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<document id="E1C417C226E687C7C558A6B7650BDF1C" ID-DOI="10.11646/zootaxa.3664.1.1" ID-GBIF-Dataset="e5ce6301-e878-4ddd-9309-c56373686f59" ID-ISSN="1175-5326" ID-Zenodo-Dep="283786" ID-ZooBank="C8136C89-7787-477D-BC64-97AA6C16057B" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460613860799" checkinUser="plazi" docAuthor="Rodriguez, Pilar &amp; Fend, Steven V." docDate="2013" docId="154FD01D4B41FFC4FF6E9766BD04CCDB" docLanguage="en" docName="zt03664p044.pdf" docOrigin="Zootaxa 3664 (1)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Rhyacodrilus propiporus Rodriguez &amp; Fend, 2013, sp. n." docType="treatment" docVersion="6" lastPageNumber="24" masterDocId="E976A8654B53FFD3FFF99271B97ACD14" masterDocTitle="New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879)" masterLastPageNumber="44" masterPageNumber="1" pageNumber="19" updateTime="1698329309631" updateUser="plazi">
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<mods:title id="552B6CF165701FA0351E7F062F10B0EA">New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879)</mods:title>
</mods:titleInfo>
<mods:name id="3ACB121E2CBDE14E9E9655B320E6E1A0" type="personal">
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<mods:namePart id="B8024B702A13EAF47994A89100EABBE4">Rodriguez, Pilar</mods:namePart>
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<mods:namePart id="23362E38D800DCF21E411E89985B42EE">Fend, Steven V.</mods:namePart>
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<mods:typeOfResource id="9727A755A2D93FE96D42366068E97B9D">text</mods:typeOfResource>
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<mods:title id="AF68336DAD955D6D8F22C77559E44F5D">Zootaxa</mods:title>
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<mods:part id="E317F456C96D1E0FE0997B5E871DB88A">
<mods:date id="67EA4FC7A84260D83C0D8AD18760A5CD">2013</mods:date>
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<mods:number id="441036C3A62E859B34E5961ADB41EE5A">3664</mods:number>
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<mods:identifier id="B90E4F7CA50F235EE79C9D9E13BD0D5E" type="DOI">10.11646/zootaxa.3664.1.1</mods:identifier>
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<treatment id="154FD01D4B41FFC4FF6E9766BD04CCDB" ID-DOI="http://doi.org/10.5281/zenodo.6160663" ID-GBIF-Taxon="127664168" ID-Zenodo-Dep="6160663" LSID="urn:lsid:plazi:treatment:154FD01D4B41FFC4FF6E9766BD04CCDB" httpUri="http://treatment.plazi.org/id/154FD01D4B41FFC4FF6E9766BD04CCDB" lastPageId="23" lastPageNumber="24" pageId="18" pageNumber="19">
<subSubSection id="D5FC32804B41FFC1FF6E9766B86DC846" pageId="18" pageNumber="19" type="nomenclature">
<paragraph id="9D59610B4B41FFC1FF6E9766BB77C825" blockId="18.[151,525,1303,1362]" box="[151,525,1303,1329]" pageId="18" pageNumber="19">
<heading id="C611D6674B41FFC1FF6E9766BB77C825" bold="true" box="[151,525,1303,1329]" fontSize="11" level="1" pageId="18" pageNumber="19" reason="1">
<emphasis id="AF92BD194B41FFC1FF6E9766BB77C825" bold="true" box="[151,525,1303,1329]" pageId="18" pageNumber="19">
<taxonomicName id="5AE61A884B41FFC1FF6E9766B8B9C825" box="[151,451,1303,1329]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="19" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B41FFC1FF6E9766B8B9C825" bold="true" box="[151,451,1303,1329]" italics="true" pageId="18" pageNumber="19">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B4A100624B41FFC1FE329766BB77C825" box="[459,525,1303,1329]" pageId="18" pageNumber="19" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="9D59610B4B41FFC1FF6E974BB86DC846" blockId="18.[151,525,1303,1362]" box="[151,279,1338,1362]" pageId="18" pageNumber="19">
(
<figureCitation id="05DD7D8E4B41FFC1FF66974BB86AC846" box="[159,272,1338,1362]" captionStart-0="FIGURE 9" captionStart-1="FIGURE 10" captionStart-2="FIGURE 11" captionStartId-0="19.[151,250,1853,1876]" captionStartId-1="20.[151,250,1609,1632]" captionStartId-2="21.[151,250,1385,1408]" captionTargetBox-0="[159,1428,782,1815]" captionTargetBox-1="[169,1422,204,1573]" captionTargetBox-2="[168,1422,213,1345]" captionTargetId-0="figure@19.[151,1436,769,1832]" captionTargetId-1="figure@20.[151,1436,193,1588]" captionTargetId-2="figure@21.[151,1436,193,1364]" captionTargetPageId-0="19" captionTargetPageId-1="20" captionTargetPageId-2="21" captionText-0="FIGURE 9. Rhyacodrilus propiporus sp. n. (A J, M N) and Rhyacodrilus subterraneus Hrabë, 1963 (K L). A J. Rhyacodrilus propiporus sp. n. A. Prostomium with coelomocytes. B E. Dorsal chaetae of segment II (B), V (C), IX (D), and postclitellar (E). F I. Ventral chaetae of segment V (F), VII (G), postclitellar (H) and very posterior (I). J. Dorsal chaetae of very posterior segments. K L. Chaetae of Rhyacodrilus subterraneus (Martínez-Ansemil's collection, see text). K. Anterior pectinates. L. Ventrals. M N. Rhyacodrilus propiporus sp. n. Ventral view of the body, whole mount. M. Spermathecal pores and male pores. N. Atrium opening in a porophore." captionText-1="FIGURE 10. Rhyacodrilus propiporus sp. n. A. Schematic outline of anterior part of the body and clitellar region. B. Schematic drawing of reproductive organs. C. Histological (sagittal) section of spermatheca. D F. Consecutive series of histological sections of atrial ampulla and vas deferens." captionText-2="FIGURE 11. Rhyacodrilus propiporus sp. n. A B Schematic drawings of reproductive organs in population from Lower Little River. C. Ventral view of spermathecae and atria showing musculature associated with the pores. D. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen)." httpUri-0="https://zenodo.org/record/283796/files/figure.png" httpUri-1="https://zenodo.org/record/283797/files/figure.png" httpUri-2="https://zenodo.org/record/283798/files/figure.png" pageId="18" pageNumber="19">Figs 911</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B41FFC1FF6E97F2B83FCA7A" pageId="18" pageNumber="19" type="materials_examined">
<paragraph id="9D59610B4B41FFC1FF6E97F2BC3CC88E" blockId="18.[151,1436,1410,2011]" box="[151,1350,1410,1435]" pageId="18" pageNumber="19">
<typeStatus id="425DDFA94B41FFC1FF6E97F2B87EC88F" box="[151,260,1411,1435]" pageId="18" pageNumber="19" type="holotype">
<emphasis id="AF92BD194B41FFC1FF6E97F2B87EC88F" bold="true" box="[151,260,1411,1435]" pageId="18" pageNumber="19">Holotype</emphasis>
</typeStatus>
. USNM 1202062, whole-mounted worm stained in hematoxylin and mounted in
<collectingCountry id="E5F1219B4B41FFC1FB6D97F3BD92C88E" box="[1172,1256,1410,1434]" name="Canada" pageId="18" pageNumber="19">Canada</collectingCountry>
balsam.
