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<document id="9C16CC2E4C852DB8E6620BE7DC337E79" ID-CLB-Dataset="29396" ID-DOI="10.11646/zootaxa.4462.3.10" ID-GBIF-Dataset="ea5e9a37-550b-41c5-be6e-c71c42945fe8" ID-ISSN="1175-5326" ID-Zenodo-Dep="1441641" ID-ZooBank="A3738754-862C-4AFD-9216-1C26D8344D1D" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1538404070486" checkinUser="plazi" docAuthor="Li, Ya-Nan, Xiao, Ning, Zhang, Lv-Ping &amp; Zhang, Haibin" docDate="2018" docId="03CD673871082E0BCBEBF9DE4768FB56" docLanguage="en" docName="zootaxa.4462.3.10.pdf" docOrigin="Zootaxa 4462 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Benthodytes marianensis Li &amp; Xiao &amp; Zhang &amp; Zhang 2018, sp. nov." docType="treatment" docVersion="7" lastPageNumber="449" masterDocId="FFF41F4071092E0DCB7CFF8F422EFFBE" masterDocTitle="Benthodytes marianensis, a new species of abyssal elasipodid sea cucumbers (Elasipodida: Psychropotidae) from the Mariana Trench area" masterLastPageNumber="450" masterPageNumber="443" pageNumber="444" updateTime="1698561095046" updateUser="ExternalLinkService">
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<mods:title id="392479DBD5C365B00575316318215716">Benthodytes marianensis, a new species of abyssal elasipodid sea cucumbers (Elasipodida: Psychropotidae) from the Mariana Trench area</mods:title>
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<mods:namePart id="6B0557E5EBDA609730707FFEA5A0FBB6">Li, Ya-Nan</mods:namePart>
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<mods:namePart id="1CD159A2D74AD6D2D0EBE6C92E4410EE">Xiao, Ning</mods:namePart>
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<mods:namePart id="FE7D89749678D0606381DC94E691A3D3">Zhang, Lv-Ping</mods:namePart>
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<mods:namePart id="0EC96543BF8393E77C5198A669B8DC56">Zhang, Haibin</mods:namePart>
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<mods:title id="B2E2ADE4D90089E86F4B3640A3F1A19D">Zootaxa</mods:title>
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<mods:date id="BBCB4ADFE0E77EF3CE975EF7FD7FF223">2018</mods:date>
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<mods:number id="EB148F7997B5A0EFE579C11DF1D6CF50">2018-08-28</mods:number>
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<mods:identifier id="A80B327AEDAC472FBCA43CD4F320C47E" type="DOI">10.11646/zootaxa.4462.3.10</mods:identifier>
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<treatment id="03CD673871082E0BCBEBF9DE4768FB56" ID-DOI="http://doi.org/10.5281/zenodo.5975694" ID-GBIF-Taxon="148424970" ID-Zenodo-Dep="5975694" LSID="urn:lsid:plazi:treatment:03CD673871082E0BCBEBF9DE4768FB56" httpUri="http://treatment.plazi.org/id/03CD673871082E0BCBEBF9DE4768FB56" lastPageId="6" lastPageNumber="449" pageId="1" pageNumber="444">
<subSubSection id="C37E85A571082E0CCBEBF9DE4300F932" pageId="1" pageNumber="444" type="nomenclature">
<paragraph id="8BDBD62E71082E0CCBEBF9DE4002F9D5" blockId="1.[151,556,1617,1677]" box="[151,556,1617,1643]" pageId="1" pageNumber="444">
<heading id="D093614271082E0CCBEBF9DE4002F9D5" bold="true" box="[151,556,1617,1643]" fontSize="11" level="1" pageId="1" pageNumber="444" reason="1">
<emphasis id="B9100A3C71082E0CCBEBF9DE4002F9D5" bold="true" box="[151,556,1617,1643]" pageId="1" pageNumber="444">
<taxonomicName id="4C64ADAD71082E0CCBEBF9DE43E8F9D5" ID-CoL="68J8C" authority="Li &amp; Xiao &amp; Zhang &amp; Zhang, 2018" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[151,454,1617,1643]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="1" pageNumber="444" phylum="Echinodermata" rank="species" species="marianensis" status="sp. nov.">
<emphasis id="B9100A3C71082E0CCBEBF9DE43E8F9D5" bold="true" box="[151,454,1617,1643]" italics="true" pageId="1" pageNumber="444">Benthodytes marianensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A223B74771082E0CCAB2F9DE4002F9D5" box="[462,556,1617,1643]" pageId="1" pageNumber="444" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BDBD62E71082E0CCBEBF9FB4300F932" blockId="1.[151,556,1617,1677]" box="[151,302,1652,1677]" pageId="1" pageNumber="444">
(
<figureCitation id="135FCAAB71082E0CCBE2F9FB430BF932" box="[158,293,1652,1677]" captionStart-0="FIGURE 1" captionStart-1="FIGURE 2" captionStart-2="FIGURE 3" captionStartId-0="2.[151,250,1844,1866]" captionStartId-1="3.[151,250,1664,1686]" captionStartId-2="4.[151,250,1965,1987]" captionTargetBox-0="[165,1423,885,1829]" captionTargetBox-1="[205,1379,188,1656]" captionTargetBox-2="[368,1236,476,1936]" captionTargetId-0="figure@2.[151,1436,879,1829]" captionTargetId-1="figure@3.[205,1379,188,1656]" captionTargetId-2="figure@4.[346,1253,442,1953]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="4" captionText-0="FIGURE 1. A: Individual in situ. The black arrow indicates flattened and serrated anterior brim. The white arrow indicates filiform tips of dorsal papillae. The yellow arrow points at anus. B: Specimen before preservation in 95% alcohol. The arrow indicates almost retracted papillae. C: Dorsal view." captionText-1="FIGURE 2. SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm." captionText-2="FIGURE 3. Benthodytes marianensis sp. nov. dorsal ossicles. ac: lateral view. d: ventral view. e: dorsal view. f: lateral view of the peculiar cross-shaped ossicle. Scale 50 µm." httpUri-0="https://zenodo.org/record/1441643/files/figure.png" httpUri-1="https://zenodo.org/record/1441645/files/figure.png" httpUri-2="https://zenodo.org/record/1441647/files/figure.png" pageId="1" pageNumber="444">Figures 13</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C37E85A571082E0FCBEBF93343FFFF6B" lastPageId="2" lastPageNumber="445" pageId="1" pageNumber="444" type="diagnosis">
<paragraph id="8BDBD62E71082E0FCBEBF93343FFFF6B" blockId="1.[151,1437,1724,2037]" lastBlockId="2.[151,1437,151,824]" lastPageId="2" lastPageNumber="445" pageId="1" pageNumber="444">
<emphasis id="B9100A3C71082E0CCBEBF933433FF96B" bold="true" box="[151,273,1724,1749]" pageId="1" pageNumber="444">Diagnosis.</emphasis>
Body long, dorsum inflated, ventrum flattened, anteriorly depressed and tapered slightly posteriorly. Skin dark violet, soft, thick, wrinkled. Dorsal papillae 9 pairs, large, conical, with filiform tips, retracted, in two uneven rows along the dorsal radii. Some minute papillae scattered anteriorly. Some equal or smaller papillae often situated outside the larger papillae. Anus dorsal terminally, mouth ventral but a certain distance from the anterior edge. Brim well developed, with dark violet channels. Tentacles, at least twelve, retracted into stalks. Dorsal ossicles cross-shaped with central bipartite, tripartite, or tetrapartite apophyses. Sparse cross-shaped ossicles with high and spinous apophyses which end in two horizontal branches with downwardly directed spines on the margin, are in the dorsum. Ventral ossicles rod-shaped with central bipartite apophyses and crosses of three or four arms with central rudimentary apophyses. Brim ossicles sturdy and spinous, tripartite with bipartite apophyses and cross-shaped with tetrapartite or pentapartite apophyses. Apophyses, when divided, present at the basal region. Tentacle ossicles large rods.
