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The discovery of deep-sea hydrothermal vents in 1977 challenged our views of ecosystem functioning and yet, the research conducted at these extreme and logistically challenging environments still continues to reveal unique biological processes. Here, we report for the first time, a unique behavior where the deep-sea skate, Bathyraja spinosissima, appears to be actively using the elevated temperature of a hydrothermal vent environment to naturally "incubate" developing egg-cases. We hypothesize that this behavior is directly targeted to accelerate embryo development time given that deep-sea skates have some of the longest egg incubation times reported for the animal kingdom. Similar egg incubating behavior, where eggs are incubated in volcanically heated nesting grounds, have been recorded in Cretaceous sauropod dinosaurs and the rare avian megapode. To our knowledge, this is the first time incubating behavior using a volcanic source is recorded for the marine environment.

Deep-sea hydrothermal vents are populated by dense communities of animals that form symbiotic associations with chemolithoautotrophic bacteria. To date, our understanding of which factors govern the distribution of host/symbiont associations (or holobionts) in nature is limited, although host physiology often is invoked. In general, the role that symbionts play in habitat utilization by vent holobionts has not been thoroughly addressed. Here we present evidence for symbiont-influenced, regional-scale niche partitioning among symbiotic gastropods (genus Alviniconcha) in the Lau Basin. We extensively surveyed Alviniconcha holobionts from four vent fields using quantitative molecular approaches, coupled to characterization of high-temperature and diffuse vent-fluid composition using gastight samplers and in situ electrochemical analyses, respectively. Phylogenetic analyses exposed cryptic host and symbiont diversity, revealing three distinct host types and three different symbiont phylotypes (one ε-proteobacteria and two γ-proteobacteria) that formed specific associations with one another. Strikingly, we observed that holobionts with ε-proteobacterial symbionts were dominant at the northern fields, whereas holobionts with γ-proteobacterial symbionts were dominant in the southern fields. This pattern of distribution corresponds to differences in the vent geochemistry that result from deep subsurface geological and geothermal processes. We posit that the symbionts, likely through differences in chemolithoautotrophic metabolism, influence niche utilization among these holobionts. The data presented here represent evidence linking symbiont type to habitat partitioning among the chemosynthetic symbioses at hydrothermal vents and illustrate the coupling between subsurface geothermal processes and niche availability.

Western Pacific hydrothermal vents will soon be subjected to deep-sea mining and peripheral sites are considered the most practical targets. The limited information on community dynamics and temporal change in these communities makes it difficult to anticipate the impact of mining activities and recovery trajectories. We studied community composition of peripheral communities along a cline in hydrothermal chemistry on the Eastern Lau Spreading Center and Valu Fa Ridge ( ELSC- VFR) and also studied patterns of temporal change. Peripheral communities located in the northern vent fields of the ELSC- VFR are significantly different from those in the southern vent fields. Higher abundances of zoanthids and anemones were found in northern peripheral sites and the symbiont-containing mussel Bathymodiolus brevior, brisingid seastars and polynoids were only present in the northern peripheral sites. By contrast, certain faunal groups were seen only in the southern peripheral sites, such as lollipop sponges, pycnogonids and ophiuroids. Taxonomic richness of the peripheral communities was similar to that of active vent communities, due to the presence of non-vent endemic species that balanced the absence of species found in areas of active venting. The communities present at waning active sites resemble those of peripheral sites, indicating that peripheral species can colonize previously active vent sites in addition to settling in the periphery of areas of venting. Growth and mortality were observed in a number of the normally slow-growing cladorhizid stick sponges, indicating that these animals may exhibit life history strategies in the vicinity of vents that differ from those previously recorded. A novel facultative association between polynoids and anemones is proposed based on their correlated distributions. [ABSTRACT FROM AUTHOR]

