Biogeochemical implications of the ubiquitous colonization of marine habitats and redox gradients by Marinobacter species
Identifieur interne : 000502 ( Pmc/Corpus ); précédent : 000501; suivant : 000503Biogeochemical implications of the ubiquitous colonization of marine habitats and redox gradients by Marinobacter species
Auteurs : Kim M. Handley ; Jonathan R. LloydSource :
- Frontiers in Microbiology [ 1664-302X ] ; 2013.
Abstract
The
Url:
DOI: 10.3389/fmicb.2013.00136
PubMed: 23734151
PubMed Central: 3660661
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Biogeochemical implications of the ubiquitous colonization of marine habitats and redox gradients by <italic>Marinobacter</italic> species</title><author><name sortKey="Handley, Kim M" sort="Handley, Kim M" uniqKey="Handley K" first="Kim M." last="Handley">Kim M. Handley</name><affiliation><nlm:aff id="aff1"><institution>Searle Chemistry Laboratory, Computation Institute, University of Chicago</institution><country>Chicago, IL, USA</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><institution>Computing, Environment and Life Sciences, Argonne National Laboratory</institution><country>Chicago, IL, USA</country></nlm:aff></affiliation></author><author><name sortKey="Lloyd, Jonathan R" sort="Lloyd, Jonathan R" uniqKey="Lloyd J" first="Jonathan R." last="Lloyd">Jonathan R. Lloyd</name><affiliation><nlm:aff id="aff3"><institution>School of Earth, Atmospheric, and Environmental Sciences, University of Manchester</institution><country>Manchester, UK</country></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">23734151</idno><idno type="pmc">3660661</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3660661</idno><idno type="RBID">PMC:3660661</idno><idno type="doi">10.3389/fmicb.2013.00136</idno><date when="2013">2013</date><idno type="wicri:Area/Pmc/Corpus">000502</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000502</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Biogeochemical implications of the ubiquitous colonization of marine habitats and redox gradients by <italic>Marinobacter</italic> species</title><author><name sortKey="Handley, Kim M" sort="Handley, Kim M" uniqKey="Handley K" first="Kim M." last="Handley">Kim M. Handley</name><affiliation><nlm:aff id="aff1"><institution>Searle Chemistry Laboratory, Computation Institute, University of Chicago</institution><country>Chicago, IL, USA</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff2"><institution>Computing, Environment and Life Sciences, Argonne National Laboratory</institution><country>Chicago, IL, USA</country></nlm:aff></affiliation></author><author><name sortKey="Lloyd, Jonathan R" sort="Lloyd, Jonathan R" uniqKey="Lloyd J" first="Jonathan R." last="Lloyd">Jonathan R. Lloyd</name><affiliation><nlm:aff id="aff3"><institution>School of Earth, Atmospheric, and Environmental Sciences, University of Manchester</institution><country>Manchester, UK</country></nlm:aff></affiliation></author></analytic><series><title level="j">Frontiers in Microbiology</title><idno type="eISSN">1664-302X</idno><imprint><date when="2013">2013</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>The <italic>Marinobacter</italic> genus comprises widespread marine bacteria, found in localities as diverse as the deep ocean, coastal seawater and sediment, hydrothermal settings, oceanic basalt, sea-ice, sand, solar salterns, and oil fields. Terrestrial sources include saline soil and wine-barrel-decalcification wastewater. The genus was designated in 1992 for the Gram-negative, hydrocarbon-degrading bacterium <italic>Marinobacter hydrocarbonoclasticus</italic>. Since then, a further 31 type strains have been designated. Nonetheless, the metabolic range of many <italic>Marinobacter</italic> species remains largely unexplored. Most species have been classified as aerobic heterotrophs, and assessed for limited anaerobic pathways (fermentation or nitrate reduction), whereas studies of low-temperature hydrothermal sediments, basalt at oceanic spreading centers, and phytoplankton have identified species that possess a respiratory repertoire with significant biogeochemical implications. Notable physiological traits include nitrate-dependent Fe(II)-oxidation, arsenic and fumarate redox cycling, and Mn(II) oxidation. There is also evidence for Fe(III) reduction, and metal(loid) detoxification. Considering the ubiquity and metabolic capabilities of the genus, <italic>Marinobacter</italic> species may perform an important and underestimated role in the biogeochemical cycling of organics and metals in varied marine habitats, and spanning aerobic-to-anoxic redox gradients.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Aguilera, M" uniqKey="Aguilera M">M. Aguilera</name></author><author><name sortKey="Jimenez Pranteda, M L" uniqKey="Jimenez Pranteda M">M. L. Jiménez-Pranteda</name></author><author><name sortKey="Kharroub, K" uniqKey="Kharroub K">K. Kharroub</name></author><author><name sortKey="Gonzalez Paredes, A" uniqKey="Gonzalez Paredes A">A. González-Paredes</name></author><author><name sortKey="Durban, J J" uniqKey="Durban J">J. J. Durban</name></author><author><name sortKey="Russell, N J" uniqKey="Russell N">N. J. 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<front><journal-meta><journal-id journal-id-type="nlm-ta">Front Microbiol</journal-id><journal-id journal-id-type="iso-abbrev">Front Microbiol</journal-id><journal-id journal-id-type="publisher-id">Front. Microbiol.</journal-id><journal-title-group><journal-title>Frontiers in Microbiology</journal-title></journal-title-group><issn pub-type="epub">1664-302X</issn><publisher><publisher-name>Frontiers Media S.A.</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">23734151</article-id><article-id pub-id-type="pmc">3660661</article-id><article-id pub-id-type="doi">10.3389/fmicb.2013.00136</article-id><article-categories><subj-group subj-group-type="heading"><subject>Microbiology</subject><subj-group><subject>Mini Review Article</subject></subj-group></subj-group></article-categories><title-group><article-title>Biogeochemical implications of the ubiquitous colonization of marine habitats and redox gradients by <italic>Marinobacter</italic> species</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Handley</surname><given-names>Kim M.</given-names></name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><xref ref-type="aff" rid="aff2"><sup>2</sup></xref><xref ref-type="author-notes" rid="fn001"><sup>*</sup></xref></contrib><contrib contrib-type="author"><name><surname>Lloyd</surname><given-names>Jonathan R.</given-names></name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref></contrib></contrib-group><aff id="aff1"><sup>1</sup><institution>Searle Chemistry Laboratory, Computation Institute, University of Chicago</institution><country>Chicago, IL, USA</country></aff><aff id="aff2"><sup>2</sup><institution>Computing, Environment and Life Sciences, Argonne National Laboratory</institution><country>Chicago, IL, USA</country></aff><aff id="aff3"><sup>3</sup><institution>School of Earth, Atmospheric, and Environmental Sciences, University of Manchester</institution><country>Manchester, UK</country></aff><author-notes><fn fn-type="edited-by"><p>Edited by: Andreas Teske, University of North Carolina at Chapel Hill, USA</p></fn><fn fn-type="edited-by"><p>Reviewed by: Purificacion Lopez-Garcia, Centre National de la Recherche Scientifique, France; Juergen Wiegel, University of Georgia, USA</p></fn><corresp id="fn001">*Correspondence: Kim M. Handley, Searle Chemistry Laboratory, Computation Institute, University of Chicago, 5735 South Ellis Avenue, Chicago, IL 60637, USA. e-mail: <email xlink:type="simple">kmhandley@uchicago.edu</email></corresp><fn fn-type="other" id="fn002"><p>This article was submitted to Frontiers in Extreme Microbiology, a specialty of Frontiers in Microbiology.</p></fn></author-notes><pub-date pub-type="epub"><day>22</day><month>5</month><year>2013</year></pub-date><pub-date pub-type="collection"><year>2013</year></pub-date><volume>4</volume><elocation-id>136</elocation-id><history><date date-type="received"><day>15</day><month>2</month><year>2013</year></date><date date-type="accepted"><day>07</day><month>5</month><year>2013</year></date></history><permissions><copyright-statement>Copyright © 2013 Handley and Lloyd.</copyright-statement><copyright-year>2013</copyright-year><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/3.0/"><license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and subject to any copyright notices concerning any third-party graphics etc.</license-p></license></permissions><abstract><p>The <italic>Marinobacter</italic> genus comprises widespread marine bacteria, found in localities as diverse as the deep ocean, coastal seawater and sediment, hydrothermal settings, oceanic basalt, sea-ice, sand, solar salterns, and oil fields. Terrestrial sources include saline soil and wine-barrel-decalcification wastewater. The genus was designated in 1992 for the Gram-negative, hydrocarbon-degrading bacterium <italic>Marinobacter hydrocarbonoclasticus</italic>. Since then, a further 31 type strains have been designated. Nonetheless, the metabolic range of many <italic>Marinobacter</italic> species remains largely unexplored. Most species have been classified as aerobic heterotrophs, and assessed for limited anaerobic pathways (fermentation or nitrate reduction), whereas studies of low-temperature hydrothermal sediments, basalt at oceanic spreading centers, and phytoplankton have identified species that possess a respiratory repertoire with significant biogeochemical implications. Notable physiological traits include nitrate-dependent Fe(II)-oxidation, arsenic and fumarate redox cycling, and Mn(II) oxidation. There is also evidence for Fe(III) reduction, and metal(loid) detoxification. Considering the ubiquity and metabolic capabilities of the genus, <italic>Marinobacter</italic> species may perform an important and underestimated role in the biogeochemical cycling of organics and metals in varied marine habitats, and spanning aerobic-to-anoxic redox gradients.