodynamic forcing ultimately constrains the capacity of this mussel species to form high biomass aggregations. This study indeed shows that the absence of these strong hydrodynamic drivers would limit the alternance of oxic and sulfidic conditions and significantly affect the growth rate of this species over time.

Non-equilibrium fractionation of stable carbon isotopes in chemosynthetic mussels

Chemosynthetic bivalves from deep-sea hydrothermal vents exploit the energy derived from chemical compounds, such as methane, sulfide or hydrogen, using symbiotic bacteria that are able to fix inorganic carbon. Available chemical resources in their habitat vary widely at various scales, from the vent field scale to the micro-habitat scale. Parallel to this environmental heterogeneity, Bathymodiolus species are considered to be flexible in their energy acquisition pathways.The goal of this study was to determine whether the isotopic compositions archived in the shells of hydrothermal vent mussels could trace chemical energy sources and their variability over spatial and temporal scales. Two different species (Bathymodiolus azoricus and Bathymodiolus thermophilus) inhabiting three vent fields with contrasted geochemical features on the Mid Atlantic Ridge (MAR; Rainbow and Menez Gwen) and the East Pacific Rise (EPR; 9°47′N), were considered for carbon isotopes and growth rate variation along the shell length. The study revealed that 13C fractionation between shells and seawater is higher than expected from calcite–bicarbonate equilibrium fractionation, suggesting a significant influence of the chemosynthetic pathway on the shell composition. Furthermore, significant differences in δ13Cshell fractionation with respect to seawater are observed between sites and habitats of the two MAR vent fields, suggesting that different chemosynthetic pathway (e.g. methanotrophic and thiotrophic) could lead to variable enrichments of the shell in 13C. Mussels supposed to rely more largely on methanotrophy (at Rainbow where free sulfide is unavailable) display a lower δ13Cshell values than mussels relying also on sulfide-oxidizing symbiosis (at Menez Gwen). Variability in δ13Cshell between habitats, or between individuals within the same assemblage, could thus reflect differences in the symbiosis activity at a micro-habitat scale. These isotopic signatures could provide useful information on the relationships between micro-habitat properties, symbiont activity and shell mineralization.

A New Barrier to Dispersal Trapped Old Genetic Clines That Escaped the Easter Microplate Tension Zone of the Pacific Vent Mussels

Comparative phylogeography of deep-sea hydrothermal vent species has uncovered several genetic breaks between populations inhabiting northern and southern latitudes of the East Pacific Rise. However, the geographic width and position of genetic clines are variable among species. In this report, we further characterize the position and strength of barriers to gene flow between populations of the deep-sea vent mussel Bathymodiolus thermophilus. Eight allozyme loci and DNA sequences of four nuclear genes were added to previously published sequences of the cytochrome c oxidase subunit I gene. Our data confirm the presence of two barriers to gene flow, one located at the Easter Microplate (between 21°33′S and 31°S) recently described as a hybrid zone, and the second positioned between 7°25′S and 14°S with each affecting different loci. Coalescence analysis indicates a single vicariant event at the origin of divergence between clades for all nuclear loci, although the clines are now spatially discordant. We thus hypothesize that the Easter Microplate barrier has recently been relaxed after a long period of isolation and that some genetic clines have escaped the barrier and moved northward where they have subsequently been trapped by a reinforcing barrier to gene flow between 7°25′S and 14°S.

Compact autonomous voltammetric sensor for sulfide monitoring in deep sea vent habitats

In situ chemical monitoring at deep-sea hydrothermal vents remains a challenge. Particularly, tools are still scarce for assessing the ranges and temporal variability of sulfide in these harsh environmental conditions. There is a particular need for compact and relatively simple devices to enlarge the capacity of in situ measurements of this major energy source in chemosynthetic ecosystems. With this objective, a voltammetric sensor based on a bare-silver working electrode was developed and tested in real conditions. In the laboratory, the sensor presented a linear response from 10 to 1000μM sulfide, together with a low pH sensitivity and moderate temperature dependence. The device was operated at 850 and 2500m depth during 3 cruises over two different vent fields. The autonomous potentiostat (290mm length, ∅ 35mm) equipped with laboratory-made electrodes was mounted on a wand, for manipulation from a submersible, or on a holder for unattended deployments. The system was applied in mussel, tubeworm and annelid worm habitats, characterized by different ranges of sulfide concentration, pH and temperature. Calibrations performed before and after each deployment confirmed the stability of the sensor response over a few hours to 11 days, with a maximum drift of 11.4% during this period. Short-term measurements in the vicinity of Riftia pachyptila and Alvinella pompejana were consistent with previous results on these habitats, with concentrations ranging from 20 to 140μM and 100 to 450μM and sulfide versus temperature ratio of 14μM°C−1 and 20μM°C−1, respectively. A continuous 4-day record on a bed of Bathymodiolus Thermophilus mussels furthermore illustrated the capacity of the sensor to capture fluctuating sulfide concentration between 0 and 70μM, in combination to temperature, and to investigate the changes in the sulfide versus temperature ratio over time. The method has a higher detection limit (<10μM) than previous in situ sulfide measurement methods, but has the advantage of selectivity to free sulfide (compared to colorimetry), low pH sensitivity (compared to amperometry) and lower reconditioning requirement for electrode surface (with respect to gold-amalgam voltammetry). This sensor is therefore a valuable complementary tool for discrete and continuous measurements within the moderate temperature environment of fauna at deep-sea vents.

