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<document id="0E679B49DAF17F698606C0D46C8D48A4" ID-DOI="10.1111/j.1096-3642.2012.00857.x" ID-ISSN="0024-4082" ID-Zenodo-Dep="5408987" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630349776171" checkinUser="marcus" docAuthor="Martínez-Ansemil, Enrique, Châtelliers, Michel Creuzé Des, Martin, Patrick &amp; Sambugar, Beatrice" docDate="2012" docId="039ADA531F5F697EFF5324F26671FC91" docLanguage="en" docName="j.1096-3642.2012.00857.x.pdf" docOrigin="Zoological Journal of the Linnean Society 166 (3)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2012.00857.x" docStyle="DocumentStyle:FBF0BF18BB2B256F1F8008F27D80BF37.3:ZoolJLinnSoc.2010-2016.journal_article" docStyleId="FBF0BF18BB2B256F1F8008F27D80BF37" docStyleName="ZoolJLinnSoc.2010-2016.journal_article" docStyleVersion="3" docTitle="Parvidrilus meyssonnieri DES CHATELLIERS &amp; MARTIN 2012, SP. NOV." docType="treatment" docVersion="4" lastPageNumber="537" masterDocId="FFA3A22B1F5B6979FFE2202C637BFF98" masterDocTitle="The Parvidrilidae - a diversified groundwater family: description of six new species from southern Europe, and clues for its phylogenetic position within Clitellata (Annelida)" masterLastPageNumber="558" masterPageNumber="530" pageNumber="534" updateTime="1699241350002" updateUser="plazi">
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<mods:title id="13E7B30D8635E8444262F2B2DEF4A12A">The Parvidrilidae - a diversified groundwater family: description of six new species from southern Europe, and clues for its phylogenetic position within Clitellata (Annelida)</mods:title>
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<mods:namePart id="B7AFE5822973412D5E5E71FEE4D944F5">Martínez-Ansemil, Enrique</mods:namePart>
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<mods:namePart id="F9287A777E15858CB498422A6EC70EE0">Châtelliers, Michel Creuzé Des</mods:namePart>
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<mods:namePart id="F27CC13163FAE234F7D456781787AAA8">Martin, Patrick</mods:namePart>
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<mods:namePart id="269D1344F44864CE9F1DA86C521690F5">Sambugar, Beatrice</mods:namePart>
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<mods:date id="99D191EAE9260C2398AD5A5991308A69">2012</mods:date>
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<treatment id="039ADA531F5F697EFF5324F26671FC91" ID-DOI="http://doi.org/10.5281/zenodo.5480034" ID-Zenodo-Dep="5480034" LSID="urn:lsid:plazi:treatment:039ADA531F5F697EFF5324F26671FC91" httpUri="http://treatment.plazi.org/id/039ADA531F5F697EFF5324F26671FC91" lastPageId="7" lastPageNumber="537" pageId="4" pageNumber="534">
<subSubSection id="C32938CE1F5F697DFF5324F262FBFA2C" pageId="4" pageNumber="534" type="nomenclature">
<paragraph id="8B8C6B451F5F697DFF5324F26105FA8B" blockId="4.[177,766,1245,1299]" pageId="4" pageNumber="534">
<heading id="D0C4DC291F5F697DFF5324F26105FA8B" centered="true" fontSize="9" level="2" pageId="4" pageNumber="534" reason="4">
<taxonomicName id="4C3310C61F5F697DFF5324F262F0FA8A" authority="DES CHATELLIERS &amp; MARTIN" authorityName="DES CHATELLIERS &amp; MARTIN" authorityYear="2012" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="534" phylum="Annelida" rank="species" species="meyssonnieri" status="sp. nov.">
<emphasis id="B947B7571F5F697DFF5324F2617CFB6A" bold="true" box="[177,519,1246,1267]" italics="true" pageId="4" pageNumber="534">PARVIDRILUS MEYSSONNIERI</emphasis>
DES CHÂTELLIERS &amp; MARTIN
</taxonomicName>
<emphasis id="B947B7571F5F697DFE77252D6281FA8A" bold="true" box="[405,506,1277,1298]" pageId="4" pageNumber="534">
<taxonomicNameLabel id="A2740A2C1F5F697DFE77252D6281FA8A" box="[405,506,1277,1298]" pageId="4" pageNumber="534" rank="species">SP. NOV.</taxonomicNameLabel>
</emphasis>
(
<figureCitation id="130877C01F5F697DFDEE24D1612DFA8A" box="[524,598,1277,1298]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="4" pageNumber="534">FIGS 1</figureCitation>
,
<figureCitation id="130877C01F5F697DFD8A24D1610DFA8B" box="[616,630,1277,1299]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="4" pageNumber="534">2</figureCitation>
)
</heading>
</paragraph>
<paragraph id="8B8C6B451F5F697DFF46250862FBFA2C" blockId="4.[164,779,1316,1674]" pageId="4" pageNumber="534">
<emphasis id="B947B7571F5F697DFF4625086391FAA1" box="[164,234,1316,1337]" italics="true" pageId="4" pageNumber="534">Types:</emphasis>
<materialsCitation id="3B5B61181F5F697DFF15250862FBFA2C" collectingDate="2000-09-28" collectionCode="MHNL" collectorName="Gallery &amp; La Martiniere'" country="Canada" elevation="350" latitude="45.680836" location="Thurins" longLatPrecision="18" longitude="4.6475" pageId="4" pageNumber="534" specimenCode="MHNL 44003358" specimenCount="1" typeStatus="holotype">
<typeStatus id="5488D5E71F5F697DFF152508621AFAA2" box="[247,353,1316,1338]" pageId="4" pageNumber="534" type="holotype">Holotype</typeStatus>
.
