Genetic comparison of two populations of the deep-sea vent shrimp Rimicaris exoculata (Decapoda: Bresiliidae) from the Mid-Atlantic Ridge

Allozyme data are presented for populations of the bresiliid shrimp Rimicaris exoculata from two hydrothermal vent fields, Trans-Atlantic Geotraverse (TAG) and Broken Spur, located along the Mid-Atlantic Ridge. These indicate that all morphotypes of R. exoculata examined, including those previously interpreted as representing separate species, are conspecific. Conversely, genetic identity between a single specimen of Chorocaris sp. and R. exoculata was high for intergeneric comparisons. Genetic variation in the populations of R. exoculata (H _o =0.034 to 0.056) was in the lower range of that estimated for other vent organisms, but similar to values obtained for other species of caridean shrimps in previous genetic studies. F-statistics were used to examine the population structure of R. exoculata. Estimates of variance of allele frequencies among populations (F _ST ) between TAG and Broken Spur were very low (mean F _ST =0.001), indicating no significant genetic differentiation between these populations although they were separated by 370 km. The number of migrants per generation was estimated from F _ST and by a private-alleles method, and indicates that migration between the two fields exceeds 100 individuals per generation. This may be because of efficient larval or adult migration or a combination of both. Estimates of the correlation between homologous alleles between individuals within local populations (F _IS ) of R. exoculata were high at two enzyme loci and indicate a heterozygote deficiency which caused a significant deviation from genotype frequencies expected under Hardy-Weinberg conditions. This deficiency was caused by the occurrence of rare homozygous genotypes in small individuals. In large individuals, rare alleles decreased in frequency or disappeared completely. This is discussed in relation to previous genetic investigations on other vent and nonvent organisms.     

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.     

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.     

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.     

Vacuolate-attached filaments: highly productive Ridgeia piscesae epibionts at the Juan de Fuca hydrothermal vents

Vacuolate sulfur bacteria with high morphological similarity to vacuolate-attached filaments previously described from shallow hydrothermal vents (White Point, CA) were found at deep-sea hydrothermal vents. These filamentous bacteria grow in dense mats that cover surfaces and potentially provide a significant source of organic carbon where they occur. Vacuolate-attached filaments were collected near vents at the Clam Bed site of the Endeavour Segment of the Juan de Fuca Ridge and from the sediment surface at Escanaba Trough on the Gorda Ridge. A phylogenetic analysis comparing their 16S rRNA gene sequences to those collected from the shallow White Point site showed that all vacuolate-attached filament sequences form a monophyletic group within the vacuolate sulfur-oxidizing bacteria clade in the gamma proteobacteria. Abundance of the attached filaments was quantified over the length of the exterior surface of the tubes of Ridgeia piscesae worms collected from the Clam Bed site at Juan de Fuca yielding a per worm average of 0.070 ± 0.018 cm³ (n = 4). In agreement with previous results for White Point filaments, anion measurements by ion chromatography showed no detectable internal nitrate concentrations above ambient seawater (n = 9). For one R. piscesae tube worm “bush” at the Easter Island vent site, potential gross epibiont productivity is estimated to be 15 to 45× the net productivity of the worms.     

深海热液喷口微生物对矿物元素行为的影响

介绍了生长在深海热液喷口周围的微生物的种类及其生长繁殖的能量来源;并就其参与的矿物沉积和浸出行为做了阐述;最后在目前国内外研究现状的基础上提出了今后应该研究的方向。     

Trophic relationships among invertebrates at the Kairei hydrothermal vent field (Central Indian Ridge)

Exploration of hydrothermal vent systems in locations remote from well-studied sites allows ecologists to determine the degree of site-specific variation in trophic relationships among communities. A preliminary outline of the trophic structure of the Kairei hydrothermal vent community on the Central Indian Ridge (25°19.23′S; 70°02.42′E) is provided here, based on analysis of collections from an April 2001 expedition. Invertebrate biomass at Kairei is dominated by organic carbon with a δ¹³C isotopic value of about –13‰, due to the abundance of primary consumers (shrimp: Rimicaris aff. exoculata) and secondary consumers (anemones: Marianactis n. sp.) with this isotopic composition. Filamentous thiotrophic episymbionts on shrimp have been interpreted to be the major diet items of the shrimp and hence are the dominant primary producers within the community. Free-living autotrophic microorganisms are implicated as the dietary base for other invertebrate species. Four trophic groups are identified within the Kairei invertebrates based on carbon- and nitrogen-isotope ratios, but these groups do not always define discrete trophic levels. Ontogenetic shifts in diet are documented for R. aff. exoculata and brachyuran crabs (Austinograea n. sp.). Diets of symbiont-bearing mussels (Bathymodiolus aff. brevior) and two species of gastropods are isotopically constant throughout the range of sizes analyzed. There is a consistent but unexplained pattern of increasing nitrogen isotopic composition with increasing carbon isotopic composition in vent communities from geographically disjunct oceanic regions. Given the assumptions associated with interpretations of isotopic data, there remains a missing pool of carbon (presumably unsampled bacterial biomass) that contributes to the maintenance of the ¹³C- and ¹⁵N-enriched primary consumers in these ecosystems.     