</paragraph>
<paragraph id="9D59610B4B41FFC1FF3097D6B852CB12" blockId="18.[151,1436,1410,2011]" pageId="18" pageNumber="19">
<emphasis id="AF92BD194B41FFC1FF3097D6B83BC8AB" bold="true" box="[201,321,1447,1471]" pageId="18" pageNumber="19">
<typeStatus id="425DDFA94B41FFC1FF3097D6B83BC8AB" box="[201,321,1447,1471]" pageId="18" pageNumber="19" type="paratype">Paratypes</typeStatus>
</emphasis>
. USNM 1202063-65: 1 dissected specimen (
<date id="E95847CB4B41FFC1FCBC97D7BD6FC8AA" box="[837,1045,1446,1470]" pageId="18" pageNumber="19" value="2007-02-14">14 February 2007</date>
), 1 sagittally sectioned (
<date id="E95847CB4B41FFC1FAD497D7B9A9C8F6" pageId="18" pageNumber="19" value="2007-03-17">17 March 2007</date>
), 1 whole mount (
<date id="E95847CB4B41FFC1FE5B97BBBB29C8F6" box="[418,595,1482,1506]" pageId="18" pageNumber="19" value="2007-03-17">17 March 2007</date>
), MNCN 16.03/3081: 1 dissected specimen (
<date id="E95847CB4B41FFC1FBA897BBBC79C8F6" box="[1105,1283,1482,1506]" pageId="18" pageNumber="19" value="2007-03-17">17 March 2007</date>
), all from the
<typeStatus id="425DDFA94B41FFC1FF6E979EB9BDCB13" box="[151,199,1519,1543]" pageId="18" pageNumber="19">type</typeStatus>
locality.
</paragraph>
<paragraph id="9D59610B4B41FFC1FF309462BC02CB3E" blockId="18.[151,1436,1410,2011]" box="[201,1400,1554,1579]" pageId="18" pageNumber="19">
<emphasis id="AF92BD194B41FFC1FF309462B825CB3F" bold="true" box="[201,351,1554,1579]" pageId="18" pageNumber="19">
<typeStatus id="425DDFA94B41FFC1FF309462B878CB3F" box="[201,258,1555,1579]" pageId="18" pageNumber="19">Type</typeStatus>
locality
</emphasis>
. Beech Swamp, Halifax Co., North Carolina,
<collectingCountry id="E5F1219B4B41FFC1FC9D9463BAE5CB3E" box="[868,927,1554,1578]" name="United States of America" pageId="18" pageNumber="19">USA</collectingCountry>
,
<geoCoordinate id="F8D207CC4B41FFC1FC539463BD5FCB3E" box="[938,1061,1554,1578]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="36.1435">N36.1435°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FBD29463BDD6CB3E" box="[1067,1196,1554,1578]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-77.5551">W77.5551°</geoCoordinate>
(
<date id="E95847CB4B41FFC1FB459463BC16CB3E" box="[1212,1388,1554,1578]" pageId="18" pageNumber="19" value="2007-03-17">17 March 2007</date>
).
</paragraph>
<paragraph id="9D59610B4B41FFC1FF309447B83FCA7A" blockId="18.[151,1436,1410,2011]" pageId="18" pageNumber="19">
<emphasis id="AF92BD194B41FFC1FF309447B8FBCB5B" bold="true" box="[201,385,1590,1615]" pageId="18" pageNumber="19">Other material</emphasis>
. 2 dissected and 1 whole mount from Lower Little River at road SR 2031, Harnett Co.,
<geoCoordinate id="F8D207CC4B41FFC1FF6E942BB868CB66" box="[151,274,1626,1650]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="35.2599">N35.2599°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FEE2942BB8E5CB66" box="[283,415,1626,1650]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-78.823">W78.8230°</geoCoordinate>
, (
<date id="E95847CB4B41FFC1FE4C942BBB3FCB66" box="[437,581,1626,1650]" pageId="18" pageNumber="19" value="2011-06-05">5 June 2011</date>
); 4 dissected, 2 whole mounts, 1 sagittally sectioned (
<date id="E95847CB4B41FFC1FB4C942BBC51CB66" box="[1205,1323,1626,1650]" pageId="18" pageNumber="19" value="2011-07">July 2011</date>
). 2 whole mounts from Lower Little River at highway NC 217,
<geoCoordinate id="F8D207CC4B41FFC1FCFD940FBA05CB82" box="[772,895,1662,1686]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="35.2637">N35.2637°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FC71940FBD76CB82" box="[904,1036,1662,1686]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-78.7406">W78.7406°</geoCoordinate>
, (
<date id="E95847CB4B41FFC1FBDB940FBDB5CB82" box="[1058,1231,1662,1687]" pageId="18" pageNumber="19" value="2011-08-05">5 August 2011</date>
). 6 whole mounts from Lower Little River at highway NC 87, Cumberland Co.,
<geoCoordinate id="F8D207CC4B41FFC1FCAB94D3BAB7CBAE" box="[850,973,1698,1722]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="35.4001">N35.4001°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FC2D94D3BD29CBAE" box="[980,1107,1698,1722]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-79.1126">W79.1126°</geoCoordinate>
(
<date id="E95847CB4B41FFC1FB9A94D3BC77CBAE" box="[1123,1293,1698,1723]" pageId="18" pageNumber="19" value="2011-08-05">5 August 2011</date>
). 2 dissected from Powell's Creek at road SR 1338, Halifax Co.,
<geoCoordinate id="F8D207CC4B41FFC1FD1194B7BA19CBCA" box="[744,867,1734,1758]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="36.1791">N36.1791°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FC9594B7BA96CBCA" box="[876,1004,1734,1758]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-77.898">W77.8980°</geoCoordinate>
(
<date id="E95847CB4B41FFC1FC0794B7BDA6CBCA" box="[1022,1244,1734,1758]" pageId="18" pageNumber="19" value="2011-12-13">13 December 2011</date>
). 1 whole mount from Friar Swamp at road SR 1740, Columbus Co.,
<geoCoordinate id="F8D207CC4B41FFC1FD11949BBA19CA16" box="[744,867,1770,1794]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="34.3695">N34.3695°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FC92949BBA91CA16" box="[875,1003,1770,1794]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-78.4572">W78.4572°</geoCoordinate>
(
<date id="E95847CB4B41FFC1FC07949BBDDACA16" box="[1022,1184,1770,1794]" pageId="18" pageNumber="19" value="1998-03-01">1 March 1998</date>
). 1 whole mount from seep at Pettiford Creek, Carteret Co., approx.
<geoCoordinate id="F8D207CC4B41FFC1FD6D957FBB89CA32" box="[660,755,1806,1830]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="555" value="34.75">N34.75°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FD00957FBA27CA32" box="[761,861,1806,1830]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="555" value="-77.02">W77.02°</geoCoordinate>
(
<date id="E95847CB4B41FFC1FC95957FBD51CA32" box="[876,1067,1806,1831]" pageId="18" pageNumber="19" value="2009-01-30">30 January 2009</date>
). 1 whole mount from Chinkapin Creek, Hertford Co.,
<geoCoordinate id="F8D207CC4B41FFC1FE7C9543BB7ACA5E" box="[389,512,1842,1866]" direction="north" orientation="latitude" pageId="18" pageNumber="19" precision="5" value="36.2534">N36.2534°</geoCoordinate>
<geoCoordinate id="F8D207CC4B41FFC1FDFE9543BBF2CA5E" box="[519,648,1842,1866]" direction="west" orientation="longitude" pageId="18" pageNumber="19" precision="5" value="-76.8491">W76.8491°</geoCoordinate>
(
<date id="E95847CB4B41FFC1FD639543BA43CA5E" box="[666,825,1842,1866]" pageId="18" pageNumber="19" value="2001-03-01">1 March 2001</date>
). All material collected by David R. Lenat from North Carolina,
<collectingCountry id="E5F1219B4B41FFC1FEFC9527B83BCA7A" box="[261,321,1878,1902]" name="United States of America" pageId="18" pageNumber="19">USA</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B41FFC1FF30950ABA2ECAA2" pageId="18" pageNumber="19" type="etymology">
<paragraph id="9D59610B4B41FFC1FF30950ABA2ECAA2" blockId="18.[151,1436,1410,2011]" pageId="18" pageNumber="19">
<emphasis id="AF92BD194B41FFC1FF30950AB832CA87" bold="true" box="[201,328,1915,1939]" pageId="18" pageNumber="19">Etymology</emphasis>
. From the Latin
<emphasis id="AF92BD194B41FFC1FDF6950DBB35CA87" box="[527,591,1916,1939]" italics="true" pageId="18" pageNumber="19">prope</emphasis>
for close and
<emphasis id="AF92BD194B41FFC1FD0E950DBA4DCA87" box="[759,823,1916,1939]" italics="true" pageId="18" pageNumber="19">porus</emphasis>
for pores. The specific name refers to the very close (nearly median) position of both spermathecal and male pores.
