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.     

Digestive enzymes in marine invertebrates from hydrothermal vents and other reducing environments

The present study demonstrates the potential hydrolytic activities in the symbiont-containing tissues of the vent invertebrates Riftia pachyptila, Bathymodiolus thermophilus (collected in 1991 at the East Pacific Rise) and the shallow-water bivalve Lucinoma aequizonata (collected in 1991 from the Santa Barbara Basin). Activities of phosphatases, esterases, -glucuronidase and leucineaminopeptidase were comparable to those of digestive tract tissues of other marine invertebrates. A lack in most glycosidases as well as in trypsin and chymotrypsin was observed. Activities of lysozyme and chitobiase were rather high. In all vent invertebrates with symbionts and in L. aequizonata, the symbiont-containing tissues and the symbiont-free tissues had similar levels of enzymatic activities, indicating that polymeric nutrients could be hydrolysed after release from the symbionts and cellular uptake. The high activities of -fucosidase in all vent invertebrates as well as in the shallow-water bivalve L. aequizonata could point to the existence of a yet undescribed substrate available to hydrolysation. The ectosymbionts-carrying polychaete Alvinella pompejana (collected in 1991 at the East Pacific Rise, EPR) shows high lysozyme activities in its gut, consistent with the proposed food source of bacteria. For the vent crab Bythogrea thermydron (also collected in 1991 at the EPR) hydrolytic activities were highest in the gut, dominated by esterase and peptidase activities which support their proposed carnivorous food source. A snail and a limpet collected from R. pachyptila tubes showed high levels of chitobiase suggesting a food source of grazed bacteria or ingested R. pachyptila tube.     

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.     

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.     

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。     

Structure and function of Halice hesmonectes (Amphipoda: Pardaliscidae) swarms from hydrothermal vents in the eastern Pacific

Swarms of the pardaliscid amphipod Halice hesmonectes were sampled at low-temperature hydrothermal vents in the Venture Hydrothermal Fields of the East Pacific Rise. In excess of 3000 individuals were collected from a range of sites/habitats, and at two points in time (December 1991 and March 1994). The structure of swarms was described in terms of growth stages, the development of primary and secondary sex characteristics and the accumulation of nutritional reserves. The results demonstrated that swarms contained a wide range of growth stages, but that small juveniles and mature males and females were absent. Swarm structure was interpreted as an equilibrium dependent on the relative proportion of time spent by each growth stage in the swarm habitat. This equilibrium swarm structure was very similar at different sites and times. Swarms dominated by small juveniles occurred at the northern sites in December 1991 following a period of eruption and new venting in April 1991. Equilibrium swarm structure had become re-established at these sites by March 1994. The function of swarming was considered. The results indicated that swarming was not associated with reproductive activity. The available evidence, though not conclusive, suggested that swarms were associated with feeding. Received: 28 May 1999 / Accepted: 16 February 2000     

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.     

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.     

Feeding biology of the shrimp Rimicaris exoculata at hydrothermal vents on the Mid-Atlantic Ridge

A newly described species of shrimp, Rimicaris exoculata Williams and Rona, 1986, dominates the megafaunal community at two hydrothermal vent sites on the Mid-Atlantic Ridge. Behavioral observations and gut-content analyses indicate, that these shrimp ingest large amounts of sulfide particles from black smoker chimneys. We found no evidence for chemoautotrophic endosymbionts in R. exoculata, based on analyses of morphology, stable isotopes, lipopolysaccharides, and ribulose-1,5-bisphosphate carboxylase (RuBPCase) activity. Instead we suggest that the shrimp, are normal heterotrophs, grazing on free-living microorganisms associated with black smoker chimneys. We infer that high bacterial productivity is required to sustain populations of R. exoculata at these vent sites.     