The potential implication of apoptosis in the control of chemosynthetic symbionts in Bathymodiolus thermophilus

The potential implication of apoptosis in the control of chemosynthetic symbionts in Bathymodiolus thermophilus

A new method for high-resolution bivalve growth rate studies in hydrothermal environments

The age and shell growth rate of deep-sea hydrothermal bivalves were investigated for the first time using in situ chemical staining combined with high-resolution micro-increment analysis. A staining chamber developed for this purpose was applied to a patch of Bathymodiolus thermophilus mussels at 2,500 m depth at the 9°47′N vent field on the East Pacific Rise (EPR) in May 2010. This approach minimizes disturbance of the mussels in their habitat. Bathymodiolus thermophilus grows according to a circalunidian rhythm, with one increment formed each day, and displays tide-related growth rate variability. Based on the von Bertalanffy growth rate model, the largest shell collected (SL = 20.5 cm) would be 10.0 year old, with a growth rate of 4.2–1.1 cm year−1 as the shell ages. This fast growth rate is consistent with the instability of the environment in this section of the EPR and observed recolonization rates and could reflect a specific adaptation of this species.

Bathymodiolamides A and B, ceramide derivatives from a deep-sea hydrothermal vent invertebrate mussel, Bathymodiolus thermophilus.

Two ceramide derivatives, bathymodiolamides A (1) and B (2), were isolated from the deep-sea hydrothermal vent invertebrate mussel Bathymodiolus thermophilus. The molecular structures of these compounds were determined using a combination of NMR spectroscopy, mass spectrometry, and chemical degradation. Biological activities were assessed in a ApopScreen cell-based screen for apoptosis induction and potential anticancer activity. To our knowledge, this is the first report of secondary metabolites from the marine hydrothermal vent mussel B. thermophilus.

Comparative analysis between protist communities from the deep‐sea pelagic ecosystem and specific deep hydrothermal habitats

Protist communities associated with deep seawater and bivalves from six hydrothermal sites in the Pacific Ocean were characterized by microscopy and molecular rRNA gene surveys (18S rRNA) and compared with planktonic communities from Pacific deep-pelagic seawater (from 500 to 3000 m in depth). Genetic libraries from larger size fractions (>3 µm) of deep-pelagic water were mainly dominated by Dinophyceae, whereas small size fractions (<3 µm) mainly revealed radiolarians and Syndiniales. In contrast, more specific opportunistic detritivores and grazers, mostly belonging to Stramenopiles and Cercozoa, were detected from water surrounding vent chimneys. Protist communities were different in the pallial cavity of the giant hydrothermal bivalves Bathymodiolus thermophilus and Calyptogena magnifica, dominated by Ciliophora (primarily belonging to Phyllopharyngea, Oligohymenophorea and Oligotrichea) and Cercozoa. Interestingly, protist communities retrieved from the pallial cavity liquid of hydrothermal bivalves were remarkably homogeneous along the Southern East Pacific Rise, in contrast to bivalves collected on the Mid-Atlantic Ridge hydrothermal vents and cold seeps from the Gulf of Mexico. Hence, complex protist communities seem to occur inside hydrothermal bivalves, and these metazoa may constitute a stable micro-niche for micro-eukaryotes, including grazers, detritivores, symbionts and potential parasites. From these communities, new lineages within the ciliates may emerge.

Pre- and post-eruption diffuse flow variability among tubeworm habitats at 9°50′ north on the East Pacific Rise

