FY 2007 PI Report - National Oceanographic Partnership Program

Investigations of Chemosynthetic Communities on the Lower Continental Slope of
the Gulf of Mexico
Project Director - James Brooks
TDI-Brooks International, 1902 Pinon, College Station, TX 77845
Phone: (979) 693-3446 FAX: (979) 693-6389 E-mail: [email protected]
CO-PI Chuck Fisher
208 Mueller Laboratory ,The Pennsylvania State University, University Park, PA 16802
Phone: (814) 865-3365 FAX: (814) 865-9131 E-mail: [email protected]
CO-PI Harry Roberts
CSI-LSU, 304 Howe-Russell Geosciences Complex, Baton Rouge, LA 70803
Phone: 225.578-2964 FAX: 225-578-2520 E-mail: [email protected]
CO-PI Bob Carney
Coastal Ecology Institute, Louisiana State University, South Stadium Road, Baton Rouge, LA 70803
Phone: 225.578-6511 FAX: 225-767-6840 E-mail: [email protected]
CO-PI Ian MacDonald
TAMUCC, 6300 Ocean Dr. ST320, Corpus Christi, TX 78412
Phone: 361.825-2234 FAX: (361) 825-2742 E-mail: [email protected]
CO-PI Erik Cordes
OEB Department, Harvard University, 3079 BioLabs, 16 Divinity Ave, Cambridge, MA 02138
Phone: 617-495-9156 FAX: 617-495-8848 E-mail: [email protected]
CO-PI Samantha Joye
Marine Sciences Bldg, University of Georgia, Athens, GA 30602-3636
Phone: 706-542-5893
FAX: (706) 542-5888 E-mail: [email protected]
CO-PI Liz Goehring
The Pennsylvania State University, 208 Mueller Laboratory, University Park PA 16802-5301
Phone: (814) 863-0278 FAX: (814) 865-9131 E-mail: [email protected]
CO-PI Bernie Bernard
TDI-Brooks International, 1902 Pinon, College Station, TX 77845
Phone: 979.690-6287 FAX: (979) 693-6389 E-mail: [email protected]
CO-PI Gary Wolff
TDI-Brooks International, 1902 Pinon, College Station, TX 77845
Phone: 979.846-7679 FAX: (979) 693-6389 E-mail: [email protected]
Award Number: 1435-01-05-39187
The long-term goals of this study are to add to the understanding of the oceanography and ecology of
the deep-sea with emphasis on cold seep communities and hard bottom communities on the Gulf of
Mexico (GoM) continental slope. Preliminary studies have shown that seep communities at the slope
base are different from those on the upper slope, in much the same way that the normal background
fauna differ. Compared to the upper-slope, there is limited understanding of seep and other hard
bottom communities below 1,000 meters in the Gulf of Mexico.
The objectives of this study are: Characterize known, or newly discovered chemosynthetic
communities at depths below 1,000 meters in the central and western Gulf of Mexico. Characterize all
other hard bottom biological communities encountered regardless of association with active
hydrocarbon seep activity or living chemosynthetic community species in the central and western Gulf
of Mexico. Determine the comparative degree of sensitivity of anthropogenic impacts for the above
through a variety of approaches such as rarity, unique taxonomy/biodiversity, or other environmental
risk assessment methodologies. Understanding how these deep communities are similar or different
from their shallower water counterparts. Further develop successful assessment methodologies for
correlation of remote sensing information such as bathymetry, seabed acoustic reflectivity, sub-bottom
structure, and other geophysical signatures obtained by non-visual techniques with the “potential”
presence of non-soft bottom biological communities at depths below 1,000 meters. The target objective
is to provide some level of predictive capability that can be used by MMS to avoid impacts to lower
slope sensitive biological communities such as presented by Roberts (2001) for upper slope
communities. Assess and explain diversity distribution and abundance of marine species at depths
below 1,000 m in the central and western Gulf, as well as improve the understanding of the functional
role of marine species in areas of active hydrocarbon seep activity or living chemosynthetic
communities. These objectives will be accomplished through a combination of both exploratory work
and more focused studies including process-based work on known communities.
In order to meet the objectives outlined above, the following scientific and technical plan is being
ƒ Compile and analyze all of the appropriate available data to predict the location of significant
chemoautotrophic or other hard-bottom communities at depths >1,000 m in the GoM. This resulted
in the selection of 10 – 20 sites for visitation during the Reconnaissance Cruise. In addition to
providing specific locations for most of the dives for the first submersible cruise, ground-truth data
collected during the Reconnaissance Cruise allowed evaluation of the predictive value of the
various criteria used for site selection and provides data on the types of communities present at the
sites. of these data (geophysical and geochemical predictors, and presence/absence of various
community types) These analyses were further enriched with multivariate analysis and when
selected sites were more intensively imaged and sampled for macro and microbiology and
chemistry. A third level of information comes from mapping community occurrence type and
density onto the high resolution maps of surficial geology and seafloor topography made at of each
of the three to four primary study sites. The“sub-goal” here is to enrich the predictive value of
these high-resolution data sets to include the occurrence of different types of communities/ habitats
on a spatial scale of meters.
Characterize types of significant hard bottom communities encountered. First order community
characterization will be identification of component taxa and descriptions of communities present
at different sites. Second order characterizations will include distributions and abundances of taxa
with respect to chemistry and surficial geology and measures of community structure and function.
Third order characterizations/analyses will include interactions with background fauna, taxonomic
relations of species from key taxa to related species at other depths and in other areas, and
community-level comparisons among sites and related communities at other depths and areas. All
sites visited by submersible or ROV are characterized with respect to surficial geology,
geochemistry of sediments and epibenthic bottom water, types of communities present, microbial
activities, and mega/macrofaunal species present. At four sites more extensive survey and
experimentation was conducted to better characterize and understand the communities, and test
hypotheses relating to community composition, tubeworms, trophic interactions, and microbiology.
During the submersible and ROV cruises we collect imagery that provides data on endemic species
occurrence, distribution and densities, and visitation by vagrant mobile megafauna. We make
quantitative collections of communities that provide the material needed for taxonomic,
biogeographic, and trophic studies, and analyze the collections in ways that provide a variety of
data on community structure and function as well as composition. In situ chemical measurements
were made to describe the microhabitat chemistry of the major community types. Faunal
distributions are mapped with respect to surficial geology and chemistry. The microbial
communities in the sediments are characterized and temporal studies of the communities initiated
(with time lapse camera, base line imaging, and growth).