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710B2E0FCBBBFF5041CDFEFE" pageId="2" pageNumber="445" type="materials_examined">
<paragraph id="8BDBD62E710B2E0FCBBBFF5041CDFEFE" blockId="2.[151,1437,151,824]" pageId="2" pageNumber="445">
<materialsCitation id="3B0CDC73710B2E0FCBBBFF5041CDFEFE" ID-GBIF-Occurrence="1913893033" collectingDate="2016-06-29" collectorName="Haibin Zhang." elevation="5567" latitude="11.799583" location="Genbank" longLatPrecision="1" longitude="142.1151" municipality="Mariana Trench area" pageId="2" pageNumber="445" specimenCount="1" specimenCount-adult="1" typeStatus="holotype">
<emphasis id="B9100A3C710B2E0FCBBBFF504395FF46" bold="true" box="[199,443,223,248]" pageId="2" pageNumber="445">Material examined.</emphasis>
<typeStatus id="54DF688C710B2E0FCAB1FF504017FF46" box="[461,569,223,248]" pageId="2" pageNumber="445" type="holotype">Holotype</typeStatus>
:
<specimenCount id="9D621DA7710B2E0FC932FF6F4088FF46" box="[590,678,223,248]" pageId="2" pageNumber="445" type="adult">1 adult</specimenCount>
specimen, catalog number IDSSE-2016-0629-HS01,
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accession numbers
<accessionNumber id="94374BCD710B2E0FCA0EFE8B43D8FEA2" box="[370,502,260,284]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MH049433" pageId="2" pageNumber="445">MH049433</accessionNumber>
,
<accessionNumber id="94374BCD710B2E0FC97EFE8B40A8FEA2" box="[514,646,260,284]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MH049434" pageId="2" pageNumber="445">MH049434</accessionNumber>
,
<accessionNumber id="94374BCD710B2E0FC9EEFE8B4138FEA3" box="[658,790,260,285]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MH049435" pageId="2" pageNumber="445">MH049435</accessionNumber>
. St. JL-Dive121-S01,
<collectingMunicipality id="6BBF4C54710B2E0FCF6DFE8B46D3FEA2" box="[1041,1277,260,285]" pageId="2" pageNumber="445">Mariana Trench area</collectingMunicipality>
,
<geoCoordinate id="EE50B0E9710B2E0FCE75FE8B47B6FEA3" box="[1289,1432,260,285]" direction="north" orientation="latitude" pageId="2" pageNumber="445" precision="1" value="11.799583">11°47.975'N</geoCoordinate>
,
<geoCoordinate id="EE50B0E9710B2E0FCBEBFEA8430DFEFE" box="[151,291,295,320]" direction="east" orientation="longitude" pageId="2" pageNumber="445" precision="1" value="142.1151">142°6.906'E</geoCoordinate>
,
<elevation id="0049311D710B2E0FCA53FEA843A8FE81" box="[303,390,295,320]" metricMagnitude="3" metricUnit="m" metricValue="5.567" pageId="2" pageNumber="445" unit="m" value="5567.0">
<quantity id="4C9C7BCB710B2E0FCA53FEA843A8FE81" box="[303,390,295,320]" metricMagnitude="3" metricUnit="m" metricValue="5.567" pageId="2" pageNumber="445" unit="m" value="5567.0">5567 m</quantity>
</elevation>
, muddy sediments,
<collectingDate id="EF9E0906710B2E0FC919FEA740D3FEFE" box="[613,765,295,320]" pageId="2" pageNumber="445" value="2016-06-29">
<date id="FFDAF0EE710B2E0FC919FEA740D3FEFE" box="[613,765,295,320]" pageId="2" pageNumber="445" value="2016-06-29">29 June 2016</date>
</collectingDate>
, coll.
<collectorName id="2691B3F8710B2E0FC83CFEA841CDFEFE" box="[832,995,295,320]" pageId="2" pageNumber="445">Haibin Zhang.</collectorName>
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710B2E08CBBBFEC3415DF8FF" lastPageId="5" lastPageNumber="448" pageId="2" pageNumber="445" type="description">
<paragraph id="8BDBD62E710B2E0FCBBBFEC34141FE13" blockId="2.[151,1437,151,824]" pageId="2" pageNumber="445">
<emphasis id="B9100A3C710B2E0FCBBBFEC3403FFEDB" bold="true" box="[199,529,332,357]" pageId="2" pageNumber="445">
Description of the
<typeStatus id="54DF688C710B2E0FCAD8FEC34023FEDB" box="[420,525,332,357]" pageId="2" pageNumber="445" type="holotype">holotype</typeStatus>
.
</emphasis>
As the specimen was seriously damaged during collection, the exact numbers of tentacles, circum-oral papillae and mid-ventral tube feet were difficult to determine. The present description was mainly derived from the living individual recorded by HD video.