</p></abstract><kwd-group><kwd><italic>Marinobacter</italic></kwd><kwd>marine</kwd><kwd>hydrothermal</kwd><kwd>biogeochemical cycling</kwd><kwd>hydrocarbon</kwd><kwd>iron</kwd><kwd>arsenic</kwd><kwd>opportunistic</kwd></kwd-group><counts><fig-count count="1"></fig-count><table-count count="2"></table-count><equation-count count="0"></equation-count><ref-count count="105"></ref-count><page-count count="10"></page-count><word-count count="8369"></word-count></counts></article-meta></front><body><sec id="s1"><title>Introduction</title><p>Marine habitats are host to a diverse range of substrates and physicochemical regimes. Among these, hydrothermal features attract particular interest owing to ore-grade concentrations of metals, physicochemical extremes, and the presence of chemolithoautotrophic macrofauna and microbiota. Bacteria and Archaea occupying marine habitats have a substantial physical presence. There are an estimated 3.9 × 10<sup>30</sup> prokaryotic cells in the open ocean and unconsolidated marine sediments, comprising ~3.1 × 10<sup>11</sup> tonnes of carbon (Whitman et al., <xref ref-type="bibr" rid="B97">1998</xref>). This is slightly more than the estimated carbon content of terrestrial prokaryotes, and just under half of that in all plant life. Prokaryotes can contribute significantly to marine ecosystem functioning and biogeochemical cycles (Jørgensen, <xref ref-type="bibr" rid="B47">2006</xref>), owing to their prevalence and enormous capacity for transforming their environments through metabolism of organic and inorganic matter (Gadd, <xref ref-type="bibr" rid="B23">2010</xref>). Yet much of the marine microbial biomass remains unexplored, and there is still much to learn about heterotrophic and autotrophic bacterial functioning in the ocean (e.g., Moran et al., <xref ref-type="bibr" rid="B68">2004</xref>; Emerson et al., <xref ref-type="bibr" rid="B20">2010</xref>; Holden et al., <xref ref-type="bibr" rid="B43">2012</xref>).</p><p><italic>Marinobacter</italic> is a heterotrophic, and in some instances mixotrophic (Dhillon et al., <xref ref-type="bibr" rid="B14">2005</xref>; Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>,<xref ref-type="bibr" rid="B39">b</xref>), metabolically flexible genus found in an exceptionally wide range of marine and saline terrestrial settings, including various low-temperature hydrothermal environments (Table <xref ref-type="table" rid="T1">1</xref>). The genus comprises Gram-negative <italic>Gammaproteobacteria</italic> within the <italic>Alteromonadales</italic> order. All known species are motile with polar flagella (excluding <italic>M. goseongensis</italic>, Roh et al., <xref ref-type="bibr" rid="B79">2008</xref>), slightly to moderately halophilic (cf. DasSarma and Arora, <xref ref-type="bibr" rid="B13">2012</xref>), aerobic heterotrophs (Table <xref ref-type="table" rid="T2">2</xref>). However, few are confirmed strict aerobes, and several are facultative anaerobes (Table <xref ref-type="table" rid="T1">1</xref>). All are rod-shaped, with the exception of the ellipsoidal <italic>M. Segnicrescens</italic> (Guo et al., <xref ref-type="bibr" rid="B33">2007</xref>), and most are neutrophilic, except the slightly alkaliphilic <italic>M. alkaliphilus</italic>, which grows optimally at pH 8.5–9.0 (Takai et al., <xref ref-type="bibr" rid="B89">2005</xref>; also see Al-Awadhi et al., <xref ref-type="bibr" rid="B2">2007</xref>; Table <xref ref-type="table" rid="T2">2</xref>). Although most species are mesophilic, many are psychrotolerant (also known as psychrotrophic) and capable of growth down to ~4°C (Table <xref ref-type="table" rid="T2">2</xref>; Moyer and Morita, <xref ref-type="bibr" rid="B69">2007</xref>). A couple of other species are either psychrophilic (with a growth optimum near 15°C) or thermotolerant (with growth up to 50°C and an optimum of 45°C). This phenotypic versatility contributes to the ubiquity of this genus, and its ability to occupy diverse physicochemical regimes.</p><table-wrap id="T1" position="float"><label>Table 1</label><caption><p><bold><italic>Marinobacter</italic> species metabolism and isolation source</bold>.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1"><bold><italic>Marinobacter</italic> species</bold></th><th align="left" rowspan="1" colspan="1"><bold>HC</bold></th><th align="left" rowspan="1" colspan="1"><bold>CH<xref ref-type="table-fn" rid="TNb"><sup>b</sup></xref></bold></th><th align="left" rowspan="1" colspan="1"><bold>NO<sup>−</sup><sub>3</sub></bold></th><th align="left" rowspan="1" colspan="1"><bold>Gluc ferm</bold></th><th align="left" rowspan="1" colspan="1"><bold>AA</bold></th><th align="left" rowspan="1" colspan="1"><bold>Anaer</bold></th><th align="left" rowspan="1" colspan="1"><bold>Env</bold></th><th align="left" rowspan="1" colspan="1"><bold>Isolation source</bold></th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>hydrocarbonoclasticus</italic><xref ref-type="table-fn" rid="TN1"><sup>1</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Oil-polluted sediment; Gulf of Fos; Mediterranean coast; France</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>aquaeolei</italic><xref ref-type="table-fn" rid="TN2"><sup>2</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Oil-producing well-head; offshore platform; Vietnam</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>excellens</italic><xref ref-type="table-fn" rid="TN3"><sup>3</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">12°C</td><td align="left" rowspan="1" colspan="1">Radionuclide-polluted sediment; 0.5 m depth; Chazhma Bay; Sea of Japan; Russia</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>lipolyticus</italic><xref ref-type="table-fn" rid="TN4"><sup>4</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">st-aer</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Saline soil; seaside city of Cádiz; Spain</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>squalenivorans</italic><xref ref-type="table-fn" rid="TN5"><sup>5</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Oil-contaminated coastal sediment; Carteau Cove; Gulf of Fos; France</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>lutaoensis</italic><xref ref-type="table-fn" rid="TN6"><sup>6</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">st-aer</td><td align="left" rowspan="1" colspan="1">43°C</td><td align="left" rowspan="1" colspan="1">Coastal hot spring water; Lutao; Taiwan</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>litoralis</italic><xref ref-type="table-fn" rid="TN7"><sup>7</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sea water; Jungdongjin beach; East Sea; Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>flavimaris</italic><xref ref-type="table-fn" rid="TN8"><sup>8</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sea water; Daepo Beach; Yellow Sea; Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>daepoensis</italic><xref ref-type="table-fn" rid="TN8"><sup>8</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sea water; Daepo Beach; Yellow Sea; Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>bryozoorum</italic><xref ref-type="table-fn" rid="TN9"><sup>9</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sediment; Bearing Sea; Russia</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>sediminum</italic><xref ref-type="table-fn" rid="TN9"><sup>9</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sediment; Peter the Great Bay; Sea of Japan; Russia</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>maritimus</italic><xref ref-type="table-fn" rid="TN10"><sup>10</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sea water; 110 km SW of subantarctic Kerguelen islands</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>alkaliphilus</italic><xref ref-type="table-fn" rid="TN11"><sup>11</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">1.7–1.9°C</td><td align="left" rowspan="1" colspan="1">Subseafloor alkaline serpentine mud; South Chamorro Seamount; Mariana Forearc</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>algicola</italic><xref ref-type="table-fn" rid="TN12"><sup>12</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Dinoflagellate <italic>Gymnodinium catenatum</italic>; Yellow Sea; Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>koreensis</italic><xref ref-type="table-fn" rid="TN13"><sup>13</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sea-shore sand at Homi Cape; Pohang; Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>vinifirmus</italic><xref ref-type="table-fn" rid="TN14"><sup>14</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">st-aer</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Wine tank decalcification wastewater-evaporation pond. Location?