Cosmopolitanism and Biogeography of the Genus Manganonema (Nematoda: Monhysterida) in the Deep Sea
Identifieur interne : 000025 ( Pmc/Corpus ); précédent : 000024; suivant : 000026Cosmopolitanism and Biogeography of the Genus Manganonema (Nematoda: Monhysterida) in the Deep Sea
Auteurs : Daniela Zeppilli ; Ann Vanreusel ; Roberto DanovaroSource :
- Animals : an Open Access Journal from MDPI [ 2076-2615 ] ; 2011.
Abstract
The deep sea comprises more than 60% of the Earth surface, and likely represents the largest reservoir of as yet undiscovered biodiversity. Nematodes are the most abundant taxon on Earth and are particularly abundant and diverse in the deep sea. Nevertheless, knowledge of their biogeography especially in the deep sea is still at its infancy. This article explores the distribution of the genus
Spatial patterns of species diversity provide information about the mechanisms that regulate biodiversity and are important for setting conservation priorities. Present knowledge of the biogeography of meiofauna in the deep sea is scarce. This investigation focuses on the distribution of the deep-sea nematode genus
Url:
DOI: 10.3390/ani1030291
PubMed: 26486501
PubMed Central: 4513462
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PMC:4513462Le document en format XML
<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Cosmopolitanism and Biogeography of the Genus <italic>Manganonema</italic> (Nematoda: Monhysterida) in the Deep Sea</title><author><name sortKey="Zeppilli, Daniela" sort="Zeppilli, Daniela" uniqKey="Zeppilli D" first="Daniela" last="Zeppilli">Daniela Zeppilli</name><affiliation><nlm:aff id="af1-animals-01-00291">Department of Marine Sciences, Polytechnic University of Marche, Via Brecce Bianche, 60131, Ancona, Italy; E-Mail:<email>d.zeppilli@univpm.it</email></nlm:aff></affiliation></author><author><name sortKey="Vanreusel, Ann" sort="Vanreusel, Ann" uniqKey="Vanreusel A" first="Ann" last="Vanreusel">Ann Vanreusel</name><affiliation><nlm:aff id="af2-animals-01-00291">Marine Biology Research Group, Ghent University, Krijgslaan, 9000 Ghent, Belgium; E-Mail:<email>ann.vanreusel@ugent.be</email></nlm:aff></affiliation></author><author><name sortKey="Danovaro, Roberto" sort="Danovaro, Roberto" uniqKey="Danovaro R" first="Roberto" last="Danovaro">Roberto Danovaro</name><affiliation><nlm:aff id="af1-animals-01-00291">Department of Marine Sciences, Polytechnic University of Marche, Via Brecce Bianche, 60131, Ancona, Italy; E-Mail:<email>d.zeppilli@univpm.it</email></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26486501</idno><idno type="pmc">4513462</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4513462</idno><idno type="RBID">PMC:4513462</idno><idno type="doi">10.3390/ani1030291</idno><date when="2011">2011</date><idno type="wicri:Area/Pmc/Corpus">000025</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Cosmopolitanism and Biogeography of the Genus <italic>Manganonema</italic> (Nematoda: Monhysterida) in the Deep Sea</title><author><name sortKey="Zeppilli, Daniela" sort="Zeppilli, Daniela" uniqKey="Zeppilli D" first="Daniela" last="Zeppilli">Daniela Zeppilli</name><affiliation><nlm:aff id="af1-animals-01-00291">Department of Marine Sciences, Polytechnic University of Marche, Via Brecce Bianche, 60131, Ancona, Italy; E-Mail:<email>d.zeppilli@univpm.it</email></nlm:aff></affiliation></author><author><name sortKey="Vanreusel, Ann" sort="Vanreusel, Ann" uniqKey="Vanreusel A" first="Ann" last="Vanreusel">Ann Vanreusel</name><affiliation><nlm:aff id="af2-animals-01-00291">Marine Biology Research Group, Ghent University, Krijgslaan, 9000 Ghent, Belgium; E-Mail:<email>ann.vanreusel@ugent.be</email></nlm:aff></affiliation></author><author><name sortKey="Danovaro, Roberto" sort="Danovaro, Roberto" uniqKey="Danovaro R" first="Roberto" last="Danovaro">Roberto Danovaro</name><affiliation><nlm:aff id="af1-animals-01-00291">Department of Marine Sciences, Polytechnic University of Marche, Via Brecce Bianche, 60131, Ancona, Italy; E-Mail:<email>d.zeppilli@univpm.it</email></nlm:aff></affiliation></author></analytic><series><title level="j">Animals : an Open Access Journal from MDPI</title><idno type="eISSN">2076-2615</idno><imprint><date when="2011">2011</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><sec><title>Simple Summary</title><p>The deep sea comprises more than 60% of the Earth surface, and likely represents the largest reservoir of as yet undiscovered biodiversity. Nematodes are the most abundant taxon on Earth and are particularly abundant and diverse in the deep sea. Nevertheless, knowledge of their biogeography especially in the deep sea is still at its infancy. This article explores the distribution of the genus <italic>Manganonema</italic> in the deep Atlantic Ocean and Mediterranean Sea providing new insights about this apparently rare deep-sea genus.</p></sec><sec><title>Abstract</title><p>Spatial patterns of species diversity provide information about the mechanisms that regulate biodiversity and are important for setting conservation priorities. Present knowledge of the biogeography of meiofauna in the deep sea is scarce. This investigation focuses on the distribution of the deep-sea nematode genus <italic>Manganonema</italic>, which is typically extremely rare in deep-sea sediment samples. Forty-four specimens of eight different species of this genus were recorded from different Atlantic and Mediterranean regions. Four out of the eight species encountered are new to science. We report here that this genus is widespread both in the Atlantic and in the Mediterranean Sea. These new findings together with literature information indicate that <italic>Manganonema</italic> is a cosmopolitan genus, inhabiting a variety of deep-sea habitats and oceans. <italic>Manganonema</italic> shows the highest diversity at water depths >4,000 m. Our data, therefore, indicate that this is preferentially an abyssal genus that is able, at the same time, to colonize specific habitats at depths shallower than 1,000 m. The analysis of the distribution of the genus <italic>Manganonema</italic> indicates the presence of large differences in dispersal strategies among different species, ranging from locally endemic to cosmopolitan. Lacking meroplanktonic larvae and having limited dispersal ability due to their small size, it has been hypothesized that nematodes have limited dispersal potential. However, the investigated deep-sea nematodes were present across different oceans covering macro-scale distances. Among the possible explanations (hydrological conditions, geographical and geological pathways, long-term processes, specific historical events), their apparent preference of colonizing highly hydrodynamic systems, could suggest that these infaunal organisms are transported by means of deep-sea benthic storms and turbidity currents over long distances.