</paragraph>
<paragraph id="8BDBD62E710B2E0FCBBBFE384149FE4B" blockId="2.[151,1437,151,824]" pageId="2" pageNumber="445">
The specimen was about
<quantity id="4C9C7BCB710B2E0FCA9CFE384030FE6E" box="[480,542,439,464]" metricMagnitude="-1" metricUnit="m" metricValue="4.5" pageId="2" pageNumber="445" unit="cm" value="45.0">45cm</quantity>
in length and
<quantity id="4C9C7BCB710B2E0FC9C1FE3840D2FE6E" box="[701,764,439,464]" metricMagnitude="-1" metricUnit="m" metricValue="1.5" pageId="2" pageNumber="445" unit="cm" value="15.0">15cm</quantity>
in width in freshly caught condition and
<quantity id="4C9C7BCB710B2E0FCFBAFE38472BFE6E" box="[1222,1285,439,464]" metricMagnitude="-1" metricUnit="m" metricValue="1.5" pageId="2" pageNumber="445" unit="cm" value="15.0">15cm</quantity>
in length and
<quantity id="4C9C7BCB710B2E0FCBEBFE5242E6FE4A" box="[151,200,477,500]" metricMagnitude="-2" metricUnit="m" metricValue="7.0" pageId="2" pageNumber="445" unit="cm" value="7.0">7cm</quantity>
in width after preservation in 95% alcohol for several days.
</paragraph>
<paragraph id="8BDBD62E710B2E0FCBBBFE704124FC86" blockId="2.[151,1437,151,824]" pageId="2" pageNumber="445">
Body about three times as long as broad, dark violet and semicircular in cross-section, flattened at the anterior end, inflated in following region, and peaked slightly at posterior end (
<figureCitation id="135FCAAB710B2E0FC896FDAB4609FD83" box="[1002,1063,548,573]" captionStart="FIGURE 1" captionStartId="2.[151,250,1844,1866]" captionTargetBox="[165,1423,885,1829]" captionTargetId="figure@2.[151,1436,879,1829]" captionTargetPageId="2" captionText="FIGURE 1. A: Individual in situ. The black arrow indicates flattened and serrated anterior brim. The white arrow indicates filiform tips of dorsal papillae. The yellow arrow points at anus. B: Specimen before preservation in 95% alcohol. The arrow indicates almost retracted papillae. C: Dorsal view." httpUri="https://zenodo.org/record/1441643/files/figure.png" pageId="2" pageNumber="445">Fig.1</figureCitation>
). Anus dorsal. Skin soft, thick, gelatinous, covered with mucus. Dorsal papillae broadly conical with filiform tips, retractile, arranged almost symmetrically in two rows along the radii, with some minute papillae confined to the anterior (
<figureCitation id="135FCAAB710B2E0FCFC9FDE3472BFD3B" box="[1205,1285,620,645]" captionStart="FIGURE 1" captionStartId="2.[151,250,1844,1866]" captionTargetBox="[165,1423,885,1829]" captionTargetId="figure@2.[151,1436,879,1829]" captionTargetPageId="2" captionText="FIGURE 1. A: Individual in situ. The black arrow indicates flattened and serrated anterior brim. The white arrow indicates filiform tips of dorsal papillae. The yellow arrow points at anus. B: Specimen before preservation in 95% alcohol. The arrow indicates almost retracted papillae. C: Dorsal view." httpUri="https://zenodo.org/record/1441643/files/figure.png" pageId="2" pageNumber="445">Fig.1C</figureCitation>
). In each row are 9 large papillae, often with equal or smaller ones situated elsewhere. Paired and smaller papillae render the dorsal skin wrinkled and uneven-coloured. Brim thin, usually bent upwards, well developed, and strongly pigmented with canals. Brim strongly expanded anteriorly, with retracted water-vascular canals forming serrated margin (
<figureCitation id="135FCAAB710B2E0FCB8BFD734364FCAB" box="[247,330,764,789]" captionStart="FIGURE 1" captionStartId="2.[151,250,1844,1866]" captionTargetBox="[165,1423,885,1829]" captionTargetId="figure@2.[151,1436,879,1829]" captionTargetPageId="2" captionText="FIGURE 1. A: Individual in situ. The black arrow indicates flattened and serrated anterior brim. The white arrow indicates filiform tips of dorsal papillae. The yellow arrow points at anus. B: Specimen before preservation in 95% alcohol. The arrow indicates almost retracted papillae. C: Dorsal view." httpUri="https://zenodo.org/record/1441643/files/figure.png" pageId="2" pageNumber="445">Fig.1A</figureCitation>
). Mouth ventral, a short distance from the anterior edge. Tentacles, at least twelve, retracted into the tentacle stalks and forming a large and flat oral disc.
</paragraph>
<caption id="DF1B86A6710B2E0FCBEBF8BB40B5F839" httpUri="https://zenodo.org/record/1441643/files/figure.png" pageId="2" pageNumber="445" startId="2.[151,250,1844,1866]" targetBox="[165,1423,885,1829]" targetPageId="2">
<paragraph id="8BDBD62E710B2E0FCBEBF8BB40B5F839" blockId="2.[151,1436,1844,1927]" pageId="2" pageNumber="445">
<emphasis id="B9100A3C710B2E0FCBEBF8BB433BF8F7" bold="true" box="[151,277,1844,1866]" pageId="2" pageNumber="445">FIGURE 1.</emphasis>
A: Individual
<emphasis id="B9100A3C710B2E0FCAD3F8BA43C1F8F4" box="[431,495,1845,1866]" italics="true" pageId="2" pageNumber="445">in situ</emphasis>
. The black arrow indicates flattened and serrated anterior brim. The white arrow indicates filiform tips of dorsal papillae. The yellow arrow points at anus. B: Specimen before preservation in 95% alcohol. The arrow indicates almost retracted papillae. C: Dorsal view.
</paragraph>
</caption>
<paragraph id="8BDBD62E710B2E0FCBBBF83B47BFF84E" blockId="2.[151,1437,1972,2032]" pageId="2" pageNumber="445">After preservation in 95% alcohol for several days, skin became reddish violet but tentacles dark violet. Skin is covered with a very superficial pigmented layer and dorsal papillae almost disappear or retract into the thick skin.</paragraph>
<caption id="DF1B86A6710A2E0ECBEBF90F415BF90A" httpUri="https://zenodo.org/record/1441645/files/figure.png" pageId="3" pageNumber="446" startId="3.[151,250,1664,1686]" targetBox="[205,1379,188,1656]" targetPageId="3">
<paragraph id="8BDBD62E710A2E0ECBEBF90F415BF90A" blockId="3.[151,1435,1664,1716]" pageId="3" pageNumber="446">
<emphasis id="B9100A3C710A2E0ECBEBF90F433DF92B" bold="true" box="[151,275,1664,1686]" pageId="3" pageNumber="446">FIGURE 2.</emphasis>
SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm.