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B41FFC5FF3095B3B8B0CEB0" lastPageId="22" lastPageNumber="23" pageId="18" pageNumber="19" type="description">
<paragraph id="9D59610B4B41FFC0FF3095B3BA86CDC0" blockId="18.[151,1436,1410,2011]" lastBlockId="19.[151,1436,151,716]" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
<emphasis id="AF92BD194B41FFC1FF3095B3B82ECACF" bold="true" box="[201,340,1986,2011]" pageId="18" pageNumber="19">Description</emphasis>
(from
<typeStatus id="425DDFA94B41FFC1FE5295B2B8A1CACF" box="[427,475,1987,2011]" pageId="18" pageNumber="19">type</typeStatus>
locality specimens). Number of segments 6374. Body diameter 440456 µm in segment VIII, 556614 µm in XI. Rounded-conical prostomium 107202 µm long. Epidermis 713 µm high, 25 45 µm at clitellum. Clitellum from the level of chaetae in X to the end of XII.
</paragraph>
<paragraph id="9D59610B4B40FFC0FF3092AEBB23CCDB" blockId="19.[151,1436,151,716]" pageId="19" pageNumber="20">
Dorsal bundles in anterior segments with 03 smooth hair chaetae (229242 µm long) and 34 bifid, nonpectinate chaetae (103107 µm long) with distal tooth longer than proximal (
<figureCitation id="05DD7D8E4B40FFC0FC0E9375BD40CC08" box="[1015,1082,260,284]" captionStart="FIGURE 9" captionStartId="19.[151,250,1853,1876]" captionTargetBox="[159,1428,782,1815]" captionTargetId="figure@19.[151,1436,769,1832]" captionTargetPageId="19" captionText="FIGURE 9. Rhyacodrilus propiporus sp. n. (A J, M N) and Rhyacodrilus subterraneus Hrabë, 1963 (K L). A J. Rhyacodrilus propiporus sp. n. A. Prostomium with coelomocytes. B E. Dorsal chaetae of segment II (B), V (C), IX (D), and postclitellar (E). F I. Ventral chaetae of segment V (F), VII (G), postclitellar (H) and very posterior (I). J. Dorsal chaetae of very posterior segments. K L. Chaetae of Rhyacodrilus subterraneus (Martínez-Ansemil's collection, see text). K. Anterior pectinates. L. Ventrals. M N. Rhyacodrilus propiporus sp. n. Ventral view of the body, whole mount. M. Spermathecal pores and male pores. N. Atrium opening in a porophore." httpUri="https://zenodo.org/record/283796/files/figure.png" pageId="19" pageNumber="20">Fig. 9</figureCitation>
BE). Hair chaetae absent from segment VIII (or IX) backwards. Posterior dorsal chaetae like ventral chaetae (69110 µm long). Ventral chaetae bifid; anterior ventral chaetae 46 per bundle (73110 µm long) with distal tooth longer than proximal; posterior ventral chaetae 24 (commonly 3) per bundle, 8899 µm long, and progressively shorter, down to 7074 µm long in the most posterior segments of complete individuals, distal tooth becoming equal to or shorter than proximal in the most posterior segments (
<figureCitation id="05DD7D8E4B40FFC0FE1993C6BB58CCDB" box="[480,546,439,463]" captionStart="FIGURE 9" captionStartId="19.[151,250,1853,1876]" captionTargetBox="[159,1428,782,1815]" captionTargetId="figure@19.[151,1436,769,1832]" captionTargetPageId="19" captionText="FIGURE 9. Rhyacodrilus propiporus sp. n. (A J, M N) and Rhyacodrilus subterraneus Hrabë, 1963 (K L). A J. Rhyacodrilus propiporus sp. n. A. Prostomium with coelomocytes. B E. Dorsal chaetae of segment II (B), V (C), IX (D), and postclitellar (E). F I. Ventral chaetae of segment V (F), VII (G), postclitellar (H) and very posterior (I). J. Dorsal chaetae of very posterior segments. K L. Chaetae of Rhyacodrilus subterraneus (Martínez-Ansemil's collection, see text). K. Anterior pectinates. L. Ventrals. M N. Rhyacodrilus propiporus sp. n. Ventral view of the body, whole mount. M. Spermathecal pores and male pores. N. Atrium opening in a porophore." httpUri="https://zenodo.org/record/283796/files/figure.png" pageId="19" pageNumber="20">Fig. 9</figureCitation>
FI).
</paragraph>
<paragraph id="9D59610B4B40FFC0FF3093ADBA54CFD8" blockId="19.[151,1436,151,716]" pageId="19" pageNumber="20">
One pair spermathecal pores at the beginning of segment X, close to the mid-ventral line of the body, usually within a shallow, common groove about 150 μm wide; the spermathecal pores surrounded by a thin, muscular ring at the body wall (similar to
<figureCitation id="05DD7D8E4B40FFC0FE379055BB67CF28" box="[462,541,548,572]" captionStart="FIGURE 11" captionStartId="21.[151,250,1385,1408]" captionTargetBox="[168,1422,213,1345]" captionTargetId="figure@21.[151,1436,193,1364]" captionTargetPageId="21" captionText="FIGURE 11. Rhyacodrilus propiporus sp. n. A B Schematic drawings of reproductive organs in population from Lower Little River. C. Ventral view of spermathecae and atria showing musculature associated with the pores. D. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen)." httpUri="https://zenodo.org/record/283798/files/figure.png" pageId="19" pageNumber="20">Fig. 11</figureCitation>
C). One pair male pores at about the posterior 1/4 of segment XI, close together near the mid-ventral line, on a prominent porophore 80120 μm wide and up to 45 μm high, lined with a thin, circular muscle layer (
<figureCitation id="05DD7D8E4B40FFC0FE65901DB89DCF90" box="[412,487,620,644]" captionStart="FIGURE 9" captionStartId="19.[151,250,1853,1876]" captionTargetBox="[159,1428,782,1815]" captionTargetId="figure@19.[151,1436,769,1832]" captionTargetPageId="19" captionText="FIGURE 9. Rhyacodrilus propiporus sp. n. (A J, M N) and Rhyacodrilus subterraneus Hrabë, 1963 (K L). A J. Rhyacodrilus propiporus sp. n. A. Prostomium with coelomocytes. B E. Dorsal chaetae of segment II (B), V (C), IX (D), and postclitellar (E). F I. Ventral chaetae of segment V (F), VII (G), postclitellar (H) and very posterior (I). J. Dorsal chaetae of very posterior segments. K L. Chaetae of Rhyacodrilus subterraneus (Martínez-Ansemil's collection, see text). K. Anterior pectinates. L. Ventrals. M N. Rhyacodrilus propiporus sp. n. Ventral view of the body, whole mount. M. Spermathecal pores and male pores. N. Atrium opening in a porophore." httpUri="https://zenodo.org/record/283796/files/figure.png" pageId="19" pageNumber="20">Figs 9</figureCitation>
M, N, 10A). No modified genital chaetae. Ventral chaetae usually absent in X at maturity; in one mated worm a single chaeta was not modified and resembled other ventral chaetae (79 µm long). Female pores in intersegment 11/12, just below lateral line.