<specimenCode id="DB95C33E1F5F697DFE902508613BFAA2" box="[370,576,1316,1338]" collectionCode="MHNL" country="France" httpUri="http://grbio.org/cool/4jmm-iazx" name="Musee Guimet d'Histoire Naturelle de Lyon" pageId="4" pageNumber="534">MHNL 44003358</specimenCode>
, slide 6, mature specimen, stained in paracarmine and wholemounted in
<collectingCountry id="F3242BD51F5F697DFECE254D62FFFAEF" box="[300,388,1377,1399]" name="Canada" pageId="4" pageNumber="534">Canada</collectingCountry>
balsam.
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of
<collectorName id="26C60E931F5F697DFD87254D6070FAEF" box="[613,779,1377,1399]" pageId="4" pageNumber="534">La Martinière</collectorName>
(
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,
<geoCoordinate id="EE070D821F5F697DFEA625AC62BDFA0D" box="[324,454,1406,1429]" degrees="04" direction="east" minutes="38" orientation="longitude" pageId="4" pageNumber="534" precision="15" seconds="51" value="4.6475">04°3851E</geoCoordinate>
,
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<quantity id="4CCBC6A01F5F697DFE382553615BFA0D" box="[474,544,1407,1429]" metricMagnitude="2" metricUnit="m" metricValue="3.5" pageId="4" pageNumber="534" unit="m" value="350.0">350 m</quantity>
asl
</elevation>
),
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(69),
<collectingCountry id="F3242BD51F5F697DFF4625B3638DFA2C" box="[164,246,1439,1460]" name="France" pageId="4" pageNumber="534">France</collectingCountry>
,
<date id="FF8D4D851F5F697DFEE625B26207FA2C" box="[260,380,1438,1460]" pageId="4" pageNumber="534" value="2000-09-28">
<collectingDate id="EFC9B46D1F5F697DFEE625B26207FA2C" box="[260,380,1438,1460]" pageId="4" pageNumber="534" value="2000-09-28">28.ix.2000</collectingDate>
</date>
.
</materialsCitation>
</paragraph>
</subSubSection>
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<materialsCitation id="3B5B61181F5F697DFF5E259161D4FA69" collectionCode="MHNL" pageId="4" pageNumber="534" specimenCode="MHNL 44003359" specimenCount="2" specimenCount-immature="1" typeStatus="paratype">
<typeStatus id="5488D5E71F5F697DFF5E25916248FA4A" box="[188,307,1469,1490]" pageId="4" pageNumber="534" type="paratype">Paratypes</typeStatus>
.
<specimenCode id="DB95C33E1F5F697DFEDC2591617FFA4B" box="[318,516,1469,1491]" collectionCode="MHNL" country="France" httpUri="http://grbio.org/cool/4jmm-iazx" name="Musee Guimet d'Histoire Naturelle de Lyon" pageId="4" pageNumber="534">MHNL 44003359</specimenCode>
, slide 1,
<specimenCount id="9D35A0CC1F5F697DFD852591607CFA4B" box="[615,775,1469,1491]" pageId="4" pageNumber="534" type="generic">six specimens</specimenCount>
,
<specimenCount id="9D35A0CC1F5F697DFF4625F76235FA69" box="[164,334,1499,1521]" pageId="4" pageNumber="534" type="immature">four immature</specimenCount>
, one mature and one fragment
</materialsCitation>
;
<materialsCitation id="3B5B61181F5F697DFD5825F062FCF9F4" collectionCode="MHNL" country="Canada" location="Canada" pageId="4" pageNumber="534" specimenCode="MHNL 44003360" specimenCount="3" specimenCount-immature="2" typeStatus="paratype">
<specimenCode id="DB95C33E1F5F697DFD5825F0626FF988" collectionCode="MHNL" country="France" httpUri="http://grbio.org/cool/4jmm-iazx" name="Musee Guimet d'Histoire Naturelle de Lyon" pageId="4" pageNumber="534">MHNL 44003360</specimenCode>
, slide 3,
<specimenCount id="9D35A0CC1F5F697DFE6525D66141F988" box="[391,570,1530,1552]" pageId="4" pageNumber="534" type="generic">four specimens</specimenCount>
, two mature and
<specimenCount id="9D35A0CC1F5F697DFF4626356236F9B6" box="[164,333,1560,1582]" pageId="4" pageNumber="534" type="immature">two immature</specimenCount>
; slide 6,
<specimenCount id="9D35A0CC1F5F697DFE5D2635611FF9B6" box="[447,612,1560,1582]" pageId="4" pageNumber="534" type="immature">one immature</specimenCount>
specimen; all specimens stained with paracarmine and wholemounted in
<collectingCountry id="F3242BD51F5F697DFECD267A62FCF9F4" box="[303,391,1622,1644]" name="Canada" pageId="4" pageNumber="534">Canada</collectingCountry>
</materialsCitation>
<materialsCitation id="3B5B61181F5F697DFE6D267A6151F912" collectingDate="2000-09-28" collectorName="La Martiniere' &amp; Thurins" country="France" location="France" pageId="4" pageNumber="534" specimenCount="1" typeStatus="paratype">
balsam;
<date id="FF8D4D851F5F697DFE12267A6113F9F4" box="[496,616,1622,1644]" pageId="4" pageNumber="534" value="2000-09-28">
<collectingDate id="EFC9B46D1F5F697DFE12267A6113F9F4" box="[496,616,1622,1644]" pageId="4" pageNumber="534" value="2000-09-28">28.ix.2000</collectingDate>
</date>
; gallery of
<collectorName id="26C60E931F5F697DFD0E267A625CF912" pageId="4" pageNumber="534">La Martinière</collectorName>
,
<collectorName id="26C60E931F5F697DFED7265862EAF912" box="[309,401,1652,1674]" pageId="4" pageNumber="534">Thurins</collectorName>
(69),
<collectingCountry id="F3242BD51F5F697DFE3526596151F912" box="[471,554,1653,1674]" name="France" pageId="4" pageNumber="534">France</collectingCountry>
</materialsCitation>
.