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.     

Reproductive traits of pioneer gastropod species colonizing deep-sea hydrothermal vents after an eruption

The colonization dynamics and life histories of pioneer species determine early succession at nascent hydrothermal vents, and their reproductive ecology may provide insight into their dispersal and population connectivity. Studies on the reproductive traits of two pioneer gastropod species, Ctenopelta porifera and Lepetodrilus tevnianus, began within a year after an eruption on the East Pacific Rise (EPR) that eliminated vent communities near 9°50′N from late 2005/early 2006. Standard histology was used to examine gamete release, instantaneous female fecundity, and time to maturation. Both species exhibited two-component oocyte size–frequency distributions indicating quasi-continuous reproduction with high fecundity. In samples collected in December 2006, both C. porifera and L. tevnianus individuals were reproductively mature. The smallest reproducing C. porifera were 4.2 mm (males) and 5.4 mm (females) in shell length, whereas reproductive L. tevnianus were smaller (2.3 and 2.4 mm in males and females, respectively). Most C. porifera were large (>6.0 mm) compared to their size at metamorphosis and reproductively mature. In contrast, most L. tevnianus were small (<1.0 mm) and immature. Reproductive traits of the two species are consistent with opportunistic colonization, but are also similar to those of other Lepetodrilus species and peltospirids at vents and do not fully explain why these particular species were the dominant pioneers. Their larvae were probably in high supply immediately after the eruption, due to oceanographic transport processes from remote source populations.     

Locomotion modes of deep-sea cirrate octopods (Cephalopoda) based on observations from video recordings on the Mid-Atlantic Ridge

The behaviour of cirrate octopods of the genera Cirroteuthis and Grimpoteuthis in their natural habitat was studied using video recordings. Sequences were filmed during the French cruise “Faranaut”, from the manned submersible “Nautile” at depths between 2702 and 4527 m, in two zones in the Fifteen-Twenty Fracture Zone on the Mid-Atlantic Ridge. Four different modes of active locomotion, namely crawling, take-off, fin-swimming, and pumping, and one apparently passive mode of locomotion, umbrella-style drifting, were observed. Jet-propulsion, a characteristic mode of locomotion typically employed by cephalopods, was not observed in the cirrate octopods filmed, although breathing movements of the mantle to aerate the gills were assumed to generate some slight propulsion through the funnel. Fin-swimming was the mode of active locomotion most frequently observed. Neutral buoyancy was confirmed in one individual after capture. This buoyancy enables passive drifting in the umbrella-style attitude observed in Cirroteuthis spp., possibly using near-bottom ocean currents. Pumping, a mode of slow locomotion generated by peristaltic waves in the primary and intermediate webs, also observed in Cirroteuthis sp., is described here for the first time. The take-off mode of locomotion, a sudden, single contraction of the brachial crown and web, is also described; it was always followed by fin-swimming. No medusoid swimming of the type previously described for opisthoteuthid cirrates was observed. A flight response to the approach of or contact with a strange object, i.e. the submersible, using the crawling, take-off, or fin-swimming modes was observed. A ballooning response was observed in a high-stress situation when a C.magna individual was captured. Received: 27 November 1996 / Accepted: 9 December 1996     

Spatial and interannual variation in the faunal distribution at Broken Spur vent field (29°N, Mid-Atlantic Ridge)

Initial ecological observations at Broken Spur in 1993 suggested a low biomass relative to other deep-water vent communities known along the Mid-Atlantic Ridge. The persistence of a low shrimp biomass over 15 mo at Broken Spur vents appears to refute the hypothesis that the community is expanding through reproduction and immigration to occupy habitat vacated during a recent period of hydrothermal quiescence or catastrophe. Although the absence of “swarms” of shrimp similar to those found at the hydrothermal mounds of TAG (26°N) and Snake Pit (23°N) is a visually striking feature of the majority of venting structures known at Broken Spur, the biomass of fauna other than shrimp may not be significantly less than that of other Mid-Atlantic sites. The discovery of “swarms” of shrimp at Bogdanov Site, visited for the first time in 1994, suggests that availability of substratum exposed to the flow of hydrothermal fluids, which is a function of the topography of venting structures, may be a prerequisite for the development of these dense aggregations. Two testable predictions arise from this hypothesis. Firstly, dense aggregations of Rimicaris exoculata should occur at any other structures with a morphology similar to Bogdanov Site that may be discovered in the Broken Spur vent field, and should not occur at other isolated chimney structures that may be found. Secondly, “swarms” of shrimp should appear at any Broken Spur chimneys that develop into structures with a morphology more similar to that of Bogdanov Site in future. Received: 2 July 1997 / Accepted: 9 July 1997     

