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- Presence of postlarval alvinocaridid
shrimps over south-west Indian Ocean hydrothermal vents, with
comparisons of the pelagic biomass at different vent sites
P. J. Herring.
Journal of the Marine Biological Association of the United
Kingdom, Vol. 86, No. 1, 2006, pp. 125-128.
Two types of alvinocaridid shrimp postlarvae were taken at
plume depth over the Kairei and Edmond hydrothermal vent fields
in the south-west Indian Ocean. These postlarvae were superficially
indistinguishable from similar postlarvae taken previously
over hydrothermal vents in the Atlantic. The micronekton invertebrate
taxa in the water column over the Kairei and Edmond sites
were similar to those in the Atlantic. The most frequently
taken deep-water fish at Kairei was an unidentified cyemid
snipe eel, whereas in the Atlantic its place was taken by
Gonostoma bathyphilum and species of the melamphaeid Scopeloberyx.
Previous sampling over East Pacific Rise hydrothermal sites
at 13 degree N failed to take any alvinocaridid postlarvae,
but the ostracod Gigantocypris agassizi was a major component
of the micronekton.
- Symbioses of methanotrophs and deep-sea
mussels (Mytilidae: Bathymodiolinae)
Eric G. DeChaine and Colleen M. Cavanaugh.
Progress in.molecular and subcellular biology, Vol. 41, 2006,
pp. 227-249.
The symbioses between invertebrates and chemosynthetic bacteria
allow both host and symbiont to colonize and thrive in otherwise
inhospitable deep-sea habitats. Given the global distribution
of the bathymodioline symbioses, this association is an excellent
model for evaluating co-speciation and evolution of symbioses.
Thus far, the methanotroph and chemoautotroph endosymbionts
of mussels are tightly clustered within two independent clades
of gamma Proteobacteria, respectively. Further physiological
and genomic studies will elucidate the ecological and evolutionary
roles that these bacterial clades play in the symbiosis and
chemosynthetic community. Due to the overall abundance of
the methanotrophic symbioses at hydrothermal vents and hydrocarbon
seeps, they likely play a significant, but as of yet unquantified,
role in the biogeochemical cycling of methane. With this in
mind, the search for methanotrophic symbioses should not be
restricted to these known deep-sea habitats, but rather should
be expanded to include methane-rich coastal marine and freshwater
environments inhabited by methanotrophs and bivalves. Our
current understanding of the bathymodioline symbioses provides
a strong foundation for future explorations into the origin,
ecology, and evolution of methanotroph symbioses, which are
now becoming possible through a combination of classical and
advanced molecular techniques.
- Characterization of Symbiont Populations
in Life-History Stages of Mussels From Chemosynthetic Environments
J. L. Salerno, S. A. Macko and S. J. Hallam, et al.
Biological Bulletin, Marine Biological Laboratory, Woods
Hole, Vol. 208, No. 2, Apr 2005, pp. 145-155.
The densities of chemoautotrophic and methanotrophic symbiont
morphotypes were determined in life-history stages (post-larvae,
juveniles, adults) of two species of mussels (Bathymodiolus
azoricus and B. heckerae) from deep-sea chemosynthetic environments
(the Lucky Strike hydrothermal vent and the Blake Ridge cold
seep) in the Atlantic Ocean. Both symbiont morphotypes were
observed in all specimens and in the same relative proportions,
regardless of life-history stage. The relative abundance of
symbiont morphotypes, determined by transmission electron
microscopy, was different in the two species: chemoautotrophs
were dominant (13:1-18:1) in B. azoricus from the vent site;
methanotrophs were dominant (2:1-3:1) in B. heckerae from
the seep site. The ratio of CH sub(4):H sub(2)S is proposed
as a determinant of the relative abundance of symbiont types:
where CH sub(4):H sub(2)S is less than 1, as at the Lucky
Strike site, chemoautotrophic symbionts dominate; where CH
sub(4):H sub(2)S is greater than 2, as at the seep site, methanotrophs
dominate. Organic carbon and nitrogen isotopic compositions
of B. azoricus ( delta super(13)C = -30ppt; delta super(15)N
= -9ppt) and B. heckerae ( delta super(13)C = -56ppt; delta
super(15)N = -2ppt) varied little among life-history stages
and provided no record of a larval diet of photosynthetically
derived organic material in the post-larval and juvenile stages.
- Composition and endemism of the deep-sea
hydrothermal vent fauna
T. Wolff.
Cahiers de Biologie Marine, Vol. 46, No. 2, 2005, pp. 97-104.
Based on enumerations from 1997/1998 and personal records,
an up-to-date survey of the hydrothermal vent fauna is presented
with number of species, genera and families of various taxa.
The number of recorded species is 712 and includes species
known exclusively from vents (71%), also occurring in cold
seeps and/or whale falls (5%), also recorded from non-vent/non-seep
environments (9%) or mainly referred to genus only (15%).
The number of genera is 373, of families 185. Molluscs, arthropods
and polychaetes are the prevailing groups (36, 34 and 18%,
respectively). Prosobranch gastropods number almost three
fourths of 257 mollusc species, copepods more than one third
of 225 crustaceans and phyllodocid polychaetes more than half
of 129 polychaetes. Calculation of vent endemism (after deduction
of 103 species of doubtful status) shows values far exceeding
those found elsewhere in the marine environment. On species
level it is 83.4%,with maximum values found in cirripeds (100%),
copepods (98%), prosobranch gastropods (89%), and polychaetes
(86%).The percentage of endemic genera is 45. All six cirriped
and most vestimentiferan genera are endemic, and other high
values are within copepods (66%), gastropods (58%), polychaetes
(51%) and decapod crustaceans (43%). Fourteen among the families
(8%) are found in vents exclusively, eight also in seeps.Original
Abstract: Fondee sur les listes publiees en 1997 et 1998 et
sur des donnees personnelles, une liste mise a jour de la
faune des sources hydrothermales est presentee. Le nombre
total d'especes signalees est 712, nombre qui inclut les especes
exclusivement infeodees aux sources hydrothermales (71%),
celles egalement presentes dans les zones de suintements froids
ou sur les carcasses de baleines (5%), celles egalement trouvees
dans d'autres types de milieux (9%), enfin celles identifiees
seulement au niveau generique (15%). Le nombre total de genres
est de 373, et celui des familles de 185. Mollusques, Arthropodes
et Polychetes sont les groupes predominants (respectivement
36, 34 et 18% du total). Le nombre de Gasteropodes prosobranches
represente a peu pres les trois quarts des 257 especes de
Mollusques, les Copepodes plus d'un tiers des 225 Crustaces
et les Phyllodocides plus de la moitie des 129 Polychetes.
Une fois soustraites les 103 especes de statut douteux, le
taux d'endemisme de la faune des sources hydrothermales atteint
des valeurs qui excedent toutes celles trouvees dans d'autres
environnements marins. Le taux d'endemisme au niveau specifique
est de 83 %, avec des valeurs maximales chez les Cirripedes
(100%), les Copepodes(98%), les Gasteropodes (89%) et les
Polychetes (86%). Le pourcentage de genres endemiques est
de 45%. Les six genres de Cirripedes et la plupart des genres
de Vestimentiferes sont endemiques; d'autres valeurs elevees
ont ete trouvees chez les Copepodes (66%), les Gasteropodes
(58%), les Polychetes (51%) et les Decapodes (43%). Quatorze
familles (8%) sont endemiques des sources hydrothermales.
- Deep-sea and shallow-water hydrothermal
vent communities; two different phenomena?; Shallow-water hydrothermal
venting
V. G. Tarasov, A. V. Gebruk, A. N. Mironov and L. I. Moskalev.
Chemical Geology, Vol. 224, No. 1-3, 15 Dec 2005, pp. 5-39.
Hydrothermal vent communities from 76 areas lying at depths
from 0 to 4100 m were split into two groups - "deep" and "shallow-water",
based on the occurrence of vent obligate (restricted to hydrothermal
vents) taxa of a high rank (genus and family). The two groups
were then compared based on a large set of environmental,
faunistic and community parameters. The degree of obligacy
of fauna sharply changes at the depth of approximately 200
m. In 21 shallow-water areas, located at depths 200 m) hydrothermal
communities differ from shallow-water ones (<200 m) in a much
higher ratio of vent obligate taxa. Deep-sea vent communities
are also distinguished by the lack of diatom and algae-bacterial
mats, phytoplankton, lower species richness, the development
of large sulphide structures, higher role of active vents
in structuring communities in space, replication of a set
of dominant taxa in communities within large regions reproduced
through the succession at newly formed vent fields, more pronounced
concentric and vertical zonality, higher biomass of macrofauna,
the important role of symbiotrophic forms dominating the biomass.
The ratio of species with a "point" species range (endemic
to small areas) is higher in the deep sea. There are no significant
differences between the vent and non-vent biogeography both
in shallow-waters and the deep ocean. Modern deep-sea hydrothermal
vent communities have a longer evolutionary history than their
shallow-water counterparts. To explain major differences between
deep and shallow-water hot vent communities, evolutionary
processes need to be analysed over long geological periods
and within global vertical zones. It is suggested that the
proportion of organic matter derived from chemosynthesis and
photosynthesis plays a crucial role in evolution. When calculated
for global zones this proportion increases with depth to a
certain critical level that separates deep and shallow-water
hot vent communities and fauna. Vertical zonation in the distribution
of obligate taxa in hydrothermal vent communities indicates
that this level is reached at approximately 200 m depth. Differences
between deep and shallow-water hydrothermal vent communities
are significant enough to distinguish the two different phenomena.
The principle differences are a higher ratio of vent obligate
taxa and a dominance of symbiotrophic forms in deep-sea hydrothermal
vent communities.
- Development of macrofaunal community
structure in mussel beds on the northern East Pacific Rise
J. C. Dreyer, K. E. Knick, W. B. Flickinger and C. L. Van
Dover.
Marine Ecology Progress Series, 121-134. 2005, pp. Vol. 302,
p.
Comparisons of macrofaunal community structure (species composition,
abundance, biovolume, rank order, species richness, diversity)
in hydrothermal vent mussel beds at 9 degree 50'N (9N) on
the East Pacific Rise were made from analysis of multiple
quantitative samples collected in 1999 and 2001 from each
of 3 mussel beds of known age (4 to 8 yr in 1999). In addition
to this time series approach, a chronosequence based on 2001
samples from 4 mussel beds at 9N and 1 mussel bed at 11 degree
25'N (11N) allowed comparison of macrofaunal community structure
for mussel beds ranging in age from 6 to >14 yr. Many aspects
of community structure (rank abundance patterns, dominant
species, and diversity) were similar in 1999 and 2001 samples.
Young (6 to 7 yr) and old (10 to 14 yr) mussel beds sampled
in 2001 could be distinguished using multivariate analyses,
based on species abundance and species biovolume matrices,
but the differences were in the relative abundance of the
dominant species rather than in any substantial changes in
the identity of the dominant species. The limpet Lepetodrilus
elevatus was the dominant species at all sites in terms of
abundance and biovolume; it and other dominant species may
have slightly greater fitness compared to other species that
allows them to persist as dominants regardless of the age
or location of mussel beds. Decreasing biovolume of macrofaunal
communities sampled at 9N and 11N in 2001 was correlated with
increasing age of the mussel beds and is postulated to be
largely a response to biological processes; including competition
for inorganic substrates by free living and symbiotic chemoautotrophs
and mussel induced mortality of larvae of macrofaunal species,
rather than the result of waning hydrothermal activity.
- Distribution, phylogenetic diversity
and physiological characteristics of epsilon-Proteobacteria
in a deep-sea hydrothermal field
Satoshi Nakagawa, Ken Takai and Fumio Inagaki, et al.