</paragraph>
<paragraph id="8B8C6B451F5F697DFF4626EC66DAFE0B" blockId="4.[164,779,1728,1903]" lastBlockId="4.[826,1441,198,403]" pageId="4" pageNumber="534">
<materialsCitation id="3B5B61181F5F697DFF4626EC66E6FE0B" collectingDate="1999-09-14" collectingDateMax="2008-01-28" collectingDateMin="1999-09-14" collectionCode="SEM" collectorName="La Martiniere'" country="France" elevation="520" latitude="45.287777" location="France" longLatPrecision="19" longitude="4.0994444" pageId="4" pageNumber="534" specimenCount="54" specimenCount-immature="1" specimenCount-juvenile="15">
<emphasis id="B947B7571F5F697DFF4626EC62B7F94D" box="[164,460,1728,1749]" italics="true" pageId="4" pageNumber="534">Other material examined:</emphasis>
Apart from the
<typeStatus id="5488D5E71F5F697DFD6C26EC61C4F94D" box="[654,703,1728,1749]" pageId="4" pageNumber="534">type</typeStatus>
material,
<specimenCount id="9D35A0CC1F5F697DFF3826F3620BF96C" box="[218,368,1758,1780]" pageId="4" pageNumber="534" type="generic">11 specimens</specimenCount>
were observed and mounted on five scanning electron microscope (
<collectionCode id="ED22F3801F5F697DFDEE26D1613DF88B" box="[524,582,1789,1811]" pageId="4" pageNumber="534">SEM</collectionCode>
) stubs, IRScNB,
<date id="FF8D4D851F5F697DFF4627306275F8AA" box="[164,270,1820,1842]" pageId="4" pageNumber="534" value="2008-01-28">
<collectingDate id="EFC9B46D1F5F697DFF4627306275F8AA" box="[164,270,1820,1842]" pageId="4" pageNumber="534" value="2008-01-28">28.i.2008</collectingDate>
</date>
; stub 1,
<specimenCount id="9D35A0CC1F5F697DFE722730612CF8AA" box="[400,599,1820,1842]" pageId="4" pageNumber="534" type="generic">three specimens</specimenCount>
,
<typeStatus id="5488D5E71F5F697DFD8F273061E5F8A9" box="[621,670,1820,1841]" pageId="4" pageNumber="534">type</typeStatus>
locality; stubs 25,
<specimenCount id="9D35A0CC1F5F697DFED527166287F8C8" box="[311,508,1850,1872]" pageId="4" pageNumber="534" type="generic">eight specimens</specimenCount>
, gallery of Navogne (
<geoCoordinate id="EE070D821F5F697DFF4E2775624BF8F6" box="[172,304,1879,1903]" degrees="45" direction="north" minutes="17" orientation="latitude" pageId="4" pageNumber="534" precision="15" seconds="16" value="45.287777">45°1716N</geoCoordinate>
,
<geoCoordinate id="EE070D821F5F697DFEA1277562BEF8F6" box="[323,453,1879,1903]" degrees="04" direction="east" minutes="05" orientation="longitude" pageId="4" pageNumber="534" precision="15" seconds="58" value="4.0994444">04°0558E</geoCoordinate>
,
<elevation id="001E8C761F5F697DFE3527756136F8F7" box="[471,589,1881,1903]" metricMagnitude="2" metricUnit="m" metricValue="5.2" pageId="4" pageNumber="534" unit="m" value="520.0">
<quantity id="4CCBC6A01F5F697DFE3527756166F8F6" box="[471,541,1881,1903]" metricMagnitude="2" metricUnit="m" metricValue="5.2" pageId="4" pageNumber="534" unit="m" value="520.0">520 m</quantity>
asl
</elevation>
), Bas-en-Basset (43),
<collectingCountry id="F3242BD51F5F697DFC6620EA60ACFF43" box="[900,983,198,219]" name="France" pageId="4" pageNumber="534">France</collectingCountry>
. Thirty-six specimens were wholemounted in
<collectingCountry id="F3242BD51F5F697DFC2A20C8675BFF62" box="[968,1056,228,250]" name="Canada" pageId="4" pageNumber="534">Canada</collectingCountry>
balsam or in Ammans lactophenol, in the collection of one of the authors (MCdC):
<collectorName id="26C60E931F5F697DFA60212F60C0FEA0" pageId="4" pageNumber="534">La Martinière</collectorName>
(
<specimenCount id="9D35A0CC1F5F697DFC29210E67F6FEA0" box="[971,1165,290,312]" pageId="4" pageNumber="534" type="generic">seven specimens</specimenCount>
, four mature and
<specimenCount id="9D35A0CC1F5F697DFA87210E60ECFECE" pageId="4" pageNumber="534" type="juvenile">three juvenile</specimenCount>
,