</paragraph>
<caption id="C99931834B40FFC0FF6E954CBBE4CAF9" httpUri="https://zenodo.org/record/283796/files/figure.png" pageId="19" pageNumber="20" targetBox="[159,1428,782,1815]" targetPageId="19">
<paragraph id="9D59610B4B40FFC0FF6E954CBBE4CAF9" blockId="19.[151,1436,1853,2029]" pageId="19" pageNumber="20">
<emphasis id="AF92BD194B40FFC0FF6E954CB861CA47" bold="true" box="[151,283,1853,1876]" pageId="19" pageNumber="20">FIGURE 9.</emphasis>
<taxonomicName id="5AE61A884B40FFC0FED0954FBB54CA40" box="[297,558,1854,1876]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B40FFC0FED0954FBB54CA40" box="[297,558,1854,1876]" italics="true" pageId="19" pageNumber="20">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B40FFC0FDC4954FBB04CA47" bold="true" box="[573,638,1854,1875]" pageId="19" pageNumber="20">
<taxonomicNameLabel id="B4A100624B40FFC0FDC4954FBB04CA47" box="[573,638,1854,1875]" pageId="19" pageNumber="20" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
(
<emphasis id="AF92BD194B40FFC0FD6C954FBBDDCA47" bold="true" box="[661,679,1854,1875]" pageId="19" pageNumber="20">A</emphasis>
<emphasis id="AF92BD194B40FFC0FD4D954FBBBBCA47" bold="true" box="[692,705,1854,1875]" pageId="19" pageNumber="20">J</emphasis>
,
<emphasis id="AF92BD194B40FFC0FD2F954FBB94CA47" bold="true" box="[726,750,1854,1875]" pageId="19" pageNumber="20">M</emphasis>
<emphasis id="AF92BD194B40FFC0FD03954FBA76CA47" bold="true" box="[762,780,1854,1875]" pageId="19" pageNumber="20">N</emphasis>
) and
<taxonomicName id="5AE61A884B40FFC0FCAE954FBC7FCA47" authority="Hrabe, 1963" authorityName="Hrabe" authorityYear="1963" box="[855,1285,1853,1876]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B40FFC0FCAE954FBD0ACA40" box="[855,1136,1854,1876]" italics="true" pageId="19" pageNumber="20">Rhyacodrilus subterraneus</emphasis>
Hrabë, 1963
</taxonomicName>
(
<emphasis id="AF92BD194B40FFC0FAE2954FBC55CA47" bold="true" box="[1307,1327,1854,1875]" pageId="19" pageNumber="20">K</emphasis>
<emphasis id="AF92BD194B40FFC0FAC5954FBC37CA47" bold="true" box="[1340,1357,1854,1875]" pageId="19" pageNumber="20">L</emphasis>
).
<emphasis id="AF92BD194B40FFC0FA93954FBC06CA47" bold="true" box="[1386,1404,1854,1875]" pageId="19" pageNumber="20">A</emphasis>
<emphasis id="AF92BD194B40FFC0FA70954FBCE6CA47" bold="true" box="[1417,1436,1854,1875]" pageId="19" pageNumber="20">J.</emphasis>
<taxonomicName id="5AE61A884B40FFC0FF6E952DB8EFCA66" box="[151,405,1884,1906]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B40FFC0FF6E952DB8EFCA66" box="[151,405,1884,1906]" italics="true" pageId="19" pageNumber="20">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B40FFC0FE62952CB897CA65" bold="true" box="[411,493,1884,1906]" pageId="19" pageNumber="20">
<taxonomicNameLabel id="B4A100624B40FFC0FE62952CB8AECA66" box="[411,468,1885,1906]" pageId="19" pageNumber="20" rank="species">sp. n.</taxonomicNameLabel>
A
</emphasis>
. Prostomium with coelomocytes.
<emphasis id="AF92BD194B40FFC0FCB0952CBA0DCA66" bold="true" box="[841,887,1885,1906]" pageId="19" pageNumber="20">BE</emphasis>
. Dorsal chaetae of segment II (B), V (C), IX (D), and postclitellar (E).
<emphasis id="AF92BD194B40FFC0FEBD950AB810CA84" bold="true" box="[324,362,1915,1936]" pageId="19" pageNumber="20">FI</emphasis>
. Ventral chaetae of segment V (F), VII (G), postclitellar (H) and very posterior (I).
<emphasis id="AF92BD194B40FFC0FB34950ABDA0CA84" bold="true" box="[1229,1242,1915,1936]" pageId="19" pageNumber="20">J</emphasis>
. Dorsal chaetae of very posterior segments.
<emphasis id="AF92BD194B40FFC0FE6595EBB8B7CABB" bold="true" box="[412,461,1946,1967]" pageId="19" pageNumber="20">KL</emphasis>
. Chaetae of
<taxonomicName id="5AE61A884B40FFC0FDAD95E8BA12CABB" box="[596,872,1945,1967]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B40FFC0FDAD95E8BA12CABB" box="[596,872,1945,1967]" italics="true" pageId="19" pageNumber="20">Rhyacodrilus subterraneus</emphasis>
</taxonomicName>
(Martínez-Ansemil's collection, see text).
<emphasis id="AF92BD194B40FFC0FADD95EBBC42CABB" bold="true" box="[1316,1336,1946,1967]" pageId="19" pageNumber="20">K</emphasis>
. Anterior pectinates.
<emphasis id="AF92BD194B40FFC0FEF095C8B860CADA" bold="true" box="[265,282,1977,1998]" pageId="19" pageNumber="20">L</emphasis>
. Ventrals.
<emphasis id="AF92BD194B40FFC0FE7195C8B8C5CADA" bold="true" box="[392,447,1977,1998]" pageId="19" pageNumber="20">MN</emphasis>
.
<taxonomicName id="5AE61A884B40FFC0FE3595C9BBB1CADA" box="[460,715,1976,1998]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B40FFC0FE3595C9BBB1CADA" box="[460,715,1976,1998]" italics="true" pageId="19" pageNumber="20">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B40FFC0FD2895C8BA71CADA" bold="true" box="[721,779,1977,1998]" pageId="19" pageNumber="20">
<taxonomicNameLabel id="B4A100624B40FFC0FD2895C8BA71CADA" box="[721,779,1977,1998]" pageId="19" pageNumber="20" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
Ventral view of the body, whole mount.
<emphasis id="AF92BD194B40FFC0FB5695C8BDBDCADA" bold="true" box="[1199,1223,1977,1998]" pageId="19" pageNumber="20">M</emphasis>
. Spermathecal pores and male pores.
<emphasis id="AF92BD194B40FFC0FEC595A6B834CAF8" bold="true" box="[316,334,2007,2028]" pageId="19" pageNumber="20">N</emphasis>
. Atrium opening in a porophore.