A prevailing paradigm of hydrothermal vent ecology is that primary consumers feed on chemoautotrophic bacteria. However, for the purposes of reconstructing vent food webs and for tracking energy flow from the generation of rock and fluid chemistry through primary/ secondary productivity and consumption to the overlying water column, it remains unclear which consumers feed on which bacteria. In paired analyses of carbon and nitrogen tissue stable isotope values with unique 16S rRNA sequences from the stomach contents, we determined that two species of gastropod grazers appear to feed on epsilon-proteobacteria, while two other species have more diverse diets, including one species that consumes alpha-proteobacteria, planctomycetes, and non-green sulfur bacteria. Different carbon fixation pathways used by epsilon- and alpha-proteobacteria may account for the variation in the carbon stable isotope values among the consumers. Furthermore, our results indicate that trophic specialization and niche partitioning may contribute to the distribution and abundance of vent-endemic gastropods and support the hypothesis that consumers in the warmer habitats commonly feed on epsilon-proteobacteria that use the rTCA cycle, while in the cooler habitats they feed on additional bacteria that use the CBB cycle. These results suggest that the phylogenetic and metabolic diversity of free-living bacteria may play an important and previously overlooked role in facilitating species coexistence among primary consumers at hydrothermal vents and other chemosynthesis-based ecosystems. [ABSTRACT FROM AUTHOR]

Patterns of succession in Lau Basin hydrothermal vent communities determined with high-resolution imagery and in situ physico--chemical data collected over 4 yr and analyzed within a Geographic Information System show that Alviniconcha snails are a pioneering group, the snail Ifremeria nautilei is a mid-successional species, and the heat-intolerant mussel Bathymodiolus brevior dominates when venting declines. The associated fauna also changes as communities progress through the successional stages, and eventually non-vent--endemic deep-sea species appear when venting has mostly subsided. This is a unique example of primary succession in which the primary producers form symbiotic associations with mobile animals, resulting in successional patterns not observed in other systems. I. nautilei dominates newly formed substrates or venting sources where both I. nautilei and Alviniconcha spp. are already established (e.g., by migration), while Alviniconcha spp. seem to be better at colonizing newly active vents (e.g., by settlement) that are remote from colonized vents. Thus, on the scale of a 5--39 m² diffuse flow area or a single edifice, the mid-successional species dominates new substrates instead of the pioneering group. These communities are remarkably stable over long time periods relative to other hydrothermal vent regions. In addition to the sequential replacements of species as sites age and overa conditions change, Lau vent animals track changes in vent fluids and relocate themselves when local hydrothermal plumbing changes over small spatial scales. [ABSTRACT FROM AUTHOR]

Like hydrothermal vents along oceanic ridges, cold seeps are patchy and isolated ecosystems along continental margins, extending from bathyal to abyssal depths. The Atlantic Equatorial Belt (AEB), from the Gulf of Mexico to the Gulf of Guinea, was one focus of the Census of Marine Life ChEss (Chemosynthetic Ecosystems) program to study biogeography of seep and vent fauna. We present a review and analysis of collections from five seep regions along the AEB: the Gulf of Mexico where extensive faunal sampling has been conducted from 400 to 3300m, the Barbados accretionary prism, the Blake ridge diapir, and in the Eastern Atlantic from the Congo and Gabon margins and the recently explored Nigeria margin. Of the 72 taxa identified at the species level, a total of 9 species or species complexes are identified as amphi-Atlantic. Similarity analyses based on both Bray Curtis and Hellinger distances among 9 faunal collections, and principal component analysis based on presence/absence of megafauna species at these sites, suggest that within the AEB seep megafauna community structure is influenced primarily by depth rather than by geographic distance. Depth segregation is observed between 1000 and 2000m, with the middle slope sites either grouped with those deeper than 2000m or with the shallower sites. The highest level of community similarity was found between the seeps of the Florida escarpment and Congo margin. In the western Atlantic, the highest degree of similarity is observed between the shallowest sites of the Barbados prism and of the Louisiana slope. The high number of amphi-atlantic cold-seep species that do not cluster according to biogeographic regions, and the importance of depth in structuring AEB cold-seep communities are the major conclusions of this study. The hydrothermal vent sites along the Mid Atlantic Ridge (MAR) did not appear as ''stepping stones'' for dispersal of the AEB seep fauna, however, the south MAR and off axis regions should be further explored to more fully test this hypothesis. [ABSTRACT FROM AUTHOR]

