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<document id="D070E7FDCA0EF421A863179FB67AF601" 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="154FD01D4B54FFD8FF6E957EBBD7CE3B" 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 quileuticus Rodriguez &amp; Fend, 2013, sp. n." docType="treatment" docVersion="6" lastPageNumber="12" 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="8" updateTime="1698329309631" updateUser="plazi">
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<mods:title id="C937B47E18568353EE8A98CA0E58ACF0">New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879)</mods:title>
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<mods:name id="E5C333A73277B75BBC31E706AEC8CFA2" type="personal">
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<mods:namePart id="2969E84DCBBA13758FE43413D5387D07">Rodriguez, Pilar</mods:namePart>
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<mods:namePart id="42915523A36E56AC204723A6B9DE4657">Fend, Steven V.</mods:namePart>
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<mods:date id="80BFBB5AC068B702C7EFC2B812D6574B">2013</mods:date>
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<treatment id="154FD01D4B54FFD8FF6E957EBBD7CE3B" ID-DOI="http://doi.org/10.5281/zenodo.6160659" ID-GBIF-Taxon="127664164" ID-Zenodo-Dep="6160659" LSID="urn:lsid:plazi:treatment:154FD01D4B54FFD8FF6E957EBBD7CE3B" httpUri="http://treatment.plazi.org/id/154FD01D4B54FFD8FF6E957EBBD7CE3B" lastPageId="11" lastPageNumber="12" pageId="7" pageNumber="8">
<subSubSection id="D5FC32804B54FFD4FF6E957EB871CA5E" pageId="7" pageNumber="8" type="nomenclature">
<paragraph id="9D59610B4B54FFD4FF6E957EBB74CA3D" blockId="7.[151,526,1807,1866]" box="[151,526,1807,1833]" pageId="7" pageNumber="8">
<heading id="C611D6674B54FFD4FF6E957EBB74CA3D" bold="true" box="[151,526,1807,1833]" fontSize="11" level="1" pageId="7" pageNumber="8" reason="1">
<emphasis id="AF92BD194B54FFD4FF6E957EBB74CA3D" bold="true" box="[151,526,1807,1833]" pageId="7" pageNumber="8">
<taxonomicName id="5AE61A884B54FFD4FF6E957EB8BECA3D" box="[151,452,1807,1833]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="7" pageNumber="8" phylum="Annelida" rank="species" species="quileuticus" status="sp. nov.">
<emphasis id="AF92BD194B54FFD4FF6E957EB8BECA3D" bold="true" box="[151,452,1807,1833]" italics="true" pageId="7" pageNumber="8">Rhyacodrilus quileuticus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B4A100624B54FFD4FE35957EBB74CA3D" box="[460,526,1807,1833]" pageId="7" pageNumber="8" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="9D59610B4B54FFD4FF6E9543B871CA5E" blockId="7.[151,526,1807,1866]" box="[151,267,1842,1866]" pageId="7" pageNumber="8">
(
<figureCitation id="05DD7D8E4B54FFD4FF669543B879CA5E" box="[159,259,1842,1866]" captionStart-0="FIGURE 4" captionStart-1="FIGURE 5" captionStartId-0="9.[151,250,1716,1739]" captionStartId-1="10.[151,250,1944,1967]" captionTargetBox-0="[165,1420,354,1684]" captionTargetBox-1="[162,1414,202,1883]" captionTargetId-0="figure@9.[151,1436,338,1695]" captionTargetId-1="figure@10.[151,1436,193,1923]" captionTargetPageId-0="9" captionTargetPageId-1="10" captionText-0="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." captionText-1="FIGURE 5. Rhyacodrilus quileuticus sp. n. A. Anterior part of the body. B. Reproductive organs. C. Spermathecal chaetae. D. Penial chaetae." httpUri-0="https://zenodo.org/record/283791/files/figure.png" httpUri-1="https://zenodo.org/record/283792/files/figure.png" pageId="7" pageNumber="8">Figs 45</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B54FFDBFF6E950AB860CC2B" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8" type="materials_examined">
<paragraph id="9D59610B4B54FFD4FF6E950AB834CACE" blockId="7.[151,1436,1914,2010]" pageId="7" pageNumber="8">
<emphasis id="AF92BD194B54FFD4FF6E950AB87ECA87" bold="true" box="[151,260,1915,1939]" pageId="7" pageNumber="8">
<typeStatus id="425DDFA94B54FFD4FF6E950AB87ECA87" box="[151,260,1915,1939]" pageId="7" pageNumber="8" type="holotype">Holotype</typeStatus>
</emphasis>
. USNM 1202053: whole mounted specimen stained in borax carmine and mounted in
<collectingCountry id="E5F1219B4B54FFD4FB36950BBC59CA86" box="[1231,1315,1914,1938]" name="Canada" pageId="7" pageNumber="8">Canada</collectingCountry>
balsam.