</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>salsuginis</italic><xref ref-type="table-fn" rid="TN15"><sup>15</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Brine-seawater interface; Shaban Deep (a brine-filled deep); Red Sea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>gudaonensis</italic><xref ref-type="table-fn" rid="TN16"><sup>16</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TN17"><sup>17</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Oil-polluted soil underlying wastewater from the coastal Shengli Oil field; China</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>segnicrescens</italic><xref ref-type="table-fn" rid="TN17"><sup>17</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Benthic sediment; 1161 m depth; South China Sea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>salicampi</italic><xref ref-type="table-fn" rid="TN18"><sup>18</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sediment; marine solar saltern; Yellow Sea; Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>pelagius</italic><xref ref-type="table-fn" rid="TN19"><sup>19</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Coastal seawater; Zhoushan Archipelago; China</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>guineae</italic><xref ref-type="table-fn" rid="TN20"><sup>20</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Marine sediment; Deception Island; Antarctica</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>psychrophilus</italic><xref ref-type="table-fn" rid="TN21"><sup>21</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Freezing</td><td align="left" rowspan="1" colspan="1">Sea-ice; Canadian Basin; Arctic Ocean</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>mobilis; zhejiangensis</italic><xref ref-type="table-fn" rid="TN22"><sup>22</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sediment; Dayu Bay; East China Sea (Lat. 27.33, Long. 120.57)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>goseongensis</italic><xref ref-type="table-fn" rid="TN23"><sup>23</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Coastal seawater; 100 m depth; East Sea of Korea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>santoriniensis</italic><xref ref-type="table-fn" rid="TN24"><sup>24</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">resp</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">25°C</td><td align="left" rowspan="1" colspan="1">Ferruginous hydrothermal marine sediment; Santorini; Greece</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>szutsaonensis</italic><xref ref-type="table-fn" rid="TN25"><sup>25</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">16–17°C</td><td align="left" rowspan="1" colspan="1">Soil; Szutsao solar saltern; southern Taiwan</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>lacisalsi</italic><xref ref-type="table-fn" rid="TN26"><sup>26</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Water; hypersaline lake; ~50 km inland; saline-wetland; Fuente de Piedra; Spain</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>zhanjiangensis</italic><xref ref-type="table-fn" rid="TN27"><sup>27</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sea water; tidal flat, Naozhou Island; South China Sea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>oulmenensis</italic><xref ref-type="table-fn" rid="TN28"><sup>28</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Brine; salt concentrator (input material?); ~60 km inland; Ain Oulmene; Algeria.</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>daqiaonensis</italic><xref ref-type="table-fn" rid="TN29"><sup>29</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Sediment; Daqiao salt pond; Yellow Sea; east coast of China</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>adhaerens</italic><xref ref-type="table-fn" rid="TN30"><sup>30</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1"><italic>Thalassiosira weissflogii</italic> diatom aggregates; Wadden sea surface; Germany</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>antarcticus</italic><xref ref-type="table-fn" rid="TN31"><sup>31</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Intertidal sandy sediment; Larsemann Hills; Antarctica</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>xestospongiae</italic><xref ref-type="table-fn" rid="TN32"><sup>32</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNa"><sup>a</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Coastal marine sponge; 8 m depth; Obhor Sharm; Red Sea; Saudi Arabia</td></tr><tr><td align="left" rowspan="1" colspan="1">Terrestrial strain MB<xref ref-type="table-fn" rid="TN33"><sup>33</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Cyanobacterial mat; saline lake; near the Red Sea</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>manganoxydans</italic><xref ref-type="table-fn" rid="TN34"><sup>34</sup></xref></td><td align="left" rowspan="1" colspan="1">G</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Heavy metal-rich sediment; hydrothermal vent; Indian Ocean (Lat. 25.32, Long. 70.04)</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Marinobacter</italic>-like isolates<xref ref-type="table-fn" rid="TN35"><sup>35</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNc"><sup>c</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">N</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">~4°C</td><td align="left" rowspan="1" colspan="1">Weathering metal sulfide rock and sediment; Main Endeavour/Middle Valley; JdFR</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Marinobacter</italic>-like clones<xref ref-type="table-fn" rid="TN36"><sup>36</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNc"><sup>d</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">≥4°C</td><td align="left" rowspan="1" colspan="1">Metal sulfides rock and sediment; Main Endeavour/Middle Valley; JdFR</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Marinobacter</italic> clones<xref ref-type="table-fn" rid="TN37"><sup>37</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNc"><sup>e</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">4°C</td><td align="left" rowspan="1" colspan="1">Relict 50 ka metal sulfide sediment; Alvin mound; TAG; Mid-Atlantic Ridge</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Marinobacter</italic> isolates<xref ref-type="table-fn" rid="TN38"><sup>38</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNc"><sup>f</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">~2°C</td><td align="left" rowspan="1" colspan="1">Lateral hydrothermal plumes; Mothra vent field and Axial Seamount; JdFR</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>Marinobacter</italic> env/enrich<xref ref-type="table-fn" rid="TN39"><sup>39</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TNc"><sup>g</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">−0.4–−0.8°C</td><td align="left" rowspan="1" colspan="1">Fresh basalt; Arctic oceanic spreading ridges; Norwegian-Greenland Sea</td></tr></tbody></table><table-wrap-foot><fn id="TNa"><label>a</label><p><italic>Validly published species names as of April 2013</italic>.</p></fn><fn id="TNb"><label>b</label><p><italic>Carbohydrates used by species are glucose, glycerol, fructose, maltose, mannitol, sucrose, cellobiose, galactose, dextrin, sorbital, trehalose, xylose, ribose, sorbose, erythritol, inositol, dulcitol, arabinose, and N-acetyl-D-glucosamine</italic>.</p></fn><fn id="TNc"><label>c–g</label><p><italic>87–94%, 89–97%, 96–99%, 99%, 96–98% 16S rRNA gene sequence similarity to Marinobacter species, respectively</italic>.</p></fn><p><italic>Abbreviations: HC, hydrocarbon utilization; CH, carbohydrate utilization; NO<sup>−</sup><sub>3</sub>, nitrate reduction; gluc ferm, glucose fermentation; AA, amino acid metabolism; Env, environment; Y, yes; N, no; –, unknown; G, genomic evidence; resp, respiratory; st-aer, strict aerobe; env/enrich, environmental/enrichment; JdFR, Juan de Fuca Ridge; TAG, Trans-Atlantic Geotransverse hydrothermal field</italic>.</p><p><sup>1–39</sup>References:</p><fn id="TN1"><label>1</label><p><italic>Gauthier et al., <xref ref-type="bibr" rid="B25">1992</xref></italic>;</p></fn><fn id="TN2"><label>2</label><p><italic>Huu et al., <xref ref-type="bibr" rid="B45">1999</xref> and Márquez and Ventosa, <xref ref-type="bibr" rid="B61">2005</xref></italic>;</p></fn><fn id="TN3"><label>3</label><p><italic>Gorshkova et al., <xref ref-type="bibr" rid="B29">2003</xref></italic>;</p></fn><fn id="TN4"><label>4</label><p><italic>Martín et al., <xref ref-type="bibr" rid="B62">2003</xref></italic>;</p></fn><fn id="TN5"><label>5</label><p><italic>Rontani et al., <xref ref-type="bibr" rid="B82">2003</xref></italic>;</p></fn><fn id="TN6"><label>6</label><p><italic>Shieh et al., <xref ref-type="bibr" rid="B83">2003</xref> and Validation List no. 94., <xref ref-type="bibr" rid="B92">2003</xref></italic>;</p></fn><fn id="TN7"><label>7</label><p><italic>Yoon et al., <xref ref-type="bibr" rid="B102">2003</xref></italic>;</p></fn><fn id="TN8"><label>8</label><p><italic>Yoon et al., <xref ref-type="bibr" rid="B103">2004</xref></italic>;</p></fn><fn id="TN9"><label>9</label><p><italic>Romanenko et al., <xref ref-type="bibr" rid="B80">2005</xref></italic>;</p></fn><fn id="TN10"><label>10</label><p><italic>Shivaji et al., <xref ref-type="bibr" rid="B84">2005</xref></italic>;</p></fn><fn id="TN11"><label>11</label><p><italic>Takai et al., <xref ref-type="bibr" rid="B89">2005</xref></italic>;</p></fn><fn id="TN12"><label>12</label><p><italic>Green et al., <xref ref-type="bibr" rid="B30">2006</xref></italic>;</p></fn><fn id="TN13"><label>13</label><p><italic>Kim et al., <xref ref-type="bibr" rid="B54">2006</xref></italic>;</p></fn><fn id="TN14"><label>14</label><p><italic>Liebgott et al., <xref ref-type="bibr" rid="B56">2006</xref></italic>;</p></fn><fn id="TN15"><label>15</label><p><italic>Antunes et al., <xref ref-type="bibr" rid="B4">2007</xref></italic>;</p></fn><fn id="TN16"><label>16</label><p><italic>Gu et al., <xref ref-type="bibr" rid="B32">2007</xref></italic>;</p></fn><fn