Abstract A seafloor eruption at 9°50′N East Pacific Rise (EPR) in 2005–2006 fundamentally changed the fluid chemical and biological community composition throughout the region. Prior to the eruption (in 2004 and 2005), this region of the EPR was dominated by siboglinid ( Riftia pachyptila ) tubeworms and bathymodiolin ( Bathymodiolus thermophilus ) mussels. Following the eruption, the tubeworm, Tevnia jerichonana , was the first observed sessile megafauna to have colonized nascent vent openings. The eruption provided the unique opportunity to compare species-specific habitat chemistry and address previously documented differences in the colonization order of tubeworm species hypothesized to be due to the different physiological tolerances to changing vent fluid chemistry. We coupled in situ electrochemical measurements of free sulfide (S free ), Σ total sulfide, O 2 , Fe(II), Mn(II), and other S(-II) species with the occurrence of tubeworm colonies to document and compare, for the first time, the chemical habitats of the dominant tubeworm fauna on the East Pacific Rise, and the pre-and post-eruptive diffuse fluid chemistry in the region. Median sulfide concentrations increased two to four fold from 2004–2005 when R. pachyptila were dominant to 2006–2008 when T. jerichonana were dominant, and median oxygen conditions decreased from 40–58 μM to below the detection limit of 3–5 μM over the same period. These data suggest fundamental differences between these tubeworm habitats, as well as the possibility of increased tolerance for microaerophilic conditions in T. jerichonana . Sulfide-to-Temperature ( S/T ) ratios increased from before (2004) to after the eruption (2006–2007). However, S/T ratios at two of the sites in 2005 increased from 2004 indicating that a part of the hydrothermal vent field may have been “heating up” prior to the eruption. The results of diffuse flow habitat chemistry provide data for the first time that R. pachyptila and T. jerichonana tubeworms may preferentially inhabit different chemical habitats.

Time-series chemical and temperature habitat characterization of diffuse flow hydrothermal sites at 9°50′N East Pacific Rise

The spatial and temporal distributions of hydrothermal vent organisms are considered to be controlled by a combination of chemical and biological interactions. However, the degree of habitat specificity imposed by fluid chemistry on hydrothermal vent-endemic siboglinid tubeworms Tevnia jerichonana and Riftia pachyptila, as well as the mussel Bathymodiolus thermophilus, remains largely unknown. A comparison of time-series sulfide and temperature data reveals that T. jerichonana primarily inhabit higher temperatures and sulfide concentrations, that B. thermophilus primarily inhabit lower temperatures and sulfide concentrations, and that R. pachyptila habitats overlap the range of temperature and sulfide concentrations observed. Based on these data in combination with previous laboratory studies, we speculate on potential in situ uptake rates of oxygen and sulfide by R. pachyptila. These estimates suggest that R. pachyptila potentially experiences both oxygen and sulfide limitations. The mean uptake rates for areas inhabited by R. pachyptila are favorable for R. pachyptila, while those inhabited by B. thermophilus and T. jerichonana appear to be sulfide and oxygen limited, respectively.

Molecular identification of differentially regulated genes in the hydrothermal-vent species Bathymodiolus thermophilus and Paralvinella pandorae in response to temperature.

BackgroundHydrothermal vents and cold seeps represent oases of life in the deep-sea environment, but are also characterized by challenging physical and chemical conditions. The effect of temperature fluctuations on vent organisms in their habitat has not been well explored, in particular at a molecular level, most gene expression studies being conducted on coastal marine species. In order to better understand the response of hydrothermal organisms to different temperature regimes, differentially expressed genes (obtained by a subtractive suppression hybridization approach) were identified in the mussel Bathymodiolus thermophilus and the annelid Paralvinella pandorae irlandei to characterize the physiological processes involved when animals are subjected to long term exposure (2 days) at two contrasting temperatures (10° versus 20°C), while maintained at in situ pressures. To avoid a potential effect of pressure, the experimental animals were initially thermally acclimated for 24 hours in a pressurized vessel.ResultsFor each species, we produced two subtractive cDNA libraries (forward and reverse) from sets of deep-sea mussels and annelids exposed together to a thermal challenge under pressure. RNA extracted from the gills, adductor muscle, mantle and foot tissue were used for B. thermophilus. For the annelid model, whole animals (small individuals) were used. For each of the four libraries, we sequenced 200 clones, resulting in 78 and 83 unique sequences in mussels and annelids (about 20% of the sequencing effort), respectively, with only half of them corresponding to known genes. Real-time PCR was used to validate differentially expressed genes identified in the corresponding libraries. Strong expression variations have been observed for some specific genes such as the intracellular hemoglobin, the nidogen protein, and Rab7 in P. pandorae, and the SPARC protein, cyclophilin, foot protein and adhesive plaque protein in B. thermophilus.ConclusionOur results indicate that mussels and worms are not responding in the same way to temperature variations. While the results obtained for the mussel B. thermophilus seem to indicate a metabolic depression (strong decrease in the level of mRNA expression of numerous genes) when temperature increased, the annelid P. pandorae mainly displayed a strong regulation of the mRNA encoding subunits and linkers of respiratory pigments and some proteins involved in membrane structure. In both cases, these regulations seem to be partly due to a possible cellular oxidative stress induced by the simulated thermal environment (10°C to 20°C). This work will serve as a starting point for studying the transcriptomic response of hydrothermal mussels and annelids in future experiments in response to thermal stress at various conditions of duration and temperature challenge.