<date id="FF8D4D851F5F697DFC41216C6761FECE" box="[931,1050,320,342]" pageId="4" pageNumber="534" value="1999-09-14">
<collectingDate id="EFC9B46D1F5F697DFC41216C6761FECE" box="[931,1050,320,342]" pageId="4" pageNumber="534" value="1999-09-14">14.ix.1999</collectingDate>
</date>
;
<specimenCount id="9D35A0CC1F5F697DFBC4216C67BAFECE" box="[1062,1217,320,342]" pageId="4" pageNumber="534" type="generic">24 specimens</specimenCount>
, ten mature and
<specimenCount id="9D35A0CC1F5F697DFA64216C60ECFEED" pageId="4" pageNumber="534" type="juvenile">14 juvenile</specimenCount>
,
<date id="FF8D4D851F5F697DFC5621736750FEED" box="[948,1067,351,373]" pageId="4" pageNumber="534" value="2000-09-28">
<collectingDate id="EFC9B46D1F5F697DFC5621736750FEED" box="[948,1067,351,373]" pageId="4" pageNumber="534" value="2000-09-28">28.ix.2000</collectingDate>
</date>
); Navogne (
<specimenCount id="9D35A0CC1F5F697DFB39217366DAFEED" box="[1243,1441,351,373]" pageId="4" pageNumber="534" type="immature">three immature</specimenCount>
specimens,
<date id="FF8D4D851F5F697DFC5F21526764FE0B" box="[957,1055,382,403]" pageId="4" pageNumber="534" value="1999-10-08">
<collectingDate id="EFC9B46D1F5F697DFC5F21526764FE0B" box="[957,1055,382,403]" pageId="4" pageNumber="534" value="1999-10-08">8.x.1999</collectingDate>
</date>
; one mature specimen,
<date id="FF8D4D851F5F697DFAC8215166EEFE0B" box="[1322,1429,381,403]" pageId="4" pageNumber="534" value="2008-01-28">
<collectingDate id="EFC9B46D1F5F697DFAC8215166EEFE0B" box="[1322,1429,381,403]" pageId="4" pageNumber="534" value="2008-01-28">28.i.2008</collectingDate>
</date>
)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8C6B451F5F697DFCD8219760E9FDF2" blockId="4.[826,1441,443,618]" pageId="4" pageNumber="534">
<emphasis id="B947B7571F5F697DFCD8219760C3FE48" box="[826,952,443,464]" italics="true" pageId="4" pageNumber="534">Etymology:</emphasis>
Named after Marcel Meyssonnier, as a tribute to this outstanding and tireless speleologist, and long-time friend and team member of one of the authors (MCdC), and whose involvement and support were instrumental in the discovery of this new species.
</paragraph>
<paragraph id="8B8C6B451F5F697DFCD822BD6740FC40" blockId="4.[826,1441,657,1903]" pageId="4" pageNumber="534">
<emphasis id="B947B7571F5F697DFCD822BD60B9FD3E" box="[826,962,657,678]" italics="true" pageId="4" pageNumber="534">Description:</emphasis>
Length
<quantity id="4CCBC6A01F5F697DFBA222BD67BAFD3E" box="[1088,1217,657,679]" metricMagnitude="-3" metricUnit="m" metricValue="1.45" metricValueMax="1.6" metricValueMin="1.3" pageId="4" pageNumber="534" unit="mm" value="1.45" valueMax="1.6" valueMin="1.3">1.3-1.6 mm</quantity>
, 2130 segments (seven complete mature specimens). Width 5585 Mm at V (mean = 59 Mm ± 11,
<emphasis id="B947B7571F5F697DFB8922E36704FD7C" box="[1131,1151,719,740]" italics="true" pageId="4" pageNumber="534">N</emphasis>
= 24), 50120 Mm at XII (mean = 72 Mm ± 19,
<emphasis id="B947B7571F5F697DFBD122C1673CFC9A" box="[1075,1095,749,770]" italics="true" pageId="4" pageNumber="534">N</emphasis>
= 24). Prostomium rounded, 3550 Mm long, 2038 Mm wide at base. Body wall not papillate, cuticle smooth. Clitellum poorly developed, limited to large, swollen, transparent cells, only visible on living animals (
<figureCitation id="130877C01F5F697DFB97234B6793FCE4" box="[1141,1256,871,893]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="4" pageNumber="534">Fig. 1C, c</figureCitation>
), budding on a mid-ventral depression of body wall (
<figureCitation id="130877C01F5F697DFAE423AA6627FC03" box="[1286,1372,902,924]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="4" pageNumber="534">Fig. 2D</figureCitation>
, vdp) that extends behind genital pores, between ventral setae in XII and XIII.