Environmental microbiology, Vol. 7, No. 10, Oct 2005, pp.
1619-1632.
Epsilon-Proteobacteria is increasingly recognized as an ecologically
significant group of bacteria, particularly in deep-sea hydrothermal
environments. In this study, we studied the spatial distribution,
diversity and physiological characteristics of the epsilon-Proteobacteria
in various microbial habitats in the vicinity of a deep-sea
hydrothermal vent occurring in the Iheya North field in the
Mid-Okinawa Trough, by using culture-dependent and -independent
approaches. The habitats studied were inside and outside hydrothermal
plume, and annelid polychaete tubes. In addition, we deployed
colonization devices near the vent emission. The polychaete
tubes harboured physiologically and phylogenetically diverse
microbial community. The in situ samplers were predominantly
colonized by epsilon-Proteobacteria. Energy metabolism of
epsilon-Proteobacteria isolates was highly versatile. Tree
topology generated from the metabolic traits was significantly
different (P = 0.000) from that of 16S rRNA tree, indicating
current 16S rRNA gene-based analyses do not provide sufficient
information to infer the physiological characteristics of
epsilon-Proteobacteria. Nevertheless, culturability of epsilon-Proteobacteria
in various microbial habitats differed among the phylogenetic
subgroups. Members of Sulfurimonas were characterized by the
robust culturability, and the other phylogenetic subgroups
appeared to lose culturability in seawater, probably because
of the sensitivity to oxygen. These results provide new insight
into the ecophysiological characteristics of the deep-sea
hydrothermal vent epsilon-Proteobacteria, which has never
been assessed by comparative analysis of the 16S rRNA genes.
- Epifaunal community structure associated
with Riftia pachyptila aggregations in chemically different
hydrothermal vent habitats
Breea Govenar, Nadine Le Bris and Sabine Gollner, et al.
Marine Ecology Progress Series, 67-77. 2005, pp. Vol. 305,
p.
The vestimentiferan tubeworm Riftia pachyptila (Polychaeta:
Sibloglinidae) often dominates early succession stages and
high productivity habitats at low-temperature hydrothermal
vents on the East Pacific Rise. We collected 8 aggregations
of R. pachyptila and the associated epifaunal community at
2 discrete sites of diffuse hydrothermal activity, in December
2001 and December 2002. Because of the high spatial and temporal
variability of the biotic and abiotic factors related to hydrothermal
vent activity, significant differences in the structure and
the composition of the community were expected to occur at
the scale of either 1 yr or 500 m distance between very different
sites. There was no significant difference in the temperature
ranges of the diffuse flow between sites or years, even though
the environmental conditions were very different at the 2
sites. At 1 site (Riftia Field), the diffuse hydrothermal
fluids had relatively low concentrations of sulfide, low pH,
and high concentrations of iron. At the other site (Tica),
the diffuse hydrothermal fluids had higher sulfide concentrations,
the pH was closer to neutral, and iron was undetectable. The
physiological condition of R. pachyptila appeared to reflect
the availability of sulfide at each site. However, the structure
and the composition of the epifaunal community were remarkably
similar between sites and years, with the exception of a few
species. Aggregations of R. pachyptila support high local
species diversity relative to the surrounding seafloor and
high community similarity in different hydrothermal vent habitats.
- Hydrothermal vent octopuses of Vulcanoctopus
hydrothermalis feed on bathypelagic amphipods of Halice hesmonectes
J. R. Voight.
Journal of the Marine Biological Association of the United
Kingdom, Vol. 85, No. 4, 2005, pp. 985-988.
A feeding frenzy of 12 octopuses of Vulcanoctopus hydrothermalis
was observed from the manned submersible Alvin at Parigo,
a 2620 m deep hydrothermal vent on the East Pacific Rise.
The aggregated benthic octopuses at the active vent used their
arms and webs to forage on bathypelagic amphipods, apparently
targeting their attacks based on contract with the swarming
amphipods. Individual octopuses wrapped their arms around
the mantles of smaller octopuses, apparently in competition
for prey. Although members of the prey species, Halice hesmonectes,
are individually small (<5 mm long), the density of their
swarms may make them attractive prey for the octopus. Inactive
sulphide spires encircled part of this vent site; octopuses
that climbed these spires had easy access to the dense prey
swarms. The presence of the spires may uniquely enable this
site to support simultaneous foraging by large numbers of
octopuses.
- Lost City Life
Antje Boetius.
Science (Washington), Vol. 307, No. 5714, 4 Mar 2005, pp.
1420-1422.
A new type of hydrothermal vent field in the Atlantic Ocean
called Lost City was discovered in 2000. As Boetius describes
in her Perspective, recent detailed mapping of this hydrothermal
vent field as well as chemical and microbial analyses (Kelley
et al.) reveal that an extraordinary array of micro- and macro-organisms
inhabit the giant white carbonate chimneys of Lost City.
- Proposed nitrate binding by hemoglobin
in Riftia pachyptila blood
Edda Hahlbeck, Mark A. Pospesel, Franck Zal, James J. Childress
and Horst Felbeck.
Deep Sea Research (Part I, Oceanographic Research Papers),
Vol. 52, No. 10, Oct 2005, pp. 1885-1895.
Riftia pachyptila lives in the unstable environment at hydrothermal
vent sites along oceanic spreading zones in the Eastern Pacific.
The tubeworm has a symbiosis with intracellular carbon-fixing
and sulfide-oxidizing bacteria. Nitrate is the main source
of nitrogen available from their habitat. This compound serves
as a substrate either for nitrate respiration or for biosynthesis
after transformation into ammonia. Very high nitrate (up to
3.2 mM) and nitrite (up to 0.8 mM) concentrations in vascular
blood of R. pachyptila indicate a novel uptake mechanism.
The dialysis experiments reported here demonstrate the binding
and transport of nitrate to the symbionts by high molecular
weight components in the blood, most likely hemoglobin. The
extent to which nitrate is bound differed markedly between
blood from different animals. In addition, a strong inverse
correlation was found between the concentrations of sulfide
and nitrate in vascular blood, as well as between the sulfur
content of trophosome and the nitrate content of vascular
blood. Specimens with low sulfur stores showed much lower
nitrate levels than those with pale green trophosome due to
high levels of elemental sulfur.
- Role of thermal conditions in habitat
selection by hydrothermal vent gastropods
Amanda E. Bates, Verena Tunnicliffe and Raymond W. Lee.
Marine Ecology Progress Series, 2005, Vol. 305, pp. 1-15
Habitat selection by 3 Juan de Fuca Ridge gastropod species
relates to their thermal environment. Both collections and
images taken along transects document the small-scale abundance
patterns of each species with respect to temperature and distance
from vent flows. Lepetodrilus fucensis and Depressigyra globulus
were most abundant at distances from vent flows of 0 to 25
cm in warm fluids with high temperature variability (10 +/-
5 degree C) over several time scales. Both species were also
abundant at 26 to 50 cm, where temperatures were lower with
less variability (4 +/- 1 degree C). Provanna variabilis was
most abundant from 51 to 75 cm, where temperatures were stable
(3 +/- 0.5 degree C). All species were absent where maximum
fluid temperatures reached 18 degree C and their substratum
coverage was related to temperature. When presented with a
choice in vent fluids from 10 to 2 degree C, L. fucensis and
D. globulus moved to areas with temperatures above 5 degree
C, while P. variabilis showed no preference. In species-specific
temperature preference experiments, L. fucensis and D. globulus
aggregated between 5 and 13 degree C, while P. variabilis
occupied areas with significantly lower temperatures from
4 to 11 degree C. These experimental temperature preferences
correspond with their thermal environments. Upper temperature
limits are moderate; extreme abiotic variability in higher
temperature fluids may constrain these 3 species. We conclude
thermal conditions are a primary determinant of habitat selection,
thereby driving gastropod abundance patterns. However, other
factors likely contribute. Space competition nearest vent
flows may result in the displacement of individuals of these
species to low quality habitats.
- Selective predation by the zoarcid
fish Thermarces cerberus at hydrothermal vents
G. Sancho, C. R. Fisher and S. Mills, et al.
Deep Sea Research (Part I, Oceanographic Research Papers),
Vol. 52, No. 5, May 2005, pp. 837-844.
This study investigates predation by the vent zoarcid fish
Thermarces cerberus through gastrointestinal analyses of 27
specimens collected with the submersible ALVIN at vents at
950'N on the East Pacific Rise. T. cerberus fed most frequently
on gastropod mollusks (mainly Lepetodrilus elevatus) and amphipod
crustaceans (mainly Ventiella sulfuris). Species found occasionally
in high abundance included the swarming amphipod Halice hesmonectes
and the snail Cyathermia naticoides. Other items also found
in gastrointestinal tracts, but in very low numbers, included
polychaete worms, crustaceans and unidentified tissue clumps.
The comparison between the size distribution of L. elevatus
limpets ingested by T. cerberus and those found attached to
vestimentiferan tubes suggest that the fish may selectively
prey on large limpets. If the selective removal of large Lepetodrilus
spp. limpets by T. cerberus does occur, then it would have
potential community-level consequences at hydrothermal vents,
since these mobile gastropods appear to inhibit the settlement
of sessile vent species, including tube-building worms. Our
results suggest possible direct and indirect effects of T.
cerberus on benthic community structure at hydrothermal vents
on the East Pacific Rise.
- A Serpentinite-Hosted Ecosystem: The
Lost City Hydrothermal Field
Deborah S. Kelley, Jeffrey A. Karson and Gretchen L. Fruh-Green,
et al.
Science (Washington), Vol. 307, No. 5714, 4 Mar 2005, pp.
1428-1434.
The serpentinite-hosted Lost City hydrothermal field is a
remarkable submarine ecosystem in which geological, chemical,
and biological processes are intimately interlinked. Reactions
between seawater and upper mantle peridotite produce methane-and
hydrogen-rich fluids, with temperatures ranging from <40 degree
to 90 degree C at pH 9 to 11, and carbonate chimneys 30 to
60 meters tall. A low diversity of microorganisms related
to methane-cycling Archaea thrive in the warm porous interiors
of the edifices. Macrofaunal communities show a degree of
species diversity at least as high as that of black smoker
vent sites along the Mid-Atlantic Ridge, but they lack the
high biomasses of chemosynthetic organisms that are typical
of volcanically driven systems.
- Sperm Storage, Internal Fertilization,
and Embryonic Dispersal in Vent and Seep Tubeworms (Polychaeta:
Siboglinidae: Vestimentifera)
A. Hilario, C. M. Young and P. A. Tyler.
Biological Bulletin, Marine Biological Laboratory, Woods
Hole, Vol. 208, No. 1, Feb 2005, pp. 20-28.
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.
- The ubiquitous mussel: Bathymodiolus
aff. brevior symbiosis at the Central Indian Ridge hydrothermal
vents
Z. P. McKiness and C. M. Cavanaugh.
Marine Ecology Progress Series, Vol. , No. 23 Jun 2005, pp.
Vol. 295, pp. 183-190
The discovery and exploration of hydrothermal vents on the
Central Indian Ridge (CIR) yielded invaluable samples for
further resolution of the biogeography of chemoautotrophic
symbioses. Mytilid mussels were collected from 2 CIR hydrothermal
vent fields and preliminary molecular analyses suggested phylogenetic
affinity with the western Pacific vent mussel Bathymodiolus
brevior. Resolving whether this mussel, designated B. aff.
brevior, hosts a dual symbiosis, as characterized for the
2 Mid-Atlantic Ridge hydrothermal vent mussel species, or
a single chemoautotrophic symbiosis, as seen in the Pacific
vent mussel species, will provide insight into the evolutionary
history of mytilid symbioses. Ultrastructural, physiological,
and molecular evidence are reported herein which support the
presence of a single endosymbiont phylotype with chemoautotrophic
metabolism. Phylogenetic analyses placed this symbiont in
the same clade as the vesicomyid clam symbionts, prompting
discussion regarding the evolutionary origin of chemoautotrophic
symbioses in vent bivalves.