id="TN17"><label>17</label><p><italic>Guo et al., <xref ref-type="bibr" rid="B33">2007</xref></italic>;</p></fn><fn id="TN18"><label>18</label><p><italic>Yoon et al., <xref ref-type="bibr" rid="B101">2007</xref></italic>;</p></fn><fn id="TN19"><label>19</label><p><italic>Xu et al., <xref ref-type="bibr" rid="B100">2008</xref></italic>;</p></fn><fn id="TN20"><label>20</label><p><italic>Montes et al., <xref ref-type="bibr" rid="B67">2008</xref></italic>;</p></fn><fn id="TN21"><label>21</label><p><italic>Zhang et al., <xref ref-type="bibr" rid="B105">2008</xref></italic>;</p></fn><fn id="TN22"><label>22</label><p><italic>Huo et al., <xref ref-type="bibr" rid="B44">2008</xref></italic>;</p></fn><fn id="TN23"><label>23</label><p><italic>Roh et al., <xref ref-type="bibr" rid="B79">2008</xref></italic>;</p></fn><fn id="TN24"><label>24</label><p><italic>Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>, <xref ref-type="bibr" rid="B37">2010</xref></italic>;</p></fn><fn id="TN25"><label>25</label><p><italic>Wang et al., <xref ref-type="bibr" rid="B94">2007</xref>, <xref ref-type="bibr" rid="B95">2009</xref></italic>;</p></fn><fn id="TN26"><label>26</label><p><italic>Aguilera et al., <xref ref-type="bibr" rid="B1">2009</xref></italic>;</p></fn><fn id="TN27"><label>27</label><p><italic>Zhuang et al., <xref ref-type="bibr" rid="B104">2009</xref> and Validation List no. 148., <xref ref-type="bibr" rid="B93">2012</xref></italic>;</p></fn><fn id="TN28"><label>28</label><p><italic>Kharroub et al., <xref ref-type="bibr" rid="B53">2011</xref></italic>;</p></fn><fn id="TN29"><label>29</label><p><italic>Qu et al., <xref ref-type="bibr" rid="B75">2011</xref></italic>;</p></fn><fn id="TN30"><label>30</label><p><italic>Kaeppel et al., <xref ref-type="bibr" rid="B48">2012</xref></italic>;</p></fn><fn id="TN31"><label>31</label><p><italic>Liu et al., <xref ref-type="bibr" rid="B57">2012</xref></italic>;</p></fn><fn id="TN32"><label>32</label><p><italic>Lee et al., <xref ref-type="bibr" rid="B55">2012</xref></italic>;</p></fn><fn id="TN33"><label>33</label><p><italic>Sigalevich et al., <xref ref-type="bibr" rid="B85">2000</xref></italic>;</p></fn><fn id="TN34"><label>34</label><p><italic>Wang et al., <xref ref-type="bibr" rid="B96">2012</xref></italic>;</p></fn><fn id="TN35"><label>35</label><p><italic>Edwards et al., <xref ref-type="bibr" rid="B19">2003</xref></italic>;</p></fn><fn id="TN36"><label>36</label><p><italic>Rogers et al., <xref ref-type="bibr" rid="B78">2003</xref></italic>;</p></fn><fn id="TN37"><label>37</label><p><italic>Müller et al., <xref ref-type="bibr" rid="B70">2010</xref></italic>;</p></fn><fn id="TN38"><label>38</label><p><italic>Kaye and Baross, <xref ref-type="bibr" rid="B50">2000</xref></italic>;</p></fn><fn id="TN39"><label>39</label><p>Lysnes et al., <xref ref-type="bibr" rid="B59">2004</xref>.</p></fn></table-wrap-foot></table-wrap><table-wrap id="T2" position="float"><label>Table 2</label><caption><p><bold><italic>Marinobacter</italic> species attributes</bold>.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1"><bold><italic>Marinobacter</italic> species</bold></th><th align="left" rowspan="1" colspan="1"><bold>Halophilic</bold></th><th align="left" rowspan="1" colspan="1"><bold>Optimum salinity (%)</bold></th><th align="left" rowspan="1" colspan="1"><bold>Salinity range (%)</bold></th><th align="left" rowspan="1" colspan="1"><bold>Mesophilic</bold></th><th align="left" rowspan="1" colspan="1"><bold>Optimum temp (°C)</bold></th><th align="left" rowspan="1" colspan="1"><bold>Temp range (°C)</bold></th><th align="left" rowspan="1" colspan="1"><bold>Optimum pH</bold></th><th align="left" rowspan="1" colspan="1"><bold>Major resp. quinone (ubiquinone)</bold></th><th align="left" rowspan="1" colspan="1"><bold>G + C (%)<xref ref-type="table-fn" rid="TN40"><sup>a</sup></xref></bold></th><th align="left" rowspan="1" colspan="1"><bold>Motility mechanism<xref ref-type="table-fn" rid="TN41"><sup>b</sup></xref></bold></th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>hydrocarbonoclasticus</italic><xref ref-type="table-fn" rid="TN43"><sup>1</sup></xref><sup>,</sup><xref ref-type="table-fn" rid="TN43"><sup>2</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">3–6</td><td align="left" rowspan="1" colspan="1">0.5–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">32</td><td align="left" rowspan="1" colspan="1">10–45</td><td align="left" rowspan="1" colspan="1">7.0–7.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">(52.7) 57.3</td><td align="left" rowspan="1" colspan="1">Polar flagellum<xref ref-type="table-fn" rid="TN42"><sup>c</sup></xref></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>aquaeolei</italic><xref ref-type="table-fn" rid="TN43"><sup>2</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">0–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30</td><td align="left" rowspan="1" colspan="1">13–50</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">55.7</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>excellens</italic><xref ref-type="table-fn" rid="TN43"><sup>3</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">28</td><td align="left" rowspan="1" colspan="1">10–41</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">56</td><td align="left" rowspan="1" colspan="1">Polarly flagellated</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>lipolyticus</italic><xref ref-type="table-fn" rid="TN43"><sup>4</sup></xref></td><td align="left" rowspan="1" colspan="1">mod</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">37</td><td align="left" rowspan="1" colspan="1">15–40</td><td align="left" rowspan="1" colspan="1">7.5</td><td rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">57</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>squalenivorans</italic><xref ref-type="table-fn" rid="TN43"><sup>5</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">>0</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">32</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">54.3</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>lutaoensis</italic><xref ref-type="table-fn" rid="TN43"><sup>6</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">3–5</td><td align="left" rowspan="1" colspan="1">0.5–12</td><td align="left" rowspan="1" colspan="1">therm</td><td align="left" rowspan="1" colspan="1">45</td><td align="left" rowspan="1" colspan="1">25–50</td><td align="left" rowspan="1" colspan="1">7.0</td><td align="left" rowspan="1" colspan="1">Q8</td><td align="left" rowspan="1" colspan="1">63.5</td><td align="left" rowspan="1" colspan="1">One–several flagella</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>litoralis</italic><xref ref-type="table-fn" rid="TN43"><sup>7</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">2–7</td><td align="left" rowspan="1" colspan="1">0.5–18</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">30–37</td><td align="left" rowspan="1" colspan="1">4–46</td><td align="left" rowspan="1" colspan="1">7.0–8.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">55</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>flavimaris</italic><xref ref-type="table-fn" rid="TN43"><sup>8</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">2–6</td><td align="left" rowspan="1" colspan="1">>0–20</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">37</td><td align="left" rowspan="1" colspan="1">≤4–45</td><td align="left" rowspan="1" colspan="1">7.0–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">58</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>daepoensis</italic><xref ref-type="table-fn" rid="TN43"><sup>8</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">2–6</td><td align="left" rowspan="1" colspan="1">>0–18</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">30–37</td><td align="left" rowspan="1" colspan="1">>4–45</td><td align="left" rowspan="1" colspan="1">7.0–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">57</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>bryozoorum</italic><xref ref-type="table-fn" rid="TN43"><sup>9</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">1.0–18</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">7–42</td><td align="left" rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">59.6</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>sediminum</italic><xref ref-type="table-fn" rid="TN43"><sup>9</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">0.5–18</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">4–42</td><td align="left" rowspan="1" colspan="1">–</td><td rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">56.5</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>maritimus</italic><xref ref-type="table-fn" rid="TN43"><sup>10</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">1–13</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">4–37</td><td align="left" rowspan="1" colspan="1">8.