Descriptions of the communities encountered and analyses of background fauna trapped, trawled
and imaged over the course of the study will contribute to the assessment of diversity, distributions,
and abundance of marine species below 1,000 m in the GoM. Correlation analyses of faunal
occurrence with geologic features and seep chemistry will further contribute to the explanation of
these patterns. Trophic analyses, time-lapse camera data, community analyses, and growth studies
greatly improve our understanding of the functional role of many of the marine species
encountered. By working under the auspices of the Census of Marine Life ChEss program and
providing all data collected to their database, we assure widespread international access to all
biodiversity and biogeography data collected.
Direct determination of sensitivity of individual species to particular potential anthropogenic
impacts is addressed through assessment of rarity and unique taxonomy/biogeography of key
species and communities, biodiversity of communities, and by interpretation made in the context of
the degree of similarity to related communities on the upper Louisiana slope and what is known
about those communities. The comparisons of community-level associations to similar
communities elsewhere, and the proposed vestimentiferan growth studies will strengthen the power
of these analyses. Existing collaborations with molecular and classical taxonomic experts will
facilitate the identification of unknown species. The molecular analyses of foundation and other
key species provide information necessary to detect significant levels of genetic isolation at any
site, analyze relations to taxa at other sites, and determine bathymetric ranges of the metapopulations.
Key individuals participating in this work and their roles are: Dr. James Brooks (TDI-BI) is the project
director and will take the lead in administration of this project and assist in the geochemical studies.
Dr. Charles Fisher (PSU) coordinates the biological studies, Dr. Harry Roberts (LSU) coordinates the
geological/geophysical studies, and Ms. Liz Goehring (PSU) coordinates the education and outreach
activities. Dr. Erik Cordes will work with Fisher’s team on studies of seep communities and synthesis
and publication of results for other hard bottom communities discovered. Dr. Stephane Schaeffer
oversees molecular phylogenetic screening of foundation species and their symbionts (tubeworms,
mussels and clams) and other potential new species (and symbioses), as needed. Dr. Robert Carney
leads the studies of interactions with background fauna and trophic exchange between seep/hard
bottom communities and larger mobile fauna. Drs. Fisher, Carney, and Cordes share responsibility for
coordination with taxonomists and molecular phylogenists and proper curation of samples. Dr. Ian
MacDonald directs the use of digital imagery in all phases of the study. Dr. Samantha Joye is
responsible for the microbial ecology and sulfide geochemistry studies. Dr. Tim Shank (WHOI) will
phylogentically characterize new species of megafaunal crustaceans and include at least the shrimp in
his ongoing biogeographic analyses. Dr. Bob Vreijenhoek (MBARI) will do the same with clams and
their symbionts and other gastropods as needed. Limpets and snails will be sent to Anders Waren
(Swedish Museum of Natural History) and chitons to Julia Sigwart (University College Dublin) for
morphological characterization. Dr. Stéphane Hourdez (Stacione Biologique de Roscoff, France) leads
the polychaete phylogenetic characterizations and descriptions of new species of polynoids and
siboglinids (using both molecular and classical approaches). He also assists with molecular
characterization of foundation species. Dr. Stephane Cairns (Smithsonian) oversees curation and
identification of cnidarians, with assistance of Daphne Fautine (University of Kansas) and Dennis
Opreska (Oak Ridge). Dr. Cheryl Morrison (USGS Leetown Science Center) will include any samples
of Lophelia pertusa collected in her ongoing studies of the phylogeography and population genetics of
this foundation coral species and also collaborate with Dr. Cairns by contributing to the molecular
systematics of other hard corals, as needed. Dr. Sabine Stohr (Swedish Museum of Natural History)
has agreed to examine all ophuiroids collected. Dr. Monika Bright and her research team (Univ.
Vienna) will sort and identify meiofauna collected with mussel and tubeworm communities and in
sediment cores. Other faunal groups will be sent to appropriate experts as needed. Additionally, two
internationally recognized research groups from the Max Planck Institute of Marine Microbiology in
Bremen will bring unique expertise and equipment to bear on the study. Dr. Nicole Dublier’s group
will use quantitative mRNA analyses to determine the relative activities of chemoautotrophic and
methanotrophic symbiont populations in the dual symbiont-containing mussels. Dr. Antje Botieus’
group will bring their in-situ seep-chemistry analysis system and expertise on the ALVIN and ROV
cruises. Dr. Bernie Bernard coordinates the isotope, hydrocarbon and ancillary measurements. Dr.
Thomas McDonald is the principal hydrocarbon chemist for the project. Dr. Gary Wolff is the project
data manager, Ms. Susan Wolff is the project’s technical editor. Ms. Suzanne Cardwell provides
financial and project administrative support.
The planned work will be primarily with analysis and interpretation of data collected from the
completed field work.
The historical data review was completed the beginning of 2006. The Reconnaissance Cruise was
conducted on the TDI-Brooks research vessel R/V GYRE from 11 to 25 March 2006, using over-theside imaging equipment and shipboard acoustic methods and was the initial cruise conducted for this
contract. The cruise was completed in two, week-long legs with an interim port call in Venice, LA.
Leg I (11-18 March) was dedicated to drift camera work to survey the seabed at selected sites. Leg II
(19-25 March) involved both drift camera and trawling/box core work efforts for isotopic
characterization of the seep-background interactions near seep sites in the deep GoM. Twenty-four
sites were studied. The cruise mobilized and embarked from Freeport, Texas. The objective was to
provide timely input for the site selection process for the subsequent ALVIN expedition (May 2006).
The Deep Chemosynthetic Community Characterization Cruise (DCCC) was conducted on the Wood’s
Hole Oceanographic Institute (WHOI) research vessel R/V ATLANTIS and the ALVIN Deep
Submergence Vehicle (DSV) from 7 May – 2 June 2006, and was the second cruise conducted for this
contract. The cruise mobilized and embarked from Key West, Florida, and de-mobilized at Galveston,
Texas. Twenty-four dives were completed on ALVIN. At some sites, multiple dives were made while
at other sites only a single dive was completed. The Deep Chemosynthetic Reconnaissance II Cruise
(DCR2) was conducted on the NOAA Ship research vessel Ronald H. Brown and the ROV JASON
from 4 June - 6 July 2007, and was the fourth cruise conducted for this contract. The cruise mobilized
and embarked from Panama City, Florida, and de-mobilized at Galveston, Texas. Post-cruise reports
were completed for all cruises and were submitted to MMS. The data from these two cruises was uploaded to a site located on the TDI-Brooks International Website. All program researchers have
password-protected access to these data.