- Vertical, lateral and temporal structure
in larval distributions at hydrothermal vents
L. S. Mullineaux, S. W. Mills, A. K. Sweetman, A. H. Beaudreau,
A. Metaxas and H. L. Hunt.
Marine Ecology Progress Series, 2 Jun 2005, pp. Vol. 293,
pp. 1-16
We examined larval abundance patterns near deep-sea hydrothermal
vents along the East Pacific Rise to investigate how physical
transport processes and larval behavior may interact to influence
larval dispersal from, and supply to, vent populations. We
characterized vertical and lateral distributions and temporal
variation of larvae of vent species using high-volume pumps
that recovered larvae in good condition (some still alive)
and in high numbers (up to 450 individuals sample super(-1)).
Moorings supported pumps at heights of 1, 20, and 175 m above
the seafloor, and were positioned directly above and at 10s
to 100s of meters away from vent communities. Sampling was
conducted on 4 cruises between November 1998 and May 2000.
Larvae of 22 benthic species, including gastropods, a bivalve,
polychaetes, and a crab, were identified unequivocally as
vent species, and 15 additional species, or species-groups,
comprised larvae of probable vent origin. For most taxa, abundances
decreased significantly with increasing height above bottom.
When vent sites within the confines of the axial valley were
considered, larval abundances were significantly higher on-vent
than off, suggesting that larvae may be retained within the
valley. Abundances of all vent species varied significantly
among sample dates; the variation was not synchronized among
taxa, except for consistently low abundances during November
1998. Lateral distributions did not vary among major larval
groups (gastropods, polychaetes and bivalves), although polychaetes
showed anomalously high abundances off-vent at 1 m above bottom.
Lateral patterns also did not vary among species of gastropods,
indicating that hydrodynamic processes may be transporting
diverse species in similar ways. However, the species-level
differences in temporal patterns indicate that there is substantial
discontinuity in the abundance of individual species at vent
communities, possibly due to timing of spawning and/or behavioral
interactions with flow.
- Metagenomic analysis of the episymbiont
community associated with Alvinella Pompejana, a deep-sea hydrothermal
vent polychaete
S. C. Cary, B. J. Campbell and A. Ghadiri, et al.
Marine Ecology Progress Series, Vol. 284, 2004, pp. 237-251
The shrimp Rimicaris exoculata dominates the megafauna of
some Mid-Atlantic Ridge hydrothermal vent fields. This species
harbours a rich bacterial epibiosis inside its gill chamber.
At the 'Rainbow' vent site (36' 14.0'N), the epibionts are
associated with iron oxide deposits. Investigation of both
bacteria and minerals by scanning electron microscopy (SEM)
and X-ray microanalysis (EDX) revealed 3 distinct compartments
in the gill chamber: (1) the lower pre-branchial chamber,
housing bacteria but devoid of minerals; (2) the 'true' branchial
chamber, containing the gills and devoid of both bacteria
and minerals; and (3) the upper pre-branchial chamber, housing
the main ectosymbiotic bacterial community and associated
mineral deposits. Our chemical and temperature data indicated
that abiotic iron oxidation appears to be kinetically inhibited
in the environment of the shrimps, which would explain the
lack of iron oxide deposits in the first 2 compartments. We
propose that iron oxidation is microbially promoted in the
third area. The discrepancy between the spatial distribution
of bacteria and minerals suggests that different bacterial
metabolisms are involved in the first and third compartments.
A possible explanation lies in the modification of physico-chemical
conditions downstream of the gills that would reduce the oxygen
content and favours the development of bacterial iron-oxidizers
in this Fell-rich environment. A potential role of such iron-oxidizing
symbionts in the shrimp diet is suggested. This would be unusual
for hydrothermal ecosystems, in which most previously described
symbioses rely on sulphide or methane as an energy source.
- Community structure in Florida Escarpment
seep and Snake Pit (Mid-Atlantic Ridge) vent mussel beds
M. Turnipseed, C. D. Jenkins and C. L. Van Dover.
Marine Biology, Vol. 145, No. 1, Jul 2004, pp. 121-132.
Comparisons between invertebrate communities hosted by similar
foundation species under different environmental conditions
permit identification of patterns of species distributions
that might be characteristic of the different ecosystems.
Similarities and differences in community structure between
two major types of chemosynthetic ecosystems were assessed
by analyzing samples of invertebrates associated with Bathymodiolus
heckerae Gustafson et al. mussel beds at the Florida Escarpment
seep (Gulf of Mexico, 26 degree 01.8N; 84 degree 54.9W; October
2000) and B. puteoserpentis von Cosel et al. mussel beds at
the Snake Pit vent (Mid-Atlantic Ridge, 23 degree 22.1N; 44
degree 56.9W; July 2001). Macrofaunal species richness was
nearly twice as high in the seep mussel bed compared to the
vent mussel bed, and only a single morphospecies, the ophiuroid
Ophioctenella acies Tyler et al., was shared between the sites.
Similarities between the two faunas at higher taxonomic levels
(genus and family) were evident for only a small percentage
of the total number of taxa, suggesting that evolutionary
histories of many of these seep and vent macrofaunal taxa
are not shared. The taxonomic distinctiveness of the seep
and vent mussel-bed macrofaunal communities supports the hypothesis
that environmental and oceanographic barriers prevent most
taxa from occupying both types of habitats. Macrofaunal community
heterogeneity among samples was similar in seep and vent mussel
beds, indicating that spatial scales of processes regulating
community variability may be similar in the two types of ecosystems.
Suspension feeders were not represented in the macrofauna
of seep or vent mussel beds. Primary consumers (deposit feeders
and grazers) contributed more to the total abundance of macrofauna
of seep mussel beds than vent mussel beds; secondary consumers
(polychaetes and shrimp) were more abundant in the vent mussel
beds.
- Composition of a One-Year-Old Riftia
pachyptila Community following a; Clearance Experiment: Insight
to Succession Patterns at Deep-Sea; Hydrothermal Vents
B. Govenar, M. Freeman, D. C. Bergquist, G. A. Johnson and
C. R. Fisher.
Biological Bulletin, Vol. 207, No. 3, December, 2004 2004,
pp. 177-182.
Following eruptions on the East Pacific Rise, tubeworms Tevnia
jerichonana and Riftia pachyptila quickly colonize the basalt-hosted
hydrothermal vents, soon growing to dominate these habitats.
Colonization by the vent mussel Bathymodiolus thermophilus
follows, with the mussels eventually overgrowing the tubeworms.
Tubeworms and mussels harbor sulfur-oxidizing chemoautotrophic
endosymbionts, which provide the bulk of their nutrition.
Mussels have maintained the ability to filter-feed, allowing
them to occupy a wider range of conditions, tolerating declining
hydrothermal vent activity better than the tubeworms. An experiment
to understand the mechanism of ecological succession in these
vent habitats was recently undertaken. A manipulative experiment
tested the hypothesis that the mussels prevent further colonization
by tubeworms. In May 1998, six aggregations of tubeworms were
cleared from a site, with location, source of diffuse flow,
and venting fluid temperatures recorded. Six hundred nearby
mussels were collected; 200 were deployed over each of three
cleared sources of diffuse flow to test the mussels' effects
on the subsequent colonization by the tubeworms. Observations
were made after 1 year; growth rates of R. pachyptila were
estimated from tube length. Colonization rates on the cleared
sources of diffuse flow were similar to those on basalt blocks.
There was an abundance of gastropods in this community, though
as grazers and scavengers, none relied on the tubeworms for
nutrition. None of the transplanted mussels persisted on the
cleared sources of diffuse flow. Situated above the venting
source, the juvenile mussels were exposed to less concentrated
hydrothermal flow, the temperatures and concentrations of
reduced chemicals were lower, and the oxygen concentration
was higher than in the fluids emanating from the basalt. Results
indicate that R. pachyptila may modify the vent habitat to
facilitate the colonization of other species in the diffuse-flow
habitat of basalt-hosted hydrothermal vents.
- Distribution of bacteria and associated
minerals in the gill chamber of the vent shrimp Rimicaris exoculata
and related biogeochemical processes
M. Zbinden, N. Le Bris, F. Gaill and P. Compere.
Marine Ecology Progress Series, 2004, pp. Vol. 284, pp. 237-251
The shrimp Rimicaris exoculata dominates the megafauna of
some Mid-Atlantic Ridge hydrothermal vent fields. This species
harbours a rich bacterial epibiosis inside its gill chamber.
At the 'Rainbow' vent site (36' 14.0'N), the epibionts are
associated with iron oxide deposits. Investigation of both
bacteria and minerals by scanning electron microscopy (SEM)
and X-ray microanalysis (EDX) revealed 3 distinct compartments
in the gill chamber: (1) the lower pre-branchial chamber,
housing bacteria but devoid of minerals; (2) the 'true' branchial
chamber, containing the gills and devoid of both bacteria
and minerals; and (3) the upper pre-branchial chamber, housing
the main ectosymbiotic bacterial community and associated
mineral deposits. Our chemical and temperature data indicated
that abiotic iron oxidation appears to be kinetically inhibited
in the environment of the shrimps, which would explain the
lack of iron oxide deposits in the first 2 compartments. We
propose that iron oxidation is microbially promoted in the
third area. The discrepancy between the spatial distribution
of bacteria and minerals suggests that different bacterial
metabolisms are involved in the first and third compartments.
A possible explanation lies in the modification of physico-chemical
conditions downstream of the gills that would reduce the oxygen
content and favours the development of bacterial iron-oxidizers
in this Fell-rich environment. A potential role of such iron-oxidizing
symbionts in the shrimp diet is suggested. This would be unusual
for hydrothermal ecosystems, in which most previously described
symbioses rely on sulphide or methane as an energy source.
- Early Jurassic Hydrothermal Vent Community
From The Franciscan Complex, California
C. T. S. Little, T. Danelian, R. J. Herrington and R. M.
Haymon.
Journal of Paleontology, Vol. 78, No. 3, May 2004, pp. 542-559.
The Figueroa sulfide deposit located in Franciscan Complex
rocks in the San Rafael Mountains, California, contains the
only known Jurassic hydrothermal vent community. Based on
radiolarian biostratigraphy it is Pliensbachian (early Jurassic)
in age. The Figueroa fossil organisms lived at a deepwater,
high temperature vent site located on a mid-ocean ridge or
seamount at an equatorial latitude. The vent site was then
translated northeastward by the motion of the Farallon Plate
and was subsequently accreted to its present location. The
vent fossils are preserved as molds of pyrite and there is
no remaining shell or tube material. The fossil assemblage
is specimen rich, but of low diversity, and comprises, in
order of decreasing abundance, vestimentiferan worm tubes,
rhynchonellide brachiopods (Anarhynchia cf. gabbi), and trochoidean
gastropods (Francisciconcha maslennikovi new genus and species).