Evolutionary relationships of deep-sea hydrothermal vent and cold-water seep clams (Bivalvia: Vesicomyidae): results from the mitochondrial cytochrome oxidase subunit I

Phylogenetic relationships among vesicomyid clams (Bivalvia: Vesicomyidae) and their placement within the order Heterodonta were examined using mitochondrial encoded cytochrome oxidase subunit I (COI) DNA sequences. The presently analyzed vesicomyids represent a recent monophyletic radiation that probably occurred within the Cenozoic. Nucleotide phylogenetic analyses resolved discrete clades that were consistent with currently recognized species: Calyptogena magnifica, C. ponderosa, Ectenagena extenta, C. phaseoliformis, Vesicomya cordata, Calyptogena n. sp. (Gulf of Mexico), C. kaikoi, C. nautilei, C. solidissima and C. soyoae (Type-A). However, specimens variously identified as: V. gigas, C. kilmeri, C. pacifica, and V. lepta comprised two “species complexes”, each composed of multiple evolutionary lineages. Most taxa are limited to hydrothermal-vent or cold-seep habitats, but the “vent” versus “seep” clams do not constitute separate monophyletic groups. Current applications of the generic names Calyptogena, Ectenagena, and Vesicomya are not consistent with phylogenetic inferences. Received: 24 July 1997 / Accepted: 22 August 1997     

Metabolic diversity in epibiotic microflora associated with the Pompeii wormsAlvinella pompejana andA. caudata (Polychaetae: Annelida) from deep-sea hydrothermal vents

Specimens of alvinellid polychaetes (Alvinella pompejana Desbruyères and Laubier, 1980 andA. caudata Desbruyères and Laubier, 1986) and their tubes were sampled from deep-sea hydrothermal vents at 13°N from the manned submersible Nautile during the Hydronaut cruise (October to November 1987) on the East Pacific Rise. Samples were subjected to bacterial analysis aboard the mother ship Nadir to detect bacteria involved in the nitrogen and sulphur cycles, in non-specific heterotrophic processes, and displaying resistance to selected heavy metals. Cultures were incubated at different temperatures under atmospheric and in situ (250 atm) pressures. Bacterial growth was observed in enrichment cultures for most metabolic types screened. Heavy-metal-resistant bacteria were also detected in many samples. No filamentous bacterial form was observed in the cultures. The results demonstrate a high metabolic diversity in episymbiotic flora, indicating that the worm (A. pompejana orA. caudata), its tube and its epiflora represent a complex micro-ecosystem.     

Response of hydrothermal vent vestimentiferan <Emphasis Type="Italic">Riftia pachyptila</Emphasis> to differences in habitat chemistry

Vestimentiferan tubeworms, which rely on intracellular sulfide-oxidizing autotrophic bacteria for organic carbon, flourish at deep-sea hydrothermal vents despite the erratic nature of their habitat. To assess the degree to which differences in habitat chemistry (sulfide, pH/CO₂) might impact host and symbiont metabolic activity, Riftia pachyptila tubeworms were collected from habitats with low (H₂S < 0.0001 mM) and high (up to 0.7 mM) sulfide concentrations. The elemental sulfur content of the symbiont-containing trophosome organ was lower in specimens collected from the low-sulfide site. Symbiont abundance, RubisCO activity, and trophosome carbon fixation rates were not significantly different for individuals collected from low- versus high-sulfide habitats. Carbonic anhydrase activities were higher in the anterior gas exchange organs of R. pachyptila from the low-sulfide habitat. Despite large differences in habitat chemistry, symbiont abundance and autotrophic potential were consistent, while the host appears to tailor carbonic anhydrase activity to environmental CO₂ availability.     