Reconnaissance Cruise - Trawls were completed at three sites, MC685, MC548 and AT209. Two box
cores were collected at site WR265, survey photos captured with the bottom in view (BIV) were
10,922, site characterization and evaluation was completed on 24 sites. DCCC Cruise – Multiple dives
were made at four sites and the geology, geochemistry, microbiology and biology of the sites
thoroughly characterized using growth rate analysis. Push cores were collected for geological,
geochemical, and microbial analyses, chemical analysis, quantitative collections of other tubeworm
communities, mussel beds, and clam beds were made. Trawls were completed, traps and cameras were
used to capture and identify the visitors to the seep and coral communities. A rotary camera was left on
the bottom for up to a year. More than 14,000 down-camera and 400 macro images were recorded.
DCR2 - Sixteen dives with the ROV JASON provided essential information on the ecology and
biodiversity of these deep-sea communities.
National Security
This program will provide critical information on the location and function of seep communities to
MMS. As manager of the nation’s seafloor mineral resources, MMS will use this information to aid in
the development of critical energy resources, which may affect domestic energy production.
Economic Development
Increased energy and mineral production will have a positive economic impact at numerous levels in
Quality of Life
Information on the location and functioning of seep communities gathered by this program will have a
positive impact on other ocean users, the natural environment, and the human environment. It will aid
in minimizing the environmental impact on sensitive habitat and mitigate any potential damage to
these communities.
Science Education and Communication
Education outreach efforts outside of the cruise website and since the last report have focused on the
merging the MMS deep slope outreach with the development of FLEXE. FLEXE (From Local to
Extreme Environments) is one of four major NSF-funded projects within the GLOBE program
(www.globe.gov), bringing the remote environments of hydrothermal vents and cold seeps for the first
time to GLOBE. GLOBE is a NASA and NSF funded web-based science education program
emphasizing K-12 Earth System Science concepts, and is currently operating in 109 countries
involving over 17,000 schools and over one million students worldwide. As the name suggests,
FLEXE guides students in understanding deep-sea extreme environments through a comparison with
analogous local environmental measures. FLEXE builds on the success of the SEAS "Classroom to
Sea" lab concept and extends it by embedding these labs in curricular units built on Earth Systems
Science essential concepts. The FLEXE project started in 2006, completed initial pilot testing in Spring
2007 and is currently testing the full system and first curricular unit this Fall 2007.
Data is provided to the ChEss database, which is a component of the Census of Marine Life (CoML)
Ocean Biogeographic Information System (OBIS) data base. All gene sequences will be submitted to
the international genetic data base, GenBank. The work proposed here will contribute significantly to
the goals of the Atlantic Equatorial Belt studies of the ChEss program, particularly the components that
will allow interpretation of our findings in the context of seeps around the world. The second
component of the CoML program relevant to this project is the CoMargE component. Dr. Carney, codirector of CoMargE and supported by the MMS Coastal Marine Program, will transfer past MMS
survey data into the CoML OBIS database system.
Chemosynthetic Ecosystems Study (MMS Report 95-0021).
Stability and Change in Gulf of Mexico Chemosynthetic Communities (MMS Report 2002036). http://www.gomr.mms.gov/homepg/regulate/environ/studies/2002-036.pdf.
The Deepwater Program: Northern Gulf of Mexico Continental Slope Habitat and Benthic
Ecology (MMS contract 1435-01-99-CT-30991).
Figure 1. Site locations of JASON dives.
Adkins, J.F., H. Cheng, E.A. Boyle, E.R.M. Druffel, and R.L. Edwards. 1998. Deep-sea coral
evidence for rapid climate change in ventilation of the deep North Atlantic 15,400 years ago.
Science 280:725-728.
Adkins, J.F., E.A. Boyle, W.B. Curry, and A. Lutringer. 2003. Stable isotopes in deep-sea corals
and a new mechanism for “vital effects”. Geochim. Cosmochim. Acta 67: 1129-1143.
Aharon, P. and B.S. Fu. 2000. Microbial sulfate reduction rates and sulfur and oxygen isotope
fractionations at oil and gas seeps in deepwater Gulf of Mexico. Geochim. Cosmochim. Acta
Aloisi, G., I. Bouloubassi, S.K. Heijs, R.D. Pancost, C. Pierre, J.S.S. Damste, J.C. gottschal, L.J.
Forney, J.M. Rouchy. 2002. CH4-consuming microorganisms and the formation of carbonate
crusts at cold seeps. Earth and Planetary Science Letters 203:195-203.
Ambrose, R.F. and T.W. Anderson. 1990. Influence of an artificial reef on the surrounding infaunal
community. Mar. Biol. 107:41-52.
Andersen, A.C., S. Hourdez, B. Marie, D. Jollivet, F.H. Lallier, and M. Sibuet. 2004. Escarpia
southwardae sp. nov., a new species of vestimentiferan tubeworm (Annelida, Siboglinidae)
from West African cold seeps. Can. J. Zool. 82: 980-999.
Andrews, A.H., E.E. Cordes, M.M. Mahoney, K. Munk, K.H. Coale, G.M. Cailliet, and J. Heifetz.
2002. Age, growth and radiometric age validation of a deep-sea, habitat-forming gorgonian
(Primnoa resedaeformis) from the Gulf of Alaska. Hydrobiologia 471:101-110.
Arvidson, R.S., J.W. Morse, and S.B. Joye. 2004. The sulfur biogeochemistry of chemosynthetic
cold seep communities, Gulf of Mexico, USA. Mar. Chem. 87: 97–119.
Bergquist, D.C., I.A. Urcuyo, and C.R. Fisher. 2002. Establishment and persistence of seep
vestimentiferan aggregations from the upper Louisiana slope of the Gulf of Mexico. Mar. Ecol.
Prog. Ser. 241: 89-98.
Bergquist, D.C., T. Ward, E.E. Cordes, T. McNelis, R. Kosoff, S. Hourdez, R. Carney, and C.R.
Fisher. 2003a. Community structure of vestimentiferan-generated habitat islands from upper
Louisiana slope cold seeps. J. Exp. Mar. Bio. Ecol. 289: 197-222.
Bergquist, D.C., J. Andras, T. McNelis, S. Howlett, M.J. van Horn, and C.R. Fisher. 2003b.
Succession in upper Louisiana slope cold seep vestimentiferan aggregations: the importance of
spatial variability. PSZN Mar. Ecol. 24:31-44.
Bergquist, D.C., C. Fleckenstein, J. Knisel, B. Begley, I. R. MacDonald, and C.R. Fisher. 2005.
