Since the first discovery of deep-sea hydrothermal vents along the Galápagos Rift in 1977, numerous vent sites and endemic faunal assemblages have been found along mid-ocean ridges and back-arc basins at low to mid latitudes. These discoveries have suggested the existence of separate biogeographic provinces in the Atlantic and the North West Pacific, the existence of a province including the South West Pacific and Indian Ocean, and a separation of the North East Pacific, North East Pacific Rise, and South East Pacific Rise. The Southern Ocean is known to be a region of high deep-sea species diversity and centre of origin for the global deep-sea fauna. It has also been proposed as a gateway connecting hydrothermal vents in different oceans but is little explored because of extreme conditions. Since 2009 we have explored two segments of the East Scotia Ridge (ESR) in the Southern Ocean using a remotely operated vehicle. In each segment we located deep-sea hydrothermal vents hosting high-temperature black smokers up to 382.8°C and diffuse venting. The chemosynthetic ecosystems hosted by these vents are dominated by a new yeti crab (Kiwa n. sp.), stalked barnacles, limpets, peltospiroid gastropods, anemones, and a predatory sea star. Taxa abundant in vent ecosystems in other oceans, including polychaete worms (Siboglinidae), bathymodiolid mussels, and alvinocaridid shrimps, are absent from the ESR vents. These groups, except the Siboglinidae, possess planktotrophic larvae, rare in Antarctic marine invertebrates, suggesting that the environmental conditions of the Southern Ocean may act as a dispersal filter for vent taxa. Evidence from the distinctive fauna, the unique community structure, and multivariate analyses suggest that the Antarctic vent ecosystems represent a new vent biogeographic province. However, multivariate analyses of species present at the ESR and at other deep-sea hydrothermal vents globally indicate that vent biogeography is more complex than previously recognised.
We examine the nature of variation in a hot vent tubeworm, Ridgeia piscesae, to determine how phenotypes are maintained and how reproductive potential is dictated by habitat. This foundation species at northeast Pacific hydrothermal sites occupies a wide habitat range in a highly heterogeneous environment. Where fluids supply high levels of dissolved sulphide for symbionts, the worm grows rapidly in a "short-fat" phenotype characterized by lush gill plumes; when plumes are healthy, sperm package capture is higher. This form can mature within months and has a high fecundity with continuous gamete output and a lifespan of about three years in unstable conditions. Other phenotypes occupy low fluid flux habitats that are more stable and individuals grow very slowly; however, they have low reproductive readiness that is hampered further by small, predator cropped branchiae, thus reducing fertilization and metabolite uptake. Although only the largest worms were measured, only 17% of low flux worms were reproductively competent compared to 91% of high flux worms. A model of reproductive readiness illustrates that tube diameter is a good predictor of reproductive output and that few low flux worms reached critical reproductive size. We postulate that most of the propagules for the vent fields originate from the larger tubeworms that live in small, unstable habitat patches. The large expanses of worms in more stable low flux habitat sustain a small, but long-term, reproductive output. Phenotypic variation is an adaptation that fosters both morphological and physiological responses to differences in chemical milieu and predator pressure. This foundation species forms a metapopulation with variable growth characteristics in a heterogeneous environment where a strategy of phenotypic variation bestows an advantage over specialization.
Vestimentiferan tubeworms are ecologically important members of deep-sea chemosynthetic communities, including hydrothermal vents and cold seeps. Some are community dominants and others are primary colonists of new vent sites; they include some of the longest living and fastest growing marine invertebrates. Their mechanisms of propagation, dispersal, and genetic exchange have been widely discussed. Direct sperm transfer from males to females has been documented in one species, Ridgeia piscesae, but others are known to discharge what are apparently primary oocytes. Brooding of embryos has never been observed in any vestimentiferan. These observations have led to the supposition that fertilization might be external in most species. Here we report sperm storage at the posterior end of the oviduct in five species, including tubeworms from both vents and seeps. We show experimentally that most eggs are inseminated internally, that fertilization rate is typically lower than 100%, that meiosis is completed after eggs are released from the female, and that the dispersal phase includes the entire embryonic period.