</p></sec></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Lambshead, P J D" uniqKey="Lambshead P">P.J.D. Lambshead</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Giere, O" uniqKey="Giere O">O. Giere</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Thiel, H" uniqKey="Thiel H">H. Thiel</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Heip, C" uniqKey="Heip C">C. 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Schüller</name></author><author><name sortKey="Ebbe, B" uniqKey="Ebbe B">B. Ebbe</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Scheltema, R S" uniqKey="Scheltema R">R.S. Scheltema</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Pearse, J S" uniqKey="Pearse J">J.S. Pearse</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Animals (Basel)</journal-id><journal-id journal-id-type="iso-abbrev">Animals (Basel)</journal-id><journal-id journal-id-type="publisher-id">Animals</journal-id><journal-title-group><journal-title>Animals : an Open Access Journal from MDPI</journal-title></journal-title-group><issn pub-type="epub">2076-2615</issn><publisher><publisher-name>MDPI</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">26486501</article-id><article-id pub-id-type="pmc">4513462</article-id><article-id pub-id-type="doi">10.3390/ani1030291</article-id><article-id pub-id-type="publisher-id">animals-01-00291</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Cosmopolitanism and Biogeography of the Genus <italic>Manganonema</italic> (Nematoda: Monhysterida) in the Deep Sea</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Zeppilli</surname><given-names>Daniela</given-names></name><xref ref-type="aff" rid="af1-animals-01-00291">1</xref></contrib><contrib contrib-type="author"><name><surname>Vanreusel</surname><given-names>Ann</given-names></name><xref ref-type="aff" rid="af2-animals-01-00291">2</xref></contrib><contrib contrib-type="author"><name><surname>Danovaro</surname><given-names>Roberto</given-names></name><xref ref-type="aff" rid="af1-animals-01-00291">1</xref><xref ref-type="corresp" rid="c1-animals-01-00291"><sup>*</sup></xref></contrib></contrib-group><aff id="af1-animals-01-00291"><label>1</label>Department of Marine Sciences, Polytechnic University of Marche, Via Brecce Bianche, 60131, Ancona, Italy; E-Mail:<email>d.zeppilli@univpm.it</email></aff><aff id="af2-animals-01-00291"><label>2</label>Marine Biology Research Group, Ghent University, Krijgslaan, 9000 Ghent, Belgium; E-Mail:<email>ann.vanreusel@ugent.be</email></aff><author-notes><corresp id="c1-animals-01-00291"><label>*</label>Author to whom correspondence should be addressed; E-Mail: <email>r.danovaro@univpm.it</email>; Tel.: +39-071-220-4654; Fax: +39-071-220-4650.</corresp></author-notes><pub-date pub-type="collection"><month>9</month><year>2011</year></pub-date><pub-date pub-type="epub"><day>05</day><month>9</month><year>2011</year></pub-date><volume>1</volume><issue>3</issue><fpage>291</fpage><lpage>305</lpage><history><date date-type="received"><day>20</day><month>7</month><year>2011</year></date><date date-type="rev-recd"><day>24</day><month>8</month><year>2011</year></date><date date-type="accepted"><day>1</day><month>9</month><year>2011</year></date></history><permissions><copyright-statement>© 2011 by the authors; licensee MDPI, Basel, Switzerland.</copyright-statement><copyright-year>2011</copyright-year><license><license-p><pmc-comment>CREATIVE COMMONS</pmc-comment>This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution license (<ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/3.0/">http://creativecommons.org/licenses/by/3.0/</ext-link>).</license-p></license></permissions><abstract><sec><title>Simple Summary</title><p>The deep sea comprises more than 60% of the Earth surface, and likely represents the largest reservoir of as yet undiscovered biodiversity. Nematodes are the most abundant taxon on Earth and are particularly abundant and diverse in the deep sea. Nevertheless, knowledge of their biogeography especially in the deep sea is still at its infancy. This article explores the distribution of the genus <italic>Manganonema</italic> in the deep Atlantic Ocean and Mediterranean Sea providing new insights about this apparently rare deep-sea genus.</p></sec><sec><title>Abstract</title><p>Spatial patterns of species diversity provide information about the mechanisms that regulate biodiversity and are important for setting conservation priorities. Present knowledge of the biogeography of meiofauna in the deep sea is scarce. This investigation focuses on the distribution of the deep-sea nematode genus <italic>Manganonema</italic>, which is typically extremely rare in deep-sea sediment samples. Forty-four specimens of eight different species of this genus were recorded from different Atlantic and Mediterranean regions. Four out of the eight species encountered are new to science. We report here that this genus is widespread both in the Atlantic and in the Mediterranean Sea. These new findings together with literature information indicate that <italic>Manganonema</italic> is a cosmopolitan genus, inhabiting a variety of deep-sea habitats and oceans. <italic>Manganonema</italic> shows the highest diversity at water depths >4,000 m. Our data, therefore, indicate that this is preferentially an abyssal genus that is able, at the same time, to colonize specific habitats at depths shallower than 1,000 m. The analysis of the distribution of the genus <italic>Manganonema</italic> indicates the presence of large differences in dispersal strategies among different species, ranging from locally endemic to cosmopolitan. Lacking meroplanktonic larvae and having limited dispersal ability due to their small size, it has been hypothesized that nematodes have limited dispersal potential. However, the investigated deep-sea nematodes were present across different oceans covering macro-scale distances. Among the possible explanations (hydrological conditions, geographical and geological pathways, long-term processes, specific historical events), their apparent preference of colonizing highly hydrodynamic systems, could suggest that these infaunal organisms are transported by means of deep-sea benthic storms and turbidity currents over long distances.