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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">2</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">ZooKeys</journal-title>
        <abbrev-journal-title xml:lang="en">ZK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1313-2989</issn>
      <issn pub-type="epub">1313-2970</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zookeys.738.22927</article-id>
      <article-id pub-id-type="publisher-id">22927</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Amphinomida</subject>
          <subject>Amphinomidae</subject>
          <subject>Animalia</subject>
          <subject>Annelida</subject>
          <subject>Invertebrata</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Identification key</subject>
          <subject>Systematics</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Japan</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>A new species of the rare, deep-sea polychaete genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> from the Sea of Kumano, Japan (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="phylum">Annelida</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Amphinomidae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Jimi</surname>
            <given-names>Naoto</given-names>
          </name>
          <email xlink:type="simple">beniimo7010@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0001-8586-3320</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kimura</surname>
            <given-names>Taeko</given-names>
          </name>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Ogawa</surname>
            <given-names>Akito</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kajihara</surname>
            <given-names>Hiroshi</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-6510-9355</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Department of Natural History Sciences, Graduate School of Science, Hokkaido University, Kita 10 Nishi 8 Kitaku, Sapporo, Hokkaido 060-0810, Japan</addr-line>
        <institution>Hokkaido University</institution>
        <addr-line content-type="city">Sapporo</addr-line>
        <country>Japan</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Graduate School of Bioresources, Mie University, 1577 Kurimamachiya-cho, Tsu, Mie 514-8507, Japan</addr-line>
        <institution>Mie University</institution>
        <addr-line content-type="city">Tsu</addr-line>
        <country>Japan</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Graduate School of Science, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-8654, Japan</addr-line>
        <institution>The University of Tokyo</institution>
        <addr-line content-type="city">Tokyo</addr-line>
        <country>Japan</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Naoto Jimi (<email xlink:type="simple">beniimo7010@gmail.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: C. Glasby</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2018</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>19</day>
        <month>02</month>
        <year>2018</year>
      </pub-date>
      <volume>738</volume>
      <fpage>81</fpage>
      <lpage>88</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/FFF7FFC1-E626-F501-5919-FF82FFA4FFEA">FFF7FFC1-E626-F501-5919-FF82FFA4FFEA</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/C0C7CC5E-B5B7-4CF3-8DF7-C52567C24819">C0C7CC5E-B5B7-4CF3-8DF7-C52567C24819</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/1222401">1222401</uri>
      <history>
        <date date-type="received">
          <day>12</day>
          <month>12</month>
          <year>2017</year>
        </date>
        <date date-type="accepted">
          <day>24</day>
          <month>01</month>
          <year>2018</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Naoto Jimi, Taeko Kimura, Akito Ogawa, Hiroshi Kajihara</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <self-uri content-type="zoobank" xlink:type="simple">http://zoobank.org/C0C7CC5E-B5B7-4CF3-8DF7-C52567C24819</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>A new species of amphinomid polychaete, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. n.</bold>, is described from the Sea of Kumano, Japan, from depths of 487–596 m. The species is distinguishable from its congeners by the following features: i) palps 1.8 times as long as lateral antennae; ii) branchiae do not reach to the tip of the notochaetae. This is the first record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> from Japan. A partial mitochondrial cytochrome <italic>c</italic> oxidase subunit I gene sequence from the holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. n.</bold> is provided for reliable species identification in the future.</p>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="order">Amphinomida</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>deep sea</kwd>
        <kwd>new genus record</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="class">Polychaeta</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>polychaetes</kwd>