These fossils represent only primary consuming organisms,
some of which may have had chemosynthetic microbial endosymbionts,
like many modern dominant vent animals. The Figueroa vent
assemblage shares vestimentiferan tube worms and gastropods
with other fossil and modern vent communities, but is unique
in having rhynchonellide brachiopods. It shares this feature
with contemporary Mesozoic cold seep communities. Many other
taxonomic groups found at modern vent sites are missing from
the Figueroa assemblage. The presence of vestimentiferan tube
worm fossils in the Figueroa deposit is at odds with the supposed
time of origin of the modern vestimentiferans ( similar to
100 Ma), based on molecular data.
- Experimental ecology at deep-sea hydrothermal
vents: a perspective
C. L. Van Dover and R. A. Lutz.
Journal of experimental marine biology and ecology, Vol.
300, No. 1-2, Mar 2004, pp. 273-307.
In situ and laboratory experiments conducted over the past
quarter of a century have greatly increased our understanding
of the ecology of deep-sea hydrothermal systems. Early experiments
suggested that chemosynthetic primary production constituted
the principal source of organic matter for biological communities
associated with vents, although subsequent studies have revealed
many complexities associated with interactions between microbes
and higher organisms inhabiting these ecosystems. A diversity
of host-microbial symbiont relationships has been identified
and experimental studies have revealed the exquisite physiological
adaptations within the giant tubeworm, Riftia pachyptila,
for the uptake, fixation, and assimilation of carbon. In vitro
experiments demonstrated the unusual sulfide binding properties
of tubeworm hemoglobin that prevent inhibition of the cytochrome-c
oxidase enzyme system during transport of sulfide to symbiont-bearing
tissues. Studies of respiration and growth of several species
of vent organisms conducted over the past two decades transformed
earlier views that low metabolism and slow growth are characteristics
of all organisms inhabiting all deep-sea environments. Results
of recent experiments suggest that metabolic rates correlate
with the degree of mobility of the organisms rather than with
any specific attribute of the deep- sea environment itself,
and growth rates of certain vent organisms (e.g. R. pachyptila)
were found to be among the highest in any marine environments.
While extreme thermal tolerance has been suggested as characteristic
of certain vent fauna (e.g. alvinellid polychaetes and alvinocarid
shrimp), the majority of vent metazoans live at temperatures
below 20 degree C and additional experiments are necessary
to reconcile field experiments documenting thermal tolerance
in situ, thermal tolerance in vivo, and thermal sensitivity
of biochemical constituents of vent organisms. Transplantation
and clearance experiments, as well as in situ characterization
of vent fluid chemistry, have greatly increased our understanding
of organism-environment interactions. Early analyses of metazoan
egg size and larval morphology, coupled with in vivo larval
culture experiments, available physical oceanographic data,
and genetic studies of gene flow, have contributed greatly
to our understanding of mechanisms of dispersal between widely
separated vent sites. The documentation of invertebrate colonization
and succession of new vents following a volcanic eruption,
and a series of manipulative field experiments, provide considerable
insights into the relative roles of abiotic conditions and
biotic interactions in structuring vent communities. Recent
and emerging technological developments, such as in situ chemical
analyzers, observatory approaches, and laboratory-based pressure
culture systems, should provide invaluable new experimental
tools for tackling many remaining questions concerning the
ecology of deep-sea hydrothermal systems.
- Feeding behaviour: Hydrothermal vent
crabs feast on sea 'snow'
M. -S Jeng, N. K. Ng and P. K. L. Ng.
Nature, Vol. 432, No. 7020, 23 Dec 2004, pp. 969.
The crab Xenograpsus testudinatus lives at enormously high
densities around the sulphur-rich hydrothermal vents found
in shallow waters off Taiwan, even though this acidic environment
is low in nutrients. Here we show that these crabs swarm out
of their crevices at slack water and feed on the vast numbers
of zooplankton that are killed by the vents' sulphurous plumes,
and that rain down like marine 'snow'. This opportunistic
feeding behaviour explains how the crabs are able to survive
in the adverse toxic environment of these shallow hydrothermal
vents.
- Novel Forms of Structural Integration
between Microbes and a Hydrothermal Vent Gastropod from the
Indian Ocean
S. K. Goffredi, A. Waren, V. J. Orphan, C. L. Van Dover and
R. C. Vrijenhoek.
Applied and Environmental Microbiology, Vol. 70, No. 5, May
2004, pp. 3082-3090.
Here we describe novel forms of structural integration between
endo- and episymbiotic microbes and an unusual new species
of snail from hydrothermal vents in the Indian Ocean. The
snail houses a dense population of gamma -proteobacteria within
the cells of its greatly enlarged esophageal gland. This tissue
setting differs from that of all other vent mollusks, which
harbor sulfur-oxidizing endosymbionts in their gills. The
significantly reduced digestive tract, the isotopic signatures
of the snail tissues, and the presence of internal bacteria
suggest a dependence on chemoautotrophy for nutrition. Most
notably, this snail is unique in having a dense coat of mineralized
scales covering the sides of its foot, a feature seen in no
other living metazoan. The scales are coated with iron sulfides
(pyrite and greigite) and heavily colonized by epsilon - and
delta -proteobacteria, likely participating in mineralization
of the sclerites. This novel metazoan-microbial collaboration
illustrates the great potential of organismal adaptation in
chemically and physically challenging deep-sea environments.
- Phylogenetic diversity of sulfate-reducing
prokaryotes in active deep-sea hydrothermal vent chimney structures
T. Nakagawa, S. Nakagawa, F. Inagaki, K. Takai and K. Horikoshi.
FEMS microbiology letters, Vol. 232, No. 2, 19 Mar 2004,
pp. 145-152.
The phylogenetic diversity of sulfate-reducing prokaryotes
occurring in active deep-sea hydrothermal vent chimney structures
was characterized based on the deduced amino acid sequence
analysis of the polymerase chain reaction-amplified dissimilatory
sulfite reductase (DSR) gene. The DSR genes were successfully
amplified from microbial assemblages of the chimney structures,
derived from three geographically and geologically distinct
deep-sea hydrothermal systems in the Central Indian Ridge
(CIR), in the Izu-Bonin Arc (IBA), and the Okinawa Trough
(OT), respectively. Phylogenetic analysis revealed seven major
phylogenetic groups. More than half of the clones from the
CIR chimney structure were related to DSR amino acid sequences
of the hyperthermophilic archaeal members of the genus Archaeoglobus,
and those of environmental DSR clones within the class Thermodesulfobacteria.
From the OT chimney structure, a different group was obtained,
which comprised a novel, deep lineage associated with the
DSRs of the thermophilic sulfate-reducing bacterium Thermodesulfovibrio.
Most of the DSR clones from the IBA chimney structure were
phylogenetically associated with the delta -proteobacterial
sulfate-reducing bacteria represented by the genus Desulfobulbus.
Sequence analysis of DSR clones demonstrated a diverse sulfate-reducing
prokaryotic community in the active deep-sea hydrothermal
chimney structures.
- Physicochemical characterization of
the microhabitat of the epibionts associated with Alvinella
pompejana, a hydrothermal vent annelid
Carol A. Di Meo-Savoie, George W. Luther III and S. Craig
Cary.
Geochimica et Cosmochimica Acta, Vol. 68, No. 9, May 2004,
pp. 2055-2066.
Alvinella pompejana is a polychaetous annelid that inhabits
narrow tubes along the walls of high-temperature hydrothermal
vent chimneys. The worm hosts a rich community of epibiotic
bacteria that coats its dorsal surface. Although the worm
tube microhabitat is a challenging environment to sample,
characterizing the thermal and geochemical regime is important
for understanding the ecology of the worm and its bacteria,
as the worm spends most of its time inside the tube. We characterized
the physicochemical conditions of diffuse hydrothermal flow
inside inhabited worm tubes using in situ analysis and wet
chemical analysis of discrete water samples. Thermistor probes
deployed inside worm tubes measured temperatures ranging from
28.6 degrees C to 84.0 degrees C, while temperatures at tube
orifices ranged from 7.5 degrees C to 40.0 degrees C. In situ
electrochemical analysis of tube fluids revealed undetectable
oxygen (<5 mu M) and surprisingly low levels of free H (sub
2) S (<0.2 mu M), with most of the sulfide existing as aqueous
FeS molecular clusters. Acid-volatile sulfide measured on
discrete samples of tube fluids ranged from 62.9 to 359.3
mu M, while free sulfide (H (sub 2) S) ranged from undetectable
(<0.2 mu M) to 46.5 mu M. The pH ranged from 5.33 to 6.40,
and sulfate ranged from 22.5 mM to 27.5 mM. Nitrate ranged
from 13.9 to 20.0 mu M, whereas ammonium ranged from 2.5 to
9.7 mu M. Total Fe ranged from 72.1 to 730.2 mu M. Mn, Zn,
Ni, V, P, and Cu were present in micromolar amounts; Pb, Cd,
Co, and Ag were present in nanomolar levels. The worm tube
fluids contained between 72% to 91% of Mg concentrations typically
found in deep seawater. Plots of Mg concentrations vs. other
fluid components showed that the tube fluid is geochemically
altered from theoretical mixing values. Values of SO (sub
4) (super 2-) were enriched inside the worm tube fluids, whereas
NO (sub 3) (super -) , Sr, Mn, Fe, Zn, and acid-volatile sulfide
were depleted. The geochemistry of the tube microhabitat likely
influences the structure of resident microbial communities.
- Spatial and temporal patterns in larval
supply at hydrothermal vents in the northeast Pacific Ocean
A. Metaxas.
Limnology and Oceanography, Vol. 49, No. 6, Nov 2004, pp.
1949-1956.
Larval supply in marine benthic invertebrates influences
patterns of settlement and recruitment to adult populations.
To successfully colonize newly formed or distant hydrothermal
vents, which are discontinuous and ephemeral habitats, larvae
of vent invertebrates must possess long-range dispersal abilities.
However, a significant proportion of locally produced larvae
must remain within the adult habitat to maintain the observed
dense assemblages of invertebrates. I compared spatial and
temporal patterns in larval availability (flux and abundance)
at hydrothermal vents within and between ridge segments in
the northeast Pacific ocean at Axial Seamount and Endeavour
Segment on the Juan de Fuca Ridge and at Magic Mountain on
Explorer Ridge. Near-bottom (within 50 cm) larval supply (individuals
m super(-2) d super(-1)) at five vents on Axial Seamount,
measured with passively collecting traps in 2000 and 2001,
varied temporally by an order of magnitude, but not spatially
at scales of tens of meters to kilometers. The most abundant
larval taxa were gastropods (particularly the limpet Lepetodrilus
fucensis) and polychaetes. Larval abundance in the water column
within the axial valley was measured with net tows by the
remotely operated vehicle ROPOS at the three ridge segments:
Axial Seamount (2000, 2001); Endeavour Segment (2001, 2002);
Magic Mountain (2002). Abundance was greater by an order of
magnitude at the Endeavour Segment than at Axial Seamount
and Magic Mountain, and it did not differ between on-vent
(within tens of meters) and off-vent (up to 5 km from the
venting source) locations within the axial valley. The uniformly
high abundance of larvae within axial valleys suggests that
larval supply within a ridge segment is most likely localized,
implying significant larval retention on the scale of vent
fields and possibly ridge segments.
- Thermal selection of PGM allozymes
in newly founded populations of the thermotolerant vent polychaete
Alvinella pompejana
Patrice Piccino, Frederique Viard, Pierre-Marie Sarradin,
Nadine Le Bris, Dominique Le Guen and Didier Jollivet.
Proceedings of the Royal Society of London, Series B: Biological
Sciences, Vol. 271, No. 1555, 22 Nov 2004, pp. 2351-2359.
Alvinella pompejana lives on the top of chimneys at deep-sea
hydrothermal vents of the East Pacific Rise. It is thought
to be one of the most thermotolerant and eurythermal metazoans.