</paragraph>
</caption>
<caption id="C99931834B47FFC7FF6E9438BBB3CB88" httpUri="https://zenodo.org/record/283797/files/figure.png" pageId="20" pageNumber="21" targetBox="[169,1422,204,1573]" targetPageId="20">
<paragraph id="9D59610B4B47FFC7FF6E9438BBB3CB88" blockId="20.[151,1436,1609,1692]" pageId="20" pageNumber="21">
<emphasis id="AF92BD194B47FFC7FF6E9438B85ECB4A" bold="true" box="[151,292,1609,1632]" pageId="20" pageNumber="21">FIGURE 10.</emphasis>
<taxonomicName id="5AE61A884B47FFC7FEC99438BB48CB4B" box="[304,562,1609,1631]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="20" pageNumber="21" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B47FFC7FEC99438BB48CB4B" box="[304,562,1609,1631]" italics="true" pageId="20" pageNumber="21">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B47FFC7FDC4943BBBE3CB4A" bold="true" box="[573,665,1609,1631]" pageId="20" pageNumber="21">
<taxonomicNameLabel id="B4A100624B47FFC7FDC4943BBB06CB4B" box="[573,636,1610,1631]" pageId="20" pageNumber="21" rank="species">sp. n.</taxonomicNameLabel>
A
</emphasis>
. Schematic outline of anterior part of the body and clitellar region.
<emphasis id="AF92BD194B47FFC7FA7D943BBCEFCB4B" bold="true" box="[1412,1429,1610,1631]" pageId="20" pageNumber="21">B</emphasis>
. Schematic drawing of reproductive organs.
<emphasis id="AF92BD194B47FFC7FD909416BB01CB6A" bold="true" box="[617,635,1639,1662]" pageId="20" pageNumber="21">C</emphasis>
. Histological (sagittal) section of spermatheca.
<emphasis id="AF92BD194B47FFC7FB859419BDD2CB69" bold="true" box="[1148,1192,1640,1661]" pageId="20" pageNumber="21">DF</emphasis>
. Consecutive series of histological sections of atrial ampulla and vas deferens.
</paragraph>
</caption>
<paragraph id="9D59610B4B47FFC7FF3094B5BB0ACAC3" blockId="20.[151,1437,1732,2007]" pageId="20" pageNumber="21">
Abundant nucleated coelomocytes (diameter 1024 µm), containing large granules (
<figureCitation id="05DD7D8E4B47FFC7FB5F94B5BD94CBC8" box="[1190,1262,1732,1756]" captionStart="FIGURE 9" captionStartId="19.[151,250,1853,1876]" captionTargetBox="[159,1428,782,1815]" captionTargetId="figure@19.[151,1436,769,1832]" captionTargetPageId="19" captionText="FIGURE 9. Rhyacodrilus propiporus sp. n. (A J, M N) and Rhyacodrilus subterraneus Hrabë, 1963 (K L). A J. Rhyacodrilus propiporus sp. n. A. Prostomium with coelomocytes. B E. Dorsal chaetae of segment II (B), V (C), IX (D), and postclitellar (E). F I. Ventral chaetae of segment V (F), VII (G), postclitellar (H) and very posterior (I). J. Dorsal chaetae of very posterior segments. K L. Chaetae of Rhyacodrilus subterraneus (Martínez-Ansemil's collection, see text). K. Anterior pectinates. L. Ventrals. M N. Rhyacodrilus propiporus sp. n. Ventral view of the body, whole mount. M. Spermathecal pores and male pores. N. Atrium opening in a porophore." httpUri="https://zenodo.org/record/283796/files/figure.png" pageId="20" pageNumber="21">Fig. 9</figureCitation>
A). Pharyngeal glands in V and poorly developed in IV and VI. Chloragogenous tissue scantily covering the gut from VII backwards, and formed by a thin layer of cells (usually &lt;20 µm high). One pair testes in segment X and one pair ovaries in segment XI. Sperm sac back to segment XV, egg sac back to segment XVI. Spermathecae composed of a short duct (4377 µm long and 5864 µm maximum diameter) and an ampulla (154246 µm long, 138215 µm wide) in the most anterior part of segment X (
<figureCitation id="05DD7D8E4B47FFC7FD6A9506BB9ECA9B" box="[659,740,1911,1935]" captionStart="FIGURE 10" captionStartId="20.[151,250,1609,1632]" captionTargetBox="[169,1422,204,1573]" captionTargetId="figure@20.[151,1436,193,1588]" captionTargetPageId="20" captionText="FIGURE 10. Rhyacodrilus propiporus sp. n. A. Schematic outline of anterior part of the body and clitellar region. B. Schematic drawing of reproductive organs. C. Histological (sagittal) section of spermatheca. D F. Consecutive series of histological sections of atrial ampulla and vas deferens." httpUri="https://zenodo.org/record/283797/files/figure.png" pageId="20" pageNumber="21">Fig. 10</figureCitation>
B, C). Duct lined with columnar epithelium, with a narrow (5 10 μm) lumen and very thin muscle layer. Ampulla with thin layer of cuboidal epithelium (621 μm); lumen with sperm typically arranged in loose clusters.
</paragraph>
<caption id="C99931834B46FFC6FF6E9718BAE4C8A9" httpUri="https://zenodo.org/record/283798/files/figure.png" pageId="21" pageNumber="22" targetBox="[168,1422,213,1345]" targetPageId="21">
<paragraph id="9D59610B4B46FFC6FF6E9718BAE4C8A9" blockId="21.[151,1436,1385,1469]" pageId="21" pageNumber="22">
<emphasis id="AF92BD194B46FFC6FF6E9718B864C86B" bold="true" box="[151,286,1385,1408]" pageId="21" pageNumber="22">FIGURE 11.</emphasis>
<taxonomicName id="5AE61A884B46FFC6FEDD971BBB5BC894" box="[292,545,1386,1408]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B46FFC6FEDD971BBB5BC894" box="[292,545,1386,1408]" italics="true" pageId="21" pageNumber="22">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B46FFC6FDDE971BBBEFC86B" bold="true" box="[551,661,1386,1407]" pageId="21" pageNumber="22">
<taxonomicNameLabel id="B4A100624B46FFC6FDDE971BBB1AC86B" box="[551,608,1386,1407]" pageId="21" pageNumber="22" rank="species">sp. n.</taxonomicNameLabel>
AB
</emphasis>
Schematic drawings of reproductive organs in population from Lower Little River.
<emphasis id="AF92BD194B46FFC6FF2497F9B995C88B" bold="true" box="[221,239,1416,1439]" pageId="21" pageNumber="22">C</emphasis>
. Ventral view of spermathecae and atria showing musculature associated with the pores.
<emphasis id="AF92BD194B46FFC6FB6E97F8BDD3C88A" bold="true" box="[1175,1193,1417,1438]" pageId="21" pageNumber="22">D</emphasis>
. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen).
</paragraph>
</caption>
<paragraph id="9D59610B4B46FFC6FF309798B8FFCBEA" blockId="21.[151,1436,1513,2007]" pageId="21" pageNumber="22">
Male duct consisting of a sperm funnel at the most ventral part of septum 10/11, and a relatively straight vas deferens which joins the atrial ampulla subapically; vas deferens of homogeneous width (diameter 2324 µm), longer than atrium (ca. 230315 µm long) (
<figureCitation id="05DD7D8E4B46FFC6FD719440BBA6CB5D" box="[648,732,1585,1609]" captionStart="FIGURE 10" captionStartId="20.[151,250,1609,1632]" captionTargetBox="[169,1422,204,1573]" captionTargetId="figure@20.[151,1436,193,1588]" captionTargetPageId="20" captionText="FIGURE 10. Rhyacodrilus propiporus sp. n. A. Schematic outline of anterior part of the body and clitellar region. B. Schematic drawing of reproductive organs. C. Histological (sagittal) section of spermatheca. D F. Consecutive series of histological sections of atrial ampulla and vas deferens." httpUri="https://zenodo.org/record/283797/files/figure.png" pageId="21" pageNumber="22">Fig. 10</figureCitation>
B). Atria formed by a spherical ampulla (diameter 5775 µm) and a duct (78125 µm long, 43 µm maximum diameter) about 1.52 times longer than the ampulla; duct tapering towards the male pore and usually towards the ampulla too. Penis absent. Atrial ampulla with a thin epithelium (7 8 µm) and very thin musculature, covered by a diffuse layer of prostate cells (up to 1014 µm high), not forming bundles of cells (
<figureCitation id="05DD7D8E4B46FFC6FEA694B0B8C8CBCD" box="[351,434,1729,1753]" captionStart="FIGURE 10" captionStartId="20.[151,250,1609,1632]" captionTargetBox="[169,1422,204,1573]" captionTargetId="figure@20.[151,1436,193,1588]" captionTargetPageId="20" captionText="FIGURE 10. Rhyacodrilus propiporus sp. n. A. Schematic outline of anterior part of the body and clitellar region. B. Schematic drawing of reproductive organs. C. Histological (sagittal) section of spermatheca. D F. Consecutive series of histological sections of atrial ampulla and vas deferens." httpUri="https://zenodo.org/record/283797/files/figure.png" pageId="21" pageNumber="22">Fig. 10</figureCitation>
DF). The relative size of the atrium is very small, total length less than one third the diameter of the body.