</paragraph>
</caption>
<paragraph id="8BDBD62E710A2E09CBBBF96E40DBFE36" blockId="3.[151,1436,1761,2038]" lastBlockId="4.[151,1436,151,392]" lastPageId="4" lastPageNumber="447" pageId="3" pageNumber="446">
Ossicles in dorsal body wall cross-shaped, of two
<typeStatus id="54DF688C710A2E0EC87CF96D4110F944" box="[768,830,1762,1786]" pageId="3" pageNumber="446">types</typeStatus>
. One
<typeStatus id="54DF688C710A2E0EC8FCF96D419EF944" box="[896,944,1762,1786]" pageId="3" pageNumber="446">type</typeStatus>
is about 190300 µm in width with central spinous, bipartite, tripartite or tetrapartite apophyses. Each arm is 90180 µm in length, slightly curved downwards tapered distally. Spines on arms commonly distributed on distal part, or throughout the arms. The other dorsal ossicle
<typeStatus id="54DF688C710A2E0ECB97F8C14335F8D8" box="[235,283,1870,1894]" pageId="3" pageNumber="446">type</typeStatus>
is peculiar, cross-shaped, and uncommon. The arms are about 100µm in length, with multiple spines, and curve downwards. The central apophysis with spines is longer than arms of the cross and ends in two horizontal arms which possess downwardly vertical spines along their edge (
<figureCitation id="135FCAAB710A2E0ECF56F81A4652F810" box="[1066,1148,1941,1966]" captionStart="FIGURE 2" captionStartId="3.[151,250,1664,1686]" captionTargetBox="[205,1379,188,1656]" captionTargetId="figure@3.[205,1379,188,1656]" captionTargetPageId="3" captionText="FIGURE 2. SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm." httpUri="https://zenodo.org/record/1441645/files/figure.png" pageId="3" pageNumber="446">Fig.2A</figureCitation>
, the ossicle with arrow;
<figureCitation id="135FCAAB710A2E0ECBEBF83642F0F86C" box="[151,222,1977,2002]" captionStart="FIGURE 3" captionStartId="4.[151,250,1965,1987]" captionTargetBox="[368,1236,476,1936]" captionTargetId="figure@4.[346,1253,442,1953]" captionTargetPageId="4" captionText="FIGURE 3. Benthodytes marianensis sp. nov. dorsal ossicles. ac: lateral view. d: ventral view. e: dorsal view. f: lateral view of the peculiar cross-shaped ossicle. Scale 50 µm." httpUri="https://zenodo.org/record/1441647/files/figure.png" pageId="3" pageNumber="446">Fig.3f</figureCitation>
). Ossicles in ventral body wall include rods, and crosses with three and four arms. The rods are about 500 µm in length with large bipartite apophyses situated slightly off the central. Crosses of three and four arms are about 230410 µm and 260320 µm, respectively, and both possess central rudimentary apophyses (
<figureCitation id="135FCAAB710D2E09CE42FF1847A1FF0E" box="[1342,1423,151,176]" captionStart="FIGURE 2" captionStartId="3.[151,250,1664,1686]" captionTargetBox="[205,1379,188,1656]" captionTargetId="figure@3.[205,1379,188,1656]" captionTargetPageId="3" captionText="FIGURE 2. SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm." httpUri="https://zenodo.org/record/1441645/files/figure.png" pageId="4" pageNumber="447">Fig.2B</figureCitation>
). Ossicles in brim comprise tripartite and cross-shaped
<typeStatus id="54DF688C710D2E09C98CFF324105FF6B" box="[752,811,189,213]" pageId="4" pageNumber="447">types</typeStatus>
with less curved arms. The tripartite
<typeStatus id="54DF688C710D2E09CFB2FF3246D0FF6B" box="[1230,1278,189,213]" pageId="4" pageNumber="447">type</typeStatus>
is about 290 375 µm across with a large and strong central bifurcate apophysis. The cross-shaped
<typeStatus id="54DF688C710D2E09CF1DFF6F46BFFF46" box="[1121,1169,224,248]" pageId="4" pageNumber="447">type</typeStatus>
is 240330µm, usually with very large, spinous central tetrapartite and sometimes pentapartite apophyses. Compared to ossicles in body wall, brim ossicles usually develop rather robust and larger spines on arms and apophyses (
<figureCitation id="135FCAAB710D2E09CFDBFEA846D7FEFE" box="[1191,1273,295,320]" captionStart="FIGURE 2" captionStartId="3.[151,250,1664,1686]" captionTargetBox="[205,1379,188,1656]" captionTargetId="figure@3.[205,1379,188,1656]" captionTargetPageId="3" captionText="FIGURE 2. SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm." httpUri="https://zenodo.org/record/1441645/files/figure.png" pageId="4" pageNumber="447">Fig.2C</figureCitation>
). The division sites of apophyses in
<taxonomicName id="4C64ADAD710D2E09CAFAFEC24000FEDA" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[390,558,333,356]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="4" pageNumber="447" phylum="Echinodermata" rank="species" species="marianensis">
<emphasis id="B9100A3C710D2E09CAFAFEC24000FEDA" box="[390,558,333,356]" italics="true" pageId="4" pageNumber="447">B. marianensis</emphasis>
</taxonomicName>
all occur at the basal region. Ossicles in tentacles are large rods about 830 µm in length with spines only in their distal part (
<figureCitation id="135FCAAB710D2E09C9E9FEE040C6FE36" box="[661,744,367,392]" captionStart="FIGURE 2" captionStartId="3.[151,250,1664,1686]" captionTargetBox="[205,1379,188,1656]" captionTargetId="figure@3.[205,1379,188,1656]" captionTargetPageId="3" captionText="FIGURE 2. SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm." httpUri="https://zenodo.org/record/1441645/files/figure.png" pageId="4" pageNumber="447">Fig.2D</figureCitation>
).