<emphasis id="AF92BD194B54FFD4FF3095EEB83BCAA3" bold="true" box="[201,321,1951,1975]" pageId="7" pageNumber="8">
<typeStatus id="425DDFA94B54FFD4FF3095EEB83BCAA3" box="[201,321,1951,1975]" pageId="7" pageNumber="8" type="paratype">Paratypes</typeStatus>
</emphasis>
. USNM 1202054-55: 2 whole mounted specimens, stained in borax carmine, from the
<typeStatus id="425DDFA94B54FFD4FAE895EEBC3BCAA3" box="[1297,1345,1951,1975]" pageId="7" pageNumber="8">type</typeStatus>
locality (
<date id="E95847CB4B54FFD4FF6695B3B838CACE" box="[159,322,1986,2010]" pageId="7" pageNumber="8" value="2004-04-29">29 April 2004</date>
).
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3092E9BB1FCDC7" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF3092E9B812CDA4" bold="true" box="[201,360,151,176]" pageId="8" pageNumber="9">
<typeStatus id="425DDFA94B5BFFDBFF3092E9B87ECDA4" box="[201,260,152,176]" pageId="8" pageNumber="9">Type</typeStatus>
locality
</emphasis>
. South Fork Calawah River, Olympic National Forest, Clallam Co., Washington,
<collectingCountry id="E5F1219B4B5BFFDBFAA492E6BCE2CDBB" box="[1373,1432,151,175]" name="United States of America" pageId="8" pageNumber="9">USA</collectingCountry>
,
<geoCoordinate id="F8D207CC4B5BFFDBFF6E92CAB86BCDC7" box="[151,273,187,211]" direction="north" orientation="latitude" pageId="8" pageNumber="9" precision="5" value="47.9627">N47.9627°</geoCoordinate>
<geoCoordinate id="F8D207CC4B5BFFDBFEE192CAB8DDCDC7" box="[280,423,187,211]" direction="west" orientation="longitude" pageId="8" pageNumber="9" precision="5" value="-124.2941">W124.2941°</geoCoordinate>
(
<date id="E95847CB4B5BFFDBFE4F92CABB23CDC7" box="[438,601,187,211]" pageId="8" pageNumber="9" value="2004-04-29">29 April 2004</date>
).
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3092AEB860CC2B" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF3092AEB8FECDEC" bold="true" box="[201,388,223,248]" pageId="8" pageNumber="9">Other material</emphasis>
. 4 mature whole-mounted worms (
<date id="E95847CB4B5BFFDBFCCA92AEBA9FCDE3" box="[819,997,223,247]" pageId="8" pageNumber="9" value="2004-04-29">29 April 2004</date>
). 1 mature whole mount, stained in hematoxylin, from West Fork Humptulips River, Washington, Grays Harbor Co., approx.
<geoCoordinate id="F8D207CC4B5BFFDBFB599372BD85CC0F" box="[1184,1279,259,283]" direction="north" orientation="latitude" pageId="8" pageNumber="9" precision="555" value="47.48">N47.48°</geoCoordinate>
<geoCoordinate id="F8D207CC4B5BFFDBFAF19372BC01CC0F" box="[1288,1403,259,283]" direction="west" orientation="longitude" pageId="8" pageNumber="9" precision="555" value="-123.68">W123.68°</geoCoordinate>
(
<date id="E95847CB4B5BFFDBFA749372B874CC2B" pageId="8" pageNumber="9" value="2003-06-02">2 June 2003</date>
).