</paragraph>
<paragraph id="9D59610B4B46FFC5FF30957BB87DCC09" blockId="21.[151,1436,1513,2007]" lastBlockId="22.[151,1437,151,2012]" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22">
<emphasis id="AF92BD194B46FFC6FF30957BBB4DCA36" bold="true" box="[201,567,1801,1826]" pageId="21" pageNumber="22">Lower Little River population</emphasis>
(supplemental description, differences from
<typeStatus id="425DDFA94B46FFC6FBC1957BBD12CA36" box="[1080,1128,1802,1826]" pageId="21" pageNumber="22">type</typeStatus>
locality population). Body dimensions generally smaller than Beech Swamp population (
<typeStatus id="425DDFA94B46FFC6FCAB955FBAFECA52" box="[850,900,1838,1862]" pageId="21" pageNumber="22">type</typeStatus>
locality): number of segments 5372, diameter 270400 µm in segment VIII and 360490 µm in XI (slide-mounted worms). Chaetae similar, but usually shorter than in Beech Swamp population: anterior dorsal bundles with 13 smooth hair chaetae (maximum 200260 µm long, shortest in II and X) and 34 bifid chaetae (85110 µm long) with distal tooth slightly or up to 2 times longer than proximal. Hair chaetae absent from segment XXII backwards. Posterior dorsal chaetae like ventral chaetae (both 85100 µm long). Anterior ventral chaetae 48 per bundle (95115 µm long) with distal tooth longer than proximal (sometimes with a thin intermediate tooth). Posterior to clitellum 25 ventral chaetae; commonly 3 per bundle in posterior half of worm, with distal tooth becoming equal or shorter than proximal in the most posterior segments.
</paragraph>
<paragraph id="9D59610B4B45FFC5FF309356BC53CC93" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
Spermathecal pores usually within a shallow, common groove 5070 μm wide, pores surrounded by a thin, circular muscle layer ectal to the ventral muscle bands of the body wall (
<figureCitation id="05DD7D8E4B45FFC5FC3D933DBD6ECC70" box="[964,1044,332,356]" captionStart="FIGURE 11" captionStartId="21.[151,250,1385,1408]" captionTargetBox="[168,1422,213,1345]" captionTargetId="figure@21.[151,1436,193,1364]" captionTargetPageId="21" captionText="FIGURE 11. Rhyacodrilus propiporus sp. n. A B Schematic drawings of reproductive organs in population from Lower Little River. C. Ventral view of spermathecae and atria showing musculature associated with the pores. D. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen)." httpUri="https://zenodo.org/record/283798/files/figure.png" pageId="22" pageNumber="23">Fig. 11</figureCitation>
C). Male pores on a low porophore about 180240 μm wide, and up to about 40 μm high; porophore lined with a thin, circular muscle layer.
</paragraph>
<paragraph id="9D59610B4B45FFC5FF3093E5BA89CF90" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
Spermathecal duct may be cylindrical and up to twice as long as wide, or short and conical, about as wide as long entally (
<figureCitation id="05DD7D8E4B45FFC5FED793C6B804CCDB" box="[302,382,439,463]" captionStart="FIGURE 11" captionStartId="21.[151,250,1385,1408]" captionTargetBox="[168,1422,213,1345]" captionTargetId="figure@21.[151,1436,193,1364]" captionTargetPageId="21" captionText="FIGURE 11. Rhyacodrilus propiporus sp. n. A B Schematic drawings of reproductive organs in population from Lower Little River. C. Ventral view of spermathecae and atria showing musculature associated with the pores. D. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen)." httpUri="https://zenodo.org/record/283798/files/figure.png" pageId="22" pageNumber="23">Fig. 11</figureCitation>
A, B) (length 5596 µm, width 3862 µm). Spermathecal ampulla ovate (125245 µm long, 100180 µm wide in mated worms), usually entirely within X. Atrial ampulla spherical to slightly ovate (60145 µm wide); duct cylindrical or tapered (55104 µm long, 3042 µm maximum diameter); total length of atrium 120225 µm, about one third to half the diameter of the body at segment XI; relative lengths of duct and ampulla variable, but usually about equal (
<figureCitation id="05DD7D8E4B45FFC5FDCF9036BBF1CF4B" box="[566,651,583,607]" captionStart="FIGURE 11" captionStartId="21.[151,250,1385,1408]" captionTargetBox="[168,1422,213,1345]" captionTargetId="figure@21.[151,1436,193,1364]" captionTargetPageId="21" captionText="FIGURE 11. Rhyacodrilus propiporus sp. n. A B Schematic drawings of reproductive organs in population from Lower Little River. C. Ventral view of spermathecae and atria showing musculature associated with the pores. D. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen)." httpUri="https://zenodo.org/record/283798/files/figure.png" pageId="22" pageNumber="23">Fig. 11</figureCitation>
A, B). Atrial ampulla with epithelium 1020 µm high, very thin musculature, and covered by a diffuse (usually single) layer of prostate cells.
</paragraph>
<paragraph id="9D59610B4B45FFC5FF3090FEBD0DCFFB" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
The other listed populations from different locations in North Carolina are represented by limited material, and have been provisionally ascribed to
<taxonomicName id="5AE61A884B45FFC5FDC890C4BBB6CFD8" box="[561,716,693,716]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus">
<emphasis id="AF92BD194B45FFC5FDC890C4BBB6CFD8" box="[561,716,693,716]" italics="true" pageId="22" pageNumber="23">R. propiporus</emphasis>
</taxonomicName>
. Specimens from Powell's Creek had very similar reproductive organs, but were unmated and had unmodified ventral chaetae in segment XI (
<figureCitation id="05DD7D8E4B45FFC5FBFA90A6BD28CFFB" box="[1027,1106,727,751]" captionStart="FIGURE 11" captionStartId="21.[151,250,1385,1408]" captionTargetBox="[168,1422,213,1345]" captionTargetId="figure@21.[151,1436,193,1364]" captionTargetPageId="21" captionText="FIGURE 11. Rhyacodrilus propiporus sp. n. A B Schematic drawings of reproductive organs in population from Lower Little River. C. Ventral view of spermathecae and atria showing musculature associated with the pores. D. Schematic drawing of reproductive organs in population from Powell's Creek (unmated specimen)." httpUri="https://zenodo.org/record/283798/files/figure.png" pageId="22" pageNumber="23">Fig. 11</figureCitation>
D).