</paragraph>
<caption id="DF1B86A6710D2E09CBEBF82240BEF85C" httpUri="https://zenodo.org/record/1441647/files/figure.png" pageId="4" pageNumber="447" startId="4.[151,250,1965,1987]" targetBox="[368,1236,476,1936]" targetPageId="4">
<paragraph id="8BDBD62E710D2E09CBEBF82240BEF85C" blockId="4.[151,1435,1965,2018]" pageId="4" pageNumber="447">
<emphasis id="B9100A3C710D2E09CBEBF822433DF87D" bold="true" box="[151,275,1965,1987]" pageId="4" pageNumber="447">FIGURE 3.</emphasis>
<taxonomicName id="4C64ADAD710D2E09CA66F8224037F87A" authority="Li &amp; Xiao &amp; Zhang &amp; Zhang, 2018" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[282,537,1965,1988]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="4" pageNumber="447" phylum="Echinodermata" rank="species" species="marianensis" status="sp. nov.">
<emphasis id="B9100A3C710D2E09CA66F8224037F87A" box="[282,537,1965,1988]" italics="true" pageId="4" pageNumber="447">Benthodytes marianensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A223B747710D2E09C95CF821405FF87D" box="[544,625,1966,1987]" pageId="4" pageNumber="447" rank="species">
<emphasis id="B9100A3C710D2E09C95CF821405FF87D" bold="true" box="[544,625,1966,1987]" pageId="4" pageNumber="447">sp. nov.</emphasis>
</taxonomicNameLabel>
dorsal ossicles. ac: lateral view. d: ventral view. e: dorsal view. f: lateral view of the peculiar cross-shaped ossicle. Scale 50 µm.
</paragraph>
</caption>
<caption id="DF1B86A6710C2E08CBEBFC1A43F6FC74" httpUri="https://zenodo.org/record/1441649/files/figure.png" pageId="5" pageNumber="448" startId="5.[151,250,917,939]" targetBox="[152,1378,199,884]" targetPageId="5">
<paragraph id="8BDBD62E710C2E08CBEBFC1A43F6FC74" blockId="5.[151,1436,917,970]" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBEBFC1A433CFC15" bold="true" box="[151,274,917,939]" pageId="5" pageNumber="448">FIGURE 4.</emphasis>
Neighbour-joining (NJ) tree based on 16S rRNA sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off.
</paragraph>
</caption>
<caption id="DF1B86A6710C2E08CBEBF90A43BFF907" httpUri="https://zenodo.org/record/1441651/files/figure.png" pageId="5" pageNumber="448" startId="5.[151,250,1669,1691]" targetBox="[237,1115,1102,1637]" targetPageId="5">
<paragraph id="8BDBD62E710C2E08CBEBF90A43BFF907" blockId="5.[151,1436,1668,1721]" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBEBF90A433CF924" bold="true" box="[151,274,1669,1691]" pageId="5" pageNumber="448">FIGURE 5.</emphasis>
Neighbour-joining (NJ) tree based on COI sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off.
</paragraph>
</caption>
<paragraph id="8BDBD62E710C2E08CBBBF96F415DF8FF" blockId="5.[151,1437,1760,2038]" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBBBF96F434BF947" bold="true" box="[199,357,1760,1785]" pageId="5" pageNumber="448">Relationship.</emphasis>
<taxonomicName id="4C64ADAD710C2E08CA10F96E40ABF946" authority="Li &amp; Xiao &amp; Zhang &amp; Zhang, 2018" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[364,645,1761,1785]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="5" pageNumber="448" phylum="Echinodermata" rank="species" species="marianensis" status="sp. nov.">
<emphasis id="B9100A3C710C2E08CA10F96E40ABF946" box="[364,645,1761,1785]" italics="true" pageId="5" pageNumber="448">Benthodytes marianensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A223B747710C2E08C9F7F96F40CBF947" box="[651,741,1760,1785]" pageId="5" pageNumber="448" rank="species">
<emphasis id="B9100A3C710C2E08C9F7F96F40CBF947" bold="true" box="[651,741,1760,1785]" pageId="5" pageNumber="448">sp. nov.</emphasis>
</taxonomicNameLabel>
is unique in the number and arrangement of dorsal papillae: 9 pairs of large broadly conical papillae with equal or smaller papillae situated outside in two zigzag rows. Dorsal deposits most closely resemble those of
<taxonomicName id="4C64ADAD710C2E08C926F8A6415EF8FF" authority="Ludwig, 1894" authorityName="Ludwig" authorityYear="1894" box="[602,880,1832,1857]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="5" pageNumber="448" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710C2E08C926F8A640E6F8FE" box="[602,712,1833,1856]" italics="true" pageId="5" pageNumber="448">B. incerta</emphasis>
<bibRefCitation id="EFF5ABDF710C2E08C9B3F8A7415EF8FF" box="[719,880,1832,1857]" pageId="5" pageNumber="448" refId="ref3689" refString="Ludwig, H. (1894) The Holothurioidea. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish Comission Streamer &quot; Albatross, during 1891. Memoirs of the Museum of Comparative Zoology, Cambridge, 17 (3), 1 - 183." type="journal article">Ludwig, 1894</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710C2E08CBBBF8C2402EF837" pageId="5" pageNumber="448" type="etymology">
<paragraph id="8BDBD62E710C2E08CBBBF8C2474CF8D8" blockId="5.[151,1437,1760,2038]" box="[199,1378,1869,1894]" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBBBF8C24360F8D8" bold="true" box="[199,334,1869,1894]" pageId="5" pageNumber="448">Etymology.</emphasis>
This species is named after the location (Mariana Trench) where the species was discovered.
</paragraph>
<paragraph id="8BDBD62E710C2E08CBBBF8FF402EF837" blockId="5.[151,1437,1760,2038]" box="[199,512,1904,1929]" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBBBF8FF4304F837" bold="true" box="[199,298,1904,1929]" pageId="5" pageNumber="448">Habitat.</emphasis>
Muddy sediments.
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710C2E08CBBBF81A47A2F813" box="[199,1420,1941,1966]" pageId="5" pageNumber="448" type="distribution">
<paragraph id="8BDBD62E710C2E08CBBBF81A47A2F813" blockId="5.[151,1437,1760,2038]" box="[199,1420,1941,1966]" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBBBF81A4373F810" bold="true" box="[199,349,1941,1966]" pageId="5" pageNumber="448">Distribution.</emphasis>
Known only from the
<typeStatus id="54DF688C710C2E08C92AF81940ABF810" box="[598,645,1942,1966]" pageId="5" pageNumber="448">type</typeStatus>
locality of the Mariana Trench, western Pacific Ocean, Depth
<quantity id="4C9C7BCB710C2E08CE4EF81A47A6F813" box="[1330,1416,1941,1966]" metricMagnitude="3" metricUnit="m" metricValue="5.567" pageId="5" pageNumber="448" unit="m" value="5567.0">5567 m</quantity>
.