        <kwd>taxonomy</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>JSPS KAKENHI, Research Institute of Marine Invertebrates</funding-statement>
      </funding-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0EQG">
        <title>Citation</title>
        <p>Jimi N, Kimura T, Ogawa A, Kajihara H (2018) A new species of the rare, deep-sea polychaete genus <italic>Benthoscolex</italic> from the Sea of Kumano, Japan (Annelida, Amphinomidae). ZooKeys 738: 81–88. <ext-link xlink:type="simple" ext-link-type="doi" xlink:href="10.3897/zookeys.738.22927">https://doi.org/10.3897/zookeys.738.22927</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="Introduction" id="SECID0EMH">
      <title>Introduction</title>
      <p>Marine annelids in the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Amphinomidae</tp:taxon-name-part></tp:taxon-name> are commonly known as fireworms, characterized by having defensive, dorsally-oriented, calcareous chaetae that are thought to be used to inject a venomous substance into predators (<xref ref-type="bibr" rid="B16">Verdes et al. 2017</xref>). The family consists of approximately 180 nominal species in 22 genera (<xref ref-type="bibr" rid="B3">Borda et al. 2012</xref>; <xref ref-type="bibr" rid="B2">Barroso et al. 2017</xref>; <xref ref-type="bibr" rid="B15">Sun and Li 2017</xref>), mostly distributed in shallow and tropical waters (<xref ref-type="bibr" rid="B2">Barroso et al. 2017</xref>).</p>
      <p>One genus, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> Horst, 1912, is rare and known mainly from deep-sea substrates. Previous collection records of the genus are limited to low latitude areas (<xref ref-type="bibr" rid="B7">Horst 1912</xref>; <xref ref-type="bibr" rid="B13">Monro 1937</xref>; <xref ref-type="bibr" rid="B6">Hartman 1942</xref>; <xref ref-type="bibr" rid="B5">Fauvel 1953</xref>; <xref ref-type="bibr" rid="B14">Salazar-Vallejo 1999</xref>; <xref ref-type="bibr" rid="B17">Wehe and Fiege 2002</xref>). The genus consists of two species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coecus">coecus</tp:taxon-name-part></tp:taxon-name></italic> Horst, 1912 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic> Hartman, 1942. Previous studies have reported <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coecus">coecus</tp:taxon-name-part></tp:taxon-name></italic> from the Red Sea to the SW Pacific Ocean (<xref ref-type="bibr" rid="B7">Horst 1912</xref>; <xref ref-type="bibr" rid="B13">Monro 1937</xref>; <xref ref-type="bibr" rid="B5">Fauvel 1953</xref>; <xref ref-type="bibr" rid="B17">Wehe and Fiege 2002</xref>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic> from the Caribbean Sea (<xref ref-type="bibr" rid="B6">Hartman 1942</xref>). The main diagnostic features of the genus are: 1) caruncle consisting of three ridges and 2) branchiae are absent at least in the first five chaetigers. In Japan, several amphinomids have been reported from the deep sea (e.g., <xref ref-type="bibr" rid="B8">Imajima 2001</xref>, <xref ref-type="bibr" rid="B9">2005</xref>, <xref ref-type="bibr" rid="B10">2006</xref>, <xref ref-type="bibr" rid="B11">2011</xref>), but there is no record of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> species from Japanese waters.</p>
      <p>During the research cruise No. 1722 by TRV <italic>Seisui-maru</italic>, we collected three specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic>. We describe the specimens here as a new species and provide a <abbrev xlink:title="cytochrome c oxidase subunit I" id="ABBRID0EPEAC">COI</abbrev> sequence as a DNA barcode of the species. This is the first report of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> from Japan.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0E1EAC">
      <title>Materials and methods</title>
      <p>Fresh specimens were collected by beam trawl from the Sea of Kumano, Japan (34°00.992'N to 33°55.258'N, 136°27.720'E to 136°26.650'E) from 487–596 m depth. The live specimens were fixed in 70% ethanol. After preservation, these specimens were observed with a Nikon SMZ1500 dissecting microscope and OLYMPUS BX51 compound microscope, and photographed with a Nikon D5200 digital camera. All of the material has been deposited in the <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">National Museum of Nature and Science, Tsukuba</named-content> (<bold><named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content></bold>). We followed the morphological terminology of <xref ref-type="bibr" rid="B2">Barroso et al. (2017)</xref> in the taxonomic description below.</p>
      <p>DNA extraction and sequencing for a partial region of mitochondrial cytochrome <italic>c</italic> oxidase subunit I (<abbrev xlink:title="cytochrome c oxidase subunit I " id="ABBRID0EOFAC">COI</abbrev>) gene were carried out following the method of <xref ref-type="bibr" rid="B12">Jimi and Fujiwara (2016)</xref>. The newly obtained sequence data has been deposited in the DNA Data Bank of Japan (<bold><abbrev content-type="institution" xlink:title="DNA Data Bank of Japan" id="ABBRID0EWFAC">DDBJ</abbrev></bold>).</p>
    </sec>
    <sec sec-type="Systematics" id="SECID0E3FAC">
      <title>Systematics</title>