Biotic interactions at hydrothermal vents: Recruitment inhibition by the mussel Bathymodiolus thermophilus

The structure and dynamics of marine communities are regulated in part by variation in recruitment. As in other ecosystems, recruitment at deep-sea hydrothermal vents is controlled by the interplay of propagule supply and behavior, gradients in physical–chemical conditions, and biotic interactions during pre- and post-settlement periods. Recent research along the East Pacific Rise indicates that inhibition of recently settled larvae by mobile predators (mainly limpets) influences patterns of recruitment and subsequent community succession. We conducted a manipulative experiment at the same sites (∼2510 m water depth) to test whether high-density assemblages of the mussel Bathymodiolus thermophilus also inhibit recruitment. In a preliminary study, recruitment of vent invertebrates within the faunal zone dominated by B. thermophilus was strikingly different at two sites, East Wall and Worm Hole. East Wall had high densities of mussels but very low total recruitment. In contrast, Worm Hole had few mussels but high recruitment. Using the submersible Alvin, we transplanted a large number of mussels from East Wall to Worm Hole and quantified recruitment on basalt blocks placed in three treatments: (1) naturally high densities of mussels at East Wall; (2) naturally low densities of mussels at Worm Hole; and (3) high densities of transplanted mussels at Worm Hole. After 11 months, a total of 24 taxa had recruited to the basalt blocks. Recruitment was 44–60% lower in the transplanted high-density mussel patch at Worm Hole and the natural high-density patch at East Wall than within the natural low-density patch at Worm Hole. Biotic processes that may have caused the pattern of recruitment observed included predation of larvae via water filtration by mussels, larval avoidance of superior competitors, interference competition, and enhanced predation by species within the mussel-bed community. Our results indicate that biotic interactions affecting recruitment must be understood to explain patterns of invertebrate community organization and dynamics at hydrothermal vents.

Hydrothermal Vent Mussel Habitat Chemistry, Pre- and Post-Eruption at 9°50′North on the East Pacific Rise

Between October 2005 and March 2006, a seafloor volcanic eruption occurred at 9°50′N East Pacific Rise (EPR), establishing a “time zero” for characterizing newly-formed hydrothermal vent habitats and comparing them to pre-eruption habitats. Before the eruption, mussels (Bathymodiolus thermophilus) formed large aggregates between 9°49.6′ and 9°50.3′N. After the eruption, the few mussels remaining were in sparsely-distributed individuals and clumps, seemingly transported via lava flows or from mass wasting of the walls of the axial trough. In situ voltammetry with solid state gold-amalgam microelectrodes was used to characterize the chemistry of vent fluids in mussel habitats from 2004 to 2007, providing data sets for comparison of oxygen, sulfide, and temperature. Posteruption fluids contained higher sulfide-to-temperature ratios (i.e., slopes of linear regressions) (10.86 μM °C−1) compared with pre-eruption values in 2004 and 2005 (2.79 μM °C−1 and −0.063 μM °C−1, respectively). These chemical differences can be attributed to the difference in geographic location in which mussels were living and physical factors arising from posteruptive fluid emissions.

Interrelationships Between Vent Fluid Chemistry, Temperature, Seismic Activity, and Biological Community Structure at a Mussel-Dominated, Deep-Sea Hydrothermal Vent Along the East Pacific Rise

In April 1991, submarine volcanic eruptions initiated the formation of numerous hydrothermal vents between 945# and 952#N along the crest of the East Pacific Rise (EPR). Dramatic changes in biological community structure and vent fluid chemistry have been documented throughout this region since the eruptive event. By April 2004, mussels (Bathymodiolus thermophilus) dominated the faunal assemblages at several of the vent sites formed during of after the 1991 eruptions, whereas other habitats within the region were dominated by the vestimentiferan Riftia pachyptila. In the present paper, we build upon the extensive data sets obtained at these sites over the past decade and describe a manipulative experiment (conducted at 949.94#N; 10414.43#W on the EPR) designed to assess interrelationships between vent fluid chemistry, temperature, biological community structure, and seismic activity. To this end, in situ voltammetric systems and thermal probes were used to measure H2S/HS - and temperature over time in a denuded region of an extensive mussel bed in which an exclusion cage was placed to inhibit the subsequent migration of mussels into the denuded area. Fluid samples were taken from the same locations to characterize the associated microbial constituents. Basalt blocks, which were placed in the cage in April 2004 and subsequently recovered in April 2005, were colonized by more than 25 different species of invertebrates, including numerous vestimentiferans and remarkably few mussels. Recorded temporal changes in vent fluid chemistry and temperature regimes, when coupled with microbiological characterization of the vent fluids and seismic activity data obtained from ocean bottom seismometers, shed considerable light on factors controlling biological community structure in these hydrothermal ecosystems.