Deep-sea hydrothermal vents host lush chemosynthetic communities, dominated by endemic fauna that cannot live in other ecosystems. Despite over 500 active vents found worldwide, the Arctic has remained a little-studied piece of vent biogeography. Though located as early as 2001, the faunal communities of the Aurora Vent Field on the ultra-slow spreading Gakkel Ridge remained unsampled until recently, owing to difficulties with sampling on complex topography below permanent ice. Here, we report an unusual cocculinid limpet abundant on inactive chimneys in Aurora (3883–3884 m depth), describing it as Cocculina aurora n. sp. using an integrative approach combining traditional dissection, electron microscopy, molecular phylogeny, and three-dimensional anatomical reconstruction. Gross anatomy of the new species was typical for Cocculina , but it has a unique radula with broad, multi-cuspid rachidian where the outermost lateral is reduced compared to typical cocculinids. A phylogenetic reconstruction using the mitochondrial COI gene also confirmed its placement in Cocculina . Only the second cocculinid found at vents following the description of the Antarctic Cocculina enigmadonta , this is currently the sole cocculinid restricted to vents. Our discovery adds to the evidence that Arctic vents host animal communities closely associated with wood falls and distinct from other parts of the world.
Abstract. Previous work in the mud volcanoes from the Gulf of Cadiz (South Iberian Margin) revealed a high number of chemosymbiotic species, namely bivalves and siboglinid polychaetes. In this study we give an overview of the distribution and life styles of these species in the Gulf of Cadiz, determine the role of autotrophic symbionts in the nutrition of selected species using stable isotope analyses (δ13C, δ15N and δ34S) and investigate the intra-specific variation of isotope signatures within and between study sites. During our studies, we identified twenty siboglinidae and nine bivalve chemosymbiotic species living in fifteen mud volcanoes. Solemyid bivalves and tubeworms of the genus Siboglinum are widespread in the study area, whereas other species were found in a single mud volcano (e.g. "Bathymodiolus" mauritanicus) or restricted to deeper mud volcanoes (e.g. Polybrachia sp., Lamelisabella denticulata). Species distribution suggests that different species may adjust their position within the sediment according to their particular needs, and to the intensity and variability of the chemical substrata supply. Tissue stable isotope signatures for selected species are in accordance with values found in other studies, with thiotrophy as the dominant nutritional pathway, and with methanotrophy and mixotrophy emerging as secondary strategies. The heterogeneity in terms of nutrient sources (expressed in the high variance of nitrogen and sulphur values) and the ability to exploit different resources by the different species may explain the high diversity of chemosymbiotic species found in the Gulf of Cadiz. This study increases the knowledge on distributional patterns and resource partitioning of chemosymbiotic species and highlights how trophic fuelling varies on spatial scales with direct implications to seep assemblages and potentially to the biodiversity of continental margin.
Abstract. Previous work in the mud volcanoes from the Gulf of Cadiz revealed a high number of chemosymbiotic species, namely bivalves and siboglinid polychaetes. In this study we give an overview of the distribution and life styles of those species in the Gulf of Cadiz, determine the role of autotrophic symbionts in the nutrition of selected species using stable isotope analyses (δ13C, δ15N and δ34S) and investigate the intra-specific variation of isotope signatures within and between study sites. Twenty siboglinid and nine bivalve chemosymbiotic species have been identified and were found living in fifteen mud volcanoes during our studies. Solemyids bivalves and tubeworms of the genus Siboglinum are the most widespread, whereas other species were found in a single mud volcano (e.g. "Bathymodiolus" mauritanicus) or restricted to deeper mud volcanoes (e.g. Polybrachia sp., Lamelisabella denticulata). Species distribution suggests that different species may adjust their position within the sediment according to their particular needs and intensity and variability of the chemical substrata supply. Isotopic values found for selected species are in accordance with values found in other studies, with thiotrophy as the dominant nutritional pathway, and with methanotrophy and mixotrophy emerging as secondary strategies. The heterogeneity in terms of nutrient sources (expressed in the high variance of nitrogen and sulphur values) and the ability to exploit different resources by the different species may explain the high diversity of chemosymbiotic species found in the Gulf of Cadiz. This study increases the knowledge of the chemosymbiotic species in the Gulf of Cadiz, highlight the relevance of seep chemoautolithotrophic production in this area and provide a starting point for future trophic ecology studies.