</p></sec></abstract><kwd-group><kwd>deep sea</kwd><kwd>biogeography</kwd><kwd>cosmopolitanism</kwd><kwd>nematodes</kwd><kwd><italic>Manganonema</italic></kwd></kwd-group></article-meta></front><body><sec sec-type="intro"><label>1.</label><title>Introduction</title><p>Nematodes are the most abundant metazoans on Earth [<xref rid="b1-animals-01-00291" ref-type="bibr">1</xref>], they are ubiquitous in marine habitats [<xref rid="b2-animals-01-00291" ref-type="bibr">2</xref>] and their numerical dominance increases in the deep sea (up to >90%) with increasing water depth [<xref rid="b3-animals-01-00291" ref-type="bibr">3</xref>-<xref rid="b6-animals-01-00291" ref-type="bibr">6</xref>]. This phylum is characterized by a very high species number: more than 20,000 have been described [<xref rid="b7-animals-01-00291" ref-type="bibr">7</xref>], and, among them, about 4,000-5,000 are marine species. Meanwhile, the number of marine nematode species is evaluated from 10,000 to 20,000 species according to the most conservative estimation [<xref rid="b8-animals-01-00291" ref-type="bibr">8</xref>]. In particular, deep-sea nematode assemblages are characterized by a very high α-diversity and evenness, which is comparable with that of the tropical sub-littoral zone [<xref rid="b9-animals-01-00291" ref-type="bibr">9</xref>]. Even if the deep sea comprises about 91% of the ocean seafloor, investigations focused on the study of nematode diversity have been so far performed on a cumulative area of only 60–70 m<sup>2</sup> [<xref rid="b10-animals-01-00291" ref-type="bibr">10</xref>]. Moreover, because of the small size, the taxonomy of deep-sea nematodes is difficult [<xref rid="b11-animals-01-00291" ref-type="bibr">11</xref>] and information on their distribution remains extremely scant [<xref rid="b12-animals-01-00291" ref-type="bibr">12</xref>].</p><p>Spatial patterns of species diversity provide information about the mechanisms that regulate biodiversity [<xref rid="b13-animals-01-00291" ref-type="bibr">13</xref>,<xref rid="b14-animals-01-00291" ref-type="bibr">14</xref>] and are important for setting conservation priorities [<xref rid="b15-animals-01-00291" ref-type="bibr">15</xref>]. Nevertheless, the present knowledge of the biogeography of deep-sea meiofauna is limited [<xref rid="b16-animals-01-00291" ref-type="bibr">16</xref>,<xref rid="b17-animals-01-00291" ref-type="bibr">17</xref>]. Cosmopolitan species are known among shallow-water nematodes [<xref rid="b18-animals-01-00291" ref-type="bibr">18</xref>]. Cosmopolitanism is known also for some deep-sea nematode genera, whilst only a few genera have been reported to inhabit a single habitat [<xref rid="b19-animals-01-00291" ref-type="bibr">19</xref>]. The cosmopolitanism of most genera does not necessarily apply to all species of the genus; and several studies suggest the presence of large differences in species distribution at both local and regional scale [<xref rid="b10-animals-01-00291" ref-type="bibr">10</xref>,<xref rid="b12-animals-01-00291" ref-type="bibr">12</xref>,<xref rid="b19-animals-01-00291" ref-type="bibr">19</xref>-<xref rid="b22-animals-01-00291" ref-type="bibr">22</xref>]. Studies on deep-sea nematode species distribution over large areas demonstrate that the distribution of some species is widespread [<xref rid="b7-animals-01-00291" ref-type="bibr">7</xref>,<xref rid="b12-animals-01-00291" ref-type="bibr">12</xref>,<xref rid="b23-animals-01-00291" ref-type="bibr">23</xref>]. For example, among the nine species of the genus <italic>Dichromadora</italic> described from the Weddell Sea, seven are found at more than one location and can cover distances in excess of 2,500 km [<xref rid="b12-animals-01-00291" ref-type="bibr">12</xref>].</p><p>This study focuses on the distribution of the deep-sea nematode genus <italic>Manganonema</italic> (Bussau 1993). This genus belong to the family Monhysteridae, recognized as one of the most under-investigated deep sea taxa [<xref rid="b10-animals-01-00291" ref-type="bibr">10</xref>]. <italic>Manganonema</italic> has been recorded in very low abundances (typically <2%), at water depths >600 m in many oceans [<xref rid="b24-animals-01-00291" ref-type="bibr">24</xref>-<xref rid="b35-animals-01-00291" ref-type="bibr">35</xref>]. As previously reported for the genera <italic>Bathyeurystomina</italic> and <italic>Bathychaetosoma</italic>, the genus <italic>Manganonema</italic> is apparently restricted to the deep sea. At present, the genus <italic>Manganonema</italic> contains six described species [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>]. Different <italic>Manganonema</italic> species are typically reported from different sites, suggesting a potentially restricted distribution [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>]. The aim of the present study is to investigate large-scale biogeography of the genus <italic>Manganonema</italic> in the deep sea. Forty-four specimens of eight different species were recorded from different Atlantic and Mediterranean regions. Four of the encountered species are new to science. The geographical and bathymetrical distributions of <italic>Manganonema</italic> genus and species were analyzed in order to test the cosmopolitan character of this genus also at species level.</p></sec><sec><label>2.