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B5BFFDBFF30933AB9A0CC92" pageId="8" pageNumber="9" type="etymology">
<paragraph id="9D59610B4B5BFFDBFF30933AB9A0CC92" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF30933AB832CC77" bold="true" box="[201,328,331,355]" pageId="8" pageNumber="9">Etymology</emphasis>
. the species has been named after the Quileute People (also called Quilayute) of the Calawah River basin.
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B5BFFDBFF3093E0BABDCBA4" pageId="8" pageNumber="9" type="description">
<paragraph id="9D59610B4B5BFFDBFF3093E0BDB4CF94" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF3093E0B82ECCBE" bold="true" box="[201,340,401,426]" pageId="8" pageNumber="9">Description</emphasis>
. Number of segments: more than 25 (all specimens incomplete). Diameter of the body in segment VIII (whole-mounted specimens) 276358 µm. Round prostomium (148217 µm long) (
<figureCitation id="05DD7D8E4B5BFFDBFB8293C7BDC5CCDA" box="[1147,1215,438,462]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
A). Epidermis 712 µm high, and up to 30 µm high at the clitellum, which extends from the level of chaetae in segment X to the end of XII (
<figureCitation id="05DD7D8E4B5BFFDBFF36938CB869CF01" box="[207,275,509,533]" captionStart="FIGURE 5" captionStartId="10.[151,250,1944,1967]" captionTargetBox="[162,1414,202,1883]" captionTargetId="figure@10.[151,1436,193,1923]" captionTargetPageId="10" captionText="FIGURE 5. Rhyacodrilus quileuticus sp. n. A. Anterior part of the body. B. Reproductive organs. C. Spermathecal chaetae. D. Penial chaetae." httpUri="https://zenodo.org/record/283792/files/figure.png" pageId="8" pageNumber="9">Fig. 5</figureCitation>
A). Clitellum formed by large glandular cells (
<figureCitation id="05DD7D8E4B5BFFDBFCD1938CBA17CF01" box="[808,877,509,533]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
C) except for the ventral side around male pores, where there are only small glandular cells. One pair male pores in XI in line of ventral chaetae. One pair spermathecal pores in X, slightly lateral to the line of ventral chaetae, close to intersegment 9/10. One pair female pores opening in intersegment 11/12, about midway between ventral chaetae and the lateral line.
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3090FDBDE7CE6E" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
Hair chaetae (0)12 per bundle anteriorly, and 0(1) in postclitellar segments, usually absent posterior to segment XII or XV, and shorter than body diameter (length 110200 µm). Dorsal pectinate chaetae 24 per bundle, nodulus 0.30.4 from distal end, 6473 µm long (5660 µm in II), distal tooth longer or about equal to proximal, intermediate teeth same length or shorter than lateral teeth (
<figureCitation id="05DD7D8E4B5BFFDBFCAA9086BAE0CE1B" box="[851,922,759,783]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
D, E). Ventral chaetae bifid; anterior ventral chaetae 36 per bundle, 7582 µm long (5870 µm in II), distal tooth twice length of proximal; 35 ventral chaetae in postclitellar segments, 5876 µm long, with distal tooth thinner, but about the same length as proximal (
<figureCitation id="05DD7D8E4B5BFFDBFAC5914EBC04CE43" box="[1340,1406,831,855]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
F H). Nodulus at 0.40.5 from distal end in anterior ventral chaetae,
<quantity id="5A1ECCEE4B5BFFDBFC799113BA8BCE6E" box="[896,1009,866,890]" metricMagnitude="-2" metricUnit="m" metricValue="1.016" metricValueMax="1.27" metricValueMin="0.762" pageId="8" pageNumber="9" unit="in" value="0.4" valueMax="0.5" valueMin="0.3">0.30.5 in</quantity>
posterior ones.