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710C2E0BCBBBF837460DFDDE" lastPageId="6" lastPageNumber="449" pageId="5" pageNumber="448" type="discussion">
<paragraph id="8BDBD62E710C2E0BCBBBF8374189FEDB" blockId="5.[151,1437,1760,2038]" lastBlockId="6.[151,1437,151,1256]" lastPageId="6" lastPageNumber="449" pageId="5" pageNumber="448">
<emphasis id="B9100A3C710C2E08CBBBF8374315F86F" bold="true" box="[199,315,1976,2001]" pageId="5" pageNumber="448">Remarks.</emphasis>
Morphologically,
<taxonomicName id="4C64ADAD710C2E08C977F836409AF86E" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[523,692,1977,2000]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="5" pageNumber="448" phylum="Echinodermata" rank="species" species="marianensis">
<emphasis id="B9100A3C710C2E08C977F836409AF86E" box="[523,692,1977,2000]" italics="true" pageId="5" pageNumber="448">B. marianensis</emphasis>
</taxonomicName>
is very close to
<taxonomicName id="4C64ADAD710C2E08C812F83646ABF86F" authority="Ludwig, 1894" authorityName="Ludwig" authorityYear="1894" box="[878,1157,1976,2001]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="5" pageNumber="448" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710C2E08C812F83641F3F86E" box="[878,989,1977,2000]" italics="true" pageId="5" pageNumber="448">B. incerta</emphasis>
<bibRefCitation id="EFF5ABDF710C2E08C898F83746ABF86F" box="[996,1157,1976,2001]" pageId="5" pageNumber="448" refId="ref3689" refString="Ludwig, H. (1894) The Holothurioidea. Reports on an exploration off the west coasts of Mexico, Central and South America, and off the Galapagos Islands, in charge of Alexander Agassiz, by the U. S. Fish Comission Streamer &quot; Albatross, during 1891. Memoirs of the Museum of Comparative Zoology, Cambridge, 17 (3), 1 - 183." type="journal article">Ludwig, 1894</bibRefCitation>
</taxonomicName>
. They share two rows of large dorsal papillae, and peculiar ossicles (
<figureCitation id="135FCAAB710C2E08C901F85240FBF84B" box="[637,725,2013,2038]" captionStart="FIGURE 2" captionStartId="3.[151,250,1664,1686]" captionTargetBox="[205,1379,188,1656]" captionTargetId="figure@3.[205,1379,188,1656]" captionTargetPageId="3" captionText="FIGURE 2. SEM images of ossicles. A: the dorsum, arrow indicates the peculiar cross-shaped ossicles. B: the ventrum, arrow represents the large rod-shaped. C: the brim. D, tentacles. Scale 100 µm." httpUri="https://zenodo.org/record/1441645/files/figure.png" pageId="5" pageNumber="448">Fig. 2A</figureCitation>
, the ossicle with arrow).
<taxonomicName id="4C64ADAD710C2E08C891F8514675F84B" authorityName="Ludwig" authorityYear="1894" box="[1005,1115,2014,2037]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="5" pageNumber="448" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710C2E08C891F8514675F84B" box="[1005,1115,2014,2037]" italics="true" pageId="5" pageNumber="448">B. incerta</emphasis>
</taxonomicName>
was described by Ludwig in 1894 based on two specimens from the eastern Pacific (one large and one small). The original description lacked illustrations of dorsal ossicles. Also, it was not clear if the
<typeStatus id="54DF688C710F2E0BC847FF324145FF6B" box="[827,875,189,213]" pageId="6" pageNumber="449">type</typeStatus>
specimens had the peculiar
<typeStatus id="54DF688C710F2E0BCFCCFF3246CEFF6B" box="[1200,1248,189,213]" pageId="6" pageNumber="449">type</typeStatus>
of ossicles. The papillae ossicles of the
<typeStatus id="54DF688C710F2E0BCAE0FF6F43E2FF46" box="[412,460,224,248]" pageId="6" pageNumber="449">type</typeStatus>
specimens are cross-shaped with high apophyses and the apophyses are bipartite at the top. In addition, there are long side branches on the arms of ventral ossicles and in the high apophyses of papillae ossicles. In contrast. in
<emphasis id="B9100A3C710F2E0BCADFFEA6407DFE81" box="[419,595,296,320]" italics="true" pageId="6" pageNumber="449">
<taxonomicName id="4C64ADAD710F2E0BCADFFEA64061FE81" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[419,591,296,320]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="marianensis">B. marianensis</taxonomicName>
,
</emphasis>
the apophyses are bipartite, tripartite and tetrapartite at the base part. The long side branches are absent in the ossicles of the Mariana specimen.