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">58</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>alkaliphilus</italic><xref ref-type="table-fn" rid="TN43"><sup>11</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">2.5–3.5</td><td align="left" rowspan="1" colspan="1">0–21</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30–35</td><td align="left" rowspan="1" colspan="1">10–45</td><td align="left" rowspan="1" colspan="1">8.5–9.0</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">57.5</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>algicola</italic><xref ref-type="table-fn" rid="TN43"><sup>12</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">3–6</td><td align="left" rowspan="1" colspan="1">1.0–12</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">25–30</td><td align="left" rowspan="1" colspan="1">5–40</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">54–55</td><td align="left" rowspan="1" colspan="1">Polar flagellum<xref ref-type="table-fn" rid="TN42"><sup>c</sup></xref></td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>koreensis</italic><xref ref-type="table-fn" rid="TN43"><sup>13</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">3–8</td><td align="left" rowspan="1" colspan="1">0.5–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">28</td><td align="left" rowspan="1" colspan="1">10–45</td><td align="left" rowspan="1" colspan="1">6.0–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">54.1</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>vinifirmus</italic><xref ref-type="table-fn" rid="TN43"><sup>14</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">3–6</td><td align="left" rowspan="1" colspan="1">0–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">20–30</td><td align="left" rowspan="1" colspan="1">15–45</td><td align="left" rowspan="1" colspan="1">6.5–8.4</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">58.7</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>salsuginis</italic><xref ref-type="table-fn" rid="TN43"><sup>15</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">1–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">35–37</td><td align="left" rowspan="1" colspan="1">10–45</td><td align="left" rowspan="1" colspan="1">7.5–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">55.9</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>gudaonensis</italic><xref ref-type="table-fn" rid="TN43"><sup>16</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">2.0–3.0</td><td align="left" rowspan="1" colspan="1">0–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">10–45</td><td align="left" rowspan="1" colspan="1">7.5–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">57.9</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>segnicrescens</italic><xref ref-type="table-fn" rid="TN43"><sup>17</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">4–8</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30–37</td><td align="left" rowspan="1" colspan="1">15–45</td><td align="left" rowspan="1" colspan="1">7.5–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">62.2</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>salicampi</italic><xref ref-type="table-fn" rid="TN43"><sup>18</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">>0–15</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">30</td><td align="left" rowspan="1" colspan="1">4–39</td><td align="left" rowspan="1" colspan="1">7.0–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">58.1</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>pelagius</italic><xref ref-type="table-fn" rid="TN43"><sup>19</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">5.0</td><td align="left" rowspan="1" colspan="1">0.5–15</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">35–30</td><td align="left" rowspan="1" colspan="1">4–48</td><td align="left" rowspan="1" colspan="1">7.0–8.0</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">59.0</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>guineae</italic><xref ref-type="table-fn" rid="TN43"><sup>20</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">4–42</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">57.1</td><td align="left" rowspan="1" colspan="1">Polar flagella</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>psychrophilus</italic><xref ref-type="table-fn" rid="TN43"><sup>21</sup></xref></td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">2–8</td><td align="left" rowspan="1" colspan="1">psyph</td><td align="left" rowspan="1" colspan="1">16–18</td><td align="left" rowspan="1" colspan="1">0–22</td><td align="left" rowspan="1" colspan="1">6.0–9.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">55.4</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>mobilis</italic><xref ref-type="table-fn" rid="TN43"><sup>22</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">3.0–5.0</td><td align="left" rowspan="1" colspan="1">0.5–10.0</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30–35</td><td align="left" rowspan="1" colspan="1">15–42</td><td align="left" rowspan="1" colspan="1">7.0–7.5</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">58.0–58.9</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>zhejiangensis</italic><xref ref-type="table-fn" rid="TN43"><sup>22</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">1.0–3.0</td><td align="left" rowspan="1" colspan="1">0.5–10.0</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30–35</td><td align="left" rowspan="1" colspan="1">15–42</td><td align="left" rowspan="1" colspan="1">7.0–7.5</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">58.4</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>goseongensis</italic><xref ref-type="table-fn" rid="TN43"><sup>23</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">4–5</td><td align="left" rowspan="1" colspan="1">1–25</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">25–30</td><td align="left" rowspan="1" colspan="1">10–37</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>santoriniensis</italic><xref ref-type="table-fn" rid="TN43"><sup>24</sup></xref></td><td align="left" rowspan="1" colspan="1">mod</td><td align="left" rowspan="1" colspan="1">5–10</td><td align="left" rowspan="1" colspan="1">0.5–16</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">35–40</td><td align="left" rowspan="1" colspan="1">15–45</td><td align="left" rowspan="1" colspan="1">7–8</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">58.1</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>szutsaonensis</italic><xref ref-type="table-fn" rid="TN43"><sup>25</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">0–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">35–40</td><td align="left" rowspan="1" colspan="1">10–50</td><td align="left" rowspan="1" colspan="1">7.5–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">56.5</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>lacisalsi</italic><xref ref-type="table-fn" rid="TN43"><sup>26</sup></xref></td><td align="left" rowspan="1" colspan="1">mod</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">3–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30–35</td><td align="left" rowspan="1" colspan="1">20–40</td><td align="left" rowspan="1" colspan="1">7.0</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">58.6</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>zhanjiangensis</italic><xref ref-type="table-fn" rid="TN43"><sup>27</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">2–4</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">25–30</td><td align="left" rowspan="1" colspan="1">4–35</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">60.6</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>oulmenensis</italic><xref ref-type="table-fn" rid="TN43"><sup>28</sup></xref></td><td align="left" rowspan="1" colspan="1">mod</td><td align="left" rowspan="1" colspan="1">5–7.5</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">37–40</td><td align="left" rowspan="1" colspan="1">30–47</td><td align="left" rowspan="1" colspan="1">6.5–7.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">57.4</td><td align="left" rowspan="1" colspan="1">–</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>daqiaonensis</italic><xref ref-type="table-fn" rid="TN43"><sup>29</sup></xref></td><td align="left" rowspan="1" colspan="1">mod</td><td align="left" rowspan="1" colspan="1">5–10</td><td align="left" rowspan="1" colspan="1">1–15</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">30</td><td align="left" rowspan="1" colspan="1">10–45</td><td align="left" rowspan="1" colspan="1">7.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">60.8</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>adhaerens</italic><xref ref-type="table-fn" rid="TN43"><sup>30</sup></xref></td><td align="left" rowspan="1" colspan="1">s-mod</td><td align="left" rowspan="1" colspan="1">2–6</td><td align="left" rowspan="1" colspan="1">0.5–20</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">34–38</td><td align="left" rowspan="1" colspan="1">4–45</td><td align="left" rowspan="1" colspan="1">7.0–8.5</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">56.9</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>antarcticus</italic><xref ref-type="table-fn" rid="TN43"><sup>31</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">3.0–4.0</td><td align="left" rowspan="1" colspan="1">0–25</td><td align="left" rowspan="1" colspan="1">psyt</td><td align="left" rowspan="1" colspan="1">25</td><td align="left" rowspan="1" colspan="1">4–35</td><td align="left" rowspan="1" colspan="1">7.0</td><td align="left" rowspan="1" colspan="1">–</td><td align="left" rowspan="1" colspan="1">55.8</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>xestospongiae</italic><xref ref-type="table-fn" rid="TN43"><sup>32</sup></xref></td><td align="left" rowspan="1" colspan="1">slighlty</td><td align="left" rowspan="1" colspan="1">2.0</td><td align="left" rowspan="1" colspan="1">0.5–6.0</td><td align="left" rowspan="1" colspan="1">Y</td><td align="left" rowspan="1" colspan="1">28–36</td><td align="left" rowspan="1" colspan="1">15–42</td><td align="left" rowspan="1" colspan="1">7.0–8.0</td><td align="left" rowspan="1" colspan="1">Q9</td><td align="left" rowspan="1" colspan="1">57.1</td><td align="left" rowspan="1" colspan="1">Polar flagellum</td></tr></tbody></table><table-wrap-foot><fn id="TN40"><label>a</label><p><italic>GC contents range from 54.0–63.5% (average, 57.6%), using the Márquez and Ventosa (<xref ref-type="bibr" rid="B61">2005</xref>) value for hydrocarbonoclasticus</italic>.