</paragraph>
<paragraph id="8B8C6B451F5F697DFCB123CD66DAF92F" blockId="4.[826,1441,657,1903]" pageId="4" pageNumber="534">
Dorsal (dorsolateral) and ventral (ventrolateral) setae present from III, posteriorly situated in each segment (
<figureCitation id="130877C01F5F697DFC482433675FFBAC" box="[938,1060,1055,1077]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="4" pageNumber="534">Figs 1A, B</figureCitation>
,
<figureCitation id="130877C01F5F697DFBD324336728FBAC" box="[1073,1107,1055,1076]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="4" pageNumber="534">2B</figureCitation>
, ds, vs). Dorsal bundles with two (three) straight, thin single-pointed needles (15 28 Mm long), distal end bluntly pointed, alternating with (one) two thin and flexible hair setae (length 150230 Mm at V, 200260 Mm at VIII, 170225 Mm at XI). Hair setae densely pilose (with soft hairlets) on one side of the shaft (pilosity mostly observed on specimens during study using SEM). Ventral bundles with (two) three (four) sigmoid, bifid crotchets with upper tooth slightly thinner and shorter than lower tooth, and accompanied by one (two) pilose hair setae in preclitellar segments (80170 Mm long); ventral hair setae absent from XII; crotchets without nodulus, 1728 Mm long, shaft out of the body wall faintly pilose, with distal half enlarged on the convex side. Within ventral bundles, the uppermost seta bifid, followed by one hair seta; these two setae separated from the other two bifid setae by a distance sometimes greater than that between them. One modified genital seta per bundle in XII, 5560 Mm long, straight with enlarged, spearhead ectal tip, the latter with a spoon-like curvature (
<figureCitation id="130877C01F5F697DFB66268D6785F92F" box="[1156,1278,1697,1719]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="4" pageNumber="534">Figs 1A, B</figureCitation>
,
<figureCitation id="130877C01F5F697DFAE8268D6615F92F" box="[1290,1390,1697,1719]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="4" pageNumber="534">2A, C, D</figureCitation>
, gs).
</paragraph>
<paragraph id="8B8C6B451F5F697CFCB126EC6195F94D" blockId="4.[826,1441,657,1903]" lastBlockId="5.[144,759,1329,1903]" lastPageId="5" lastPageNumber="535" pageId="4" pageNumber="534">
No eyes. Brain long, extending into IV, posteriorly indented. Ventral nerve cord in contact with epidermis, beneath muscles of the body wall. Digestive tract complete; eversible pharynx with dorsal pad, sinuous, ciliated oesophagus, with pharyngeal glands present in IV- VI, followed by a simple intestine, slowly dilating from X- XI and ending in a terminal anus. Layer of chloragogen cells surrounding digestive tract possibly present; details not observable in specimens thus far examined. Coelomocytes absent. Dorsal glandular body-wall pouches very small and difficult to observe, but observed in VIII and XII in
<specimenCount id="9D35A0CC1F5E697CFD6025E763BFFA67" pageId="5" pageNumber="535" type="generic">one specimen</specimenCount>
), opening mid-dorsally at about the intersection with setal line. Voluminous glands associated with genital setae (2048 Mm long; 2032 Mm wide), bulging at the body surface; fine internal ducts running inside the glands and converging on the shaft of genital setae; setal glands asymmetrically arranged on each lateral side of the body, the left gland anteriad to the right one (
<figureCitation id="130877C01F5E697CFDEC26EC61F3F94D" box="[526,648,1728,1750]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535">Figs 1AC</figureCitation>
,
<figureCitation id="130877C01F5E697CFD7426EC61C3F94D" box="[662,696,1728,1749]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="5" pageNumber="535">2B</figureCitation>
, sg).
</paragraph>
<caption id="DF4C3BCD1F5E697CFF72244D60EDFB72" ID-DOI="http://doi.org/10.5281/zenodo.5408989" ID-Zenodo-Dep="5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535" startId="5.[144,223,1121,1140]" targetBox="[147,1421,200,1089]" targetPageId="5">
<paragraph id="8B8C6B451F5E697CFF72244D60EDFB72" blockId="5.[144,1421,1121,1258]" pageId="5" pageNumber="535">
<emphasis id="B947B7571F5E697CFF72244D61F0FBED" bold="true" box="[144,651,1121,1141]" pageId="5" pageNumber="535">
Figure 1.
<taxonomicName id="4C3310C61F5E697CFEE6244D6157FBED" authority="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar, 2012" authorityName="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar" authorityYear="2012" box="[260,556,1121,1141]" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="5" pageNumber="535" phylum="Annelida" rank="species" species="meyssonnieri" status="sp. nov.">
<emphasis id="B947B7571F5E697CFEE6244D6157FBED" bold="true" box="[260,556,1121,1141]" italics="true" pageId="5" pageNumber="535">Parvidrilus meyssonnieri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2740A2C1F5E697CFDD6244E61F0FBED" box="[564,651,1122,1141]" pageId="5" pageNumber="535" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
A, left lateral view of genitalia in segments X- XII (holotype). B, ventral view of genitalia in segments XI- XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm.