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3091F7BBCEC98C" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
Modified spermathecal chaetae in segment X, 12 per bundle, in line with ventral chaetae, in the posterior third of the segment, well-separated from spermathecal pores (7597 µm long and ca. 2 µm shaft diameter). The spermathecal chaetae have a very long, sharply-pointed distal tooth (717 µm long, most commonly 12 µm), with a short (3 µm), rounded proximal tooth at the base, nodulus at 0.3 from the distal end (
<figureCitation id="05DD7D8E4B5BFFDBFB909180BDCBC91D" box="[1129,1201,1009,1033]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Figs 4</figureCitation>
J, 5C). A ventral pad of thickened (to about 30 µm) epidermis surrounds spermathecal chaetae; cells columnar but not distinctly glandular (
<figureCitation id="05DD7D8E4B5BFFDBFEEB9648B822C945" box="[274,344,1081,1105]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
K). Modified penial chaetae (7284 µm long), 2 per bundle (sometimes an additional, partially developed one growing laterally), arranged in parallel, adjacent to male pores in XI, bifid with long, distinct teeth, and nodulus at 0.2 from distal end (
<figureCitation id="05DD7D8E4B5BFFDBFDDA96F1BB16C98C" box="[547,620,1152,1176]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">
Figs
<date id="E95847CB4B5BFFDBFDA596F1BB16C98C" box="[604,620,1152,1176]" pageId="8" pageNumber="9">4</date>
</figureCitation>
I, 5D).
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3096D5B819C85F" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
Nucleated coelomocytes with granulated cytoplasm (diameter 1022 µm) (
<figureCitation id="05DD7D8E4B5BFFDBFBE396D5BD1AC9A8" box="[1050,1120,1188,1212]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
B). Pharyngeal glands in IV (ventro-laterally) and V (ventrally), in some specimens also ventrally in VI. Chloragogenous tissue from the back part of segment IV and densely covering the gut from V backwards. No glands observed behind ventral chaetae. Sperm sac may extend anteriorly to segment VIII and back to XV. Egg sac back to XVI. One pair testes in X, one pair ovaries in XI.
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF309726BD55C8A2" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
Spermathecal duct short and bulbous (6096 µm long, 5570 µm maximum diameter), formed by a thick wall of columnar epithelium and a muscular layer (
<figureCitation id="05DD7D8E4B5BFFDBFD52970BBB8AC886" box="[683,752,1402,1426]" captionStart="FIGURE 5" captionStartId="10.[151,250,1944,1967]" captionTargetBox="[162,1414,202,1883]" captionTargetId="figure@10.[151,1436,193,1923]" captionTargetPageId="10" captionText="FIGURE 5. Rhyacodrilus quileuticus sp. n. A. Anterior part of the body. B. Reproductive organs. C. Spermathecal chaetae. D. Penial chaetae." httpUri="https://zenodo.org/record/283792/files/figure.png" pageId="8" pageNumber="9">Fig. 5</figureCitation>
B). Oval spermathecal ampullae occupy most of segment X (188217 µm maximum diameter). Sperm free in lumen of spermathecal ampulla.
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3097B3B8D7CB98" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
One pair sperm funnels on 10/11. Vas deferens 2026 µm wide and about 150200 µm long, coiled, and joining the atrial ampulla subapically (
<figureCitation id="05DD7D8E4B5BFFDBFD999797BBD2C8EA" box="[608,680,1510,1534]" captionStart="FIGURE 5" captionStartId="10.[151,250,1944,1967]" captionTargetBox="[162,1414,202,1883]" captionTargetId="figure@10.[151,1436,193,1923]" captionTargetPageId="10" captionText="FIGURE 5. Rhyacodrilus quileuticus sp. n. A. Anterior part of the body. B. Reproductive organs. C. Spermathecal chaetae. D. Penial chaetae." httpUri="https://zenodo.org/record/283792/files/figure.png" pageId="8" pageNumber="9">Fig. 5</figureCitation>
B). Atrial ampulla globular (84111 µm long, 5781 µm wide), completely covered by a diffuse layer of prostate cells up to 4057 µm high. Atrial duct (3945 µm long) shorter than the ampulla and associated with a few muscular strands, its diameter widening from the simple male pore towards the ental end, and not clearly separated from the ampulla (
<figureCitation id="05DD7D8E4B5BFFDBFC539420BA8BCB7D" box="[938,1009,1617,1641]" captionStart="FIGURE 5" captionStartId="10.[151,250,1944,1967]" captionTargetBox="[162,1414,202,1883]" captionTargetId="figure@10.[151,1436,193,1923]" captionTargetPageId="10" captionText="FIGURE 5. Rhyacodrilus quileuticus sp. n. A. Anterior part of the body. B. Reproductive organs. C. Spermathecal chaetae. D. Penial chaetae." httpUri="https://zenodo.org/record/283792/files/figure.png" pageId="8" pageNumber="9">Fig. 5</figureCitation>
B). Atrial lumen irregularly filled by epithelial cells (
<figureCitation id="05DD7D8E4B5BFFDBFEB09405B8F6CB98" box="[329,396,1652,1676]" captionStart="FIGURE 4" captionStartId="9.[151,250,1716,1739]" captionTargetBox="[165,1420,354,1684]" captionTargetId="figure@9.[151,1436,338,1695]" captionTargetPageId="9" captionText="FIGURE 4. Rhyacodrilus quileuticus sp. n. A. Anterior body end. B. Coelomocytes in body cavity. C. Epithelium in segment X in front of, and at the clitellum. D E. Dorsal chaetae from segments IV (D) and XIII (E). F H. Ventral chaetae from segments III (F), V (G) and posterior (H). I. Penial chaetae in XI. J. Spermathecal chaetae in X. K. Thickened epidermal pad surrounding the spermathecal chaeta. L. Atrium." httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="8" pageNumber="9">Fig. 4</figureCitation>
L).