Variations in seep mussel bed communities along physical and chemical environmental
gradients. Mar. Ecol. Prog. Ser 293:89-97.
Bett, B.J. 1997. Atlantic Margin Environmental Survey: Seabed Survey of the Shelf Edge and
Slope west of Shetland. Published by Challenger Division for Seafloor Processes: Southampton
Oceanography Centre, NERC/ University of Southampton. Pp.165.
Boetius, A., K. Ravenschlag, C. Schubert, D. Rickert, F. Widdel, A. Gieseke, R. Amann, B.B.
Jørgensen, U. Witte, and O. Pfannkuche. 2000. A marine microbial consortium apparently
mediating anaerobic oxidation of methane. Nature 407:623-626.
Brooks, J.M., D.A. Wiesenburg, H. Roberts, R.S. Carney, I.R. MacDonald, C.R. Fisher, N.L.
Guinasso Jr., W.W. Sager, S.J. McDonald, R.A. Burke Jr., P. Aharon, and T.J. Bright. 1990.
Salt, seeps and symbiosis in the Gulf of Mexico. EOS 71(45):1772-1773
Cairns, S.D. 1978. A checklist of the ahermatypic scleractinia of the Gulf of Mexico, with the
description of a new species. Gulf Res. Rep.6: 9-15.
Cairns, S.D. 2001. A brief history of taxonomic research on azooxanthellate Scleractinia (Cnidaria:
Anthozoa). Bull. Biol. Soc. Wash. 10:191-203.
Cairns, S.D., D.M. Opresko, T.S. Hopkins, and W.W. Schroeder. 1993. New records of deep-water
Cnidaria (Scleractinia & Antipatharia) from the Gulf of Mexico. Northeast Gulf Sci. 13:1-11.
Carney, R.S. 1994. Consideration of the oasis analogy for chemosynthetic communities at Gulf of
Mexico hydrocarbon vents. Geo-Mar. Let. 14:149-159.
Carney, R.S. 1997. Basing conservation policies for the deep-sea floor on current diversity
concepts: a consideration of rarity. Biodiv. Conserv. 6:1463-1485
Carney, R.S. 2005. Zonation of deep biota on continental margins. Oceanogr. Mar. Biol. Ann.
Rev. 43:211-278.
Cary, S.C. 1989. Multiple trophic resources for a chemoautotrophic community at a cold water
brine seep at the base of the Florida escarpment. Mar. Biol. 100:411-418
Cary, S.C., C.R. Fisher, and H. Felbeck. 1988. Mussel growth supported by methane as sole carbon
and energy source. Science 240:78-80.
Childress, J.J., C.R. Fisher, J.M. Brooks, M.C. Kennicutt II, R. Bidigare, and A.E. Anderson. 1986.
A methanotrophic marine molluscan symbiosis: Mussels fueled by gas. Science 233: 13061308.
Clarke, K.R., and R.M. Warwick. 2001. Change in marine communities: An
statistical analysis. Primer-E, Plymouth, UK. 170 pp.
approach to
Cordes, E.E., D.C. Bergquist, K. Shea, and C.R. Fisher. 2003. High hydrogen sulfide demand of
long-lived vestimentiferan tube worm aggregations modifies the chemical environment at deepsea hydrocarbon seeps. Ecology Letters 6: 212-219.
Cordes, E.E., M.A. Arthur, K. Shea, and C.R. Fisher. 2005. Modeling the mutualistic interactions
between tubeworms and microbial consortia. PLoS Biol. 3: e77.
Cordes, E.E., S. Hourdez, B.L. Predmore, M.L. Redding, and C.R. Fisher. In press. Succession of
hydrocarbon seep communities associated with the long-lived foundation species
Lamellibrachia luymesi. Mar. Ecol. Prog. Ser.
Craddock, C., W.R. Hoeh, R.G. Gustafson, R.A. Lutz, J. Hashimoto, and R.J. Vrijenhoek 1995.
Evolutionary relationships among deep-sea mytilids (Bivalvia: Mytilidae) from hydrothermal
vents and cold water methane sulfide seeps. Mar. Biol. 121(3):477-485
De Beukelaer, S.M., I.R. MacDonald, N.L. Guinnasso, and J. A. Murray. 2003. Distinct side-scan
sonar, RADARSAT SAR, and acoustic profiler signatures of gas and oil seeps on the Gulf of
Mexico slope. Geo-Marine Letters 23(3-4):177-186.
Dhillon, A., A. Teske, J. Dillon, D.A. Stahl, and M.L. Sogin. 2003. Molecular characterization of
sulfate-reducing bacteria in the Guaymas Basin. App. Env. Microbiol. 69: 2765-2772.
Dodge, R.E. and J. Thomson. 1974. The natural radiochemical and growth records in contemporary
hermatypic corals from the Atlantic to the Caribbean. Earth Planet Sci 36:339-356.
Dodge, R.E. and G.W. Brass. 1984. Skeletal extension, density, and calcification of the reef coral,
Montastrea annularis: St. Criox, U.S. Virgin Islands. Bull Mar Sci 34:288-307.
Doerries, M.B. and C.L. Van Dover. 2003. Higher-taxon richness as a surrogate for species
richness in chemosynthetic communities. Deep-Sea Res. I 50:749-755.
Druffel, E.R.M., L.L. King, R.A. Belastock, and K.O. Buesseler. 1990. Growth rates of a deep-sea
coal using 210Pb and other isotopes.
Dunbar, R.B., G.M. Wellington, M.W. Colgan, and P.W. Glynn. 1994. Eastern Pacific sea surface
temperature since 1600 A.D.: The 18O record of climate variability in Galapagos corals.
Paleoceanography 9:291-315.
Espedal, H. and T. Wahl. 1999. Satellite SAR oil spill detection using wind history information.
Int. J. Remote Sensing 20(1):49-65.
Fisher, C.R. 1996. Ecophysiology of primary production at deep-sea vents and seeps. In: Deep-sea
and extreme shallow-water habitats: affinities and adaptations. R. Uiblein, J. Ott, and M.
Stachowtish (eds.) Biosystematics and Ecology Series 11: 311-334.
Fisher, C.R., J.J. Childress, S.A. Macko, and J.M. Brooks. 1994. Nutritional interactions at
Galapagos hydrothermal vents: Inferences from stable carbon and nitrogen isotopes. Mar. Ecol.
Prog. Ser. 103: 45-55.
Frazer, T.K., W.J. Lindberg, and G.R. Stanton. 1991. Predation on sand dollars by gray triggerfish,
Ballistes capriscus, in the northeastern Gulf of Mexico. Bull. Mar. Sci. 48:159-164.