Abstract Deep-sea mining in the Pacific Clarion-Clipperton Fracture Zone (CCZ), a low-energy sedimentary habitat with polymetallic nodules, is expected to have considerable and long-lasting environmental impact. The CCZ hosts extraordinarily high species diversity across representatives from all Domains of Life. Data on species biology and ecology remain scarce, however. The current study describes the reproductive biology of Ophiosphalma glabrum (Lütken & Mortensen, 1899) (Ophiosphalmidae) and Ophiacantha cosmica (Lyman, 1878) (Ophiacanthidae), two ophiuroids frequently found in the CCZ. Specimens collected in Spring 2015 and 2019 in four contract areas were examined morphologically and histologically. Size-class frequencies (disc diameter and oocytes feret diameters), sex ratios, gametogenic status, putative reproductive mode and a simple proxy for fecundity are presented. Habitat use differs in each. While Ophiosphalma glabrum is epibenthic, occurring as single individuals, Ophiacantha cosmica often form size-stratified groups living on stalked sponges, suggesting gregarious settlement or retention of offspring (though no brooding individuals were found). Further molecular analyses are needed to establish whether O. cosmica groups are familial. In Ophiosphalma glabrum , for which sample sizes were larger, sex ratios approximated a 1:1 ratio with no size-structuring. In both species, individuals were at various stages of gametogenic maturity but no ripe females were identified. Based on this, O. glabrum is most probably gonochoric. Reproductive mode remains inconclusive for Ophiacantha cosmica . Both species are presumptively lecithotrophic, with vitellogenic-oocyte feret diameters exceeding 250 µm. Oocyte feret diameters at times exceeded 400 µm in Ophiosphalma glabrum , indicating substantial yolk reserves. Estimates of instantaneous fecundity (vitellogenic specimens of O. glabrum only) were confounded by interindividual variability in gonad characteristics. The well-furnished lecithotrophic larvae of O. glabrum would be capable of dispersing even under food-impoverished conditions. The current study examines ophiuroid reproductive biology over multiple localities in the CCZ concurrently for the first time, at sites characterised by differing productivity regimes. The reproductive biology of each species is thus discussed with reference to past evolutionary (habitat stability), contemporary (food supply) and future environmental drivers (potential impacts of deep-sea mining).
Abstract Motivation Traits are increasingly being used to quantify global biodiversity patterns, with trait databases growing in size and number, across diverse taxa. Despite growing interest in a trait‐based approach to the biodiversity of the deep sea, where the impacts of human activities (including seabed mining) accelerate, there is no single repository for species traits for deep‐sea chemosynthesis‐based ecosystems, including hydrothermal vents. Using an international, collaborative approach, we have compiled the first global‐scale trait database for deep‐sea hydrothermal‐vent fauna – sFDvent ( s Div‐funded trait database for the F unctional D iversity of vent s). We formed a funded working group to select traits appropriate to: (a) capture the performance of vent species and their influence on ecosystem processes, and (b) compare trait‐based diversity in different ecosystems. Forty contributors, representing expertise across most known hydrothermal‐vent systems and taxa, scored species traits using online collaborative tools and shared workspaces. Here, we characterise the sFDvent database, describe our approach, and evaluate its scope. Finally, we compare the sFDvent database to similar databases from shallow‐marine and terrestrial ecosystems to highlight how the sFDvent database can inform cross‐ecosystem comparisons. We also make the sFDvent database publicly available online by assigning a persistent, unique DOI. Main types of variable contained Six hundred and forty‐six vent species names, associated location information (33 regions), and scores for 13 traits (in categories: community structure, generalist/specialist, geographic distribution, habitat use, life history, mobility, species associations, symbiont, and trophic structure). Contributor IDs, certainty scores, and references are also provided. Spatial location and grain Global coverage (grain size: ocean basin), spanning eight ocean basins, including vents on 12 mid‐ocean ridges and 6 back‐arc spreading centres. Time period and grain sFDvent includes information on deep‐sea vent species, and associated taxonomic updates, since they were first discovered in 1977. Time is not recorded. The database will be updated every 5 years. Major taxa and level of measurement Deep‐sea hydrothermal‐vent fauna with species‐level identification present or in progress. Software format .csv and MS Excel (.xlsx).
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