</label><title>Experimental Section</title><p>Specimens were collected from 20 distinct deep-sea sites from the Atlantic Ocean and the Mediterranean Sea at water depth ranging from 567 to 4,987 (<xref ref-type="table" rid="t1-animals-01-00291">Table 1</xref>). Sediment samples were collected from several oceanographic cruises in the North Atlantic at Rockall Through (R/V Pelagia, 2006), and off the Portugal coast (R/V Pelagia, 2006), in the Gulf of Guinea (R/V Pourquoi pas?, 2008), in the Eastern Atlantic and in the Western and in the Central Mediterranean Sea (R/V Urania, 2007, 2008 and 2009) and in the Eastern Mediterranean Sea (R/Vs Aegaeo and Universitatis, 2006 and 2007 respectively). Sediment samples were collected by multiple-, box- and interface-corers and preserved in buffered 4% formalin solution and stained with Rose Bengal.</p><p>Sediment samples were pre-screened through a 1,000-μm mesh net, and the organisms retained on a 20-μm mesh net. The fraction remaining on the latter sieve was re-suspended and centrifuged three times with Ludox HS40 (density 1.31 g cm<sup>3</sup>) and nematodes were mounted on slides after formalin-ethanol-glycerol treatment following the detailed protocols reported by Danovaro [<xref rid="b36-animals-01-00291" ref-type="bibr">36</xref>].</p><p>Drawings and photos were made on Leica DMLS microscope. Cobb formula, showing the distance of each character from anterior end was calculated for each specimen. Ratio a represents the body length divided by the body maximum width, ratio b represents the body length divided by the pharynx length, and ratio c represents body length divided by the tail length.</p></sec><sec><label>3.</label><title>Results and Discussion</title><sec><label>3.1.</label><title>Distribution of the Genus Manganonema</title><p>We encountered the genus <italic>Manganonema</italic> in 20 different deep-sea sites from the Atlantic Ocean and the Mediterranean Sea (<xref ref-type="table" rid="t1-animals-01-00291">Table 1</xref>). The presence of the genus <italic>Manganonema</italic> was recorded in the Atlantic Ocean (Rockall Through, Portuguese Margin, Gulf of Cadiz and Gulf of Guinea) and in the Mediterranean Sea (Western, Central and Eastern). <xref ref-type="fig" rid="f1-animals-01-00291">Figure 1</xref>(A) shows the distribution of <italic>Manganonema</italic> at the global scale, including the records available in literature. The new findings of <italic>Manganonema ssp.</italic> specimens together with literature data confirm the cosmopolitan character of this genus, which can be found in all oceans (Atlantic, Indian, Pacific, Arctic, and Antarctic) and at latitudes spanning from 70°N to 60°S. We discovered <italic>Manganonema</italic> in 16 specimens from the Mediterranean Sea (<xref ref-type="fig" rid="f1-animals-01-00291">Figure 1</xref>(B)). Only one study previously documented the presence of this genus in the Mediterranean Sea [<xref rid="b31-animals-01-00291" ref-type="bibr">31</xref>]. <italic>Manganonema</italic> was recorded in three sites of the Eastern Mediterranean Sea, in three sites of the Southern Adriatic Sea and in two sites in the Western Mediterranean Sea. This demonstrates that the genus <italic>Manganonema</italic> is widespread in the deep-sea sediments of the Mediterranean Sea, from the Eastern part of the basin to the Atlantic Ocean. In addition half of the <italic>Manganonema</italic> species identified in the present study were reported in the Mediterranean Sea, including one species new to science. This supports the hypothesis that the deep Mediterranean basin provides a variety of environmental conditions that might favor the colonization of several rare species [<xref rid="b22-animals-01-00291" ref-type="bibr">22</xref>].</p></sec><sec><label>3.2.</label><title>Bathymetric Distribution of the Genus Manganonema</title><p>Our data show that <italic>Manganonema</italic> is present at depths ranging from 567 to 4,997 m. The highest diversity (7 species) was found at a water depth >4,000 m (<xref ref-type="fig" rid="f2-animals-01-00291">Figure 2</xref>). <italic>Manganonema</italic> is considered a deep-sea genus as for <italic>Bathyeurystomina</italic> and <italic>Bathychaetosoma</italic> [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>]. However, two species encountered at water depths <1,000 m in the North-east Atlantic Ocean (Rockall Trough, 567 m) and in the Central Mediterranean Sea (Southern Adriatic Sea, between 590–824 m) indicate that this genus can be piezotolerant. The only other study that described the presence of <italic>Manganonema</italic> at shallow depths (600 m) was in the Weddell Sea [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>], this was explained by the fact that the peculiar conditions of this marine system could be compared with those of the deepest ocean, and thus offer the opportunity to this genus to colonize shallow sediments. Accordingly, here we discovered <italic>Manganonema</italic> in shallow sites that are characterized by habitats such as cold-water corals, seamounts, erosional features and different physical conditions (e.g., bottom temperature of the Mediterranean Sea and Atlantic Ocean), which share the presence of high hydrodynamics (<italic>i.e.</italic>, strong bottom currents).</p></sec><sec><label>3.3.</label><title>Habitats Inhabited by Manganonema</title><p>The genus <italic>Manganonema</italic> was previously reported from diverse deep-sea habitats, from the Arctic to the Antarctic Ocean, in reduced ecosystems (hydrothermal vents and mud volcanoes), in submarine canyons and in polymetallic nodule deposits [<xref rid="b24-animals-01-00291" ref-type="bibr">24</xref>,<xref rid="b26-animals-01-00291" ref-type="bibr">26</xref>-<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>,<xref rid="b32-animals-01-00291" ref-type="bibr">32</xref>-<xref rid="b35-animals-01-00291" ref-type="bibr">35</xref>]. Here we discovered <italic>Manganonema</italic> in cold-water coral habitats and in seamounts. Furthermore, we recorded <italic>Manganonema</italic> also in different morphologies and sedimentary features related to submarine landslides, unstable slopes or seafloor erosion (e.g., mud waves and furrows). This study reinforces the idea that the genus <italic>Manganonema</italic> is able to colonize a very different variety of deep-sea habitats.</p></sec><sec><label>3.4.