      <sec sec-type="Family Amphinomidae Lamarck, 1818" id="SECID0EAGAC">
        <title>Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Amphinomidae</tp:taxon-name-part></tp:taxon-name> Lamarck, 1818</title>
        <p>[Japanese name: umikemushi-ka]</p>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Amphinomida</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Amphinomidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>Genus</label>
            <tp:taxon-name><object-id content-type="arpha">7E8B42D1-7E6B-5B75-ADF3-906365697E15</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>Horst, 1912</tp:taxon-authority>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0EMHAC">
            <title>Diagnosis.</title>
            <p>Body fusiform, flat. Eyes absent. Caruncle consisting of three ridges without ornamentation. Branchiae absent at least in first five chaetigers, dendritically branched. Dorsal and ventral cirri occur singly on the notopodium and neuropodium.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Amphinomida</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Amphinomidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">FBC36141-2EA8-5530-8D48-9EB0B3ADEFD5</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/BC2C42AA-5761-44F6-89AE-F1D2B7260BBC</object-id>
            </tp:taxon-name>
            <tp:taxon-status>sp. n.</tp:taxon-status>
            <xref ref-type="fig" rid="F1">Figs 1</xref>
            <xref ref-type="fig" rid="F2">, 2 [New Japanese name: Seisui-mitsu-one-umikemushi]</xref>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="material" id="SECID0ECJAC">
            <title>Material examined.</title>
            <p>Holotype: <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content>-Pol H-676, 21 mm long, 5 mm wide (without chaetae, at widest chaetiger), 29 chaetigers, female, the Sea of Kumano, 487–596 m depth, 8 November 2017, collected by NJ (left parapodium of chaetiger 15 was dissected for DNA extraction). Paratypes: <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content>-Pol P-677, two specimens, 19–28 mm long, 4–7 mm wide (without chaetae, at widest chaetiger), 29 chaetigers, female, collection data is the same as that of the holotype.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Sequence" id="SECID0ESJAC">
            <title>Sequence.</title>
            <p>LC360809, <abbrev xlink:title="cytochrome c oxidase subunit I" id="ABBRID0EYJAC">COI</abbrev> gene, 507 bp, determined from holotype.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0E3JAC">
            <title>Description.</title>
            <p>Body flat, tapered in anterior and posterior regions, whitish both in life and after fixation; pair of brown longitudinal lines on ventral middle line; no pigmentation on dorsal surface (Fig. <xref ref-type="fig" rid="F1">1A</xref>). Body surface smooth.</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.738.22927.figure1</object-id>
              <object-id content-type="zenodo_dep_id">3486316</object-id>
              <object-id content-type="arpha">891662F1-9B0F-56E3-9234-7C1EBF1D7D62</object-id>
              <label>Figure 1.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n., holotype (<named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content>-Pol H-676). <bold>A</bold> whole body, dorsal view <bold>B</bold> anterior end, dorsal view <bold>C</bold> median body, dorsal view <bold>D</bold> posterior end, dorsal view. Scale bars: 5 mm (<bold>A</bold>); 1 mm (<bold>B</bold>); 3 mm (<bold>C–D</bold>).</p>
              </caption>
              <graphic xlink:href="zookeys-738-081_article-22927__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_186771.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/186771</uri>
              </graphic>
            </fig>
            <p>Prostomium triangular; eyes absent. Pairs of lateral antennae and palps present, conical, smooth; palps 1.8 times as long as lateral antennae. Median antenna present, conical, as long as lateral antennae (Figs <xref ref-type="fig" rid="F1">1B</xref>, <xref ref-type="fig" rid="F2">2A</xref>). Caruncle consists of three longitudinal ridges, without ornamentation, extends to chaetigers 1–2 (depending on fixation), unattached in posterior part (Fig. <xref ref-type="fig" rid="F2">2A</xref>). Mouth composed of chaetigers 1–2. Pharynx eversible with black pigmentation.</p>
            <p>Parapodia biramous, notopodia and neuropodia clearly separated (Fig. <xref ref-type="fig" rid="F2">2B</xref>). Dorsal and ventral cirri occur singly on notopodium and neuropodium, conical, whitish, arising from body wall, present in all chaetigers. Branchiae present on chaetiger 6 or 7 and succeeding posterior chaetigers: anterior ones simple, conical lobes; gradually increasing in number and size posteriorly (Fig. <xref ref-type="fig" rid="F1">1B, C</xref>), branched from base; filaments digitiform, 8–10 filaments per branchia in middle body chaetigers, 15–18 filaments <!--PageBreak-->per branchia in posterior chaetigers; branchiae in posterior chaetigers differ in size between specimens, but never reaching to tip of notochaetae (Fig. <xref ref-type="fig" rid="F1">1D</xref>).</p>