Our experimental approach combines methods of population genetics
and biochemistry, considering temperature as a potential selective
factor. Phosphoglucomutase (Pgm-1 locus) is one of the most
polymorphic loci of A. pompejana and exhibits four alleles,
from which alleles 90 and 100 dominate with frequencies of
approximately 0.5 in populations. Results from previous studies
suggested that allele 90 might be more thermostable than allele
100. Significant genetic differentiation was found by comparing
contrasted microhabitats, especially the young, still hot,
versus older and colder chimneys, with allele 90 being at
highest frequency on young chimneys. Moreover the frequency
of allele 90 was positively correlated with mean temperature
at the opening of Alvinella tubes. In parallel, thermostability
and thermal optimum experiments demonstrated that allele 90
is more thermostable and more active at higher temperatures
than allele 100. This dataset supports an additive model of
diversifying selection in which allele 90 is favored on young
hot chimneys but counterbalanced over the whole metapopulation
by the dynamics of the vent ecosystem.
- Chemosynthesis-based ecosystems in
the deep Atlantic - what we do know and we don't
D. Desbruyeres and Commission Internationale pour l'Exploration
de la Mer Mediterranee - CIESM, Monaco.
Monaco (Monaco): CIESM, 2003
The discovery of hydrothermal vents, followed by that of
cold seeps, has been one of the great interdisciplinary successes
of oceanography in at least the last 25 years. One can hardly
imagine a greater contrast to the typical soft-sediment deep
sea than the exuberant oases of life at hydrothennal vents
and cold seeps areas. With the steep gradient of chemistry
and the high biological primary production, the ecology of
vent invertebrates is probably more closely similar to that
of intertidal communities than to the "regular" deep sea ones.
These oases of life are small-sized and discontinuous in space
and cannot be observed and studied without the use of deep-sea
submersibles. The direct linkage between fluid emission and
bacterial chemo-autotrophic production makes them also discontinuous
in time at different scales.
- Constrained circulation at Endeavour
ridge facilitates colonization by vent larvae
R. E. Thomson, S. F. Mihaly, A. B. Rabinovich, R. E. Mcduff,
S. R. Veirs and F. R. Stahr.
Nature, Vol. 424, No. 6948, 31 Jul 2003, pp. 545-549.
Understanding how larvae from extant hydrothermal vent fields
colonize neighbouring regions of the mid-ocean ridge system
remains a major challenge in oceanic research. Among the factors
considered important in the recruitment of deep-sea larvae
are metabolic lifespan, the connectivity of the seafloor topography,
and the characteristics of the currents. Here we use current
velocity measurements from Endeavour ridge to examine the
role of topographically constrained circulation on larval
transport along-ridge. We show that the dominant tidal and
wind-generated currents in the region are strongly attenuated
within the rift valley that splits the ridge crest, and that
hydrothermal plumes rising from vent fields in the valley
drive a steady near- bottom inflow within the valley. Extrapolation
of these findings suggests that the suppression of oscillatory
currents within rift valleys of mid-ocean ridges shields larvae
from cross-axis dispersal into the inhospitable deep ocean.
This effect, augmented by plume-driven circulation within
rift valleys having active hydrothermal venting, helps retain
larvae near their source. Larvae are then exported preferentially
down-ridge during regional flow events that intermittently
over-ride the currents within the valley.
- Current challenges in the study of
biological communities at deep-sea hydrothermal vents
A. Metaxas and Commission Internationale pour l'Exploration
de la Mer Mediterranee - CIESM, Monaco.
Monaco (Monaco): CIESM, 2003
The discovery of hydrothermal vents in 1977 impacted profoundly
our preconceptions of structure and function of marine biological
communities. These extreme habitats harbour communities of
exceedingly high biomass but relatively low diversity, consisting
mostly of endemic, newly discovered species. The luxuriant
communities at vents are sustained by chemoautotrophic production
by microbes, using CO sub(2) as substrate and energy derived
from the oxidation of hydrogen sulfide (H sub(2)S). Over the
last 25 years, research on the geological, physical, chemical
and biological properties of hydrothermal vents has expanded
significantly our understanding of these systems. Presently
we are faced with our next challenge: to move forward in our
understanding, we must adopt a combination of traditional
approaches, which have been used in other systems for several
decades, and innovative technologies, which currently are
under development.
- Deep Submergence Synergy: Alvin and
ABE Explore the Galapagos Rift at 86 degree W
T. Shank, D. Fornari and D. Yoerger, et al.
EOS, Transactions, American Geophysical Union, Vol. 84, No.
41, 14 Oct 2003, pp. 430-432.
For over 25 years, hydrothermal vent communities discovered
at the Galapagos Rift near 86 degree W have provided the foundation
of deep-sea vent biology as their study has led to fundamental
discoveries of chemoautorophy and novel symbioses in the deep
sea. Since 1979, numerous physiological and geochemical investigations
of the Rose Garden vent community have been made possible
through routine access to this deep sea floor site, provided
by the deep submergence vehicle Alvin. This research revolutionized
our understanding of basic biological and chemical processes
in the deep ocean.
- Dispersal at hydrothermal vents: a
summary of recent progress
P. A. Tyler and C. M. Young.
Hydrobiologia, Vol. 503, No. 1-3, Aug 2003, pp. 9-19.
The discovery of hydrothermal vents along the Galapagos Rift
in 1977 opened up one of the most dynamic and productive research
themes in marine biology. In the intervening 25 years, hydrothermal
vent faunas have been described from the eastern, northeastern
and western Pacific, the mid-Atlantic Ridge and the Indian
Ocean in the region of the Rodriguez Triple Junction. In addition,
there is evidence of hydrothermal signals from the Gakkel
Ridge in the Arctic, the central and southwest Indian Ridges
and the Scotia Arc in Antarctica. Although often perceived
as a continuous linear structure, there are many discontinuities
that have given rise to separate biogeographic provinces.
In addition, the intervening 25 years have seen a massive
increase in our understanding of the biological processes
at hydrothermal vents. However, how vents are maintained,
and how new vents are colonised has been relatively poorly
understood until recently. This review addresses the known
larval development of vent-endemic invertebrates. The distribution
of larvae in relation to the hydrothermal plume, and the ocean
ridge in general, are discussed and the experimental evidence
of larval longevity and transport are discussed using such
variables as gene flow and larval development rates. The concept
of larval dispersal along the mid-ocean ridge is discussed
in relation to dispersal barriers and relates the known biogeography
of hydrothermal vent systems to both local and evolutionary
processes.
- Distributional and biological aspects
of copepods at hydrothermal vents on the Juan de Fuca Ridge,
north-east Pacific ocean
Maia Tsurumi, R. C. De Graaf and V. Tunnicliffe.
Journal of the Marine Biological Association of the United
Kingdom, Vol. 83, No. 3, 2003, pp. 469-477.
The abundance patterns of copepods on the Juan de Fuca Ridge
was examined. One species was studied in detail. Twelve non-parasitic
species are recorded from the Juan de Fuca, but only three
dirivultid species and some unidentified harpacticoids are
abundant in collections. Densities are estimated at 0.5 copepod
cm super(-2) on vestimentiferan tubes to over 8 cm super(-2)
on chimney surfaces. Aphotopontius forcipatus is most abundant
at new vents and Benthoxynus spiculifer is most abundant at
mature vents. Vents with reduced or undetectable fluid flow
have higher diversity of copepod fauna. The life cycle of
the siphonostome Stygiopontius quadrispinosus begins with
a centrolecithal egg brooded singly or doubly on the female.
Hatching and naupliar stages are unknown in benthic samples.
The pre-adult stage (copepodite V) recruits to the vent habitat.
Pre-adult males attach to pre-adult females and fertilize
at the final copepodite VI moult. As the sex ratio is highly
skewed in favour of females, males probably inseminate many
females and there may be mate competition in populations where
males are rare. Reproduction is probably continuous or semi-continuous.
Abundance is greatest on sulphide edifices near the points
of hot water egress. This copepod co-occurs with the alvinellid
polychaete Paralvinella sulfincola.
- Diversity at hydrothermal vents
M. Tsurumi.
Global Ecology and Biogeography, Vol. 12, No. 3, May 2003,
pp. 181-190.
To describe patterns of hydrothermal vent community diversity
and dispersion at the intersegment scale (> 100 km). The area
discussed is an approximately 170 km portion of the Juan de
Fuca Ridge, a mid-ocean ridge in the north-east Pacific Ocean.
Samples of benthic invertebrates from hydrothermal vents on
three segments of the Juan de Fuca are examined for community
characteristics such as diversity, abundance and distribution.
Species richness (55 species) and evenness are low. If the
macrofauna only are considered, species richness is about
30% lower than when meiofauna are also considered. The geometric
series describes the species-abundance distribution. The relationship
between vent species' distribution and abundance is significantly
positive (r super(2) = 0.818; P < 0.001). Alpha diversity
is lower in patchy habitat than continuous habitat and gamma
diversity is similar for both habitat types. Beta diversity
is higher in patchy habitat. Local diversity is linearly related
to regional diversity. Species richness is comparable to other
highly disturbed systems. The geometric series species abundance
model implies some degree of niche pre-emption in the vent
community and is consistent with the suggestion that the geometric
series distribution can be found in species-poor environments
that experience harsh conditions and are structured by relatively
few environmental factors. Species distribution and abundance
are highly correlated. The regional species pool affects local
vent diversity. Vent diversity studies should be conducted
on at least the ridge scale.
- Evidence of Chemolithoautotrophy in
the Bacterial Community Associated with Alvinella pompejana,
a Hydrothermal Vent Polychaete
B. J. Campbell, J. L. Stein and S. C. Cary.
Applied and Environmental Microbiology, Vol. 69, No. 9, Sep
2003, pp. 5070-5078.
The deep-sea polychaete Alvinella pompejana colonizes tubes
on the sides of black smoker chimneys along the East Pacific
Rise. A diverse, yet phylogenetically constrained episymbiotic
community is obligately associated with its dorsal surface.
The morphologically and phylogenetically distinct dominant
episymbionts have not yet been cultured, and there are no
clearly defined roles for these bacteria in this symbiosis.
A large insert fosmid library was screened for the presence
of the two dominant phylotypes. Two fosmids, 35.2 and 38 kb,
containing phylotype-specific 16S ribosmal DNA sequences were
fully sequenced. Each fosmid had a gene encoding ATP citrate
lyase, a key enzyme in the reverse tricarboxylic acid (rTCA)
cycle, a CO sub(2) fixation pathway. A selection of episymbiont
communities from various geographic locations and vent sites
were screened for the presence, diversity, and expression
(via reverse transcription-PCR) of the ATP citrate lyase gene.
Our results indicate that the ATP citrate lyase gene is not
only a consistent presence in these episymbiont communities
but is also expressed. Phylogenetically distinct forms of
ATP citrate lyase were also found associated with and expressed
by bacteria extracted from the tubes of A. pompejana. Utilizing
PCR with degenerate primers based on a second key enzyme in
the rTCA cycle, 2-oxoglutarate:acceptor oxidoreductase, we
also demonstrated the persistent presence and expression of
this gene in the episymbiont community. Our results suggest
that members of both the episymbiont and the surrounding free-living
communities display a chemolithoautotrophic form of growth
and therefore contribute fixed carbon to other organisms in
the vent community.