</label><title>Distribution of the Manganonema Species</title><p>Some meiofaunal species (including nematodes) display a wide spatial distribution [<xref rid="b37-animals-01-00291" ref-type="bibr">37</xref>] and a trans-oceanic dispersal of free-living marine nematodes is not rare [<xref rid="b18-animals-01-00291" ref-type="bibr">18</xref>]. Most dominant deep-sea nematode genera are considered to be cosmopolitan, but at the species level little is known about the geographic distribution of their species.</p><p>The first exhaustive analysis of the <italic>Manganonema</italic> species hypothesised a restricted distribution of this genus [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>]. However, subsequent studies revealed that some species of <italic>Manganonema</italic> are not endemic [<xref rid="b35-animals-01-00291" ref-type="bibr">35</xref>]. In the present study 44 specimens of <italic>Manganonema</italic> belonging of eight species were analysed. Although a description of the new <italic>Manganonema</italic> species is beyond the scope of this work, a pictorial key is provided in <xref ref-type="fig" rid="f3-animals-01-00291">Figure 3</xref>, <xref ref-type="fig" rid="f4-animals-01-00291">Figure 4</xref> and <xref ref-type="fig" rid="f5-animals-01-00291">Figure 5</xref>. Species represented only by females and juveniles were considered as putative species (<italic>Manganonema sp.2</italic> and <italic>Manganonema sp.3</italic>).</p><p><italic>Manganonema pitilica</italic> (Fonseca, Decraemer, Vanreusel, 2006) was previously found in the Pacific Ocean and in the South-west Atlantic Ocean (<xref ref-type="fig" rid="f3-animals-01-00291">Figure 3</xref>(A,B,C)) [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>,<xref rid="b35-animals-01-00291" ref-type="bibr">35</xref>]. We found this species in the North-east Atlantic (Rockall Trough, Portuguese Margin, Gulf of Cadiz, and in Gulf of Guinea) and in the Western, Central and Eastern Mediterranean Sea at all investigated depths. <italic>M. pitilica</italic> is widely distributed and able to colonize different habitats from manganese nodule areas [<xref rid="b35-animals-01-00291" ref-type="bibr">35</xref>] to seamounts and cold-water coral ecosystems.</p><p><italic>Manganonema robustus</italic> (Fonseca, Decraemer, Vanreusel, 2006) was discovered in the North-east Atlantic Ocean (Portuguese Margin) at water depths between 2,000 and 5,000 m. This species was previously reported only in the Southwest Atlantic, off the Brazilian coasts (<xref ref-type="fig" rid="f3-animals-01-00291">Figure 3</xref>(D,E,F)) [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>].</p><p><italic>Manganonema bussuaensis</italic> (Fonseca, Decraemer, Vanreusel, 2006) was found in the North-east Atlantic Ocean (Gulf of Cadiz and Gulf of Guinea) and in Western, Central and Eastern Mediterranean Sea from 714 to 4,400 m water depth. This species was previously reported only in the North Atlantic at 2,000 m water depth (<xref ref-type="fig" rid="f3-animals-01-00291">Figure 3</xref>(G,H,I)) [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>].</p><p><italic>Manganonema media</italic> (Fonseca, Decraemer, Vanreusel, 2006) was encountered in the North-east Atlantic Ocean (Portuguese Margin, Gulf of Cadiz and Gulf of Guinea) and in Western Mediterranean Sea at water depth between 2000 and 5000 m. This species was previously reported only in the South-west Atlantic, off the Brazilian coasts and in the North-east Atlantic, at Goban Spur (<xref ref-type="fig" rid="f4-animals-01-00291">Figure 4</xref>(A,B,C)) [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>].</p><p><italic>Manganonema sp. 1</italic> was found in the North-east Atlantic Ocean (Portuguese Margin) at 2,000 and 4,000 m water depth (<xref ref-type="fig" rid="f4-animals-01-00291">Figure 4</xref>(D,E,F)).</p><p><italic>Manganonema sp. 2</italic> (putative species) was encountered in the North-east Atlantic Ocean (Portuguese Margin) at 3475 m water depth (<xref ref-type="fig" rid="f4-animals-01-00291">Figure 4</xref>(G,H,I)).</p><p><italic>Manganonema sp. 3</italic> (putative species) was reported in the North-east Atlantic Ocean (Portuguese Margin and Gulf of Guinea) at water depth between 1,700 and 5,000 m (<xref ref-type="fig" rid="f5-animals-01-00291">Figure 5</xref>(A,B,C)).</p><p><italic>Manganonema sp. 4</italic> was encountered in the Eastern Mediterranean Sea at 3000 m water depth (<xref ref-type="fig" rid="f5-animals-01-00291">Figure 5</xref>(D,E,F)).</p><p>Finally, in order to provide a complete review on biogeography and distribution of the <italic>Manganonema</italic> genus, the distribution of the remaining described species (not encountered in this study) are presented.</p><p><italic>Manganonema antarctica</italic> was reported in two different sites of the Southern Ocean: the Weddell Sea and the South Sandwich Trench at water depth between 600 and 4,000 m [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>].</p><p><italic>Manganonema microchepalum</italic> was reported in the Peru Basin, Central Pacific Ocean at 4,000 m water depth [<xref rid="b26-animals-01-00291" ref-type="bibr">26</xref>,<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>].</p><p>The results of this study suggest that <italic>Manganonema</italic> species are not isolated as previously supposed [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>]. Based on distribution patterns of single species we observed that <italic>Manganonema</italic> species can be either apparently restricted or widespread and one <italic>Manganonema</italic> species (<italic>M. pitilica</italic>) is a possible cosmopolitan species. Furthermore, at some sites, up to 3 different species were found to coexist, supporting the hypothesis of a trophic specialization and resulting in an important contribution to the local species richness [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>].</p></sec><sec><label>3.5.</label><title>Dispersal Mechanisms of Deep-Sea Nematodes</title><p>Some <italic>Manganonema</italic> species are apparently confined to specific areas, while others have a much wider spatial distribution. These contrasting dispersal strategies may be linked to their behaviour or life cycle. Adult nematodes are expected to have limited dispersal capabilities and hence a reduced gene flow, making restricted species distributions more plausible [<xref rid="b38-animals-01-00291" ref-type="bibr">38</xref>]. However some deep-sea nematodes have been demonstrated to have important dispersal capabilities [<xref rid="b7-animals-01-00291" ref-type="bibr">7</xref>]. The continental and cosmopolitan distributions of many meiofaunal taxa appear to result from a variety of dispersive mechanisms. Several geological, geographical and hydrodynamic pathways (<italic>i.e.