            <p>Notochaetae contain three types: i) harpoon chaetae, with serrations limited to one side (Fig. <xref ref-type="fig" rid="F2">2C</xref>); ii) bifurcate chaetae with weakly serrated or non-serrated short tip (Fig. <xref ref-type="fig" rid="F2">2D</xref>); iii) bifurcate chaetae, with long serrated tip (Fig. <xref ref-type="fig" rid="F2">2E</xref>). Neurochaetae contain two types: i) bifurcate chaetae, with weakly serrated or non-serrated short tip (Fig. <xref ref-type="fig" rid="F2">2F</xref>); ii) bifurcate chaetae, with long serrated tip (Fig. <xref ref-type="fig" rid="F2">2G</xref>). Neurochaetae longer than notochaetae.</p>
            <p>Anus opening dorsally on terminal chaetiger; anal papilla absent (Fig. <xref ref-type="fig" rid="F1">1D</xref>).</p>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.738.22927.figure2</object-id>
              <object-id content-type="zenodo_dep_id">3486318</object-id>
              <object-id content-type="arpha">41D2178A-FC64-5868-BD09-1C23CD92DFED</object-id>
              <label>Figure 2.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n., holotype (<named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content>-Pol H-676). <bold>A</bold> anterior end, dorsal view <bold>B</bold> left parapodium of chaetiger 15, posterior view <bold>C</bold> harpoon notochaeta <bold>D</bold> bifurcate notochaeta with short tip <bold>E</bold> bifurcate notochaeta with long tip <bold>F</bold> bifurcate neurochaeta with short tip <bold>G</bold> bifurcate neurochaeta with long tip. Scale bars: 1 mm (<bold>A–B</bold>); 100 μm (<bold>C–G</bold>).</p>
              </caption>
              <graphic xlink:href="zookeys-738-081_article-22927__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_186772.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/186772</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0E1OAC">
            <title>Etymology.</title>
            <p>The species is named after the TRV <italic>Seisui-maru</italic>. The type specimens from the Sea of Kumano were collected by beam trawl gear of the ship. The specific name is a noun in the genitive case.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0ECPAC">
            <title>Confirmed distribution.</title>
            <p>Only known from the type locality, the Sea of Kumano, Japan, 487–596 m depth.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0EHPAC">
            <title>Remarks.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n. can be discriminated from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coecus">coecus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic> by the following features: i) palps 1.8 times as long as lateral antennae (vs. same length as lateral antennae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic>; 2.0 times as long as lateral antennae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coecus">coecus</tp:taxon-name-part></tp:taxon-name></italic>), and ii) branchiae do not reach to tip of notochaetae (vs. extending beyond tip of notochaetae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coecus">coecus</tp:taxon-name-part></tp:taxon-name></italic>; they also do not reach to tip of notochaetae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic>). In addition, the tip of the bifurcate neurochaetae is reportedly serrated in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic>, whereas it is only weakly serrated, or not serrated at all, in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n., although chaetal serration is known to be variable in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eurythoe">Eurythoe</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B1">Barroso and Paiva 2007</xref>).</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic> is reported to be endocommensal in the body cavity of the bathyal irregular sea urchin <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Heterobrissus">Heterobrissus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hystrix">hystrix</tp:taxon-name-part></tp:taxon-name></italic> (A. Agassiz, 1880) (<xref ref-type="bibr" rid="B6">Hartman 1942</xref>; <xref ref-type="bibr" rid="B4">Emson et al. 1993</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n. was collected by a beam trawl and found free living. In the same haul, 49 specimens representing five species of irregular sea urchins [<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brisaster">Brisaster</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="latifrons">latifrons</tp:taxon-name-part></tp:taxon-name></italic> (A. Agassiz, 1898) (n = 13, <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10723–10724), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brissopsis">Brissopsis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="luzonica">luzonica</tp:taxon-name-part></tp:taxon-name></italic> (Gray, 1851) (n = 6, <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10721–10722), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brissopsis">Brissopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. (n = 1, <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10727), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lovenia">Lovenia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gregalis">gregalis</tp:taxon-name-part></tp:taxon-name></italic> Alcock, 1893 (n = 22, <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10719–10720), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Schizaster">Schizaster</tp:taxon-name-part></tp:taxon-name></italic> sp. (n = 7, <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10725–10726)] were present and some were broken in the net. However, examination of body cavity in all but one specimen (used for spe<!