Frederiksen, R., A. Jensen, and H. Westerberg. 1992. The distribution of the scleractinian coral
Lophelia pertusa around the Faroe Islands and the relation to internal mixing. Sarsia 77:157171.
Freidwald, A., R. Heinrich, and J. Patzold. 1997. Anatomy of a deep water coral reef mound from
Sternsund, west Finnmark, northern Norway: James, N.P. and J.A.D. Clarke Eds. Cool-water
carbonates. Society for Sedimentary Geology, Special Volume 56:144-146.
Freytag, J.K., P. Girguis, D.C. Bergquist, J.P. Andras, J.J. Childress, and C.R. Fisher. 2001. Sulfide
acquisition by roots of seep tubeworms sustains net chemoautotrophy. Proc. Nat. Acad. Sci. 98;
Gardiner, S.L. and S. Hourdez. 2003. On the occurrence of the vestimentiferan tube worm
Lamellibrachia luymesi van der Land and Nørrevang, 1975 (Annelida: Pogonophora) in
hydrocarbon seep communities in the Gulf of Mexico. Proc. Biol. Soc. Wash. 116(2):380-394.
Gilliam, J.F. 1989. Strong effects of a foraging minnow on a stream benthic community. Ecology
Glover, A.G. and C.R. Smith. 2003. The deep-sea floor ecosystem: current status and prospects of
anthropogenic change by the year 2025. Conserv. Biol. 30:219-141.
Heifetz, J. 2002. Coral in Alaska: distribution, abundance, and species associations. Hydrobiologia
Heikoop, J.M., J.J. Dunn, M.J. Risk, H.P. Schwarcz, T.A. McConnaughey, and I.M. Sandeman.
2000. Separation of kinetic and metabolic isotope effects in carbon-13 records preserved in reef
coral skeletons. Geochem Cosmochim Acta 64:975-987.
Helmle, K.P., K.E. Kohler, and R.E. Dodge. 2002. Relative Optical Densitometry and The Coral
X-radiograph Densitometry System: CoralXDS. Presented Poster, Int. Soc. Reef Studies 2002
European Meeting. Cambridge, England. Sept. 4-7.
Highsmith R.C. 1979. Coral growth rates and environmental control of density banding. J. Exp.
Mar. Bio. Ecol. 37:105-125.
Hovland, M. and E. Thomsen. 1997. Cold-water corals- are they hydrocarbon seep related? Mar.
Geol. 137:159-164.
Hovland, M., P.B. Mortensen, T. Brattegard, P. Strass, and K. Rokoengen. 1998. Ahermatypic
coral banks off mid-Norway: Evidence of a link with seepage of light hydrocarbons. Palaios
Hudson J.H., E.A. Shinn, R.B. Halley, and B. Lidz. 1976. Sclerochronology: a tool for interpreting
past environments. Geology 4:361-364.
Ivany, L.C., W.P. Patterson, and K.C. Lohmann. 2000. Increase in seasonality across the EoceneOligocene boundary inferred from fossil otoliths. Nature 407:887-890.Jensen, A., and R.
Frederiksen. 1992. The fauna associated with the bank-forming deepwater coral Lophelia
pertusa (Scleractinia) on the Faroe shelf. Sarsia 77:53-69.
Ivany, L.C., K.C. Lohmann, and W.P. Patterson. 2003. Paleogene temperature history of the U.S.
Gulf Coastal Plain inferred from 18O of fossil otoliths, pp. 232-251. In:D.R. Prothero, L.C.
Ivany and E.A. Nesbitt Eds. From Greenhouse to Icehouse, The Marine Eocene-Oligocene
Transition. Columbia Univ. Press, New York.
Joye, S.B., A. Boetius, B.N. Orcutt, J.P. Montoya, H.N. Schulz, M.J. Erickson, and S.K. Lugo.
2004. The anaerobic oxidation of methane and sulfate reduction in sediments from Gulf of
Mexico cold seeps. Chem. Geol. 205: 219–238.
Kennicutt II, M.C., R.H. Green, P. Montagna, and P.F. Roscigno. 1996. Gulf of Mexico Offshore
Operations Monitoring Experiment (GOOMEX), Phase I: Sublethal responses to contaminant
exposure-introduction and overview. Can. J. Fish. Aquat. Sci. 53:2540-2553.
Knittel, K., A. Boetius, A. Lemke, H. Eilers, K. Lochte, O. Pfannkuche, P. Linke, and R. Amann.
2003. Activity, distribution, and diversity of sulfate reducers and other bacteria in sediments
above gas hydrate (Cascadia Margin, OR). Geomicrobiol. J. 20:269-294.
Knittel, K., T. Lösekann, A. Boetius, R. Kort, and R. Amann. 2005. Diversity and distribution of
methanotrophic Archaea at cold seeps. Applied and Environmental Microbiology, pp. 467–479.
Kornacki, A.S., J.W. Kendrick and J.L. Berry. 1994. The impact of oil and gas vents and slicks on
petroleum exploration in the deepwater Gulf of Mexico. Geo-Marine Letters 14(2/3): 160-169.
Koslow, J.A., K. Gowlett-Jones, J.K. Lowry, T. O’Hara, G.C.B. Poore, and A. Williams. 2001.
Seamount benthic macrofauna off southern Tasmania: community structure and impacts of
trawling. Mar. Ecol. Prog. Ser. 213:111-125.
Kulm, L.D., E. Suess, J.C. Moore, B. Carson, B.T. Lewis, S.D. Ritger, D.C. Kadko, T.M.
Thornburg, R.W. Embley, W.D. Rugh, G.J. Massoth, M.G. Langseth, G.R. Cochrane, and R.L.
Scamman, 1986, Oregon subduction zone: Venting, fauna, and carbonates, Science, v. 231, p.
Lanoil, B.D., R. Sassen, M.T. La Duc, S.T. Sweet, and K.H. Nealson. 2001. Bacteria and Archaea
physically associated with Gulf of Mexico gas hydrates. Appl. Environ. Microbiol.
Levin, L.A., W. Ziebis, G.F. Mendoza, V.A. Growney, M.D. Tryon, K.M. Brown, C. Mahn, J.M.
Gieskes, and A.E. Rathburn. 2003. Spatial heterogeneity of macrofauna at northern California
methane seeps: Influence of sulfide concentration and fluid flow. Mar. Ecol. Prog. Ser. 256:
LGL Ecological Research. 1987. Northern Gulf of Mexico Continental Slope Study Technical
Volume, Minerals Management Service, Gulf of Mexico OCS Regional Office: 632.