</paragraph>
<paragraph id="9D59610B4B5BFFDBFF3E94E9BABDCBA4" blockId="8.[151,1437,151,2034]" box="[199,967,1688,1713]" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF3E94E9B836CBA5" bold="true" box="[199,332,1688,1713]" pageId="8" pageNumber="9">Anomalies.</emphasis>
In one specimen, genitalia shifted one segment forward.
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B5BFFDBFF3094CDB99ACABE" pageId="8" pageNumber="9" type="distribution">
<paragraph id="9D59610B4B5BFFDBFF3094CDB99ACABE" blockId="8.[151,1437,151,2034]" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF3094CDB896CBC1" bold="true" box="[201,492,1724,1749]" pageId="8" pageNumber="9">Distribution and habitat</emphasis>
. Known only from two sites in northwestern Washington. The Calawah River (
<typeStatus id="425DDFA94B5BFFDBFA9394CDBCE6CBC0" box="[1386,1436,1724,1748]" pageId="8" pageNumber="9">type</typeStatus>
locality) and tributaries are low gradient, streams; surface flow varies seasonally, with low flows in July-August. Areas of groundwater upwellings result in localized cold spots. Sediment is dominated by cobble-size stones. Dissolved oxygen levels are high (
<quantity id="5A1ECCEE4B5BFFDBFDE19556BBD3CA2B" box="[536,681,1831,1855]" metricMagnitude="-5" metricUnit="kg" metricValue="1.02" metricValueMax="1.0699999999999998" metricValueMin="0.97" pageId="8" pageNumber="9" unit="mg" value="10.2" valueMax="10.7" valueMin="9.7">9.710.7 mg</quantity>
/L), and the river is rated Class AA (extraordinary water quality) by the Washington Department of Ecology (Hook 2004). There are no domestic water systems or hatcheries located within the South Fork Calawah, and the watershed has substantial populations of several salmonid species (Hook 2004).
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B5BFFD8FF3095C6BE2ECF91" lastPageId="11" lastPageNumber="12" pageId="8" pageNumber="9" type="discussion">
<paragraph id="9D59610B4B5BFFDAFF3095C6BB37CC0F" blockId="8.[151,1437,151,2034]" lastBlockId="9.[151,1436,151,283]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
<emphasis id="AF92BD194B5BFFDBFF3095C6B84CCADB" bold="true" box="[201,310,1975,1999]" pageId="8" pageNumber="9">Remarks</emphasis>
. The species is closely related to
<taxonomicName id="5AE61A884B5BFFDBFD4A95C9BA4DCADB" box="[691,823,1975,1999]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="8" pageNumber="9" phylum="Annelida" rank="species" species="saelonae">
<emphasis id="AF92BD194B5BFFDBFD4A95C9BA4DCADB" box="[691,823,1975,1999]" italics="true" pageId="8" pageNumber="9">R. saelonae</emphasis>
</taxonomicName>
based on the presence of both penial and spemathecal chaetae, and the prostate gland covering all of the atrial ampulla. It is, however, clearly distinguished from it and other related species (see
<tableCitation id="D06454B04B5AFFDAFE4392E6BB71CDBB" box="[442,523,151,175]" captionStart="TABLE 1" captionStartId="11.[125,208,551,573]" captionText="TABLE 1. Rhyacodrilus species with both modified spermathecal and penial chaetae. nd: no data." pageId="9" pageNumber="10">Table 1</tableCitation>