</p></fn><fn id="TN41"><label>b</label><p><italic>All species are motile, excluding M. goseongensis. M. lutaoensis also has bipolar pili. The number of flagella on M. guineae cells is unknown</italic>.</p></fn><fn id="TN42"><label>c</label><p><italic>Unsheathed flagellum</italic>.</p></fn><p>Abbreviations: Haloph, halophile; Mesoph, mesophile; temp, temperature; resp, respiratory; Y, yes; N, no; –, unknown; s-mod, slightly-moderately; psyt, psychrotolerant; psyph, psychrophile; therm, thermotolerant.</p><fn id="TN43"><label>1–32</label><p>References: refer to Table <xref ref-type="table" rid="T1">1</xref>.</p></fn></table-wrap-foot></table-wrap></sec><sec><title><italic>Marinobacter hydrocarbonoclasticus</italic>, denitrification and hydrocarbons</title><p>The genus was created for <italic>M. hydrocarbonoclasticus</italic>, which was isolated from hydrocarbon-polluted sediment, collected from the mouth of an oil refinery outlet along the French Mediterranean coast (Gauthier et al., <xref ref-type="bibr" rid="B25">1992</xref>), and includes the later heterotypic synonym, <italic>M. aquaeolei</italic> (Huu et al., <xref ref-type="bibr" rid="B45">1999</xref>; Márquez and Ventosa, <xref ref-type="bibr" rid="B61">2005</xref>). The species has an obligate requirement for sodium. It grows readily on complex organic media containing yeast extract and peptone, and aerobically on a range of organic acids (acetate, butyrate, caproate, fumarate, adipate, lactate, citrate), and the amino acids L-glutamate, L-glutamine, and L-proline. Under anaerobic conditions <italic>M. hydrocarbonoclasticus</italic> can perform denitrification using a membrane-bound respiratory NarGHI complex to reduce nitrate (Correia et al., <xref ref-type="bibr" rid="B11">2008</xref>). The nitrite formed is reduced to N<sub>2</sub><italic>via</italic> nitrite reductase cytochrome cd1 (Besson et al., <xref ref-type="bibr" rid="B8">1995</xref>), nitric oxide reductase NorBC (EMBL-EBI ABM20188.1, ABM20189.1), and nitrous oxide reductase (Prudêncio et al., <xref ref-type="bibr" rid="B74">2000</xref>). <italic>M. hydrocarbonoclasticus</italic> is most notable, however, for its ability to aerobically degrade liquid and solid, aliphatic (pristane, heneicosane, eicosane, hexadecane, tetradecane) and aromatic (phenanthrene, phenyldecane) hydrocarbons. It uses each hydrocarbon as a sole energy source, and produces large quantities of bioemulsifier. Bioemulsifiers (biosurfactants), are thought to aid in bacterial adhesion to hydrophobic surfaces, water-immiscible material breakdown, and competitor inhibition, and are attracting increasing interest for various industrial applications (Banat et al., <xref ref-type="bibr" rid="B7">2000</xref>; Nerurkar et al., <xref ref-type="bibr" rid="B71">2009</xref>; Williams, <xref ref-type="bibr" rid="B98">2009</xref>; Soberón-Chávez and Maier, <xref ref-type="bibr" rid="B88">2011</xref>).</p><p>Of the 34 species named since the genus was created, several exhibit hydrocarbonoclastic activity, while others remain untested (Table <xref ref-type="table" rid="T1">1</xref>). Additional hydrocarbons utilized by <italic>Marinobacter</italic> species include squalene, which is metabolized under denitrifying conditions (Rontani et al., <xref ref-type="bibr" rid="B82">2003</xref>), polycyclic aromatic hydrocarbons (PAHs) (Cui et al., <xref ref-type="bibr" rid="B12">2008</xref>), hexane, heptane, petroleum ether (Shivaji et al., <xref ref-type="bibr" rid="B84">2005</xref>; Antunes et al., <xref ref-type="bibr" rid="B4">2007</xref>), <italic>n</italic>-pentadecane, <italic>n</italic>-tridecane, <italic>n</italic>-undecane, <italic>n</italic>-decane, <italic>n</italic>-nonane, butane, and kerosene (Takai et al., <xref ref-type="bibr" rid="B89">2005</xref>).</p><p>This hydrocarbonoclastic capacity in <italic>Marinobacter</italic> has attracted attention owing to the potential for these bacteria to remediate crude oil contamination in environments as diverse as the Arabian Gulf (Al-Awadhi et al., <xref ref-type="bibr" rid="B2">2007</xref>) and Artic sea ice (Gerdes et al., <xref ref-type="bibr" rid="B26">2005</xref>). Nitrate reduction by <italic>Marinobacter</italic> species has also been exploited for potential use in oilfield maintenance. Dunsmore et al. (<xref ref-type="bibr" rid="B15">2006</xref>) showed reduction of added nitrate prevented deleterious growth of sulfate-reducing bacteria in produced water from a North Sea oilfield oil reservoir, controlling microbial souring reactions. The beneficial reduction of nitrate was largely attributed to indigenous <italic>Marinobacter</italic> species.</p></sec><sec><title>Expanded functional traits of the genus</title><p>Following the characterization of <italic>M. hydrocarbonoclasticus</italic>, the functional range of the genus has been further expanded to include (non-exhaustively) fermentation; the ability to respire at least 19 different carbohydrates (Table <xref ref-type="table" rid="T1">1</xref>) and several extra amino (e.g., L-alanine, D-glutamate, L-phenylalanine; Antunes et al., <xref ref-type="bibr" rid="B4">2007</xref> and Green et al., <xref ref-type="bibr" rid="B30">2006</xref>) and organic acids [e.g., malonate, formate, pyruvate, alpha-ketoglutarate; Kim et al. (<xref ref-type="bibr" rid="B54">2006</xref>) and Kharroub et al. (<xref ref-type="bibr" rid="B53">2011</xref>)]; degradation of the isoprenoid ketone 6,10,14-trimethylpentadecan-2-one (Rontani et al., <xref ref-type="bibr" rid="B81">1997</xref>); growth on ethanol (Gu et al., <xref ref-type="bibr" rid="B32">2007</xref>; Huo et al., <xref ref-type="bibr" rid="B44">2008</xref>), phenol (Liebgott et al., <xref ref-type="bibr" rid="B56">2006</xref>), and various Tweens (e.g., Takai et al., <xref ref-type="bibr" rid="B89">2005</xref>; Green et al., <xref ref-type="bibr" rid="B30">2006</xref>) following enzymatic evidence in <italic>M. hydrocarbonoclasticus</italic>; utilization of fumarate as an electron acceptor (Takai et al., <xref ref-type="bibr" rid="B89">2005</xref>; Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>); and oxidation/reduction of arsenic, iron or manganese (Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>,<xref ref-type="bibr" rid="B39">b</xref>; Wang et al., <xref ref-type="bibr" rid="B96">2012</xref>).</p><p>As for <italic>M. hydrocarbonoclasticus</italic>, all subsequently described species are able to grow aerobically on complex organic matter, and oxidize organic acids. Many, but not all are enzymatically able to reduce nitrate (Table <xref ref-type="table" rid="T1">1</xref>). Lack of fermentation by <italic>M. hydrocarbonoclasticus</italic> was initially proposed as a distinctive feature of the genus; however, a number of subsequently isolated type strains exhibit both fermentative and respiratory metabolisms, owing to their ability to ferment glucose (Table <xref ref-type="table" rid="T1">1</xref>), lactate (Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>) and other substrates (Lee et al., <xref ref-type="bibr" rid="B55">2012</xref>). Evaluation of recently available genome sequences also suggests certain <italic>Marinobacter</italic> species may exhibit enzymatic resistance to arsenic and heavy metals (e.g., Wang et al., <xref ref-type="bibr" rid="B96">2012</xref>).</p></sec><sec><title>Biogeography and phylogeny</title><p><italic>Marinobacter</italic> colonize diverse saline habitats, e.g., sea ice and hydrothermal sediments, facilitated by psychrophilic to thermotolerant physiologies, and an ability to metabolize an array of (in)organic compounds under aerobic or anaerobic conditions. However, evaluation of phylogenetic trees, constructed using 16S rRNA gene sequences, suggest the genus is monophyletic, forming a single clade distinct from other closely related epsilonproteobacterial lineages (Figure <xref ref-type="fig" rid="F1">1</xref>).</p><fig id="F1" position="float"><label>Figure 1</label><caption><p><bold>16S rRNA gene phylogenetic maximum-likelihood tree comparing <italic>Marinobacter</italic> species and their nearest neighbors within the epsilonproteobacterial orders, <italic>Alteromonadales, Pseudomonadales</italic>, and <italic>Oceanospirillales</italic>.</bold> The tree indicates the genus is monophyletic, despite the three orders being non-monophyletic (Williams et al., <xref ref-type="bibr" rid="B99">2010</xref>). The same result was obtained using the neighbor-Joining method. Trees were constructed using MEGA v5.0 (Tamura et al., <xref ref-type="bibr" rid="B90">2011</xref>), Clustal W alignments (Thompson et al., <xref ref-type="bibr" rid="B91">1994</xref>), and 1000 bootstrap replicates. Bootstrap values ≥50 are shown. Sequences used were ≥1350 bp long. <italic>Marinobacter</italic> isolates are in dark font with type species bolded, and closely related <italic>Gammaproteobacteria</italic> are in pale font. GenBank accession numbers are given in parentheses. The symbols indicate <italic>Marinobacter</italic> isolate sources.