- Food resource partitioning and competition
among alvinellid polychaetes of Juan de Fuca Ridge hydrothermal
vents
C. Levesque, S. K. Juniper and J. Marcus.
Marine Ecology Progress Series, 2003, pp. Vol. 246, pp. 173-182
Biotic communities in highly variable, frequently perturbed
habitats are usually expected to be structured mainly by abiotic
factors. In the highly variable deep-sea hydrothermal vent
environment, physical and chemical factors are known to play
an important role in limiting the distribution and abundance
of species, but the importance of biotic interactions remains
largely unresolved. The high density and biomass attained
by the vent macrofauna suggest that resource partitioning
and competition may be significant in these communities. This
study of food resource utilisation at northeast Pacific deep-sea
hydrothermal vents uses an approach based on stable carbon
and nitrogen isotope analyses to characterise trophic interactions
between the 3 dominant alvinellid polychaetes, Paralvinella
palmiformis, P. sulfincola, and P. pandorae. We also examine
size structure in sympatric and allopatric populations of
P. palmiformis and P. pandorae. Results indicate that food
resource partitioning occurs both intra- and interspecifically
in P. palmiformis and P. sulfincola, and we advocate that
this process contributes to explaining their co-existence
at very high densities. In contrast, P. pandorae has a much
more restricted trophic niche, overlaps P. palmiformis in
diet and is much smaller in size when found in sympatry with
P. palmiformis. P. pandorae is the most likely of the 3 species
to be affected by intra- and/or interspecific competition
for food, and this may explain the drastic change in the population
size structure observed between successive years. Our work
indicates that within the limits imposed by environmental
conditions, biotic interactions such as food resource partitioning
and competition can be significant factors structuring deep-sea
hydrothermal vent communities.
- Functional morphology and feeding strategy
of the hydrothermal-vent polychaete Archinome rosacea (family
Archinomidae)
M. E. Ward, C. D. Jenkins and C. L. M. Dover.
Canadian Journal of Zoology/Revue Canadienne de Zoologie,
Vol. 81, No. 4, 2003, pp. 582-590.
Polychaetes are a large component of deep-sea hydrothermal-vent
invertebrate faunas, yet little is known about the trophic
ecology of most species. In this study, the feeding mode of
a common and widespread vent polychaete, Archinome rosacea
(order Amphinomida), is investigated through morphological
studies and gut-content analyses. The foregut of this polychaete
is similar in structure to that of other shallow-water Amphinomida.
Prey capture is facilitated by a ventral muscular eversible
proboscis with two glandular lateral-lip structures. Extracellular
digestion is suggested by packets of digestive enzymes observed
to be discharged from the apical ends of the midgut epithelial
cells into the lumen. Midgut tissues also contain inclusions
resembling spherocrystals, possibly sites where toxic materials
are sequestered. Gut-content analyses indicate a preference
for mobile prey, including polychaetes and crustaceans. We
propose that predation and scavenging are important trophic
strategies for A. rosacea.Original Abstract: Les polychetes
sont une composante importante de la faune des cheminees hydrothermales
dans la zone profonde des oceans et pourtant, on connait peu
de choses au sujet de l'ecologie trophique de la plupart des
especes. Nous etudions ici le mode d'alimentation d'un polychete
commun dans les cheminees hydrothermales, Archinome rosacea
(ordre Amphinomida), par examen morphologique et par analyse
des contenus stomacaux. La partie anterieure du tube digestif
est de structure semblable a celle d'autres Amphinomida d'eaux
peu profondes. La capture des proies est assuree par le proboscis
ventral, musculeux, reversible, qui porte deux structures
laterales glandulaires en forme de levres. La digestion semble
extracellulaire, vu la presence de paquets d'enzymes digestives
liberes dans la lumiere aux extremites apicales des cellules
epitheliales de la partie mediane du tube digestif. Les tissus
de cette partie du tube contiennent aussi des inclusions ressemblant
a des cristaux spheriques qui servent peut-etre a l'entreposage
de matieres toxiques. Les analyses de contenus stomacaux ont
revele une preference pour les proies mobiles, notamment des
polychetes et des crustaces. Il semble qu'A. rosacea soit
a la fois predatrice et detritivore, deux strategies alimentaires
importantes pour cette espece.
- Habitat, growth and physiological ecology
of a basaltic community of Ridgeia piscesae from the Juan de
Fuca Ridge
I. A. Urcuyo, G. J. Massoth, D. Julian and C. R. Fisher.
Deep-Sea Research (Part I, Oceanographic Research Papers),
Vol. 50, No. 6, Jun 2003, pp. 763-780.
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 super(-1)). Although the incidence of plume
damage from partial predation was very high (>95%), mortality
was very low (<4% yr super(-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 mu M), while their
posterior sections were consistently exposed to sulfide concentrations
in the 100 mu M 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.
- Incidence and Diversity of Microorganisms
within the Walls of an Active Deep-Sea Sulfide Chimney
M. O. Schrenk, D. S. Kelley, J. R. Delaney and J. A. Baross.
Applied and Environmental Microbiology, Vol. 69, No. 6, Jun
2003, pp. 3580-3592.
A large, intact sulfide chimney, designated Finn, was recovered
from the Mothra Vent Field on the Juan de Fuca Ridge in 1998.
Finn was venting 302 degree C fluids on the seafloor and contained
complex mineralogical zones surrounding a large open central
conduit. Examination of microorganisms within these zones,
followed by community analysis with oligonucleotide probes,
showed that there were variations in the abundance and diversity
of eubacteria and archaea from the exterior to the interior
of the chimney. The microbial abundance based upon epifluorescence
microscopy and quantitative fatty acid analyses varied from
>10 cells/g of sulfide 2 to 10 cm within the chimney wall
to <10 cells/g in interior zones. Direct microscopic observation
indicated that microorganisms were attached to mineral surfaces
throughout the structure. Whole-cell hybridization results
revealed that there was a transition from a mixed community
of eubacteria and archaea near the cool exterior of the chimney
to primarily archaea near the warm interior. Archaeal diversity
was examined in three zones of Finn by cloning and sequencing
of the 16S rRNA gene. The majority of sequences from the exterior
of the chimney were related to marine group I of the Crenarchaeota
and uncultured Euryarchaeota from benthic marine environments.
In contrast, clone libraries from interior regions of the
chimney contained sequences closely related to methanogens,
Thermococcales, and Archaeoglobales, in addition to uncultured
crenarchaeal phylotypes obtained from deep subsurface sites.
These observations of microbial communities within an active
hydrothermal chimney provide insight into the microbial ecology
within such structures and may facilitate follow-up exploration
into expanding the known upper temperature limits of life.
- Striking succession of mussels at newly
formed deep-sea hydrothermal vents
R. A. Lutz, T. M. Shank and D. J. Fornari.
Journal of Shellfish Research, Vol. 22, No. 1, Jun 2003,
pp. 341-342.
In April, 1991 a volcanic eruption obliterated existing biological
communities within extensive regions along the crest of the
East Pacific Rise between 9 degree 45'N and 9 degree 52'N
(depth 2520 m). The initiation of hydrothermal venting during
the eruptive process afforded the unique opportunity to follow,
over a 10+ year period, temporal changes in biological community
structure from the "birth" of numerous hydrothermal vents.
Vestimentiferan tubeworms, amphipods, copepods, octopods,
and galatheid and brachyruan crabs gradually colonized the
vents during the first 2 years. Noticeably absent during this
time was any evidence of the mussel Bathymodiolus thermophilus.
Small mussels (5 cm were common in the region and had begun
to colonize the tubes of Riftia pachyptila, concomitant with
declining concentrations of H2S in the venting diffuse flow
fluids. Over the next 5-6 years, the abundance of mussels
increased dramatically until most of the existing communities
previously dominated (in biomass) by tubeworms were now dominated
by extensive populations of mussels.
- Thermal Tolerances of Deep-Sea Hydrothermal
Vent Animals From the Northeast Pacific
R. W. Lee.
Biological Bulletin, Marine Biological Laboratory, Woods
Hole, Vol. 205, No. 2, 1 Oct 2003, pp. 98-101.
- Tubeworm-associated communities at
hydrothermal vents on the Juan de Fuca Ridge, northeast Pacific
M. Tsurumi and V. Tunnicliffe.
Deep Sea Research (Part I, Oceanographic Research Papers),
Vol. 50, No. 5, May 2003, pp. 611-629.
Hydrothermal vent communities on a mid-ocean ridge crest
can be separated by large distances on separate segments.
Heat sources, vent character, fluid chemistry and current
patterns may differ markedly. This study examines whether
vent community characteristics on three of the four southern
segments of the Juan de Fuca Ridge are significantly different.
Taxonomic composition and relative abundance of the fauna
over 1 mm in size associated with vestimentiferan tubeworm
bushes are examined from fifty-one collections. Among nearly
350, 000 specimens, 37 taxa are recognized, most to species
level. Another 14 taxa are meiofaunal in size classification.
Species richness and selected diversity indicators are highest
on Axial Volcano while animal density within the bushes does
not differ significantly. Cluster analysis does not group
collections by location, year of collection or vent temperature;
collection substratum--basalt or sulphide--may influence clustering.
The architecture of the tubes of tubeworm bushes appears to
affect the numbers of species present and the resultant clusters.
The tightly interwoven, knotted Ridgeia piscesae tubes found
on Axial host twice as many species as tubeworm bushes with
a less complex structure. Four species dominate most of the
collections: two gastropods (Lepetodrilus fucensis and Depressigyra
globulus) and two polychaetes (Paralvinella pandorae and Amphisamytha
galapagensis). Other vent species are low in abundance (<1%
relative abundance) and patchy in distribution. Four collections
with no visible flow had markedly different assemblages representing
a transition state from vent assemblages to normal deep-sea
fauna. There are differences in community structure among
the segments, but the causes for these differences are unclear.
Higher diversity on Axial Volcano may be supported by a greater
time of sustained venting, a larger venting area, water circulation
contained within the caldera, or flow conditions that sponsor
growth of more complex habitat.
- Variation in community structure within
hydrothermal vent mussel beds of the East Pacific Rise
C. L. Van Dover.
Marine Ecology Progress Series, 2003, pp. Vol. 253, pp. 55-66
Patterns in invertebrate community structure associated with
mussel beds at 3 hydrothermal vents on the northern East Pacific
Rise (NEPR) were explored using quantitative, replicate sampling
methods and were compared to those of southern East Pacific
Rise (SEPR) mussel beds (~63000 km apart). Univariate measures
of diversity (H super('), J super(')) did not differ among
3 NEPR mussel beds. Diversity by most estimates was lower
at NEPR mussel beds than at SEPR mussel beds. Invertebrate
faunas of NEPR and SEPR mussel beds belong to the same biogeographic
province, and the numerically dominant species at NEPR mussel
beds were also numerical dominants at SEPR mussel beds. Patterns
of community structure within and among NEPR mussel beds,
between NEPR and SEPR mussel beds, and between 'young' 8 yr)
mussel beds could be differentiated using multivariate techniques
based on species-abundance matrices. Overall, these observations
suggest that NEPR and SEPR mussel-bed communities are remarkably
similar, differing primarily in the relative abundances of
their shared, numerically dominant species and in the composition
of the rare species.
- Colonization by pioneer populations
of epsilon -proteobacteria and community succession at mid-ocean
ridge hydrothermal vents as determined by T-RFLP analysis; AGU
2002 fall meeting
Craig L. Moyer and Jeffrey J. Engebretson.
EOS, Transactions, American Geophysical Union, Vol. 83, No.
47, Suppl, 19 Nov 2002, pp. 1472.
- Diversity of anaerobic heterotrophic
thermophiles isolated from deep-sea hydrothermal vents of the
Mid-Atlantic Ridge
N. Wery, M. A. Cambon-Bonavita, F. Lesongeur and G. Barbier.
FEMS microbiology ecology, Vol. 41, No. 2, Aug 2002, pp.
105-114.