</italic>, passive erosive suspension, active emergence, seamounts, sediment transports) and natural/anthropogenic rafting should be considered [<xref rid="b2-animals-01-00291" ref-type="bibr">2</xref>]. Adult benthic organisms can be transported also within moving sediment, for example benthic storms, submarine landslides and turbidity currents can carry large amounts of sediment over long distances [<xref rid="b39-animals-01-00291" ref-type="bibr">39</xref>,<xref rid="b40-animals-01-00291" ref-type="bibr">40</xref>], with an additional effect of defaunating large areas that are then free for re-colonisation. In addition to the adult macrofaunal organisms, larvae and buoyant eggs can stay in the water column for a long time, extending their potential dispersal [<xref rid="b41-animals-01-00291" ref-type="bibr">41</xref>]. The lack of meroplanktonic larvae or buoyant eggs does not allow nematodes to use these strategies for their dispersal, but sediment resuspension due to bottom currents or benthic storms can certainly contribute to the dispersal of the nematodes. This could apply particularly to the genus <italic>Manganonema</italic> that apparently prefers habitats characterized by high hydrodynamic conditions.</p></sec></sec><sec sec-type="conclusions"><label>4.</label><title>Conclusions</title><p><italic>Manganonema</italic> is a cosmopolitan genus, inhabiting all oceans (Atlantic, Indian, Pacific, Arctic, and Antarctic) and spanning across a large variety of deep-sea habitats. This genus can be found at latitudes ranging from 70°N to 60°S. We report here that the genus <italic>Manganonema</italic> is widespread also in both basins of the Mediterranean Sea. Fifty percent of the species encountered were present also in the Mediterranean Sea, and one of these is new to science. <italic>Manganonema</italic> is a deep-sea genus characterized by the highest diversity at depths >4,000 m. However, this genus is able to colonize also sediments at a water depth <1,000 m, where peculiar environmental conditions exist, for example the presence of strong near bottom currents.</p><p>Finally, the results of this study show that <italic>Manganonema</italic> showed remarkable differences among species, spanning the entire range between endemic to cosmopolitan. This genus shows the co-presence of several species, indicating a potential local diversity. Moreover, half of the species encountered are new to science, revealing the potential and hidden biodiversity of deep-sea nematodes. In spite of the evidence that adult nematodes have limited dispersal capabilities, the deep-sea nematodes revealed a high dispersion. A possible explanation is that infaunal organisms, and thus also nematodes, can be transported along with surface sediments by turbidity currents and benthic storms over long distances.</p></sec></body><back><ack><title>Acknowledgements</title><p>This research was supported by a research grant provided by “Fondazione Luigi e Francesca Brusarosco”, in the framework of the projects funded by the European Commission HERMES (Hot Spot Ecosystem Research on the Margins of European Seas FPVI, Contract No. GOCE-CT-2005-511234-1) and HERMIONE (Hotspot Ecosystem Research and Man's Impact on European Seas FP7-ENV-2008-1, Contract No. 226354), the project funded by ESF BIOFUN (BIOdiversity and Ecosystem FUNctioning Contrasting Southern European Deep-Sea Environments), and the Italian project OBAMA (COFIN-PRIN). The authors are indebted to Marianna Mea and Cristina Gambi (DISMAR, Italy) for providing specimens of <italic>Manganonema</italic>, to Michele Voisset and Antonio Cattaneo (cruise ERIG3D, IFREMER, France) for support in the sampling activities, and to the crews of the R/Vs Urania and Universitatis (Italy), Pelagia (The Netherlands), Aegeao (Greece) and Pourquoi pas? (France) for their valuable help during the sea-going activities.</p></ack><notes><title>Conflict of Interest</title><p>The authors declare no conflict of interest.</p></notes><ref-list><title>References</title><ref id="b1-animals-01-00291"><label>1.</label><element-citation publication-type="book"><person-group person-group-type="author"><name><surname>Lambshead</surname><given-names>P.J.D.</given-names></name></person-group><article-title>Marine nematode biodiversity</article-title><source>Nematode Morphology, Physiology and Ecology</source><person-group person-group-type="editor"><name><surname>Chen</surname><given-names>Z.X.</given-names></name><name><surname>Chen</surname><given-names>S.Y.</given-names></name><name><surname>Dickson</surname><given-names>D.W.</given-names></name></person-group><publisher-name>University Press</publisher-name><publisher-loc>Beijing, China</publisher-loc><year>2004</year><volume>1</volume><fpage>438</fpage><lpage>492</lpage></element-citation></ref><ref id="b2-animals-01-00291"><label>2.</label><element-citation publication-type="book"><person-group person-group-type="author"><name><surname>Giere</surname><given-names>O.</given-names></name></person-group><source>Meiobenthology. 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Red circles indicate locations of <italic>Manganonema</italic> records in the present study, green circles indicate <italic>Manganonema</italic> records available in literature (Antarctic Ocean [<xref rid="b28-animals-01-00291" ref-type="bibr">28</xref>,<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>], Arctic Ocean [<xref rid="b33-animals-01-00291" ref-type="bibr">33</xref>], Atlantic Ocean [<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>,<xref rid="b30-animals-01-00291" ref-type="bibr">30</xref>,<xref rid="b32-animals-01-00291" ref-type="bibr">32</xref>], Barents Sea [<xref rid="b34-animals-01-00291" ref-type="bibr">34</xref>], Pacific Ocean [<xref rid="b24-animals-01-00291" ref-type="bibr">24</xref>,<xref rid="b26-animals-01-00291" ref-type="bibr">26</xref>,<xref rid="b27-animals-01-00291" ref-type="bibr">27</xref>,<xref rid="b29-animals-01-00291" ref-type="bibr">29</xref>,<xref rid="b35-animals-01-00291" ref-type="bibr">35</xref>], Mediterranean Sea [<xref rid="b31-animals-01-00291" ref-type="bibr">31</xref>], Weddell Sea [<xref rid="b25-animals-01-00291" ref-type="bibr">25</xref>]).