</paragraph>
<paragraph id="8BDBD62E710F2E0BCBBBFEFF460DFDDE" blockId="6.[151,1437,151,1256]" pageId="6" pageNumber="449">
<bibRefCitation id="EFF5ABDF710F2E0BCBBBFEFF435FFE36" box="[199,369,367,392]" pageId="6" pageNumber="449" refId="ref3413" refString="Hansen, B. (1975) Scientific results of the Danish deep-sea expedition round the world 1950 - 52. Systematics and biology of the deep-sea holothurians. Part. l. Elasipoda. Galathea Report, 13, 1 - 262." type="journal article">Hansen (1975)</bibRefCitation>
re-examined
<typeStatus id="54DF688C710F2E0BC969FEFF406BFE36" box="[533,581,368,392]" pageId="6" pageNumber="449">type</typeStatus>
specimens and assigned his
<taxonomicName id="4C64ADAD710F2E0BC8E3FEFF4629FE36" box="[927,1031,368,392]" class="Malacostraca" family="Galatheidae" genus="Galathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="6" pageNumber="449" phylum="Arthropoda" rank="genus">
<emphasis id="B9100A3C710F2E0BC8E3FEFF4629FE36" box="[927,1031,368,392]" italics="true" pageId="6" pageNumber="449">Galathea</emphasis>
</taxonomicName>
specimens to
<taxonomicName id="4C64ADAD710F2E0BCFC9FEFE4706FE39" authorityName="Ludwig" authorityYear="1894" box="[1205,1320,368,392]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710F2E0BCFC9FEFE4706FE39" box="[1205,1320,368,392]" italics="true" pageId="6" pageNumber="449">B. incerta</emphasis>
</taxonomicName>
based on external morphology. The ossicles in the
<taxonomicName id="4C64ADAD710F2E0BC90DFE1B40F7FE12" box="[625,729,404,428]" class="Malacostraca" family="Galatheidae" genus="Galathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="6" pageNumber="449" phylum="Arthropoda" rank="genus">
<emphasis id="B9100A3C710F2E0BC90DFE1B40F7FE12" box="[625,729,404,428]" italics="true" pageId="6" pageNumber="449">Galathea</emphasis>
</taxonomicName>
specimens are larger and more regular than those in the
<typeStatus id="54DF688C710F2E0BCE17FE1A47B2FE13" box="[1387,1436,405,429]" pageId="6" pageNumber="449">type</typeStatus>
specimens. The Mariana specimen and Hansens
<taxonomicName id="4C64ADAD710F2E0BC9A3FE374169FE6E" box="[735,839,440,464]" class="Malacostraca" family="Galatheidae" genus="Galathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="6" pageNumber="449" phylum="Arthropoda" rank="genus">
<emphasis id="B9100A3C710F2E0BC9A3FE374169FE6E" box="[735,839,440,464]" italics="true" pageId="6" pageNumber="449">Galathea</emphasis>
</taxonomicName>
specimens differ in the following features: (1) In
<taxonomicName id="4C64ADAD710F2E0BCBEBFE5342D1FE4A" box="[151,255,476,500]" class="Malacostraca" family="Galatheidae" genus="Galathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="6" pageNumber="449" phylum="Arthropoda" rank="genus">
<emphasis id="B9100A3C710F2E0BCBEBFE5342D1FE4A" box="[151,255,476,500]" italics="true" pageId="6" pageNumber="449">Galathea</emphasis>
</taxonomicName>
specimens, the crosses are much larger than those of Mariana specimen. (2) In
<taxonomicName id="4C64ADAD710F2E0BCFFAFE5346C0FE4A" box="[1158,1262,476,500]" class="Malacostraca" family="Galatheidae" genus="Galathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="6" pageNumber="449" phylum="Arthropoda" rank="genus">
<emphasis id="B9100A3C710F2E0BCFFAFE5346C0FE4A" box="[1158,1262,476,500]" italics="true" pageId="6" pageNumber="449">Galathea</emphasis>
</taxonomicName>
specimens, the apophyses of crosses are bipartite at least half the length; in Mariana specimen, the apophyses are bipartite, tripartite or tetrapartite at the base part. (3) In
<taxonomicName id="4C64ADAD710F2E0BC9C1FDAB410BFD82" box="[701,805,548,572]" class="Malacostraca" family="Galatheidae" genus="Galathea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Decapoda" pageId="6" pageNumber="449" phylum="Arthropoda" rank="genus">
<emphasis id="B9100A3C710F2E0BC9C1FDAB410BFD82" box="[701,805,548,572]" italics="true" pageId="6" pageNumber="449">Galathea</emphasis>
</taxonomicName>
specimens, the apophyses of the peculiar ossicles are smooth; in Mariana specimen, the apophyses of the peculiar ossicles are spinous.
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710F2E0BCBBBFDE34053FC76" pageId="6" pageNumber="449" type="reference_group">
<paragraph id="8BDBD62E710F2E0BCBBBFDE34694FCAA" blockId="6.[151,1437,151,1256]" pageId="6" pageNumber="449">
<taxonomicName id="4C64ADAD710F2E0BCBBBFDE34064FD3B" ID-CoL="68J6V" authority="Ohshima, 1915" authorityName="Ohshima" authorityYear="1915" box="[199,586,620,645]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="gotoi">
<emphasis id="B9100A3C710F2E0BCBBBFDE343BDFD3A" box="[199,403,620,644]" italics="true" pageId="6" pageNumber="449">Benthodytes gotoi</emphasis>
<bibRefCitation id="EFF5ABDF710F2E0BCAE0FDE34064FD3B" box="[412,586,620,645]" pageId="6" pageNumber="449" refId="ref3818" refString="Ohshima, H. (1915) Report on the holothurians collected by the United States Steamer &quot; Albatross &quot; in the northwestern Pacific during the summer of 1906. Proceedengs of the United States National Museum, 48, 213 - 291. https: // doi. org / 10.5479 / si. 00963801.48 - 2073.213" type="journal article">Ohshima, 1915</bibRefCitation>
</taxonomicName>
is known from four specimens, taken in the Okhotsk Sea.
<bibRefCitation id="EFF5ABDF710F2E0BCF88FDE247B2FD3B" box="[1268,1436,620,645]" pageId="6" pageNumber="449" refId="ref3413" refString="Hansen, B. (1975) Scientific results of the Danish deep-sea expedition round the world 1950 - 52. Systematics and biology of the deep-sea holothurians. Part. l. Elasipoda. Galathea Report, 13, 1 - 262." type="journal article">Hansen (1975)</bibRefCitation>
considered this species to be a synonym of
<taxonomicName id="4C64ADAD710F2E0BC9F0FD1E40D2FD19" ID-CoL="5WJ6Z" authorityName="Ludwig" authorityYear="1894" box="[652,764,656,680]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710F2E0BC9F0FD1E40D2FD19" box="[652,764,656,680]" italics="true" pageId="6" pageNumber="449">B. incerta</emphasis>
</taxonomicName>
. The crosses of Ohshimas specimens differ from those of Mariana specimen in their large size. The arms of the crosses are usually 300800 µm, and the apophyses of crosses are usually bipartite for one-half of its length. In the Mariana specimen, the arms of the crosses are usually 90180 µm, and the apophyses of crosses are bipartite, tripartite or tetrapartite at the base part.