Molecular systematics of six Calanus and three Metridia species (Calanoida: Copepoda)

The discrimination of species of the copepod genus, Calanus (Copepoda; Calanoida), is problematical-especially in regions of sympatry. Although the species of Calanus exhibit exceptional morphological similarity, they are quite distinct in genetic character. The DNA base sequences of the mitochondrial large subunit (16S) ribosomal RNA (rRNA) gene unambiguously discriminated C. finmarchicus (Gunnerus 1765), C. glacialis (Jaschnov 1955), C. marshallae (Frost 1974), C. helgolandicus (Claus 1863), C. pacificus (Brodsky 1948), C. sinicus (Brodsky 1965), and C. hyperboreus (Kroyer 1838). Sequence differences among Calanus species for this gene portion range from 7.3% (between C. glacialis and C. marshallae) to 23.9% (between C. glacialis and C. sinicus). Differences among conspecific individuals were approximately 1 to 2%. [These sequence data were determined between April and November 1993; the sequenced domain is similar to that published previously in Bucklin et al. (1992) but are derived from analysis of additional individuals.] Statistical analysis of the sequence data using a variety of tree-building algorithms separated the taxa into one group of species corresponding to the C. finmarchicus group (C. finmarchicus, C. marshallae, and C. glacialis) and another ungrouped set of species corresponding to the C. helgolandicus group (C. helgolandicus, C. pacificus, and C. sinicus). The C. helgolandicus group may be older than the C. finmarchicus group, making the tree topology less reliable in this area. Calanus hyperboreus was an outlier; Nannocalanus minor (Claus 1863) was the outgroup. Similar analysis of Metridia species confirmed that M. lucens (Boeck 1864) and M. pacifica (Brodsky 1948) are distinct species; M. longa (Lubbock 1854) was still more divergent. These sequence data will allow the design of simple, molecular tools for taxonomic identifications. Diagnostic characters, assayed by rapid molecular protocols, will enable biological oceanographers to answer important questions about the distribution and abundance of all life stages (as well as patterns of reproduction) of morphologically similar species, such as those of Calanus.     

Sulfur contents and sulfur-isotope compositions of thiotrophic symbioses in bivalve molluscs and vestimentiferan worms

Total sulfur (S_TOT), elemental sulfur (S°) and sulfur-isotope compositions (δ³⁴S) of marine animals were analyzed to determine whether these chemical characteristics could help distinguish animals with a sulfur-based, thiotrophic nutrition from animals whose nutrition is based on methanotrophy or on more normal consumption of phytoplankton-derived organic matter. The presence of S° was almost entirely confined to the symbiont-containing tissues of thiotrophs, but was sometimes undetectable in thiotrophic species where sulfide availability was probably low. When S° contents were subtracted, the remaining tissue-sulfur concentrations were similar for all nutritional groups. δ³⁴S values were typically lower for thiotrophs than for other groups, although there was overlap in methanotroph and thiotroph values at some sites. Field evidence supported the existence of small to moderate (1 to 10‰)³⁴S fractionations in the uptake of sulfides and metabolism of thiosulfate. In general, a total sulfur content of >3% dry weight, the presence of elemental sulfur, and δ³⁴S values less than +5‰ can be used to infer a thiotrophic mode of nutrition. Received: 2 October 1997 / Accepted: 8 June 1998     

Feeding and swimming of lysianassid amphipods in a shallow cold-water bay

The potential for dispersal by lysianassid amphipods and their localization to carrion in a shallow cold-water bay in the Middle Saint Lawrence Estuary were assessed by means of endobenthic sampling, SCUBA observations, measures of swimming speeds, and by exposure of bait (50–100 g of fish) in traps. Seventy-five to 99.9% of animals attracted to traps were lysianassid amphipods belonging to five species. Lysianassid species were spatially segregated in the Bay at low tide but all were more or less dispersed at high tide. Second cohortAnonyx sarsi Steele and Brunel,Boeckosimus edwardsi andOnisimus littoralis (Krøyer) were more dispersed than the small first cohort individuals. Second cohortA. sarsi were crawlers or low (0–0.5 m off the bottom) suprabenthic swimmers in the day, but upper (0.5–2 m) suprabenthic swimmers at night. In contrast, first cohortA. sarsi were crawlers or low suprabenthic swimmers day-and-night, whileOrchomenella pinguis (Boeck) followed this swimming pattern at night but were generally akinetic in the day. Mean swimming speeds ofA. sarsi (13.6 cm s^-1) andOn. littoralis (12.1 cm s^-1) were 2 to 3 times greater than those ofOr. pinguis (7.4 cm s^-1) andPsammonyx nobilis (Stimpson) (4.4 cm s^-1). Catchability coefficients (i.e. ratio number of individuals per trap:endobenthic abundance) were 74 (A. sarsi), 8 (On. littoralis), 7 (Or. pinguis), and 0.7 (P. nobolis) m² of bottom. Gut content analysis indicated thatA. sarsi fed mostly on large carrion, whileOn. littoralis were markedly opportunistic, andOr. pinguis andP. nobilis relied on detritus, algae, and small crustaceans.     