During the 'MARVEL' oceanographical cruise performed in September
1997, samples were collected from the deep-sea vents of hydrothermal
sites on the Mid-Atlantic Ridge. Eighty-four thermophilic
and hyperthermophilic heterotrophic microorganisms were isolated
using different culture media containing cellobiose, xylan,
starch, lipidic or proteic substrates. These isolates were
obtained in anaerobic conditions, at 65 degree C, 85 degree
C and 95 degree C. Fifty of them were classified using amplified
ribosomal DNA restriction analysis, random amplified polymorphic
DNA and 16S rDNA sequencing. The strains classified have been
assigned to the archaeal order Thermococcales and to the bacterial
orders Thermotogales and Clostridiales. Variations in growth
temperature and carbon sources were efficient enough to generate
taxonomic diversity within enrichment cultures. Presumptive
new genera and new species were isolated. Two isolates were
confirmed as type strains of new species of new genera recently
described: Marinitoga camini and Caloranaerobacter azorensis.
- Ecology of hydrothermal vents on three
segments of the Juan de Fuca Ridge, northeast Pacific
M. Tsurumi.
Dissertation Abstracts International Part B: Science and
Engineering, Vol. 62, No. 10, Apr 2002, pp. 4337.
- Merging Genomes with Geochemistry in
Hydrothermal Ecosystems
A. Reysenbach and E. Shock.
Science (Washington), Vol. 296, No. 5570, 10 May 2002, pp.
1077-1082.
Thermophilic microbial inhabitants of active seafloor and
continental hot springs populate the deepest branches of the
universal phylogenetic tree, making hydrothermal ecosystems
the most ancient continuously inhabited ecosystems on Earth.
Geochemical consequences of hot water-rock interactions render
these environments habitable and supply a diverse array of
energy sources. Clues to the strategies for how life thrives
in these dynamic ecosystems are beginning to be elucidated
through a confluence of biogeochemistry, microbiology, ecology,
molecular biology, and genomics. These efforts have the potential
to reveal how ecosystems originate, the extent of the subsurface
biosphere, and the driving forces of evolution.
- A novel landscape ecology approach
for determining habitat correlations and macrofaunal patchiness
in extreme environments: pilot study for the Southern East Pacific
Rise at 17-18oS
AM Wargo Rub, DJ Wright and JA Jones.
Roscoff (France): Station Biologique de Roscoff, 2002, 307-311.
[Cah. Biol. Mar.]
- Nutritional relations of deep-sea hydrothermal
fields at the Mid-Atlantic Ridge: a stable isotope approach
A. Colaco, F. Dehairs and D. Desbruyeres.
Deep-Sea Research (Part I, Oceanographic Research Papers),
Vol. 49, No. 2, Feb 2002, pp. 395-412.
Nutritional relations among invertebrates from the hydrothermal
vent fields at the Mid Atlantic Ridge (MAR) were studied via
the carbon and nitrogen stable isotope approach. A large number
of specimens of different vent species from different MAR
vent fields were analysed, providing a general picture of
the community structure. The isotopic composition at each
vent field presents the same general trend. There is an obvious
dichotomy of the trophic structure, with the mussels being
significantly depleted in super(13)C and shrimps being significantly
enriched in super(13)C. MAR and Pacific vent fields present
the same picture, despite a different species composition.
Primary consumers are divided into main groups according to
their delta super(13)C signature: > -15 (shrimps) and < -20ppt
(mussels). Vent predators are tightly linked to one or the
other group, but a mixed diet cannot be excluded. Bathyal
species are top predators, making incursions into the vent
fields to profit from the large biomass. Taking into account
the above associations, a descriptive trophic model was elaborated.
At the base of the food chain the chemolithotrophic bacteria
predominate. Four trophic levels were then distinguished:
primary consumers, feeding only on bacteria; mixotrophs feeding
on bacteria and small invertebrates; vent predators feeding
only on small invertebrates; and finally top predators that
are mainly constituted by deep-sea fauna.
- Phylogenetic characterization and biomass
estimation of bacterial endosymbionts associated with invertebrates
dwelling in chemosynthetic communities of hydrothermal vent
and cold seep fields
H. Yamamoto, K. Fujikura, A. Hiraishi, K. Kato and Y. Maki.
Marine Ecology Progress Series, 2002, Vol. 245, pp. 61-67
Molecular phylogenetics for endosymbiotic bacteria recovered
from vestimentiferan tubeworm Lamellibrachia satsuma from
Kagoshima Bay, vesicomyid clam Calyptogena laubieri from the
Nankai Trough and mytilid mussel Bathymodiolus from the Mariana
Back-arc Basin, were examined by PCR-aided 16S rDNA cloning
and sequencing, and quinone profiling. The 16SrRNA clones
of the endosymbionts from the 3 organisms fall within gamma
-Proteobacteria and showed distinct lines of descent specific
to their respective host. The 16S rRNA gene phylogeny confirms
the host endosymbiont specificity in the coevolutionary process.
Ubiquinones with 9 isoprene units (Q-9) or Q-10 were found
as the major quinones in all test tissues of the host. Larger
amounts of Q-8 were detectable only in those host body parts
(gill or trophosome) harboring the endosymbiotic bacteria.
These observations suggest that Q-8 is the major quinone of
the endosymbionts. Based on the bacterial quinone concentration,
the population densities of the bacteria present were estimated
to be 10 super(10) to 10 super(11) cells/g wet wt of host
tissue. The quantitative determination of quinones may provide
information about physiological activity of the chemosynthetic
communities as well as their biomass production.
- Predation structures communities at
deep-sea hydrothermal vents
F. Micheli, C. H. Peterson and G. A. Johnson, et al.
Ecological Monographs, Vol. 72, No. 3, Aug 2002, pp. 365-382.
The structure and dynamics of natural communities result
from the interplay of abiotic and biotic factors. We used
manipulative field experiments to determine the relative roles
of abiotic conditions and biotic interactions in structuring
deep-sea (2500 m depth) communities along environmental gradients
around hydrothermal vents of the eastern tropical Pacific
Ocean (East Pacific Rise, at 9 degree 50 degree N). We tested
(1) whether predation by crabs and fishes affects the recruitment
of benthic species and subsequent community structure and
(2) whether the effects of predation vary along the steep
gradients of temperature, oxygen, sulfide, and metal concentrations
near vents. Recruitment substrates (basalt cubic blocks, roughly
10 cm on a side), both uncaged and caged to exclude predators
(crabs, fishes, whelks, and octopi), were deployed along a
decreasing vent fluid-flux gradient. The exclusion of predators
for 8 mo increased the abundance of small mobile gastropods
and amphipod crustaceans but decreased the abundance of sessile
invertebrates, including juvenile vestimentiferan worms, tubiculous
polychaetes, and mussels. Effects of predation were strongest
nearest to hydrothermal vents, where abiotic environmental
conditions were most extreme but productivity and the overall
abundances of benthic invertebrates and mobile predators were
the greatest. Additional 5-mo experiments conducted at three
different locations showed similar trends at all sites, indicating
that these effects of predation on benthic community structure
are repeatable. Stomach-content analyses of the most abundant
predators found at vents indicated that the zoarcid fish (Thermarces
cerberus) primarily feeds on the vent snail Cyathermia naticoides,
the limpet Lepetodrilus elevatus, and the amphipod crustacean
Ventiella sulfuris, the very species that showed the greatest
increase following predator exclusion. In contrast, brachyuran
(Bythograea thermydron) and galatheid (Munidopsis subsquamosa)
crab stomachs did not contain small mobile grazers, and crabs
presented with arrays of the most common vent invertebrate
species preferred mussels and vestimentiferans over limpets.
Our results indicate that predation by large mobile predators
influences the structure of hydrothermal vent communities,
directly by reducing the abundance of gastropod prey species,
and indirectly by reducing gastropod grazing and by bulldozing
of recruits of sessile invertebrates.
- Species-specific variation in sulfide
physiology between closely related vesicomyid clams
S. K. Goffredi and J. P. Barry.
Marine Ecology Progress Series, Vol. 225, Jan 2002, pp. 227-238.
Symbioses involving sulfide-oxidizing bacteria and metazoan
phyla dominate invertebrate assemblages at cold seeps and
hydrothermal vents worldwide. The predominant species inhabiting
cold seeps in Monterey Bay are the vesicomyid clams Calyptogena
kilmeri and C. pacifica. The growth and survival of these
clams depend directly upon the productivity of their chemoautotrophic
endosymbionts, which is fueled by the oxidation of sulfide.
For this reason, sulfide availability and sulfide-related
physiology are thought to be the most influential factors
governing the productivity of these associations. Both species
inhabit sulfide-rich sediments and depend nutritionally on
their symbionts, yet many aspects of their life histories
differ considerably. Our results indicate that C. pacifica,
which inhabits areas with lower environmental sulfide levels,
is physiologically poised for the uptake and transport of
sulfide, indicated by increased sulfide consumption rates,
sulfide-binding ability, and internal sulfide levels. C. pacifica
also has a greater potential for symbiont energy turnover,
supported by increased sulfide oxidation potential, enzymes
involved in sulfur metabolism, and bacterial densities. Conversely,
C. kilmeri demonstrates a less effective sulfide uptake mechanism
and, therefore, a specific need for higher environmental sulfide
levels. It appears that the abilities of these 2 species to
process sulfide differ greatly and reflect not only the environments
in which they are found but also the capabilities of their
symbionts. This research represents the first comparative
investigation of the physiological functioning of closely
related species in chemosynthetic symbioses and elucidates
the constraints and advantages posed by different modes of
sulfide (energy) uptake and assimilation in these, and perhaps
other, symbiotic organisms.
- Three dimensional view of the Atlantic
abyssal benthopelagic vent community
AL Vereshchaka and ME Vinogradov.
Roscoff (France): Station Biologique de Roscoff, 2002, pp.
303-305. [Cah. Biol. Mar.]
- Three Ridgeia piscesae assemblages
from a single Juan de Fuca Ridge sulphide edifice: structurally
different and functionally similar
BW Govenar, DC Bergquist, IA Urcuyo, JT Eckner and CR Fisher.
Roscoff (France): Station Biologique de Roscoff, 2002, pp.
247-252. [Cah. Biol. Mar.]
- Trophic relationships among invertebrates
at the Kairei hydrothermal vent field (Central Indian Ridge)
C. L. Van Dover.
Marine Biology, Vol. 141, No. 4, 2002, pp. 761-772.
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 degree 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 delta super(13)C isotopic value of about - 13 ppt,
due to the abundance of primary consumers (shrimp: Rimicaris
aff. exoculata) and secondary consumers (anemones: Marianactis)
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). 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 super(13)C- and super(15)N-enriched primary consumers
in these ecosystems. Electronic supplemen- tary material to
this paper can be obtained by using the Springer LINK server
located at http://dx.doi.org/ 10. 1007/s00227-002-0865-y.
- Understanding the biogeography of chemosynthetic
ecosystems
P. A. Tyler, C. R. German, E. Ramirez-Llodra and C. L. Van
Dover.
Oceanologica Acta, Vol. 25, No. 5, 2002, pp. 227-241.
ChEss is a recently-funded Census of marine life programme
aimed at improving our knowledge of the biogeography of deepwater
chemosynthetically driven ecosystems by promoting an international
field phase of discovery and exploration. The main objectives
are to assess and explain the diversity, distribution and
abundance of hydrothermal vent and cold seep species. With
the global mid-ocean ridge system extending ~65 000 km, it
is unlikely that its entire length would be examined in detail.
The ChEss programme proposes to select a limited number of
target areas chosen for the discovery of new vents and seeps.