</paragraph>
</caption>
<paragraph id="8B8C6B451F5E697FFF4A26F26745F8F7" blockId="5.[144,759,1329,1903]" lastBlockId="6.[826,1441,1575,1903]" lastPageId="6" lastPageNumber="536" pageId="5" pageNumber="535">
Clearly delimited testes not present; however, a lump of cells with small nuclei observed dorsally in XI, in posterior end of coelom of XI, is probably unpaired male protogonia (
<figureCitation id="130877C01F5E697CFE2E27166142F8C8" box="[460,569,1850,1872]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535">Fig. 1A, t</figureCitation>
). Diffuse formation of male gametes in XI with many morulae (or cysts according to
<bibRefCitation id="EFA216B41F5E697CFBE0251E67CCFAD0" author="Ferraguti M" box="[1026,1207,1330,1352]" pageId="5" pageNumber="535" pagination="125 - 182" refId="ref19578" refString="Ferraguti M. 1997. Euclitellata. In: Jamieson BGM, Adiyodi RG, eds. Progress in male gamete ultrastructure and phylogeny. New Delhi, Calcutta: Oxford &amp; IBH Publishing Co. Pvt. Ltd, 125 - 182." type="book chapter" year="1997">Ferraguti, 1997</bibRefCitation>
), 2.84.0 Mm wide, freely floating inside the coelomic cavity although usually lying ventrally in XI (
<figureCitation id="130877C01F5E697CFB422543678EFA1C" box="[1184,1269,1391,1413]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535">Fig. 1A</figureCitation>
, mo); a few globular cytophores in a more dorsal and posterior position in XI, 8.011.3 Mm wide, sometimes in a stage of disintegration with the detachment of spermatozoa in the coelom. Sperm funnels and vasa deferentia not seen. Atria elongate (90190 Mm long, 710 Mm wide), either extending into XIII or restricted to XII; in the latter case, atria folded in different configurations: one atrium bent anteriad, with distal end folded up posteriad, the other atrium (or if present, both atria), bent posteriad, with distal end folded up anteriad (
<figureCitation id="130877C01F5E697CFAD6268D602DF94D" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535">Fig. 1A, B, a</figureCitation>
). Atria merging below the nerve cord (
<figureCitation id="130877C01F5E697CFAF126EC66F3F94D" box="[1299,1416,1728,1750]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535">Fig. 1A, B</figureCitation>
, nc) into a common ejaculatory duct, and opening at the tip of a mid-ventral, conical porophore (
<figureCitation id="130877C01F5E697CFAF226D166F3F88A" box="[1296,1416,1789,1811]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="5" pageNumber="535">Figs 1A, B</figureCitation>
,
<figureCitation id="130877C01F5E697CFCC427306016F8A9" box="[806,877,1820,1841]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="5" pageNumber="535">2A, C</figureCitation>
, mp), in posterior part of XII, between setal glands and in front of a median ventral depression of body wall in XII- XIII (
<figureCitation id="130877C01F5E697CFBD0277567F3F8F6" box="[1074,1160,1881,1903]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="5" pageNumber="535">Fig. 2D</figureCitation>
, vdp). Atrial ampullae longitudinally striated, with large lumen filled with sperm, mature spermatozoa more concentrated near the distal part of ampullae (
<figureCitation id="130877C01F5D697FFE3D26486128F9E1" box="[479,595,1636,1658]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="6" pageNumber="536">Fig. 1A, B</figureCitation>
, sp); no peculiar aggregate of sperm in the ampullae. An (unpaired?) ovary, apparently developing on one side, observed in anterior part of XII, with a ventral attachment on 11/12 (
<figureCitation id="130877C01F5D697FFF1326F26213F96C" box="[241,360,1758,1780]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="6" pageNumber="536">Fig. 1A, B</figureCitation>
, ov). Specimen ovigerous with large mature eggs filling coelom in XII, and extending through XIII. Female funnels not observed. No typical spermatheca observed. However, an unpaired, spherical ampulla in XII, 10 Mm in diameter, ventrally located, ahead of the point of union of atrial ampullae, usually on the right side of the ventral nervous chord but sometimes on the left side in some specimens (
<figureCitation id="130877C01F5D697FFCA026AF60CCF900" box="[834,951,1667,1689]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="6" pageNumber="536">Fig. 1AC</figureCitation>
, am); ampulla opening, through a duct, 14 Mm long, mid-ventrally, close to the basis of male porophore (
<figureCitation id="130877C01F5D697FFC5C26EC6769F94D" box="[958,1042,1728,1750]" captionStart="Figure 2" captionStartId="6.[164,243,1329,1348]" captionTargetBox="[164,1444,196,1300]" captionTargetId="figure-182@6.[164,1444,196,1300]" captionTargetPageId="6" captionText="Figure 2. Parvidrilus meyssonnieri sp. nov.; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="6" pageNumber="536">Fig. 2C</figureCitation>
, amo); duct surrounded by a muscular ring near opening. Spherical ampulla filled with cell material (sperm?) and surrounded by a cell cluster, clearly delimitated, noticeable on living worms but difficult to resolve on mounted slides, possibly glandular (
<figureCitation id="130877C01F5D697FFC592775674AF8F7" box="[955,1073,1881,1903]" captionStart="Figure 1" captionStartId="5.[144,223,1121,1140]" captionTargetBox="[147,1421,200,1089]" captionTargetId="figure-342@5.[144,1424,197,1092]" captionTargetPageId="5" captionText="Figure 1. Parvidrilus meyssonnieri sp. nov. A, left lateral view of genitalia in segments X-XII (holotype). B, ventral view of genitalia in segments XI-XIII (paratype MHNL 3.2). C, hand drawings of the unpaired spherical ampulla and associated cell cluster (from living animal). Abbreviations: a, atrium; am, spherical ampulla; c, clitellum; ccl, cell cluster; ds, dorsal seta; gs, genital seta; mo, morula; mp, male pore; nc, nerve cord; ov, oocyte; sg, setal gland; sp, spermatozoa; st, septum; t, testis (or germ cells); vs, ventral seta. Scale bars = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.5408989" httpUri="https://zenodo.org/record/5408989/files/figure.png" pageId="6" pageNumber="536">Fig. 1B, C</figureCitation>
).