</p></caption><graphic xlink:href="fmicb-04-00136-g0001"></graphic></fig><p>Despite the physiological versatility of the genus, and the ability of some strains to grow (non-optimally) without salt (e.g., Huu et al., <xref ref-type="bibr" rid="B45">1999</xref>; Sigalevich et al., <xref ref-type="bibr" rid="B85">2000</xref>; Liebgott et al., <xref ref-type="bibr" rid="B56">2006</xref>), <italic>Marinobacter</italic> appear to be geographically restricted to marine or terrestrial environments rich in sodium salts. This observation is consistent with the hypothesis that microorganisms exhibit non-random biogeographical differentiation and distribution, due in part to environmental selection (Martiny et al., <xref ref-type="bibr" rid="B64">2006</xref>).</p><p>In marine environments, dispersal does not appear to be a limiting factor for <italic>Marinobacter</italic>. Strains have been isolated from, and phylogenetically detected in, oceans (Pacific, Atlantic, Indian, Arctic, and Antarctic) and seas, spanning the globe from pole to equator (Table <xref ref-type="table" rid="T1">1</xref>; Kaye et al., <xref ref-type="bibr" rid="B51">2011</xref>). They display both attached and planktonic lifestyles, and the distribution of the genus extends from deep-ocean (hydrothermal) benthic sediment and exposed basalt to surface water, or coastal (hydrothermal) sediment, hot spring water and sand (Table <xref ref-type="table" rid="T1">1</xref>; Figure <xref ref-type="fig" rid="F1">1</xref>).</p><p>In many instances, terrestrial isolation sources can be clearly linked to the ocean (e.g., coastal solar salterns and hot springs, and polluted soil from a coastal oil field; Table <xref ref-type="table" rid="T1">1</xref>). Isolation of species from terrestrial sources, up to 50–60 km inland, implies a greater degree of terrestrial dispersal (Table <xref ref-type="table" rid="T1">1</xref>; Figure <xref ref-type="fig" rid="F1">1</xref> and Table S2 in Kaye et al., <xref ref-type="bibr" rid="B51">2011</xref>). However, there is insufficient information regarding the nature of terrestrial isolation sources, and too few isolate and phylogenetic data, to judge how well-dispersed this genus is on land, or whether terrestrial sources are strictly independent from marine influences.</p></sec><sec><title>Lifesytles</title><p>In many respects <italic>Marinobacter</italic> species are generalists like their marine and terrestrial <italic>Alteromonadales</italic> cousins in the <italic>Shewanella</italic> genus. <italic>Shewanella</italic> species are respiratory generalists (e.g., Heidelberg et al., <xref ref-type="bibr" rid="B42">2002</xref>), and at least one species (<italic>S. baltica</italic>) has been described as “very close to the ultimate [marine] <italic>r-strategist,”</italic> starkly contrasting with genomically streamlined K-strategist (or oligotrophic) marine bacteria like <italic>Prochlorococcus</italic> (Caro-Quintero et al., <xref ref-type="bibr" rid="B10">2011</xref>). In the presence of surplus organic carbon, <italic>Marinobacter</italic> can grow rapidly, out-competing other bacteria in enrichment cultures (e.g., Handley et al., <xref ref-type="bibr" rid="B37">2010</xref>). This <italic>r</italic>-strategist (or copiotrophic) behavior renders them weed-like and relatively easy to cultivate, even compared with other heterotrophic marine bacteria (Kaye and Baross, <xref ref-type="bibr" rid="B50">2000</xref>). <italic>Marinobacter</italic> can also excel under aerobic-to-anaerobic conditions with no added substrate, while in the presence of Fe(II) (Edwards et al., <xref ref-type="bibr" rid="B19">2003</xref>; Handley et al., <xref ref-type="bibr" rid="B34">2013a</xref>).</p><p>This type of lifestyle exhibited by <italic>Marinobacter</italic> strains has been described as opportunistic or “opportunitrophic” (Singer et al., <xref ref-type="bibr" rid="B86">2011</xref>), following the definition given by Moran et al. (<xref ref-type="bibr" rid="B68">2004</xref>) in describing the ability of the marine bacterium <italic>Silicibacter pomeroyi</italic> to switch rapid between lithoheterotrophy to heterotrophy in response to nutrient pulses. Use of “-troph” in this context describes the mode of obtaining nourishment (as for the term “psychrotroph”) rather than the source of the nourishment (as in “organotroph”), and as such may be considered a misnomer. The term was applied in order to differentiate between types of fast growing and nominally <italic>r</italic>-strategist bacteria, specifically between specialists (e.g., <italic>Geobacter</italic>, Mahadevan and Lovley, <xref ref-type="bibr" rid="B60">2008</xref>), and generalists like <italic>Marinobacter, Shewanella, Pseudomonas, Vibrio</italic> and <italic>Roseobacter</italic> (Singer et al., <xref ref-type="bibr" rid="B86">2011</xref>)—with the latter two genera already having been deemed opportunistic based on their versatile lifestyles (“opportunitrophs”; Polz et al., <xref ref-type="bibr" rid="B73">2006</xref>). Singer et al. (<xref ref-type="bibr" rid="B86">2011</xref>) also identified other potential commonalities shared among the genomes of opportunistic bacteria and <italic>M. aquaeolei</italic> VT8, including a large genomic toolkit for responding to environmental stimuli and for defense (cf. Polz et al., <xref ref-type="bibr" rid="B73">2006</xref>).</p><p>There are few phylogenetic studies of the environments from which <italic>Marinobacter</italic> species have been isolated that evaluate their <italic>in situ</italic> relative abundance. Nevertheless, the studies that have been published show two different scenarios for <italic>Marinobacter</italic>. Strains may be characterized as <italic>r</italic>-strategists or opportunistic (Polz et al., <xref ref-type="bibr" rid="B73">2006</xref>; Singer et al., <xref ref-type="bibr" rid="B86">2011</xref>), and dominate communities sporadically when stimulated by high nutrient loads, encountered, for example, in marine aggregates or enrichment cultures (Balzano et al., <xref ref-type="bibr" rid="B6">2009</xref>; Handley et al., <xref ref-type="bibr" rid="B37">2010</xref>). In contrast, relatively high <italic>in situ</italic> abundances of <italic>Marinobacter</italic> (Müller et al., <xref ref-type="bibr" rid="B70">2010</xref>) and uncultured organisms closely related to <italic>Marinobacter</italic> (Rogers et al., <xref ref-type="bibr" rid="B78">2003</xref>; Edwards et al., <xref ref-type="bibr" rid="B17">2004</xref>) have been observed in some hydrothermal systems, implying these organisms may play an important and sustained role in post-depositional mineral alteration.</p></sec><sec><title>Hydrothermal settings</title><p>Marine hydrothermal systems are dispersed throughout the world's oceans (Martin et al., <xref ref-type="bibr" rid="B63">2008</xref>), and support abundant psychrophilic-to-mesophilic life even in close proximity to high-temperature venting (Reysenbach and Cady, <xref ref-type="bibr" rid="B77">2001</xref>; Edwards et al., <xref ref-type="bibr" rid="B19">2003</xref>). A number of studies suggest <italic>Marinobacter</italic> may be significant in ‘low-temperature’ hydrothermal systems, defined by low-temperature hydrothermalism (e.g., ~8 to ~40°C, McCollom and Shock, <xref ref-type="bibr" rid="B65">1997</xref>) or ambient seawater temperatures (e.g., ~2° in the deep ocean). This is due to their documented association with several different hydrothermal features (Table <xref ref-type="table" rid="T1">1</xref>), and to their ability to heterotrophically or mixotrophically respire inorganic compounds abundant in hydrothermal systems.</p><p>A common hydrothermal feature found at plate boundaries, and with which <italic>Marinobacter</italic> or near-relatives have been associated (Edwards et al., <xref ref-type="bibr" rid="B19">2003</xref>; Rogers et al., <xref ref-type="bibr" rid="B78">2003</xref>; Müller et al., <xref ref-type="bibr" rid="B70">2010</xref>), are massive sulfides, which comprise an estimated 6 × 10<sup>8</sup> tonnes of material globally (Hannington et al., <xref ref-type="bibr" rid="B40">2011</xref>), and adjunct metalliferous sediments. While much of the material for massive sulfides originates from high-temperature hydrothermal fluids (>350°C) emanating from black smoker chimneys (Hannington et al., <xref ref-type="bibr" rid="B40">2011</xref>), particulates distributed locally by plumes and talus from mound and chimney collapse can equilibrate with ambient temperatures (Edwards et al., <xref ref-type="bibr" rid="B19">2003</xref>), or entire mounds can be inactive (Müller et al., <xref ref-type="bibr" rid="B70">2010</xref>). Adjacent to massive sulfide deposits are low-temperature iron- and manganese-rich metalliferous sediments, derived from distal plume fallout with contributions from mound mass wasting (Jannasch and Mottl, <xref ref-type="bibr" rid="B46">1985</xref>; Mills et al., <xref ref-type="bibr" rid="B66">1993</xref>; Hannington et al., <xref ref-type="bibr" rid="B41">1998</xref>)—possibly of the type from which <italic>M. manganoxydans</italic> was isolated (Wang et al., <xref ref-type="bibr" rid="B96">2012</xref>).</p><p>Deposits consisting of iron oxyhydroxides, nontronite (a ferric iron-rich clay) and iron-manganese crusts can form independently at plate boundaries or at places of intra-plate volcanism (e.g., Alt, <xref ref-type="bibr" rid="B3">1988</xref>; Karl et al., <xref ref-type="bibr" rid="B49">1988</xref>; Boyd and Scott, <xref ref-type="bibr" rid="B9">2001</xref>; Kennedy et al., <xref ref-type="bibr" rid="B52">2004</xref>; Edwards et al., <xref ref-type="bibr" rid="B18">2011</xref>). They form from diffuse low-temperature venting (Karl et al., <xref ref-type="bibr" rid="B49">1988</xref>; Edwards et al., <xref ref-type="bibr" rid="B18">2011</xref>), and can span areas >100 m<sup>2</sup> (Boyd and Scott, <xref ref-type="bibr" rid="B9">2001</xref>). Similar deposits exist in shallow marine settings, such as the ferruginous arsenic-rich sediments found in Papua New Guinea and Santorini (Smith and Cronan, <xref ref-type="bibr" rid="B87">1983</xref>; Pichler and Veizer, <xref ref-type="bibr" rid="B72">1999</xref>). <italic>M. santoriniensis</italic> was isolated from the Santorini sediment (Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>).