) by the smaller number of genital chaetae, the bifid penial chaetae, the structure of the male duct, and the absence of ventral glands in segment X and adjacent segments. The atrial duct is shorter than the ampulla, gradually narrowing towards the male pore in
<taxonomicName id="5AE61A884B5AFFDAFC9F9291BD7ACDE3" box="[870,1024,223,247]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="9" pageNumber="10" phylum="Annelida" rank="species" species="quileuticus" status="sp. nov.">
<emphasis id="AF92BD194B5AFFDAFC9F9291BD7ACDE3" box="[870,1024,223,247]" italics="true" pageId="9" pageNumber="10">R. quileuticus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B5AFFDAFBFE9291BD3CCDEC" bold="true" box="[1031,1094,224,248]" pageId="9" pageNumber="10">
<taxonomicNameLabel id="B4A100624B5AFFDAFBFE9291BD3CCDEC" box="[1031,1094,224,248]" pageId="9" pageNumber="10" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
, while it is well-separated and longer than the ampulla in
<taxonomicName id="5AE61A884B5AFFDAFE3D9375BB3DCC0F" box="[452,583,259,283]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="9" pageNumber="10" phylum="Annelida" rank="species" species="saelonae">
<emphasis id="AF92BD194B5AFFDAFE3D9375BB3DCC0F" box="[452,583,259,283]" italics="true" pageId="9" pageNumber="10">R. saelonae</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="C99931834B5AFFDAFF6E94C5BB79CA33" httpUri="https://zenodo.org/record/283791/files/figure.png" pageId="9" pageNumber="10" targetBox="[165,1420,354,1684]" targetPageId="9">
<paragraph id="9D59610B4B5AFFDAFF6E94C5BB79CA33" blockId="9.[151,1436,1716,1831]" pageId="9" pageNumber="10">
<emphasis id="AF92BD194B5AFFDAFF6E94C5B869CBDE" bold="true" box="[151,275,1716,1739]" pageId="9" pageNumber="10">FIGURE 4.</emphasis>
<taxonomicName id="5AE61A884B5AFFDAFEE394C4BB6CCBDF" box="[282,534,1717,1739]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="9" pageNumber="10" phylum="Annelida" rank="species" species="quileuticus" status="sp. nov.">
<emphasis id="AF92BD194B5AFFDAFEE394C4BB6CCBDF" box="[282,534,1717,1739]" italics="true" pageId="9" pageNumber="10">Rhyacodrilus quileuticus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B5AFFDAFDE494C4BB0CCBDE" bold="true" box="[541,630,1717,1738]" pageId="9" pageNumber="10">
<taxonomicNameLabel id="B4A100624B5AFFDAFDE494C4BB2DCBDE" box="[541,599,1717,1738]" pageId="9" pageNumber="10" rank="species">sp. n.</taxonomicNameLabel>
A.
</emphasis>
Anterior body end.
<emphasis id="AF92BD194B5AFFDAFCBD94C4BA21CBDE" bold="true" box="[836,859,1717,1738]" pageId="9" pageNumber="10">B.</emphasis>
Coelomocytes in body cavity.
<emphasis id="AF92BD194B5AFFDAFB6F94C5BDD5CBDF" bold="true" box="[1174,1199,1716,1739]" pageId="9" pageNumber="10">C.</emphasis>
Epithelium in segment X in front of, and at the clitellum.
<emphasis id="AF92BD194B5AFFDAFE1494A5BB67CBFD" bold="true" box="[493,541,1748,1769]" pageId="9" pageNumber="10">DE</emphasis>
. Dorsal chaetae from segments IV(D) and XIII (E).
<emphasis id="AF92BD194B5AFFDAFBD394A5BD23CBFD" bold="true" box="[1066,1113,1748,1769]" pageId="9" pageNumber="10">FH</emphasis>
. Ventral chaetae from segments III (F), V (G) and posterior (H).
<emphasis id="AF92BD194B5AFFDAFE259483B89CCA13" bold="true" box="[476,486,1778,1799]" pageId="9" pageNumber="10">I</emphasis>
. Penial chaetae in XI.