</paragraph>
<paragraph id="8BDBD62E710F2E0BCBBBFCAF4053FC76" blockId="6.[151,1437,151,1256]" pageId="6" pageNumber="449">
In addition, the Mariana specimen differs from the descriptions of
<taxonomicName id="4C64ADAD710F2E0BC8ABFCAE4664FC89" ID-CoL="5WJ6Z" authorityName="Ludwig" authorityYear="1894" box="[983,1098,800,824]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710F2E0BC8ABFCAE4664FC89" box="[983,1098,800,824]" italics="true" pageId="6" pageNumber="449">B. incerta</emphasis>
</taxonomicName>
and its synonym
<taxonomicName id="4C64ADAD710F2E0BCE62FCAE4755FC89" ID-CoL="68J6V" authorityName="Ohshima" authorityYear="1915" box="[1310,1403,800,824]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="gotoi">
<emphasis id="B9100A3C710F2E0BCE62FCAE4755FC89" box="[1310,1403,800,824]" italics="true" pageId="6" pageNumber="449">B. gotoi</emphasis>
</taxonomicName>
in number and arrangement of dorsal large papillae: In
<taxonomicName id="4C64ADAD710F2E0BC993FCCA4171FCE2" ID-CoL="5WJ6Z" authorityName="Ludwig" authorityYear="1894" box="[751,863,837,860]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="species" species="incerta">
<emphasis id="B9100A3C710F2E0BC993FCCA4171FCE2" box="[751,863,837,860]" italics="true" pageId="6" pageNumber="449">B. incerta</emphasis>
</taxonomicName>
, there are 69 (58) large papillae in a single row along each dorsal ambulacrum, while in Mariana specimen there are 9 papillae with equal or smaller ones situated elsewhere, forming a zigzag row. Therefore, based on the external morphology and internal ossicles, the Mariana specimen can be assigned to a new species.
</paragraph>
</subSubSection>
<subSubSection id="C37E85A5710F2E0BCBBBFC5B4768FB56" pageId="6" pageNumber="449" type="description">
<paragraph id="8BDBD62E710F2E0BCBBBFC5B4768FB56" blockId="6.[151,1437,151,1256]" pageId="6" pageNumber="449">
<geoCoordinate id="EE50B0E9710F2E0BCBBBFC5B4336FC53" box="[199,280,980,1005]" direction="south" orientation="latitude" pageId="6" pageNumber="449" precision="925" value="-16.266666">16 16S</geoCoordinate>
rRNA gene sequences, 14 COI and 14 H3 gene sequences were selected from GenBank, including almost all species in the
<taxonomicName id="4C64ADAD710F2E0BCACCFC78401AFBAE" authorityName="Theel" authorityYear="1882" box="[432,564,1015,1040]" class="Holothuroidea" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="order">Elasipodida</taxonomicName>
especially those in the
<taxonomicName id="4C64ADAD710F2E0BC83DFC7841DFFBAE" authorityName="Theel" authorityYear="1882" box="[833,1009,1015,1040]" class="Holothuroidea" family="Psychropotidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="family">Psychropotidae</taxonomicName>
. The topological structures of NJ and ML tree were all highly similar to each other based on these genes separately (
<figureCitation id="135FCAAB710F2E0BCF75FB93464AFB8B" box="[1033,1124,1052,1077]" captionStart-0="FIGURE 4" captionStart-1="FIGURE 5" captionStart-2="FIGURE 6" captionStartId-0="5.[151,250,917,939]" captionStartId-1="5.[151,250,1669,1691]" captionStartId-2="6.[151,250,1970,1992]" captionTargetBox-0="[152,1378,199,884]" captionTargetBox-1="[237,1115,1102,1637]" captionTargetBox-2="[152,1346,1316,1903]" captionTargetId-0="figure@5.[151,1435,193,896]" captionTargetId-1="figure@5.[151,1436,1020,1647]" captionTargetId-2="figure@6.[151,1435,1311,1948]" captionTargetPageId-0="5" captionTargetPageId-1="5" captionTargetPageId-2="6" captionText-0="FIGURE 4. Neighbour-joining (NJ) tree based on 16S rRNA sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off." captionText-1="FIGURE 5. Neighbour-joining (NJ) tree based on COI sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off." captionText-2="FIGURE 6. Neighbour-joining (NJ) tree based on H3 sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off." httpUri-0="https://zenodo.org/record/1441649/files/figure.png" httpUri-1="https://zenodo.org/record/1441651/files/figure.png" httpUri-2="https://zenodo.org/record/1441653/files/figure.png" pageId="6" pageNumber="449">Fig.46</figureCitation>
). The phylogenetic analysis based on mitochondrial (16S and COI) and nuclear genes (H3) were all consistent with the result of morphological identification, embedding the new species within the
<taxonomicName id="4C64ADAD710F2E0BC83CFBEB41C3FBC3" authorityName="Theel" authorityYear="1882" box="[832,1005,1124,1149]" class="Holothuroidea" family="Psychropotidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="family">Psychropotidae</taxonomicName>
clade. However,
<taxonomicName id="4C64ADAD710F2E0BCFA8FBEB4773FBC2" authorityName="Li &amp; Xiao &amp; Zhang &amp; Zhang" authorityYear="2018" box="[1236,1373,1124,1148]" class="Holothuroidea" family="Psychropotidae" genus="Benthodytes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="genus">
<emphasis id="B9100A3C710F2E0BCFA8FBEB4773FBC2" box="[1236,1373,1124,1148]" italics="true" pageId="6" pageNumber="449">Benthodytes</emphasis>
</taxonomicName>
was paraphyletic according to mitochondrial genes (
<figureCitation id="135FCAAB710F2E0BC9C5FB0840D9FB1E" box="[697,759,1159,1184]" captionStart="FIGURE 4" captionStartId="5.[151,250,917,939]" captionTargetBox="[152,1378,199,884]" captionTargetId="figure@5.[151,1435,193,896]" captionTargetPageId="5" captionText="FIGURE 4. Neighbour-joining (NJ) tree based on 16S rRNA sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off." httpUri="https://zenodo.org/record/1441649/files/figure.png" pageId="6" pageNumber="449">Fig.4</figureCitation>
,
<figureCitation id="135FCAAB710F2E0BC879FB08413AFB1E" box="[773,788,1159,1184]" captionStart="FIGURE 5" captionStartId="5.[151,250,1669,1691]" captionTargetBox="[237,1115,1102,1637]" captionTargetId="figure@5.[151,1436,1020,1647]" captionTargetPageId="5" captionText="FIGURE 5. Neighbour-joining (NJ) tree based on COI sequences. The bootstrap values are NJ/ML at each node and the values below 50% were cut off." httpUri="https://zenodo.org/record/1441651/files/figure.png" pageId="6" pageNumber="449">5</figureCitation>
), which is inconsistent with the traditional classification system. The results showed poor support, likely due to the limited sequences available for the
<taxonomicName id="4C64ADAD710F2E0BCF95FB2347B7FB7B" authorityName="Theel" authorityYear="1882" box="[1257,1433,1196,1221]" class="Holothuroidea" family="Psychropotidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Elasipodida" pageId="6" pageNumber="449" phylum="Echinodermata" rank="family">Psychropotidae</taxonomicName>
. Sequences obtained in the present study will enrich the database and provide more data for future research.
</paragraph>
</subSubSection>
</treatment>
</document>