</paragraph>
<paragraph id="9D59610B4B45FFC5FF30908AB8B0CEB0" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
<emphasis id="AF92BD194B45FFC5FF30908AB83DCE00" bold="true" box="[201,327,763,788]" pageId="22" pageNumber="23">Anomalies</emphasis>
. One Lower Little River specimen had paired spermathecae in VII and VIII, paired atria in VIII, and a single atrium in IX; only the second male duct had sperm on the funnel. Another specimen had normal reproductive organs, but they were shifted forward by two segments. Six mated specimens had 1 or 2 unmodified chaetae in XI, but these were presumably remnant somatic chaetae, as they were on the line of ventral chaetae, and were not medially directed.
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B45FFC5FF3091DEB981C818" pageId="22" pageNumber="23" type="distribution">
<paragraph id="9D59610B4B45FFC5FF3091DEB8EFC968" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
<emphasis id="AF92BD194B45FFC5FF3091DEB895CEDC" bold="true" box="[201,495,943,968]" pageId="22" pageNumber="23">Distribution and habitat</emphasis>
.
<taxonomicName id="5AE61A884B45FFC5FE0691C1BA63CEDC" box="[511,793,944,968]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B45FFC5FE0691C1BA63CEDC" box="[511,793,944,968]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B45FFC5FCD891C1BA18CEDC" bold="true" box="[801,866,944,968]" pageId="22" pageNumber="23">
<taxonomicNameLabel id="B4A100624B45FFC5FCD891C1BA18CEDC" box="[801,866,944,968]" pageId="22" pageNumber="23" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
was collected in diverse habitats in eastern North Carolina. Beech Swamp (the
<typeStatus id="425DDFA94B45FFC5FE2791A4BB74CEF9" box="[478,526,981,1005]" pageId="22" pageNumber="23">type</typeStatus>
locality) and Powell's Creek are in the Tar River basin, in the northeastern part of the State.
<taxonomicName id="5AE61A884B45FFC5FEFD9189B8E3C904" box="[260,409,1016,1040]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="AF92BD194B45FFC5FEFD9189B8E3C904" box="[260,409,1016,1040]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus</emphasis>
</taxonomicName>
habitats were small (
<quantity id="5A1ECCEE4B45FFC5FD7E9189BBC8C904" box="[647,690,1016,1040]" metricMagnitude="0" metricUnit="m" metricValue="2.0" pageId="22" pageNumber="23" unit="m" value="2.0">2 m</quantity>
wide), sandy streams that typically go dry during summer months. Both streams appear to have good water quality based on the macroinvertebrate composition. Beech Swamp has a diverse oligochaete community, and is the only known location for the monotypic
<taxonomicName id="5AE61A884B45FFC5FBAD9631B8EBC968" authority="Fend and Lenat, 2007" authorityName="Fend and Lenat" authorityYear="2007" class="Clitellata" family="Lumbriculidae" genus="Pilaridrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="AF92BD194B45FFC5FBAD9631BDADC94C" box="[1108,1239,1088,1112]" italics="true" pageId="22" pageNumber="23">Pilaridrilus</emphasis>
Fend and Lenat, 2007 (Lumbriculidae)
</taxonomicName>
.
</paragraph>
<paragraph id="9D59610B4B45FFC5FF3096F9B981C818" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
The Little River is in southeastern North Carolina, in the Cape Fear River basin. It is a large (
<quantity id="5A1ECCEE4B45FFC5FAF696F9BC36C9B4" box="[1295,1356,1160,1184]" metricMagnitude="1" metricUnit="m" metricValue="2.0" pageId="22" pageNumber="23" unit="m" value="20.0">20 m</quantity>
wide), permanent stream, with low gradient and excellent water quality based on the macroinvertebrate community (D.R. Lenat, pers. comm. 2012).
<taxonomicName id="5AE61A884B45FFC5FE3C96A1BBA7C9FC" box="[453,733,1232,1256]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus">
<emphasis id="AF92BD194B45FFC5FE3C96A1BBA7C9FC" box="[453,733,1232,1256]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
was usually collected from sand and gravel near the middle of the river.
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B45FFC4FF309769BD04CCDB" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23" type="discussion">
<paragraph id="9D59610B4B45FFC5FF309769BB0DCBA8" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
<emphasis id="AF92BD194B45FFC5FF309769B84CC824" bold="true" box="[201,310,1304,1328]" pageId="22" pageNumber="23">Remarks</emphasis>
.
<taxonomicName id="5AE61A884B45FFC5FEBD9769BB27C824" box="[324,605,1304,1328]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B45FFC5FEBD9769BB27C824" box="[324,605,1304,1328]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B45FFC5FD9D9769BBDEC824" bold="true" box="[612,676,1304,1328]" pageId="22" pageNumber="23">
<taxonomicNameLabel id="B4A100624B45FFC5FD9D9769BBDEC824" box="[612,676,1304,1328]" pageId="22" pageNumber="23" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
belongs to a small group of
<taxonomicName id="5AE61A884B45FFC5FC1F9769BD06C824" box="[998,1148,1304,1328]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="AF92BD194B45FFC5FC1F9769BD06C824" box="[998,1148,1304,1328]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus</emphasis>
</taxonomicName>
species with hair chaetae in dorsal bundles and a layer of prostate cells covering the atrial ampulla, and lacking both modified spermathecal and penial chaetae. This group is formed by
<taxonomicName id="5AE61A884B45FFC5FD649710BAA4C86C" authority="Malevich, 1949" authorityName="Malevich" authorityYear="1949" box="[669,990,1376,1400]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="lepnevae">
<emphasis id="AF92BD194B45FFC5FD649710BA58C86C" box="[669,802,1376,1400]" italics="true" pageId="22" pageNumber="23">R. lepnevae</emphasis>
Malevich, 1949
</taxonomicName>
emend. Hrabë (1974) (
<collectingCountry id="E5F1219B4B45FFC5FB15972EBC46C863" box="[1260,1340,1375,1399]" name="Russia" pageId="22" pageNumber="23">Russia</collectingCountry>
), and
<taxonomicName id="5AE61A884B45FFC5FA7D9710B8F4C888" authority="Hrabe, 1963" authorityName="Hrabe" authorityYear="1963" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="lindbergi">
<emphasis id="AF92BD194B45FFC5FA7D9710B987C888" italics="true" pageId="22" pageNumber="23">R. lindbergi</emphasis>
Hrabë, 1963
</taxonomicName>
a (described from one immature specimen from a cave in
<collectingCountry id="E5F1219B4B45FFC5FBE197F5BD06C888" box="[1048,1148,1412,1436]" name="Portugal" pageId="22" pageNumber="23">Portugal</collectingCountry>
). Martínez-Ansemil
<emphasis id="AF92BD194B45FFC5FA9D97F4BCE6C888" box="[1380,1436,1412,1436]" italics="true" pageId="22" pageNumber="23">et al.</emphasis>
(1997) attributed a single specimen with penial chaetae from northwestern
<collectingCountry id="E5F1219B4B45FFC5FBFA97D6BD39C8AB" box="[1027,1091,1447,1471]" name="Spain" pageId="22" pageNumber="23">Spain</collectingCountry>
to
<taxonomicName id="5AE61A884B45FFC5FB9497D8BD8FC8D4" box="[1133,1269,1448,1472]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="lindbergi">
<emphasis id="AF92BD194B45FFC5FB9497D8BD8FC8D4" box="[1133,1269,1448,1472]" italics="true" pageId="22" pageNumber="23">R. lindbergi</emphasis>
</taxonomicName>
, although this character was not verified in the
<typeStatus id="425DDFA94B45FFC5FDDE97BCBB2DC8F1" box="[551,599,1485,1509]" pageId="22" pageNumber="23">type</typeStatus>
specimen (E. Martínez-Ansemil, pers. comm. 2011). Both species are distinguished externally from
<taxonomicName id="5AE61A884B45FFC5FE1E9780BBFBCB1C" box="[487,641,1521,1544]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus" status="sp. nov.">
<emphasis id="AF92BD194B45FFC5FE1E9780BBFBCB1C" box="[487,641,1521,1544]" italics="true" pageId="22" pageNumber="23">R. propiporus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B45FFC5FD719781BBB2CB1C" bold="true" box="[648,712,1520,1544]" pageId="22" pageNumber="23">
<taxonomicNameLabel id="B4A100624B45FFC5FD719781BBB2CB1C" box="[648,712,1520,1544]" pageId="22" pageNumber="23" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
by the presence of ventral bundles with both simple-pointed and bifid chaetae. Externally, the near-median positions of the spermathecal and male pores distinguish the new species from other Nearctic members of the genus. The new species is also well-separated from other Nearctic
<taxonomicName id="5AE61A884B45FFC5FF6E942DB857CB60" box="[151,301,1628,1652]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="AF92BD194B45FFC5FF6E942DB857CB60" box="[151,301,1628,1652]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus</emphasis>
</taxonomicName>
by the small size of the atrium and relative lengths of the atrial ampulla and duct (the latter being 1.5 to 2 times the length of the former), and also by the very short, one cell layer prostate covering the ampulla (unique among the Nearctic
<taxonomicName id="5AE61A884B45FFC5FE8094D5BB75CBA8" box="[377,527,1700,1724]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="AF92BD194B45FFC5FE8094D5BB75CBA8" box="[377,527,1700,1724]" italics="true" pageId="22" pageNumber="23">Rhyacodrilus</emphasis>
</taxonomicName>
species).