<emphasis id="AF92BD194B5AFFDAFD309483BBAFCA13" bold="true" box="[713,725,1778,1799]" pageId="9" pageNumber="10">J</emphasis>
. Spermathecal chaetae in X.
<emphasis id="AF92BD194B5AFFDAFC059483BD6ACA13" bold="true" box="[1020,1040,1778,1799]" pageId="9" pageNumber="10">K</emphasis>
. Thickened epidermal pad surrounding the spermathecal chaeta.
<emphasis id="AF92BD194B5AFFDAFE6C9560B8DCCA32" bold="true" box="[405,422,1809,1830]" pageId="9" pageNumber="10">L</emphasis>
. Atrium.
</paragraph>
</caption>
<caption id="C99931834B59FFD9FF6E95E9B84ACAD9" httpUri="https://zenodo.org/record/283792/files/figure.png" pageId="10" pageNumber="11" targetBox="[162,1414,202,1883]" targetPageId="10">
<paragraph id="9D59610B4B59FFD9FF6E95E9B84ACAD9" blockId="10.[151,1435,1944,1997]" pageId="10" pageNumber="11">
<emphasis id="AF92BD194B59FFD9FF6E95E9B868CABA" bold="true" box="[151,274,1944,1967]" pageId="10" pageNumber="11">FIGURE 5.</emphasis>
<taxonomicName id="5AE61A884B59FFD9FEE095E9BB6FCABA" box="[281,533,1944,1966]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="10" pageNumber="11" phylum="Annelida" rank="species" species="quileuticus" status="sp. nov.">
<emphasis id="AF92BD194B59FFD9FEE095E9BB6FCABA" box="[281,533,1944,1966]" italics="true" pageId="10" pageNumber="11">Rhyacodrilus quileuticus</emphasis>
</taxonomicName>
<emphasis id="AF92BD194B59FFD9FDE295E8BB16CABA" bold="true" box="[539,620,1945,1966]" pageId="10" pageNumber="11">
<taxonomicNameLabel id="B4A100624B59FFD9FDE295E8BB2ECABA" box="[539,596,1945,1966]" pageId="10" pageNumber="11" rank="species">sp. n.</taxonomicNameLabel>
A
</emphasis>
. Anterior part of the body.
<emphasis id="AF92BD194B59FFD9FC7995E8BAEBCABA" bold="true" box="[896,913,1945,1966]" pageId="10" pageNumber="11">B</emphasis>
. Reproductive organs.
<emphasis id="AF92BD194B59FFD9FB8595E9BDF4CABB" bold="true" box="[1148,1166,1944,1967]" pageId="10" pageNumber="11">C</emphasis>
. Spermathecal chaetae.
<emphasis id="AF92BD194B59FFD9FA7A95E8BCEFCABA" bold="true" box="[1411,1429,1945,1966]" pageId="10" pageNumber="11">D</emphasis>
. Penial chaetae.
</paragraph>
</caption>
<caption id="C99931834B58FFD8FF849056BD3ECF29" box="[125,1092,551,573]" pageId="11" pageNumber="12">
<paragraph id="9D59610B4B58FFD8FF849056BD3ECF29" blockId="11.[125,1891,551,815]" box="[125,1092,551,573]" pageId="11" pageNumber="12">
<emphasis id="AF92BD194B58FFD8FF849056B998CF29" bold="true" box="[125,226,551,573]" pageId="11" pageNumber="12">TABLE 1</emphasis>
.
<taxonomicName id="5AE61A884B58FFD8FF169056B808CF28" box="[239,370,551,572]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="11" pageNumber="12" phylum="Annelida" rank="genus">
<emphasis id="AF92BD194B58FFD8FF169056B808CF28" box="[239,370,551,572]" italics="true" pageId="11" pageNumber="12">Rhyacodrilus</emphasis>
</taxonomicName>
species with both modified spermathecal and penial chaetae. nd: no data.