</p><p>Further examples of low-temperature hydrothermal habitats, with which <italic>Marinobacter</italic> or near relatives are associated, include those created by sharp temperature gradients that form across high-temperature chimney walls (Rogers et al., <xref ref-type="bibr" rid="B78">2003</xref>), or buoyant plumes (Kaye and Baross, <xref ref-type="bibr" rid="B50">2000</xref>) that rise 200–300 m up from these vents and spread laterally (German et al., <xref ref-type="bibr" rid="B27">1991</xref>). Exposed, iron-rich basalt, delivered by oceanic spreading centers, provides another environment associated with many deep-sea hydrothermal systems (Lysnes et al., <xref ref-type="bibr" rid="B59">2004</xref>), whereas <italic>M. alkaliphilus</italic> was isolated from alkaline serpentine mud (Takai et al., <xref ref-type="bibr" rid="B89">2005</xref>) from a setting peculiar to mud volcanoes on the non-accretionary Mariana forearc (Fryer et al., <xref ref-type="bibr" rid="B22">1999</xref>).</p></sec><sec><title>Function, biogeochemistry and hydrothermal systems</title><p>The various low-temperature hydrothermal settings <italic>Marinobacter</italic>(-like) species inhabit are rich in metals/metalloids, such as iron, manganese, arsenic, copper and zinc (Smith and Cronan, <xref ref-type="bibr" rid="B87">1983</xref>; Hannington et al., <xref ref-type="bibr" rid="B41">1998</xref>) that certain <italic>Marinobacter</italic> strains can transform enzymatically. Moreover, oxygen gradients established in these sediments may be exploited by <italic>Marinobacter</italic> species able to grow heterotrophically under anaerobic/aerobic conditions and mixotrophically under aerobic conditions.</p><p>Among the functions <italic>Marinobacter</italic> may perform in these environments is ferrous iron oxidation. The potential for <italic>Marinobacter</italic> Fe(II) oxidation was first suggested by Edwards et al. (<xref ref-type="bibr" rid="B19">2003</xref>) after isolating iron-oxidizing bacteria, phylogenetically resembling <italic>M. aquaeolei</italic>, from low-temperature hydrothermal metal sulfides. The isolates were able to grow chemoautotrophically on pyrite, basalt glass and siderite under micro-aerobic conditions. This promoted subsequent study of <italic>M. aquaeolei</italic>, including genome sequencing, and identification of its ability to anaerobically oxidize Fe(II) under mixotrophic conditions (Dhillon et al., <xref ref-type="bibr" rid="B14">2005</xref>; Edwards et al., <xref ref-type="bibr" rid="B16">2006</xref>; Singer et al., <xref ref-type="bibr" rid="B86">2011</xref>). Subsequently, <italic>M. santoriniensis</italic>, which was isolated from iron-rich hydrothermal sediment, was also shown to perform nitrate-dependent Fe(II) oxidation when supplemented with a small amount of organic carbon (Handley et al., <xref ref-type="bibr" rid="B38">2009a</xref>).</p><p>Interestingly, <italic>M. santoriniensis</italic> was isolated from sediment rife with stalk-like cells and bacteria phylogenetically resembling iron-oxidizing <italic>Zetaproteobacteria</italic> (Handley et al., <xref ref-type="bibr" rid="B37">2010</xref>). Other <italic>Marinobacter</italic> (or near relatives) were also cultivated from environments containing stalks (Edwards et al., <xref ref-type="bibr" rid="B19">2003</xref>; Lysnes et al., <xref ref-type="bibr" rid="B59">2004</xref>) that speculatively belong to this increasingly characteristic phylum of marine iron-oxidizers—the <italic>Zetaproteobacteria</italic> (Emerson et al., <xref ref-type="bibr" rid="B21">2007</xref>, <xref ref-type="bibr" rid="B20">2010</xref>; Edwards et al., <xref ref-type="bibr" rid="B18">2011</xref>).</p><p>As <italic>Marinobacter</italic> are reputedly more versatile than <italic>Mariprofundus ferrooxydans</italic> strains (the sole representatives of the <italic>Zetaproteobacteria</italic>) it is possible they perform other functions in these environments instead of, or in addition to, Fe(II) oxidation. For instance, <italic>Marinobacter</italic> and <italic>Marinobacter</italic>-like isolates have been implicated in Fe(III) reduction, but only in complex or simple co-cultures with other bacteria (Lysnes et al., <xref ref-type="bibr" rid="B59">2004</xref>; Balzano et al., <xref ref-type="bibr" rid="B6">2009</xref>; Handley et al., <xref ref-type="bibr" rid="B37">2010</xref>). This metabolic trait remains to be demonstrated in definitively anexic cultures. <italic>M. santoriniensis</italic> has the genetic potential to reductively detoxify arsenate and mercury using proteins encoded by an <italic>Escherichia coli</italic>-like <italic>arsC</italic> and <italic>merRTA</italic> genes (Handley et al., <xref ref-type="bibr" rid="B36">2013c</xref>), in addition to being able to conserve energy for growth <italic>via</italic> arsenate [As(V)] respiration using an unidentified mechanism, and mixotrophically oxidize arsenite [As(III)] using the <italic>aro</italic> gene cluster—making it one of a handful of bacteria currently known to completely redox-cycle arsenic (Handley et al., <xref ref-type="bibr" rid="B39">2009b</xref>). This is particularly relevant given that the bacterium was isolated from sediment containing ~400 ppm of arsenic.</p><p>It remains to be explored whether other <italic>Marinobacter</italic> species share this ability to respire arsenic. However, there is cursory evidence for non-respiratory arsenate reductase (plus/minus putative respiratory arsenite oxidase) genes in several publically available <italic>Marinobacter</italic> genomes (namely, <italic>M. hydrocarbonoclasticus</italic> ATCC49840, GenBank FO203363.1, Grimaud et al., <xref ref-type="bibr" rid="B31">2012</xref>; <italic>M. aquaeolei</italic> VT8, GenBank CP000514.1, Singer et al., <xref ref-type="bibr" rid="B86">2011</xref>; <italic>M. adhaerens</italic> HP15, GenBank CP001978.1, Gärdes et al., <xref ref-type="bibr" rid="B24">2010</xref>; <italic>M. algicola</italic> DG893, GenBank ABCP00000000.1; <italic>Marinobacter</italic> spp. BSs20148 and ELB17, GenBank CP003735.1 and AAXY00000000.1). Likewise, in a recent genome announcement Wang et al. (<xref ref-type="bibr" rid="B96">2012</xref>) described a <italic>Marinobacter</italic> candidate, <italic>M. manganoxydans</italic> MnI7-9 that has not only a putative <italic>arsC</italic> gene for arsenic detoxification (GenBank YP_005884959.1), but also a host of other genes that may be used for nickel, mercury, copper, chromate, zinc, cobalt, and cadmium resistance. This bacterium adsorbs and tolerates high levels of metals/metalloids, alongside a demonstrated ability to oxidize manganese, Mn(II), to a mixed-valency Mn(III)/Mn(IV) product <italic>via</italic> an unidentified genetic mechanism. Bacterial manganese oxidation is not thought to be an energy conserving process, but it is considered significant in environmental Mn(IV) oxide formation (Geszvain et al., <xref ref-type="bibr" rid="B28">2012</xref>).</p></sec><sec><title>Conclusions and future directions</title><p>Although the genus is widespread in marine settings, and dozens of cultivated representatives and several sequenced genomes exist, the functional breath of <italic>Marinobacter</italic> species remains largely unexplored. The ability to metabolize hydrocarbons and inorganic elements (e.g., iron, arsenic, manganese) has been tested in relatively few species. Information, based on cultures and isolation source characteristics, suggests species within the genus are able to contribute, for example, to the degradation of hydrocarbons in oil-polluted sediment, and the oxidation of Fe(II) in ferruginous sediment or basalt. However, we know little about the nature and magnitude of their actual function in the environment. High-throughput omics (genomics, transcriptomics, proteomics) techniques promise to expand our knowledge into the uncultivated black box that encompasses much of the microbiome, and to facilitate <italic>in situ</italic> investigations of communities (e.g., Ram et al., <xref ref-type="bibr" rid="B76">2005</xref>; Lo et al., <xref ref-type="bibr" rid="B58">2007</xref>; Baker et al., <xref ref-type="bibr" rid="B5">2012</xref>; Handley et al., <xref ref-type="bibr" rid="B35">2013b</xref>), but are limited in part by the large number of genes of unknown function. Much can still be achieved from cultivation experiments. In moving forward, a combination of omics, functional gene expression studies, isotope tracer and cultivation techniques will provide a powerful complement of tools for characterizing both the real and potential function of microorganisms in marine settings and elsewhere, and elucidating the (opportunistic?) role of <italic>Marinobacter</italic> species in environmental biogeochemical cycles.</p><sec><title>Conflict of interest statement</title><p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p></sec></sec></body><back><ack><p>We acknowledge funding support from the European Union through the BIOtransformations of TRace elements in AquatiC Systems (BIOTRACS) EST programme, Marie Curie Actions.</p></ack><ref-list><title>References</title><ref id="B1"><mixed-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Aguilera</surname><given-names>M.</given-names></name><name><surname>Jiménez-Pranteda</surname><given-names>M. 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