Lough, J.M. and D.J. Barnes. 2000. Environmental controls on growth of the massive coral Porites.
J. Exp. Mar. Bio. Eco. 245:225-243.
Loya, Y. and B. Rinkevich. 1980. Effects of oil pollution on coral reef communities. Mar. Ecol.
Prog. Ser. 3: 167-180.
MacAvoy, S.E., R.S. Carney, C.R. Fisher, and S.A. Macko. 2002. Use of chemosynthetic biomass
by large, mobile, benthic predators in the Gulf of Mexico. Mar. Ecol. Prog.Ser. 225: 65-78.
MacAvoy, S.E., C.R. Fisher, R.S. Carney, and S.A. Macko. 2005. Nutritional associations among
fauna at hydrocarbon seep communities in the Gulf of Mexico. Mar. Ecol. Prog. Ser. 225: 6578.
MacDonald, I.R., N.L. Guinasso, Jr., S.G. Ackleson, J.F. Amos, R. Duckworth, R. Sassen, and
J.M. Brooks. 1993. Natural oil slicks in the Gulf of Mexico visible from space. J. Geophys.
Res. 98 C9:16351-16364.
MacDonald, I.R., N.L. Guinasso, Jr., R. Sassen, J.M. Brooks, L. Lee, and K.T. Scott. 1994. Gas
hydrate that breaches the sea floor on the continental slope of the Gulf of Mexico. Geology
MacDonald, I.R., J.F. Reilly Jr., S.E. Best, R. Venkataramaiah, R. Sassen, J. Amos, and N.L.
Guinasso, Jr. 1996. A remote-sensing inventory of active oil seeps and chemosynthetic
communities in the northern Gulf of Mexico. Hydrocarbon migration and its near-surface
expression. D. Schumacher and M. A. Abrams, American Association of Petroleum Geologists.
Memoir 66:27-37.
MacDonald, I.R., D. Buthman, W.W. Sager, M.B. Peccini, and N.L. Guinasso Jr. 2000. Pulsed oil
discharge from a mud volcano. Geology 28(10):907-910.
MacDonald, I.R., R. Arvidson, R.S. Carney, C. F. Fisher, N.L. Guinasso Jr., S. Joye, P. Montagna,
J.W. Morse, D.C. Nelson, E. Powell, W. Sager, R. Sassen, S. Schaeffer, and G.A. Wolff. 2002.
Stability and Change in Gulf of Mexico Chemosynthetic Communities: Final Report. New
Orleans, LA, U.S. Dept. Interior, Minerals Management Service, Gulf of Mexico OCS Region,
Contract 14-35-001-31813.
MacDonald, I.R., I. Leifer, R. Sassen, P. Stine, R. Mitchell, and N.L. Guinasso Jr. 2002. Transfer
of hydrocarbons from natural seeps to the water column and atmosphere. Geofluids 5: 95-107.
MacDonald, I.R., W.W. Sager,and M.B. Peccin. 2003. Association of gas hydrate and
chemosynthetic fauna in mounded bathymetry at mid-slope hydrocarbon seeps: northern Gulf
of Mexico. Mar. Geol. 198:133-158.
MacDonald, I.R., G. Bohrmann , E. Escobar , F. Abegg , P. Blanchon , V. Blinova , W.
Brückmann, M. Drews , A. Eisenhauer, X. Han , K. Heeschen , F. Meier , C. Mortera , T.
Naehr , B. Orcutt , B. Bernard , J. Brooks, and M. de Faragó. 2004. Asphalt volcanism and
chemosynthetic life, Campeche Knolls, Gulf of Mexico. Science 304: 999-1002.
MacDonald, I.R., L.C. Bender, M. Vardaro, B. Bernard, and J.R. Brooks. 2005. Thermal and visual
time-series at a seafloor gas hydrate deposit on the Gulf of Mexico Slope. Earth and Planetary
Science Letters 233:45-59.
Masson, D.G, B.J. Bett, D.S.M. Billett, C.L. Jacobs, A.J. Wheeler, and R.B. Wynn. 2003. The
origin of deep-water, coral-topped mounds in the northern Rockall Trough, Northeast Atlantic.
Mar. Geol. 194: 159-180.
McConnaughey, T. 1989. 13C and 18O isotopic disequilibrium in biological carbonates: I. Patterns.
Geochim Cosmochim Acta 53:151-162.
McCulloch, M., S. Fallon, T. Wyndham, E. Hendy, J. Lough, and D. Barnes. 2003. Coral record of
increased sediment flux to the inner Great Barrier Reef since European settlement. Nature
McMullin, E.R., S. Hourdez, S.W. Schaeffer, and C.R. Fisher. 2003. Phylogenetics and
biogeography of deep sea vestimentiferan tubeworms and their bacterial symbionts. Symbiosis.
Mikkelsen, N., H. Erlenkauser, J.S. Killingley, and W.H. Berger. 1982. Norwegian corals:
radiocarbon and stable isotopes in Lophelia pertusa. Boreas 5:163-171.
Milkov, A.V. and R. Sassen. 2000. Thickness of the gas hydrate stability zone, Gulf of Mexico
continental slope. Marine and Petroleum Geology 17:981.
Mills, H. J., C. Hodges, K. Wilson, I.R. MacDonald, and P.A. Sobecky. 2003. Microbial diversity
in sediments associated with surface-breaching gas hydrate mounds in the Gulf of Mexico.
FEMS Microbiol. Ecol. 46: 39-52.
Mills, H. J., R.J. Martinez, S. Story, and P.A. Sobecky. 2005. Characterization of the microbial
community structure in Gulf of Mexico gas hydrates: comparative analysis of DNA- and RNAderived clone libraries. Applied and Environmental Microbiology 71: 3235-3247.
Mitchell, R., I.R. MacDonald, and K. Kvenvolden. 1999. Estimates of total hydrocarbon seepage
into the Gulf of Mexico based on satellite remote sensing images. EOS Supplement 80(49):
Moore, D.R. and H.R. Bullis. 1960. A deep-water coral reef in the Gulf of Mexico. Bull. Mar. Sci.
Mortensen, P.B., M. Hovland, T. Brattegard, and R. Farestveit. 1995. Deep-water bioherms of the
scleractinian coral Lophelia pertusa L. at 64o on the Norwegian shelf: structure and associated
megafauna. Sarsia. 80:145-158.