Identification of archaeogastropod larvae from a hydrothermal vent community

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

深海热液喷口周围微氧耐压细菌的培养研究

研究采用析因实验设计,探讨了培养基的水源、培养温度和生长pH等因素对喷口细菌生长繁殖的影响,初步探索了常压下东太平洋某海底喷口周围微氧耐压细菌的生长条件。样品培养10d后,对培养液中细菌量进行显微计数,实验数据用SPSS11.0统计软件中的方差分析程序进行处理。统计结果显示:在标准大气压下,从研究样品中获得最多细菌数的较优培养条件为:培养基水源为单蒸水;生长pH为7.60;培养温度为50℃。在该条件的培养液中细菌平均浓度达2.596×10个/mL。研究结果为进一步认识、鉴定与开发利用深海喷口周围细菌资源打下了基础。     

Elemental concentrations of hydrothermal vent organisms from the Galápagos Rift

Concentrations of 40 elements were determined in tissues and whole bodies of mussels (Bathymodiolus thermophilus) and/or whole bodies of limpets (Neomphalus fretterae) collected from the submarine thermal springs on the Galápagos Rift in 1977. Concentrations of Ag, As, Fe, Mn, Mo, Se, Sr, and Zn were comparable or elevated relative to those of intertidal mollusks, which have been utilized as indicators of anthropogenic trace-metal contamination. Both species contained concentrations of some elements (e.g. Ag, As, Fe, Mn, Mo, Se, Zn) which corresponded with the elevated levels of those elements discharged from hydrothermal vents. Many elements were most enriched in the gills and digestive glands of the mussels. Iron:manganese ratios of those tissues were similar to ratios of pelagic and transitional sediments, indicating that metal-rich suspended particulates had been absorbed onto mucous membranes. Comparisons with other benthopelagic organisms were precluded, since the present data represent essentially the only trace-element analyses of deep-sea macroinvertebrates.     

Biochemical systematics of fishes of the genus Stegastes (Pomacentridae) from the Southern Marianas

The biochemical systematics of the subset of species of Stegastes (Pisces: Pomacentridae) available from coral reefs surrounding Guam, USA, was studied using three species of Pomacentrus as the outgroup. Analyses of allozymic variation at 14 presumptive loci (10 of which were polymorphic) using various evolutionary tree-construction techniques identified S. fasciolatus as the sister species of an assemblage comprised of S. albifasciatus, S. nigricans, and S. lividus: within this assemblage, S. lividus is the sister species of S. albifasciatus and S. nigricans, the pair of species which shared the most synapomorphic alleles and consistently exhibited the lowest degree of genetic distance relative to estimates for the other pairs of species studied. The consistency of ingroup topologies produced by parsimony analyses using Manhattan rectilinear distances computed from allele frequencies with those produced using the distance Wagner procedure and UPGMA (unweighted pair-group method of arithmetic averages) clustering, substantiates strongly the proposed phylogeny. The systematic scheme inferred from molecular data is concordant with predictions based on morphologic differences, i.e., differences in numbers of dorsal spines, that have been used as evidence in support of the proposal to partition Stegastes into at least two subgenera.