The article presents the highlights of the exploration of the hydrothermal vents along the Galápagos Spreading Center (GSC) in the Galápagos Islands by scientists aboard the exploration vessel (E/V) Nautilus as of early 2016. Among the goals of the research are to collect fauna that are yet to be recorded and to visit the Iguanas black smoker vents. The vents that were explored are the Rosebud and Tempus Fugit vents.

Lamellibrachia luymesi and Seepiophila jonesi are co-occurring species of vestimentiferan tubeworms found at hydrocarbon seepage sites on the upper Louisiana slope of the Gulf of Mexico. Like all vestimentiferans, they rely on internal sulfide-oxidizing symbiotic bacteria for nutrition. These symbionts produce hydrogen ions as a byproduct of sulfide oxidation, which the host tubeworm needs to eliminate to prevent acidosis. The hydrothermal vent tubeworm Riftia pachyptila uses a high activity of P- and V-type H+-ATPases located in its plume epithelium to excrete protons. Unlike R. pachyptila, the seep species grow a posterior root, which they can use in addition to their plumes as a nutrient exchange surface. In this study we measured the ATPase activities of plume and root tissues collected from L. luymesi and S. jonesi, and used a combination of inhibitors to determine the relative activities of P- and V-type H+-ATPases. We found that the total H+-ATPase activity of their plumes was approximately 14 μmol h−1 g−1 wet weight, and that of their roots was between 5 and 7 μmol h−1 g−1 wet weight. These activities were more than ten times lower than those measured in R. pachyptila. We suggest that seep tubeworms might use passive channels to eliminate protons across their roots, in addition to ATP-dependant proton pumps located in their plumes and roots. In addition, we found strong differences between the types of ATPase activities in the plumes of L. luymesi and S. jonesi. While the H+-ATPase activity of L. luymesi plumes is dominated by P-type ATPases, S. jonesi has an unusually high activity of V-type H+-ATPases. We suggest that S. jonesi relies on its high V-type H+-ATPase activity to drive carbon dioxide uptake across its plume surface. L. luymesi, on the other hand, might rely partially on bicarbonate uptake across its root. [ABSTRACT FROM AUTHOR]

The ability of an organism to respond to changes in its environment depends upon its short- term physiological plasticity and the constraints of its genetic makeup. At hydrothermal vents and cold seeps, the spatially variable physiological characteristics of symbiont-bearing animals are often assumed to reflect short- term physiological adjustments to a patchy and dynamic chemical environment. However, the extent to which these spatially variable responses represent fixed characteristics unique to animals inhabiting the different environments (such as might arise from genetic differentiation) has not been tested. The seep mussel Bathymodiolus childressi depends upon methanotrophic bacteria for the bulk of its nutrition and inhabits a range of environments where it displays varying growth and body condition. In this study, we first investigated the multiscale environmental and physiological variability of B. childressi by measuring dissolved gas concentrations and mussel body condition in 12 mussel beds at four geographically distinct sites. Brine-dominated seeps tended to have higher methane and sulfide concentrations and host mussels of better body condition than petroleum-dominated sites. Then, using two transplant experiments, we evaluated whether local environmental conditions or stock effects determined the observed differences in growth and body condition of B. childressi. In all cases, mussels transplanted to new sites acquired or nearly acquired the characteristics of their host population, illustrating the primary role of the environment in determining the physiological characteristics of resident mussels. However, mussels from different sites sometimes responded differently to the same environment, suggesting stock-related effects also play a role in the spatial variation observed in the physiology of chemosynthetic fauna. [ABSTRACT FROM AUTHOR]