--PageBreak-->cies identification and photography, Fig. <xref ref-type="fig" rid="F3">3</xref>) for each species revealed no commensal <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> worms (A. Ogawa pers. obs.); <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brissopsis">Brissopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. was not examined because it was represented by only one specimen. Therefore, whether the new species is also endocommensal in sea urchins or not cannot be ascertained at the moment. Future studies are required to confirm the present observations of a free-living lifestyle in the new species.</p>
            <fig id="F3" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zookeys.738.22927.figure3</object-id>
              <object-id content-type="zenodo_dep_id">3486320</object-id>
              <object-id content-type="arpha">9E995144-8EAD-5073-B953-2F563DAAA9B1</object-id>
              <label>Figure 3.</label>
              <caption>
                <p>Four of the five irregular sea urchin species that were contained in the same haul with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n., aboral view. <bold>A</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brisaster">Brisaster</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="latifrons">latifrons</tp:taxon-name-part></tp:taxon-name></italic> (A. Agassiz, 1898), <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10723 <bold>B</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Brissopsis">Brissopsis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="luzonica">luzonica</tp:taxon-name-part></tp:taxon-name></italic> (Gray, 1851), <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10721 <bold>C</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lovenia">Lovenia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gregalis">gregalis</tp:taxon-name-part></tp:taxon-name></italic> Alcock, 1893, <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10719 <bold>D</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Schizaster">Schizaster</tp:taxon-name-part></tp:taxon-name></italic> sp., <named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content> E-10725. Scale bars 1 cm.</p>
              </caption>
              <graphic xlink:href="zookeys-738-081_article-22927__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_186773.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/186773</uri>
              </graphic>
            </fig>
            <!--PageBreak-->
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="Key to species of Benthoscolex Horst, 1912" id="SECID0ECKAE">
        <title>Key to species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">Benthoscolex</tp:taxon-name-part></tp:taxon-name></italic> Horst, 1912</title>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table id="TID0EENAC" rules="all">
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1">Posterior branchiae extend beyond tip of notochaetae</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="coecus">coecus</tp:taxon-name-part></tp:taxon-name></italic> Horst, 1912</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Posterior branchiae do not reach to tip of notochaetae</td>
                <td rowspan="1" colspan="1">
                  <bold>2</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">Palps 1.8 times as long as lateral antennae</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="seisuiae">seisuiae</tp:taxon-name-part></tp:taxon-name></italic> sp. n.</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Palps as long as lateral antennae</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Benthoscolex">B.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cubanus">cubanus</tp:taxon-name-part></tp:taxon-name></italic> Hartman, 1942</bold>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We thank the captain and crew of the TRV <italic>Seisui-maru</italic>, and research members of the research cruise No. 1722 for generous help in sampling. We are also grateful to Dr. Toshihiko Fujita (<named-content content-type="dwc:institutional_code" xlink:title="National Museum of Nature and Science, Tsukuba" xlink:href="http://grbio.org/institution/national-science-museum-natural-history">NSMT</named-content>) for providing photos of sea urchins. Drs. Rômulo Barroso and Chris Glasby carefully read a previous draft and made positive recommendations to improve this final version. This study was supported by JSPS KAKENHI (No. 17J05066) and Research Institute of Marine Invertebrates (No.2016IKU-1) to NJ.</p>
    </ack>
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