</paragraph>
<paragraph id="9D59610B4B45FFC5FF3094B6BB97CA64" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
The population from Lower Little River has been provisionally ascribed to
<taxonomicName id="5AE61A884B45FFC5FBCF94B8BDAFCBF4" box="[1078,1237,1737,1760]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus">
<emphasis id="AF92BD194B45FFC5FBCF94B8BDAFCBF4" box="[1078,1237,1737,1760]" italics="true" pageId="22" pageNumber="23">R. propiporus</emphasis>
</taxonomicName>
based on similar chaetal characteristics and the peculiar near-median position of male and spermathecal pores. The larger atrium, a relative length of atrial ampulla to duct of about 1, a cylindrical spermathecal duct, and ventral chaetae often present in segment X, distinguish this population from that of the
<typeStatus id="425DDFA94B45FFC5FC6D9544BABECA59" box="[916,964,1845,1869]" pageId="22" pageNumber="23">type</typeStatus>
locality, and its status may be subject to revision with new findings in eastern North
<collectingCountry id="E5F1219B4B45FFC5FD7D9529BB92CA64" box="[644,744,1880,1904]" name="United States of America" pageId="22" pageNumber="23">America</collectingCountry>
.
</paragraph>
<paragraph id="9D59610B4B45FFC5FF30950DBCE6CAC8" blockId="22.[151,1437,151,2012]" pageId="22" pageNumber="23">
This species may have a broader distribution than indicated here. The combination of dorsal hairs limited to anterior segments and anterior dorsal bifid chaetae with the distal tooth longer than the proximal would likely attribute immature
<taxonomicName id="5AE61A884B45FFC5FE7995B4BB5ECAC8" box="[384,548,1989,2012]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="propiporus">
<emphasis id="AF92BD194B45FFC5FE7995B4BB5ECAC8" box="[384,548,1989,2012]" italics="true" pageId="22" pageNumber="23">R. propiporus</emphasis>
</taxonomicName>
to
<taxonomicName id="5AE61A884B45FFC5FDA095B4BA6ECAC8" box="[601,788,1988,2012]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B45FFC5FDA095B4BA6ECAC8" box="[601,788,1988,2012]" italics="true" pageId="22" pageNumber="23">R. subterraneus</emphasis>
</taxonomicName>
using the keys by Kathman and Brinkhurst (1998).
</paragraph>
<paragraph id="9D59610B4B44FFC4FF6E92E6BD04CCDB" blockId="23.[151,1436,151,464]" pageId="23" pageNumber="24">
Examination of material collected by D. Strayer from several sites in eastern
<collectingCountry id="E5F1219B4B44FFC4FBE692E6BD20CDBB" box="[1055,1114,151,175]" name="United States of America" pageId="23" pageNumber="24">USA</collectingCountry>
, and cited by Kathman and Brinkhurst (1998) as
<taxonomicName id="5AE61A884B44FFC4FE7792CCBB38CDC0" box="[398,578,188,212]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B44FFC4FE7792CCBB38CDC0" box="[398,578,188,212]" italics="true" pageId="23" pageNumber="24">R. subterraneus</emphasis>
</taxonomicName>
, revealed no mature specimens, so the presence/absence of penial chaetae cannot be verified. All 6 individuals examined from the USNM collection (#
<date id="E95847CB4B44FFC4FBFB92AEBD23CDE3" box="[1026,1113,223,247]" pageId="23" pageNumber="24" value="32-10-28">102832</date>
, #187041 and #187042, leg. Strayer), plus 2 additional specimens from the Strayer collection, were immature, with upper tooth of dorsal chaetae thicker and much longer than lower (as in
<taxonomicName id="5AE61A884B44FFC4FD189358BAEFCC54" box="[737,917,296,320]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B44FFC4FD189358BAEFCC54" box="[737,917,296,320]" italics="true" pageId="23" pageNumber="24">R. subterraneus</emphasis>
</taxonomicName>
), and with short intermediate teeth visible in some individuals but not in others. In the absence of mature worms attributable to
<taxonomicName id="5AE61A884B44FFC4FBB2933CBD84CC70" box="[1099,1278,332,356]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B44FFC4FBB2933CBD84CC70" box="[1099,1278,332,356]" italics="true" pageId="23" pageNumber="24">R. subterraneus</emphasis>
</taxonomicName>
, we conclude that at least some of the North American records of
<taxonomicName id="5AE61A884B44FFC4FCFD9300BAC3CC9C" box="[772,953,368,392]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B44FFC4FCFD9300BAC3CC9C" box="[772,953,368,392]" italics="true" pageId="23" pageNumber="24">R. subterraneus</emphasis>
</taxonomicName>
based on chaetal morphology may be
<taxonomicName id="5AE61A884B44FFC4FA7D9300B869CCB8" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="propiporus">
<emphasis id="AF92BD194B44FFC4FA7D9300B869CCB8" italics="true" pageId="23" pageNumber="24">R. propiporus</emphasis>
</taxonomicName>
, and that an examination of the male ducts and/or molecular analyses are required to confirm the presence of
<taxonomicName id="5AE61A884B44FFC4FEE793C8B8B5CCC4" box="[286,463,440,464]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="subterraneus">
<emphasis id="AF92BD194B44FFC4FEE793C8B8B5CCC4" box="[286,463,440,464]" italics="true" pageId="23" pageNumber="24">R. subterraneus</emphasis>
</taxonomicName>
in North
<collectingCountry id="E5F1219B4B44FFC4FDC593C6BBE6CCDB" box="[572,668,439,463]" name="United States of America" pageId="23" pageNumber="24">America</collectingCountry>
(see also remarks for
<taxonomicName id="5AE61A884B44FFC4FC6D93C8BAA7CCC4" box="[916,989,440,464]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="clio" status="sp. nov.">
<emphasis id="AF92BD194B44FFC4FC6D93C8BAA7CCC4" box="[916,989,440,464]" italics="true" pageId="23" pageNumber="24">R. clio</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B44FFC4FC1D93C9BD59CCC4" bold="true" box="[996,1059,440,464]" pageId="23" pageNumber="24">
<taxonomicNameLabel id="B4A100624B44FFC4FC1D93C9BD59CCC4" box="[996,1059,440,464]" pageId="23" pageNumber="24" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
above).
</paragraph>
</subSubSection>
</treatment>
</document>