We present an investigation of the effects of disturbance and fecundity–tolerance strategies on community composition. We develop a theoretical model and apply it to macrofaunal communities at deep‐sea hydrothermal vents. We characterize community outcomes and find that dominance, coexistence, and alternative stable equilibria can all result from the interplay of disturbance regimes and fecundity–tolerance interactions. We show that fecundity–tolerance trade‐offs can permit coexistence under disturbance, but a strict fecundity–tolerance trade‐off is not required for coexistence to arise. We further describe how coexistence depends on habitat availability and disturbance regimes. Generally, our model elaborates on fecundity–tolerance strategies as a new axis of trait variation in coexistence theory. Natural disturbance regimes vary considerably across regions, and anthropogenic disturbance to vent communities will escalate with the advent of deep‐sea mineral extraction. We demonstrate how anthropogenic changes to disturbance regimes may impact species diversity, pushing communities over thresholds leading to local species extinction. Recommendations for Resource Managers Conventional wisdom suggests that disturbance‐adapted communities will not be strongly affected by novel perturbations associated with resource extraction or other human activities.We show that differing fecundity–tolerance strategies can mediate coexistence in disturbance‐prone deep‐sea environments, but that small changes to the disturbance regime can alter community composition and result in species extirpation.Natural disturbance should not be used as a rationale for expected low impacts of anthropogenic disturbance, for example, mining activities in management of deep‐sea mineral extraction. Appropriate impact studies should be performed for all communities where disturbances, such as mining, are planned. [ABSTRACT FROM AUTHOR]

Reports effects of deep-sea hydrothermal vents on the growth rates of the marine tubeworm Lamellibrachia. Details of the slow growth rate of the tubeworm in relation to other invertebrates near the vents; Use of a blue chitin stain in measuring growth; Suggestion that growth in individual tubeworms occurs episodically; Conclusions.

Abstract: Hydrothermal vent sulfide edifices contain some of the most extreme thermal and chemical conditions in which animals are able to live. As a result, sulfide edifices in the East Pacific Rise, Juan de Fuca Ridge, and Mid Atlantic Ridge vent systems often contain distinct faunal assemblages. In this study, we used high-resolution imagery and in-situ physico-chemical measurements within the context of a Geographic Information System (GIS) to examine community structure and niche differentiation of dominant fauna on sulfide edifices in the Eastern Lau Spreading Center (ELSC) and Valu Fa Ridge (VFR) in the Western Pacific Ocean. Our results show that ELSC and VFR sulfide edifices host two distinct types of communities. One type, that covers the majority of sulfide edifice faces, is overall very similar to nearby lava communities and biomass is dominated by the same chemoautotrophic symbiont-containing molluscs that dominate lava communities, namely the provannid gastropods Alviniconcha spp. and Ifremeria nautilei and the mytilid bivalve Bathymodiolus brevior. The spatial distribution of the dominant molluscs is often a variation of the pattern of concentric rings observed on lavas, with Alviniconcha spp. at the tops of edifices where exposure to vent flow is the highest, and I. nautilei and B. brevior below. Our physico-chemical measurements indicate that because of rapid dispersion of vent fluid, habitable area for symbiont-containing fauna is quite limited on sulfide edifices, and the realized niches of the mollusc groups are narrower on sulfide edifices than on lavas. We suggest that competition plays an important role in determining the realized distributions of the mollusc groups on edifices. The other habitat, present in small patches of presumably hot, new anhydrite, is avoided by the dominant symbiont-containing molluscs and inhabited by crabs, shrimp and polynoids that are likely more heat tolerant. The ratio of sulfide concentration to temperature anomaly of vent fluids was significantly different between sulfide edifice sites and lava sites in the southern vent fields but not in the northern vent fields. We suggest that this is due to increased sulfide consumption by a large microbial consortium associated with the more friable andesitic lava substrates in the south. [Copyright &y& Elsevier]