</p></caption><graphic xlink:href="animals-01-00291-g001"></graphic></fig><fig id="f2-animals-01-00291" position="float"><label>Figure 2</label><caption><p>Bathymetric species richness of the genus <italic>Manganonema</italic>.</p></caption><graphic xlink:href="animals-01-00291-g002"></graphic></fig><fig id="f3-animals-01-00291" position="float"><label>Figure 3</label><caption><p>Pictorial key for <italic>Manganonema pitilica</italic> (<bold>A</bold>–<bold>C</bold><italic>), Manganonema robustus</italic> (<bold>D</bold>–<bold>F</bold>) and <italic>Manganonema bussuaensis</italic> (<bold>G</bold>–<bold>I</bold>) with the representation of the anterior and posterior ends, geographical distribution (red circles indicate records in the present study, green circles indicate records available in literature) and short diagnosis of each species. Reported are ratio a = body length divided by maximum body diameter, b = body length divided by pharyngeal length, c = body length divided by tail length, c′ = tail length divided by anal body diameter, cbd = corresponding body diameter.</p></caption><graphic xlink:href="animals-01-00291-g003"></graphic></fig><fig id="f4-animals-01-00291" position="float"><label>Figure 4</label><caption><p>Pictorial key for <italic>Manganonema media</italic> (<bold>A</bold>–<bold>C</bold>), <italic>Manganonema sp. 1</italic> (<bold>D</bold>–<bold>F</bold>) and <italic>Manganonema sp. 2</italic> (<italic>putative species</italic>) (<bold>G</bold>–<bold>I</bold>) with the representation of the anterior and posterior ends, geographical distribution (red circles indicate records in the present study, green circles indicate records available in literature) and short diagnosis of each species. Reported are ratio a = body length divided by maximum body diameter, b = body length divided by pharyngeal length, c = body length divided by tail length, c′ = tail length divided by anal body diameter, cbd = corresponding body diameter.</p></caption><graphic xlink:href="animals-01-00291-g004"></graphic></fig><fig id="f5-animals-01-00291" position="float"><label>Figure 5</label><caption><p>Pictorial key for <italic>Manganonema sp. 3</italic> (<italic>putative species</italic>) (<bold>A</bold>–<bold>C</bold>) and <italic>Manganonema sp. 4</italic> (<bold>D</bold>–<bold>F</bold>) with the representation of the anterior and posterior ends, geographical distribution (red circles indicate records in the present study, green circles indicate records available in literature) and short diagnosis of each species. Reported are ratio a = body length divided by maximum body diameter, b = body length divided by pharyngeal length, c = body length divided by tail length, c′ = tail length divided by anal body diameter, cbd = corresponding body diameter.</p></caption><graphic xlink:href="animals-01-00291-g005"></graphic></fig><table-wrap id="t1-animals-01-00291" position="float"><label>Table 1</label><caption><p>List of the sampling sites. Reported are latitude longitude, water depth, region, habitat and number of investigated <italic>Manganonema</italic> specimens for each station.</p></caption><table frame="box" rules="all"><thead><tr><th align="center" valign="middle" rowspan="1" colspan="1"><bold>Station</bold></th><th align="center" valign="middle" rowspan="1" colspan="1"><bold>Latitude</bold></th><th align="center" valign="middle" rowspan="1" colspan="1"><bold>Longitude</bold></th><th align="center" valign="middle" rowspan="1" colspan="1"><bold>Water depth (m)</bold></th><th align="center" valign="middle" rowspan="1" colspan="1"><bold>Region</bold></th><th align="center" valign="middle" rowspan="1" colspan="1"><bold>Habitat</bold></th><th align="center" valign="middle" rowspan="1" colspan="1"><bold><italic>Manganonema</italic> specimens</bold></th></tr></thead><tbody><tr><td align="center" valign="bottom" rowspan="1" colspan="1">1</td><td align="center" valign="bottom" rowspan="1" colspan="1">55°29.71′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">15°48.56′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">567</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Rockall Through)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Cold-water corals</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">2</td><td align="center" valign="bottom" rowspan="1" colspan="1">41°52.54′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">17°00.47′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">590</td><td align="center" valign="bottom" rowspan="1" colspan="1">Central Mediterranean Sea (Southern Adriatic Sea)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Erosional structure (mud wave)</td><td align="center" valign="bottom" rowspan="1" colspan="1">2</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">3</td><td align="center" valign="bottom" rowspan="1" colspan="1">41°47.34′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">17°01.85′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">714</td><td align="center" valign="bottom" rowspan="1" colspan="1">Central Mediterranean Sea (Southern Adriatic Sea)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Erosional structure (furrow)</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">4</td><td align="center" valign="bottom" rowspan="1" colspan="1">41°31.88′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">17°25.15′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">824</td><td align="center" valign="bottom" rowspan="1" colspan="1">Central Mediterranean Sea (Southern Adriatic Sea)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Dauno seamount</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">5</td><td align="center" valign="bottom" rowspan="1" colspan="1">02°57.26′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">06°49.51′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">1,671</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Gulf of