Feeding biology of the brackish-water oncholaimid nematode Adoncholaimus thalassophygas

The brackish-water inhabiting nematode Adoncholaimus thalassophygas (De Man, 1876) may be classified as an omnivore and predator according to the gut contents of the adults. A series of ¹⁴C experiments was conducted to investigate the feeding potential of labelled microorganisms and dissolved ¹⁴C glucose. The observed consistent failure to incorporate ¹⁴C from labelled microorganisms, and its consistent success with dissolved ¹⁴C glucose suggests that dissolved organic matter may be more important than microorganisms as a food source for his nematode. This corresponds also with direct observations of living worms which showed very little uptake of particulate food other than that originating from predation. Defecation and mean gulping rates were low and, therefore, ¹⁴C glucose uptake from solution can hardly be explained by the assumption that large amounts of water are swallowed and passing through the gut. By means of mucus secretion, A. thalassophygas forms compact agglutinations of detrital sediment which are sites of enhanced microbial activity. We suggest that hatched juveniles feed primarily on dissolved organic matter released by the intensive microbial activity within this habitat. Adults retain the ability to utilize dissolved organic matter, and supplement their diet by scavenging on carcasses and occasional predation. Thus, the biology of oncholaimid nematodes may be characterized by an intimate linkage to microbial metabolism, although they do not appear to feed directly upon microorganisms.     

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.     

Extensive gene flow among mytilid (Bathymodiolus thermophilus) populations from hydrothermal vents of the eastern Pacific

Prior studies of the hydrothermal vent mussel Bathymodiolus thermophilus (Bivalvia: Mytilidae), provided conflicting predictions about the dispersal ability and population structure of this highly specialized species. Analyses of morphological features associated with its larval shells revealed a feeding larval stage that might facilitate dispersal between ephemeral vent habitats. In contrast, an allozyme study revealed substantial genetic differentiation between samples taken from populations 2370 km apart on Galápagos Rift (Latitude 0°N) and the East Pacific Rise (13°N). To resolve the discrepancy between these studies, we examined allozyme and mitochondrial (mt) DNA variation in new samples from the same localities plus more recently discovered sites (9° and 11°N) along the East Pacific Rise. Although analysis of 26 enzyme-determining loci revealed relatively low levels of genetic variation within the five populations, no evidence existed for significant barriers to dispersal among populations. We estimated an average effective rate of gege flow (Nm) of 8 migrants per population per generation. Two common mtDNA variants predominated at relatively even frequencies in each population, and similarly provided no evidence for barriers to gene flow or isolation-by-distance across this species' known range. Larvae of this species appear to be capable of dispersing hundreds of kilometers along a continuous ridge system and across gaps separating non-contiguous spreading centers.     

Gene flow among vestimentiferan tube worm (Riftia pachyptila) populations from hydrothermal vents of the eastern Pacific

Allozyme data are presented for six discrete populations of the giant hydrothermal vent tube worm Riftia pachyptila Jones, 1981 collected throughout the species' known range along mid-ocean spreading ridges of the eastern Pacific Ocean. Contrary to an earlier report, levels of genetic variation are relatively high in this species. Estimates of gene flow based on F-statistics revealed that dispersal throughout the surveyed region is sufficiently high to counter random processes that would lead to losses of genetic diversity and significant population differentiation. R. pachyptila, like other species of tube worms, displays considerable morphologic variation among populations, but this diversity is not reflected in allozyme variation. Vestimentifera, in general, appear to show extensive phenotypic plasticity. In the light of the available genetic data, caution is warranted when making inferences about the taxonomic status of collections based on morphological variation alone. A general decrease in estimated rates of gene flow between geographically more distant populations supports the hypothesis that dispersal in this species follows a stepping-stone model, with exchange between neighboring populations in great excess of long-distance dispersal. High levels of gene flow have been recorded in a variety of vent fauna and may be a prerequisite for success of species found in the ephemeral habitats associated with regions of sea-floor hydrothermal activity.     

Feeding strategies of some demersal fishes of the continental slope and rise off the Mid-Atlantic Coast of the USA

Stomach contents of 729 fishes comprising 16 species were examined from the continental slope and rise off the Middle Atlantic States of the USA. Two main feeding modes among demersal deep-sea fishes were evident: those feeding primarily on pelagic food items, and those feeding on benthic invertebrates. Both pelagic and benthic predators were euryphagous. Most pelagic predators also fed on the epibenthos. These findings support Dayton and Hessler's (1972) suggestion that benthic predators should have a generalized diet which may be responsible for the high diversity found in the deep-sea infauna. The mesopelagic fauna is an important food source for some demersal fishes on the continental slope. Pelagic prey, which are also important to ecologically dominant demersal fishes on the lower slope and continental rise, may be nutritionally supported by suspended particulate organic matter in a nepheloid layer close to the bottom, and they may exist in much higher concentrations than in the bathypelagic zone above.Virginia Institute of Marine Science Contribution No. 835.