Abstract: Hydrothermal vent environments are characterized by large gradients of toxic chemicals and high temperatures, which play a significant role in defining species’ distributions. We used high-resolution imagery and spatially explicit in-situ physico-chemical measurements analyzed within a Geographic Information System (GIS) in order to characterize the spatial relations among different groups of megafauna, temperature, and chemistry within two discrete vent communities (40 and 50m2) on the Eastern Lau Spreading Center (ELSC). Chemical (sulfide and O2 concentrations) and temperature data were obtained from approximately 75 different locations within each community using in-situ instruments. All data were integrated into a GIS, which served as a visualization tool and enabled the data to be analyzed in a spatial context. Our results confirm the importance of abiotic variables in defining the distributions of some fauna and elucidate several biological associations that are consistent between the two communities. The provannid snail, Alviniconcha spp., appears to actively avoid temperatures above 32–46°C and/or sulfide concentrations exceeding approximately 260μM. Slightly higher average sulfide concentrations and temperatures were measured among aggregations of Ifremeria nautilei compared to aggregations of the mussel Bathymodiolus brevior; however, the presence of mixed aggregations of the two species indicates an overlap in requirements. The brachyuran crab, Austinograea spp., was consistently observed directly on symbiont-containing species, particularly Alviniconcha spp. The solitary snail, Eosipho desbruyeresi, was rarely observed on biological substrata, but was often (60% of its population at the most active site) within 5cm of symbiont-containing fauna, indicating a tolerance and preference for proximity to areas of high productivity. Densities and coverage of species differed substantially between the two communities despite high species overlap. Symbiont-containing species covered much larger areas at the more hydrothermally active site, ABE1, while shrimp and anemones occurred in relatively higher densities within the less-active site, TM1. This is the first study to thoroughly characterize realized distributions of megafauna at vent sites along the ELSC. [Copyright &y& Elsevier]

The vestimentiferan tubeworm Ridgeia piscesae is an ecosystem-structuring organism in the hydrothermal vent environments of the Northeast Pacific. During this study, a single representative aggregation of the long-skinny morphotype of R. piscesae from the main endeavor segment was monitored for 3 yr before being collected in its entirety with a hydraulically actuated collection device manipulated in situ by a research vehicle. Vestimentiferan growth rates in this aggregation were determined by staining the exterior of the tubes and measuring newly deposited tube sections. The average growth rate of R. piscesae in this aggregation was very low in both years of the growth study (3.2 mm yr−1). Although the incidence of plume damage from partial predation was very high (>95%), mortality was very low (<4% yr−1). The distribution and the very tight clustering of recently recruited individuals indicated gregarious settlement behavior that is hypothesized to be partly due to biotic cues from settled larvae. Coupled measurements of vent fluid sulfide concentration and temperature were used to calculate the exposure of the vestimentiferans to sulfide from short- and long-term temperature monitoring. Plume-level temperature records indicate that most of the time individuals in this aggregation were exposed to extremely low levels of vent fluid, and therefore sulfide (<0.1 μM), while their posterior sections were consistently exposed to sulfide concentrations in the 100 μM range. A rootball-like structure formed the common base of the aggregation. In contrast to the anterior sections of the tubeworm tubes, the portions of the tubes within the “rootball” were freely permeable to sulfide. The results of this study show that R. piscesae, unlike vestimentiferans from the East Pacific Rise, can survive and grow in areas of low diffuse vent flow with very low plume-level exposure to sulfide. We propose that this morphotype of R. piscesae has the ability to acquire sulfide from sources near their posterior ends, similar to some species of cold seep vestimentiferans from the Gulf of Mexico. The ability of this single species of vestimentiferan to survive low exposure to vent flow with low mortality coupled with sulfide uptake across posterior tube sections may help explain the occurrence of a single vent vestimentiferan species in a wide variety of habitat conditions at hydrothermal vent sites in the Northeast Pacific. [Copyright &y& Elsevier]