The intention is to identify the maximum scientific return
that could be achieved from detailed investigation of the
minimum number of sites at key locations. To narrow the field
for exploration, a number of starting hypotheses and goals
have been identified. A bio- and geo-referenced database for
hydrothermal vent and cold seep species will be created. This
database will be integrated with the Ocean Biogeographic Information
System (OBIS). An international scientific committee will
coordinate the programme, facilitate collaboration between
participants, promote ship-time applications at national level
and stimulate scientific innovation from a wider community.Original
Abstract: ChEss est un des programmes les plus recents du
Recensement de la vie marine (CoML). Il doit accroitre notre
connaissance de la biogeographie des ecosystemes chemosynthetiques
profonds en mettant en place une recherche internationale
de terrain. L'objectif est de definir et d'expliquer la diversite,
la distribution et l'abondance des especes des events hydrothermaux
et des suintements froids. Avec un systeme de ride medio-oceanique
s'etendant sur 65000 kilometres, une etude fine est hors de
propos. Le programme ChEss propose de selectionner un nombre
limite d'aires-cibles choisies pour la decouverte de nouveaux
events ou sources. L'objectif est de tirer un benefice maximum
d'une etude detaillee d'un nombre reduit de sites choisis
dans des positions-clefs. Pour focaliser le champ de recherche,
quelques hypotheses et themes ont ete identifies. Une base
de donnees biologiques et geologiques pour les especes de
ces aires sera creee ; elle sera integree a Obis (Ocean buiogeographic
information system). Un Comite scientifique international
coordonnera le programme, facilitera les cooperations, appuiera
l'attribution de temps-bateau au niveau national et stimulera
l'innovation scientifique pour une communaute elargie.
- The use of video surveys, a geographic
information system and sonar backscatter data to study faunal
community dynamics at Juan de Fuca Ridge hydrothermal vents
S. Durand, M. Le Bel and KS Juniper.
Roscoff (France): Station Biologique de Roscoff, 2002, 235-240.
[Cah. Biol. Mar.]
- Biogeography and ecological setting
of Indian Ocean hydrothermal vents
C. L. Van Dover, S. E. Humphris and D. Fornari, et al.
Science (Washington), Vol. 294, No. 5543, 26 Oct 2001, pp.
818-823.
Within the endemic invertebrate faunas of hydrothermal vents,
five biogeographic provinces are recognized. Invertebrates
at two Indian Ocean vent fields (Kairei and Edmond) belong
to a sixth province, despite ecological settings and invertebrate-bacterial
symbioses similar to those of both western Pacific and Atlantic
vents. Most organisms found at these Indian Ocean vent fields
have evolutionary affinities with western Pacific vent faunas,
but a shrimp that ecologically dominates Indian Ocean vents
closely resembles its Mid-Atlantic counterpart. These findings
contribute to a global assessment of the biogeography of chemosynthetic
faunas and indicate that the Indian Ocean vent community follows
asymmetric assembly rules biased toward Pacific evolutionary
alliances.
- Distribution of Archaea in a Black
Smoker Chimney Structure
K. Takai, T. Komatsu, F. Inagaki and K. Horikoshi.
Applied and Environmental Microbiology, Vol. 67, No. 8, Aug
2001, pp. 3618-3629.
Archaeal community structures in microhabitats in a deep-sea
hydrothermal vent chimney structure were evaluated through
the combined use of culture-independent molecular analyses
and enrichment culture methods. A black smoker chimney was
obtained from the PACMANUS site in the Manus Basin near Papua
New Guinea, and subsamples were obtained from vertical and
horizontal sections. The elemental composition of the chimney
was analyzed in different subsamples by scanning electron
microscopy and energy-dispersive X-ray spectroscopy, indicating
that zinc and sulfur were major components while an increased
amount of elemental oxygen in exterior materials represented
the presence of oxidized materials on the outer surface of
the chimney. Terminal restriction fragment length polymorphism
analysis revealed that a shift in archaeal ribotype structure
occurred in the chimney structure. Through sequencing of ribosomal
DNA (rDNA) clones from archaeal rDNA clone libraries, it was
demonstrated that the archaeal communities in the chimney
structure consisted for the most part of hyperthermophilic
members and extreme halophiles and that the distribution of
such extremophiles in different microhabitats of the chimney
varied. The results of the culture-dependent analysis supported
in part the view that changes in archaeal community structures
in these microhabitats are associated with the geochemical
and physical dynamics in the black smoker chimney.
- First Hydrothermal Vent Communities
from the Indian Ocean Discovered
Jun Hashimoto, Suguru Ohta and Toshitaka Gamo, et al.
Zoological Science, Vol. 18, No. 5, Jul 2001, pp. 717-721.
Thriving chemosynthetic communities were located for the
first time in the Indian Ocean between 2420 and 2450 m, on
a volcanic knoll at the eastern crest of an axial valley,
approximately 22 km north of the Rodriguez Triple Junction.
The communities were distributed in a 40m by 80m field around
the knoll. At least seven active vent sites, including black
smoker complexes that were emitting superheated water at 360
degree C, were observed at the field. The faunal composition
of the Indian Ocean hydrothermal vent communities had links
to both Pacific and Atlantic vent assemblages. This discovery
supports the hypothesis that there is significant communication
between vent faunas in the Pacific and Atlantic Oceans via
active ridges in the Indian Ocean.
- Larval dispersal potential of the tubeworm
Riftia pachyptila at deep-sea hydrothermal vents
A. G. Marsh, L. S. Mullineaux, C. M. Young and D. T. Manahan.
Nature, Vol. 411, No. 6833, 3 May 2001, pp. 77-80.
Hydrothermal vents are ephemeral because of frequent volcanic
and tectonic activities associated with crust formation. Although
the larvae of hydrothermal vent fauna can rapidly colonize
new vent sites separated by tens to hundreds of kilometres,
the mechanisms by which these larvae disperse and recruit
are not understood. Here we integrate physiological, developmental
and hydrodynamic data to estimate the dispersal potential
of larvae of the giant tubeworm Riftia pachyptila. At in situ
temperatures and pressures (2 degree C and 250 atm), we estimate
that the metabolic lifespan for a larva of R. pachyptila averages
38 days. In the measured flow regime at a fast-spreading ridge
axis (9 degree 50'N; East Pacific Rise), this lifespan results
in potential along-ridge dispersal distances that rarely exceed
100 km. This limited dispersal results not from the physiological
performance of the embryos and larvae, but instead from transport
limitations imposed by periodic reversals in along-ridge flows
and sustained episodes of across-ridge flow. The lifespan
presented for these larvae can now be used to predict dispersal
under current regimes at other hydrothermal vent sites.
- Life After Death in the Deep Sea
R. A. Lutz, T. M. Shank and R. Evans.
American Scientist, Vol. 89, No. 5, Sep-Oct 2001, pp. 422-431.
The first examples of life forms not dependent on solar energy
were discovered by scientists using towed cameras and the
submersible Alvin in 1977 along hydrothermal vents of the
Galapagos Rift. Since then, investigators have made hundreds
of dives aboard Alvin to learn more about these unusual ecological
communities. In the spring of 1991, Alvin and its tender,
Atlantis II, happened to be on station above the East Pacific
Rise between 9 and 10 degrees north latitude only a few days
after the axial summit trough 2,550 meters below the surface
erupted, obliterating a thriving vent community. The authors
made numerous dives on the 9N Biotransect over the ensuing
10 years. Their article describes the return of life to the
vents and the ecological succession they witnessed.
- An off-axis hydrothermal vent field
near the Mid-Atlantic Ridge at 30 degree N
D. S. Kelley, J. A. Karson and D. K. Blackman, et al.
Nature, Vol. 412, No. 6843, 12 Jul 2001, pp. 145-149.
Evidence is growing that hydrothermal venting occurs not
only along mid-ocean ridges but also on old regions of the
oceanic crust away from spreading centres. Here we report
the discovery of an extensive hydrothermal field at 30 degree
N near the eastern intersection of the Mid-Atlantic Ridge
and the Atlantis fracture zone. The vent field - named "Lost
City" - is distinctly different from all other known sea-floor
hydrothermal fields in that it is located on 1.5-Myr-old crust,
nearly 15 km from the spreading axis, and may be driven by
the heat of exothermic serpentinization reactions between
sea water and mantle rocks. It is located on a dome-like massif
and is dominated by steep-sided carbonate chimneys, rather
than the sulphide structures typical of "black smoker" hydrothermal
fields. We found that vent fluids are relatively cool (40-75
degree C) and alkaline (pH 9.0-9.8), supporting dense microbial
communities that include anaerobic thermophiles. Because the
geological characteristics of the Atlantis massif are similar
to numerous areas of old crust along the Mid-Atlantic, Indian
and Arctic ridges, these results indicate that a much larger
portion of the oceanic crust may support hydrothermal activity
and microbial life than previously thought.
- Submersible Observations of the Hydrothermal
Vent Communities on the Iheya Ridge, Mid Okinawa Trough, Japan
S. Ohta and D. Kim.
Journal of Oceanography, Vol. 57, No. 6, Dec 2001, pp. 663-677.
During the Dives Nos. 409, 410, 480 and 481 of the Japanese
submersible Shinkai 2000, conducted on June 10 and 11, 1989
and on May 16 and 17, 1990, several hydrothermal vents and
prosperous vent associated biological communities were found
on the northern slope of the Iheya Ridge in the Mid-Okinawa
Trough (27 degree 32.5' N, 126 degree 58.5' E: depth 1,400
m). The first site we found, the "Calyptogena Site", was characterized
by a relatively thick blanket of sediments, pleated and/or
laminated lava flows, with occasional lobate pillows and white
and yellow stains. Although no remarkable shimmering water
and thermal anomalies were detected during the observations,
the extraordinarily dense community must be related to hydrothermal
activities. The community was dominated by the giant white
clam, Calyptogena okutanii, in biomass, and by Neolepas-type
primitive scalpellids and slender vestimentiferan tube-worms
in number. The second site, the "Pyramid Site", situated only
200 m northward from the Calyptogena Site has typical clear
smokers emitting hot water over 200 degree C, and is characterized
by a hard substratum of volcanic rocks and hydrothermal slabs.
No noteworthy succession was perceived at the Calyptogena
Site over a year. Many of the vent members occurred at both
sites. However, Calyptogena okutanii, which were confined
to the sediment bottom, Neolepas spp. and larger vestimentiferan
tube-worms were found to thrive only at the Calyptogena Site,
being only minor elements in the Pyramid Site. The global
distribution of several groups of organisms is discussed preliminarily
in zoogeographical terms based on comparison with other submersible
missions and surveys done by surface vessels.
- Temperature resistance of Hesiolyra
bergi, a polychaetous annelid living on deep-sea vent smoker
walls
B. Shillito, D. Jollivet and P. -M Sarradin, et al.
Marine Ecology Progress Series, Vol. 216, 6 Jul 2001, pp.
141-149.
For the first time, in vivo heat-exposure experiments were
conducted on the hydrothermal vent polychaete Hesiolyra bergi
from the hottest part of the vent biotope. Using a pressurised
incubator equipped with video-facilities, we found that H.
bergi, which forages around and in the tubes of the thermophilic
Alvinella sp., became hyperactive once temperature exceeded
35 degree C and further lost co-ordination in the 41 to 46
degree C interval, just before death occurred. Another exposure
experiment at 39 degree C for 3 to 4 h led to 80% mortality
(max) 9 h after heat shock, and 100% thereafter. In view of
the much higher temperatures recorded in this organism's habitat,
these results suggest that tolerance to high temperatures
(exceeding 40 degree C) is not a pre-requisite for life among
alvinellid tubes. Behavioural responses (escape from heat)
may suffice.
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