Mortensen, P.B. and H.T. Rapp. 1998. Oxygen and carbon isotope ratios related to growth line
patterns in ckeletons of Lophelia pertusa (L) (Anthozoa, Scleractinia): Implications for
determination of linear extension rates. Sarsia 83:433-446.
Nikolaus, R., J.W. Ammerman, and I.R. MacDonald. 2003. Distinct pigmentation and trophic
modes in Beggiatoa from hydrocarbon seeps in the Gulf of Mexico. Aquat. Micr. Ecol. 32: 8593.
Nix, E., C.R. Fisher, K.M. Scott, and J. Vodenichar. 1995. Physiological ecology of a mussel with
methanotrophic symbionts at three hydrocarbon seep sites in the Gulf of Mexico. Mar. Biol.
122: 605-617.
Olu, K., S. Lance, M. Sibuet, P. Henry, A. Fiala-Medioni, and A. Dinet. 1997. Cold seep
communities as indicators of fluid expulsion patterns through mud volcanoes seaward of the
Barbados accretionary prism. Deep-Sea Res. 44(5):811-841.
Olu, K., V. Rigaud, A. Fifis, M.C. Fabri, P. Cochonat, H. Ondréas, and M. Sibuet. 2001. Spatial
distribution of chemosynthetic fauna from video records and mosaic analysis at a new cold seep
site in the Gulf of Guinea. Program and abstracts volume. Second Internatinal Symposium on
Deep-sea Hydrothermal Vent Biology. October 8-12, Brest, France. Pp. 208.
Orange, D.L., M.M. Angell, J.R. Brand, J. Thompson, T. Buddin, M. Williams, W. Hart, and W.
Berger. 2003. Geological and shallow salt tectonic setting of the Mad Dog and Atlantis fields:
Relationship between salt, faults, and seafloor geomorphology: Proceedings of the Offshore
Technology conference, Houston, Texas, May 5-8, 2003, OTC 15157, 16 p.
Orphan, V.J., L.T. Taylor, D. Hafenbradl, and E.F. Delong. 2000. Culture-dependent and cultureindependent characterization of microbial assemblages associated with high-temperature
petroleum reservoirs. App. Env. Microbiol. 66: 700-711.
Parrish, J.D. 1989. Fish communities of interaction shallow water habitats in tropical oceanic
regions. Mar. Ecol. Prog. Ser. 58:143-160.
Paull, C.K., B. Hecker, R. Commeau, R.P. Freeman-Lynde, C. Neumann, W.P. Corso, S. Golubic,
J.E. Hook, E. Sikes, and J. Curray. 1984. Biological communities at the Florida escarpment
resemble hydrothermal vent taxa. Science 226:965-967
Paull, C.K., A.J.T. Jull, L.J. Toolinand, and T. Linick. 1985. Stable isotope evidence for
chemosynthesis in an abyssal seep community. Nature 317:709-711
Posey, M.H. and W.G. Ambrose Jr. 1994. Effects of proximity to an offshore hard bottom reef on
infaunal abundances. Mar. Biol. 118:745-753.
Reed, J.K. 2002. Comparison of deep-water reefs and lithoherms off southeastern USA.
Hydrobiologia 471:57-69.Ricciardi, A. and E. Bourget. 1998. Weight to weight conversion
factors for marine benthic invertebrates. Mar. Ecol. Prog. Ser. 163:245-251.
Risk, M.J., J.M. Heikoop, M.G. Snow, and R. Beukens. 2002. Lifespans and growth patterns of
two deep-sea corals: Primnoa resedaeformis and Desmophyllum cristagalli. Hydrobiologia
Ritger, S., B. and Carson, E. Seuss. 1987. Methane-derived authigenic carbonates formed by
subduction-induced pore-water expulsion along the Oregon-Washington margin: GSA Bulletin
Roberts, H.H. 1996. Surface amplitude data: 3D-seismic for interpretation of sea floor geology
(Louisiana slope): Transactions of the Gulf Coast Association of Geological Societies 46:353362.
Roberts, H.H. 2001. Improved geohazards and benthic habitat evaluations: digital acoustic data
with ground truth calibrations. U.S. Dept. of the Interior, Minerals Management Service, Gulf
of Mexico OCS Region, New Orleans, LA. OCS Study MMS 2001-050. 116 pp + appendices.
Roberts, H.H. 2001. Fluid and gas expulsion on the northern Gulf of Mexico continental slope:
Mud-prone to mineral-prone responses: American Geophysical Union, Geophysical
Monograph 124:145-161.
Roberts, H.H., P. Aharon, R. Carney, J. Larkin, and R. Sassen. 1990. Responses to hydrocarbon
seeps, Louisiana continental slope: Geo-Marine Letters 10:232-243.
Roberts, H.H., and P Aharon. 1994. Hydrocarbon-derived carbonate buildups of the northern Gulf
of Mexico continental slope: A review of submersible investigations, Geo-Marine Letters
Roberts, H.H., and R. Carney. 1997. Evidence of episodic fluid, gas and sediment venting on the
northern Gulf of Mexico continental slope, Economic Geology 92: 863-879.
Roberts, H.H. and J.M. Coleman. (In Preparation). Improving the predictive capability of 3-D
seismic surface amplitude data for identifying chemosynthetic communities: Contract no. 14301-99-CA-30951, task order 17801.
Robinson, C.A., J.M. Bernhard, L.A. Levin, G.F. Mendoza, and J.K. Blanks. 2004. Surficial
hydrocarbon seep infauna from the Blake Ridge (Atlantic Ocean, 2150m) and Gulf of mexico
(690-2240m). PSZNI Mar. Ecol. 25:313-336.
Rogers, A.D. 1999. The biology of Lophelia pertusa (Linnaeus 1758) and other deep-water reef
forming corals and impacts from human activities. Int. Rev. Hydrobiol. 844:315-406.
Rowan, M.G., M.P.A. Jackson, and B.D. Trudgill. 1999. Salt-related fault families and fault welds
in the northern Gulf of Mexico: AAPG Bulletin 83:1454-1484.
Sager, W.W., C.S. Lee, I.R. MacDonald, and W.W. Schroeder. 1999. High-frequency near-bottom
acoustic reflection signatures of hydrocarbon seeps on the northern Gulf of Mexico continental
slope. Geo-Mar. Lett. 18:267-276.
Sahling, H., D. Rickert, R.W. Lee, P. Linke, and E. Suess. 2002. Macrofaunal community structure
and sulfide flux at gas hydrate deposits from the Cascadia convergent margin. Mar. Ecol. Prog.