</paragraph>
<caption id="DF4C3BCD1F5D697FFF46251D66F5FA6C" ID-DOI="http://doi.org/10.5281/zenodo.5408991" ID-Zenodo-Dep="5408991" httpUri="https://zenodo.org/record/5408991/files/figure.png" pageId="6" pageNumber="536" startId="6.[164,243,1329,1348]" targetBox="[164,1444,196,1300]" targetPageId="6">
<paragraph id="8B8C6B451F5D697FFF46251D66F5FA6C" blockId="6.[164,1442,1329,1524]" pageId="6" pageNumber="536">
<emphasis id="B947B7571F5D697FFF46251D61DFFADC" bold="true" box="[164,676,1329,1348]" pageId="6" pageNumber="536">
Figure 2.
<taxonomicName id="4C3310C61F5D697FFEFB251D6139FADC" authority="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar, 2012" authorityName="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar" authorityYear="2012" box="[281,578,1329,1348]" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="536" phylum="Annelida" rank="species" species="meyssonnieri" status="sp. nov.">
<emphasis id="B947B7571F5D697FFEFB251D6139FADC" bold="true" box="[281,578,1329,1348]" italics="true" pageId="6" pageNumber="536">Parvidrilus meyssonnieri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2740A2C1F5D697FFDA9251D61DFFADC" box="[587,676,1329,1348]" pageId="6" pageNumber="536" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
; scanning electron micrographs. A, left lateral view of anterior part of body showing dorsal and ventral setal bundles, and the absence of setae in II. B, ventral and dorsal setae. C, left lateral view of genital region showing a genital seta and its associated gland (damaged), the single male pore, and the opening of the spherical ampulla. D, left lateral view of genital region showing the left voluminous setal gland, the median ventral depression behind the porophore (not visible on the picture), and the left genital seta in XII (gs). Specimens from Navogne (AC) and La Martinière (D) sites. Abbreviations: amo, spherical ampulla opening; ds, dorsal setae; gs, genital seta; mp, male pore; sg, setal gland; vdp, ventral depression; vs, ventral setae. Scale bars: A = 100 Mm; BD = 20 Mm.
</paragraph>
</caption>
<paragraph id="8B8C6B451F5C697EFF7220EA6276FE81" blockId="7.[144,759,198,281]" pageId="7" pageNumber="537">
<emphasis id="B947B7571F5C697EFF7220EA6240FF44" box="[144,315,198,220]" italics="true" pageId="7" pageNumber="537">COI sequences:</emphasis>
<materialsCitation id="3B5B61181F5C697EFEA620EA6271FE81" accessionNumber="HE800204, HE800205" collectionCode="EMBL" collectorName="La Martiniere' &amp; Navogne'" pageId="7" pageNumber="537" specimenCount="1">
<collectionCode id="ED22F3801F5C697EFEA620EA62F4FF43" box="[324,399,198,219]" pageId="7" pageNumber="537">EMBL</collectionCode>
accession numbers:
<accessionNumber id="9460F6A61F5C697EFD9C20EA618DFF43" box="[638,758,198,219]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/HE800204" pageId="7" pageNumber="537">HE800204</accessionNumber>
(gallery of
<collectorName id="26C60E931F5C697EFEC320C962AEFF62" box="[289,469,228,250]" pageId="7" pageNumber="537">La Martinière</collectorName>
);
<accessionNumber id="9460F6A61F5C697EFE1320C86112FF62" box="[497,617,228,250]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/HE800205" pageId="7" pageNumber="537">HE800205</accessionNumber>
(gallery of
<collectorName id="26C60E931F5C697EFF76212F6279FE81" box="[148,258,259,281]" pageId="7" pageNumber="537">Navogne</collectorName>
)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8C6B451F5C697EFF72216367DCFECE" blockId="7.[144,759,335,1213]" lastBlockId="7.[806,1421,198,342]" pageId="7" pageNumber="537">
<emphasis id="B947B7571F5C697EFF7221636380FEFC" box="[144,251,335,356]" italics="true" pageId="7" pageNumber="537">Remarks:</emphasis>
Within the parvidrilid species known so far (including all new species being described herein plus the reassigned
<taxonomicName id="4C3310C61F5C697EFEAB21A06161FE39" authorityName="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar" authorityYear="2012" baseAuthorityName="Juget" baseAuthorityYear="1959" box="[329,538,396,417]" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="537" phylum="Annelida" rank="species" species="gineti">
<emphasis id="B947B7571F5C697EFEAB21A06161FE39" box="[329,538,396,417]" italics="true" pageId="7" pageNumber="537">Parvidrilus gineti</emphasis>
</taxonomicName>
see below), the presence of genital setae is unique to
<taxonomicName id="4C3310C61F5C697EFDAA2187618CFE58" authority="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar, 2012" authorityName="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar" authorityYear="2012" box="[584,759,427,448]" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="537" phylum="Annelida" rank="species" species="meyssonnieri" status="sp. nov.">
<emphasis id="B947B7571F5C697EFDAA2187618CFE58" box="[584,759,427,448]" italics="true" pageId="7" pageNumber="537">P. meyssonnieri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2740A2C1F5C697EFF7221E56397FE46" box="[144,236,457,478]" pageId="7" pageNumber="537" rank="species">sp. nov.</taxonomicNameLabel>
clearly separating this species from its congeners. The most outstanding feature in