Guinea)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">3</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">6</td><td align="center" valign="bottom" rowspan="1" colspan="1">02°56.87′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">06°49.44′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">1,701</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Gulf of Guinea)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Submarine landslides</td><td align="center" valign="bottom" rowspan="1" colspan="1">3</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">7</td><td align="center" valign="bottom" rowspan="1" colspan="1">34°40.26′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">24°07.66′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">1,998</td><td align="center" valign="bottom" rowspan="1" colspan="1">Eastern Mediterranean Sea</td><td align="center" valign="bottom" rowspan="1" colspan="1">Unstable continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">8</td><td align="center" valign="bottom" rowspan="1" colspan="1">37°50.00′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">09°45.00′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">2,130</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Portuguese Margin)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">2</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">9</td><td align="center" valign="bottom" rowspan="1" colspan="1">36°47.38′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">00°29.06′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">2,689</td><td align="center" valign="bottom" rowspan="1" colspan="1">Western Mediterranean Sea</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">2</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">10</td><td align="center" valign="bottom" rowspan="1" colspan="1">35°00.34′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">08°16.27′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">2,788</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Gulf of Cadiz)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">6</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">11</td><td align="center" valign="bottom" rowspan="1" colspan="1">39°18.80′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">06°04.25′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">2,855</td><td align="center" valign="bottom" rowspan="1" colspan="1">Western Mediterranean Sea</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">12</td><td align="center" valign="bottom" rowspan="1" colspan="1">35°08.33′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">20°50.88′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">3,000</td><td align="center" valign="bottom" rowspan="1" colspan="1">Eastern Mediterranean Sea</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">6</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">13</td><td align="center" valign="bottom" rowspan="1" colspan="1">35°57.64′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">28°17.63′E</td><td align="center" valign="bottom" rowspan="1" colspan="1">3,009</td><td align="center" valign="bottom" rowspan="1" colspan="1">Eastern Mediterranean Sea</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">2</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">14</td><td align="center" valign="bottom" rowspan="1" colspan="1">40°10.00′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">09°59.99′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">3,475</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Portuguese Margin)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">15</td><td align="center" valign="bottom" rowspan="1" colspan="1">39°13.74′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">10°59.00′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">4,060</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Portuguese Margin)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Unstable continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">3</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">16</td><td align="center" valign="bottom" rowspan="1" colspan="1">34°37.49′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">09°17,00′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">4,335</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Gulf of Cadiz)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">17</td><td align="center" valign="bottom" rowspan="1" colspan="1">34°32.75′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">09°45.38′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">4,381</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Gulf of Cadiz)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">1</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">18</td><td align="center" valign="bottom" rowspan="1" colspan="1">34°08.02′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">09°33.28′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">4,385</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Gulf of Cadiz)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">3</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">19</td><td align="center" valign="bottom" rowspan="1" colspan="1">40°10.01′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">10°59.99′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">4,902</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Portuguese Margin)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open continental slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">2</td></tr><tr><td align="center" valign="bottom" rowspan="1" colspan="1">20</td><td align="center" valign="bottom" rowspan="1" colspan="1">37°50.01′N</td><td align="center" valign="bottom" rowspan="1" colspan="1">11°00.01′W</td><td align="center" valign="bottom" rowspan="1" colspan="1">4,987</td><td align="center" valign="bottom" rowspan="1" colspan="1">North-east Atlantic Ocean (Portuguese Margin)</td><td align="center" valign="bottom" rowspan="1" colspan="1">Open Continental Slope</td><td align="center" valign="bottom" rowspan="1" colspan="1">2</td></tr></tbody></table></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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