Ser. 231:121–138
Sassen, R., J.M. Brooks, I.R. MacDonald, M.C.Kennicutt, N.L. Guinasso, and A. G. Requejo.
1993. Association of oil seeps and chemosynthetic communities with oil discoveries, upper
continental slope, Gulf of Mexico. Trans GCAGS 43:349-355.
Schroeder, W.W. 2002. Observations of Lophelia pertusa and the surficial geology at a deep-water
site in the northeastern Gulf of Mexico. Hydrobiologia 471:29-33.
Sharp, Z.D. and T.E. Cerling. 1996. A laser GC-IRMS technique for in situ stable isotope analyses
of carbonates and phosphates. Geochim. Cosmochim. Acta 60:2909-2916.
Sibuet, M. and K. Olu. 1998. Biogeography, biodiversity and fluid dependence of deep-sea coldseep communities at active and passive margins. Deep-Sea Res. 45:517-567.
Smith, J.E., H.P. Schwarcz, M.J. Risk, T.A. McConnaughey, and N. Keller. 2000.
Paleotemperatures from deep-sea corals: overcoming ‘vital effects’. Palaios. 15:25-32.
Snelgrove, R.V.R. and C.R. Smith. 2002. A riot of species in an environmental calm. Oceanogr.
Mar. Biol. Ann. Rev. 40:311-342.
Southward, E.C. 1988. Development of the gut and segmentation of newly settled stages of Ridgeia
(Vestimentifera): implications for relationship between Vestimentifera and Pogonophora. J.
Mar. Biol. Ass. UK. 68: 465-487.
Stohr, S. and M. Segonzac. 2005. Deep- sea ophiuroids (Echinodermata) from reducing and nonreducing environments in the North Atlantic Ocean. J. Mar. Biol. Ass. U.K. 85:383-402.
Suess, E., B. Carson, S. Ritger, J.C. Moore, M. Jones, L.D. Kulm and G. Cochrane, 1985,
Biological communities at vent sites along the subduction zones off Oregon, Bulletin of the
Biological Society of Washington, v. 6, p. 475-484.
Summerson, H.C. and C.H. Peterson. 1984. Role of predation in organizing benthic communities
of a temperate zone seagrass bed. Mar. Ecol. Prog. Ser. 15:63-77.
Swart P.K. 1983. Carbon and oxygen isotope fractionation in scleractinian corals. Earth Sci Rev.
Thiel, H. and J.A. Koslow, eds. 2001. Managing Risks to Biodiversity and the Environment on
the High Sea, Including Tools such as Marine Protected Areas. Scientific Requirements and
Legal Aspects, Proceedings of the Expert Workshop held at the International Academy for
Nature Conservation, Isle of Vilm Germany, 27 February- 4 March 2001.
Turnipseed, M., K.E. Knick, R.N. Lipcius, J. Dreyer, and C.L. Van Dover. 2003. Diversity in
mussel beds at deep-sea hydrothermal vents and cold seeps. Ecology Letters 6: 518-523.
Turnipseed, M., C.D. Jenkins, and C.L. Van Dover. 2004. Community structure in Florida
Escarpment seep and Snake Pit vent mussel beds. Mar. Biol. 145(1):121-132
Tyler, P.A. (ed). 2003. Ecosystems of the deep oceans. Ecosystems of the World Vol. 28.
Elsevier, Amsterdam, 569p.
Tyler, P.A., C.R. German, E. Ramirez-Llodra , and C.L. Van Dover. 2003. Understanding the
biogeography of chemosynthetic ecosystems. Oceanol. Acta 25:227–241
Van Dover, C.L. 2002a. Community structure of mussel beds at deep-sea hydrothermal vents. Mar.
Ecol. Prog. Ser. 230: 137-158.
Van Dover, C.L. 2002b. Trophic relationships among invertebrates at the Kairei hydrothermal vent
field (Central Indian Ridge). Mar. Biol. 141:761-772.
Van Dover, CL. 2003. Variation in community structure within
of the East Pacific Rise. Mar. Ecol. Prog. Ser. 253:55-66.
hydrothermal vent mussel beds
Van Dover, C.L. and B. Fry. 1994. Microorganisms as food resources at deep-sea hydrothermal
vents. Limnol. Ocean. 39:51-57.
Van Dover, C.L. and J.L. Trask. 2000. Diversity at deep-sea hydrothermal vent and intertidal
mussel beds. Mar. Ecol. Prog. Ser. 195:169-178.
Van Dover, C.L., C.R. German, K.G. Speer, L.M. Parson, and R.C. Vrijenhoek . 2002. Evolution
and biogeography of deep-sea vent and seep invertebrates. Science 295:1253-1257.
Van Dover, C.L., P. Aharon, J.M. Bernhard, M. Doerries, W. Flickinger, W. Gilhooly, K. Knick, S.
Macko, S. Rapoport, C. Ruppel, J. Salerno, R. Seitz, B.K. Sen Gupta, T.M. Shank, M.
Turnipseed, R. Vrijenhoek, and E. Watkins. 2003. Blake Ridge methane seeps: characterization
of a soft-sediment, chemosynthethically based ecosystem. Deep-Sea Res. 50:281-300.
Vardaro, M., I.R. MacDonald, L.C. Bender, and N.L. Guinasso Jr. Accepted. Dynamic biological
and physical processes observed at a gas hydrate outcropping on the continental slope of the
Gulf of Mexico. Geo-Marine Letters.
von Cosel, R. and Olu K. 1998. Gigantism in Mytilidae. A new Bathymodiolus from cold seep
areas on the Barbados Accretionary Prism. C. R. Acad. Sci (Ser. 3) 321:655-663.
Voordouw, G., S.M. Armstrong, M.F. Reimer, B. Fouts, A.J. Telang, Y. Shen, and D. Gevertz.
1996. Characterization of 16S rRNA genes from oil field microbial communities indicates the
presence of a variety of sulfate-reducing, fermentative, and sulfide-oxidizing bacteria. App.
Env. Microbiol. 62:1623-1629.
Wainright, S.A. 1964. Studies of the mineral phase of coral skeleton. Exp Cell Research, 34:213230.
Weinstein, M.P. and K.L. Heck. 1979. Ichthyofauna of seagrass meadows along the Caribbean
coast of Panama and in the Gulf of Mexico-composition, structure, and community ecology.
Mar. Biol. 50:97-107.
Wilson, J.B. 1979. ‘Patch’ development of the deep-water coral Lophelia pertusa (L.) on Rockall
Bank. J. Mar. Biol. Ass. U.K. 59:165-177.