<taxonomicName id="4C3310C61F5C697EFD7921C46397FD84" authority="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar, 2012" authorityName="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar" authorityYear="2012" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="537" phylum="Annelida" rank="species" species="meyssonnieri" status="sp. nov.">
<emphasis id="B947B7571F5C697EFD7921C46397FD84" italics="true" pageId="7" pageNumber="537">P. meyssonnieri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2740A2C1F5C697EFF19222B6222FD84" box="[251,345,519,540]" pageId="7" pageNumber="537" rank="species">sp. nov.</taxonomicNameLabel>
is the presence of an enigmatic unpaired, spherical ampulla in XII the function of which remains obscure. There is a possibility that this structure is used for storing sperm from a concopulant, which would imply that it is a spermatheca. However, the ampulla is surrounded by a cell cluster, which seems more glandular (prostate-like cells) than shaped in order to play the role of a bag for sperm storage. In this respect, such a structure is somewhat similar to the glandular, atrium-like organ in the pre-atrial segment of some
<taxonomicName id="4C3310C61F5C697EFE3F231463BAFCF4" authority="(Rhynchelmis)" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" kingdom="Animalia" order="Lumbriculida" pageId="7" pageNumber="537" phylum="Annelida" rank="subGenus" subGenus="Rhynchelmis">
<emphasis id="B947B7571F5C697EFE3F2314610BFCD5" box="[477,624,824,845]" italics="true" pageId="7" pageNumber="537">Rhynchelmis</emphasis>
(
<emphasis id="B947B7571F5C697EFD66231463C2FCF4" italics="true" pageId="7" pageNumber="537">Rhynchelmis</emphasis>
)
</taxonomicName>
species (
<taxonomicName id="4C3310C61F5C697EFEC5237B62A9FCF5" box="[295,466,855,877]" class="Clitellata" family="Lumbriculidae" kingdom="Animalia" order="Lumbriculida" pageId="7" pageNumber="537" phylum="Annelida" rank="family">Lumbriculidae</taxonomicName>
), referred to as accessory organ (formerly rudimentary atrium), whose function and derivation are unknown (
<bibRefCitation id="EFA216B41F5C697EFDF623B863B0FC51" author="Fend SV &amp; Brinkhurst RO" pageId="7" pageNumber="537" pagination="1 - 27" refId="ref19548" refString="Fend SV, Brinkhurst RO. 2010. Contributions towards a review of the genus Rhynchelmis Hoffmeister (Clitellata: Lumbriculidae). Zootaxa 2407: 1 - 27." type="journal article" year="2010">Fend &amp; Brinkhurst, 2010</bibRefCitation>
). Some material has been observed in the spherical ampulla but it does not resemble mature spermatozoa. Hence, the absence of spermathecae in this species seems plausible. Conversely, considering the ampulla and its surrounding cell cluster as an atrium and a prostate is another possibility implying that the structures described above as atria are in fact the spermathecae and the porophore is spermathecal. However, prostates seem to be absent in
<taxonomicName id="4C3310C61F5C697EFD96248B618CFB24" authorityName="Erseus" authorityYear="1999" box="[628,759,1191,1212]" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="537" phylum="Annelida" rank="genus">
<emphasis id="B947B7571F5C697EFD96248B618CFB24" box="[628,759,1191,1212]" italics="true" pageId="7" pageNumber="537">Parvidrilus</emphasis>
</taxonomicName>
(see below). In addition, when copulatory organs are present in microdriles whether in the form of porophores or penes they are always associated with the male aperture (
<bibRefCitation id="EFA216B41F5C697EFC3D210E67CAFEA0" author="Stephenson J" box="[991,1201,290,312]" pageId="7" pageNumber="537" refId="ref21083" refString="Stephenson J. 1930. The Oligochaeta. Oxford: Oxford University Press." type="book" year="1930">Stephenson, 1930</bibRefCitation>
), thus making the latter hypothesis less convincing.
</paragraph>
<paragraph id="8B8C6B451F5C697EFCC421A96671FC91" blockId="7.[806,1422,389,778]" pageId="7" pageNumber="537">
<emphasis id="B947B7571F5C697EFCC421A966F6FE02" box="[806,1421,389,410]" italics="true" pageId="7" pageNumber="537">
Distribution and habitat:
<taxonomicName id="4C3310C61F5C697EFB8521A966F6FE02" authorityName="Martínez-Ansemil &amp; Châtelliers &amp; Martin &amp; Sambugar" authorityYear="2012" box="[1127,1421,389,410]" class="Clitellata" family="Parvidrilidae" genus="Parvidrilus" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="537" phylum="Annelida" rank="species" species="meyssonnieri">Parvidrilus meyssonnieri</taxonomicName>
</emphasis>
is known only from the La Martinière and Navogne galleries in the Rhône department,
<collectingCountry id="F3242BD51F5C697EFB0D21EE663AFE4F" box="[1263,1345,450,471]" name="France" pageId="7" pageNumber="537">France</collectingCountry>
, in a region lying on metamorphic bedrock. The La Martinière gallery opens in the upper alteration layer of a crystalline craton, which forms granitic sand with scarce, particulate organic matter. A rivulet runs along the gallery and passes slowly through a thin layer of coarse sand. Historically (between 1846 and 1919) the Navogne gallery was dug in arkoses and ferruginous sandstone for iron ore exploitation. The water is present in the gallery as isolated puddles with gravelly soil, rich in organic matter.
</paragraph>
</subSubSection>
</treatment>
</document>