</paragraph>
</caption>
<paragraph id="9D59610B4B58FFD8FF849022BE2ECF91" blockId="11.[125,1891,551,815]" pageId="11" pageNumber="12">
<emphasis id="AF92BD194B58FFD8FF849022BA7BCF7D" bold="true" box="[125,769,595,617]" pageId="11" pageNumber="12">
Character
<emphasis id="AF92BD194B58FFD8FE679022BA7BCF7D" bold="true" box="[414,769,595,617]" italics="true" pageId="11" pageNumber="12">
<taxonomicName id="5AE61A884B58FFD8FE679022BB79CF7D" box="[414,515,595,617]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="11" pageNumber="12" phylum="Annelida" rank="species" species="alcyoneus">alcyoneus</taxonomicName>
<taxonomicName id="5AE61A884B58FFD8FD559022BA7BCF7D" box="[684,769,595,617]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="11" pageNumber="12" phylum="Annelida" rank="species" species="ardierae">ardierae</taxonomicName>
</emphasis>
</emphasis>
<taxonomicName id="5AE61A884B58FFD8FC889023BD32CF7D" authority="Kosel, 1980" authorityName="Kosel" authorityYear="1980" box="[881,1096,594,617]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="11" pageNumber="12" phylum="Annelida" rank="species" species="carsticus">
<emphasis id="AF92BD194B58FFD8FC889023BAB1CF7C" bold="true" box="[881,971,594,616]" pageId="11" pageNumber="12">carsticus</emphasis>
Košel, 1980
</taxonomicName>
<taxonomicName id="5AE61A884B58FFD8FB7A9023BDA0CF7C" box="[1155,1242,594,616]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="11" pageNumber="12" phylum="Annelida" rank="species" species="gasparoi">
<emphasis id="AF92BD194B58FFD8FB7A9023BDA0CF7C" bold="true" box="[1155,1242,594,616]" italics="true" pageId="11" pageNumber="12">gasparoi</emphasis>
</taxonomicName>
Martínez-
<taxonomicName id="5AE61A884B58FFD8FA7D9023BF4ECF7D" authority="Giani" authorityName="Giani" box="[1412,1588,594,617]" class="Clitellata" family="Randiellidae" genus="Rhyacodrilus" kingdom="Animalia" order="Enchytraeida" pageId="11" pageNumber="12" phylum="Annelida" rank="species" species="gernikensis">
<emphasis id="AF92BD194B58FFD8FA7D9023BC82CF7C" bold="true" box="[1412,1528,594,616]" italics="true" pageId="11" pageNumber="12">gernikensis</emphasis>
Giani
</taxonomicName>
&amp;
<taxonomicName id="5AE61A884B58FFD8F9839023BF8ECF7C" box="[1658,1780,594,616]" class="Clitellata" family="Tubificidae" genus="Stochidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="11" pageNumber="12" phylum="Annelida" rank="species" species="glandulosus">
<emphasis id="AF92BD194B58FFD8F9839023BF8ECF7C" bold="true" box="[1658,1780,594,616]" italics="true" pageId="11" pageNumber="12">glandulosus</emphasis>
</taxonomicName>
(
<emphasis id="AF92BD194B58FFD8F8F89022B896CF91" bold="true" pageId="11" pageNumber="12">Martínezsp. n.</emphasis>
Lafont &amp; Juget, 1993 Ansemil
<emphasis id="AF92BD194B58FFD8FB23901EBC76CF90" box="[1242,1292,623,644]" italics="true" pageId="11" pageNumber="12">et al.</emphasis>
1997 Rodriguez, 1988 Ansemil
<emphasis id="AF92BD194B58FFD8F91A901EBE6FCF90" box="[1763,1813,623,644]" italics="true" pageId="11" pageNumber="12">et al.</emphasis>
1997)
</paragraph>
</subSubSection>
<subSubSection id="D5FC32804B58FFD8FF8490FCBBD7CE3B" pageId="11" pageNumber="12" type="description">
<paragraph id="9D59610B4B58FFD8FF8490FCB8C0CFE3" blockId="11.[125,1891,551,815]" pageId="11" pageNumber="12">
<emphasis id="AF92BD194B58FFD8FF8490FCB86DCFB7" bold="true" box="[125,279,653,675]" pageId="11" pageNumber="12">Dorsal chaetae</emphasis>
Hairs and pectinates. No hairs, bifids Bifids, teeth equal Hairs and pectinates. No hairs, bifids with Hairs and pectinates. Hairs absent from variable (modified in XI) equal teeth Hairs absent from XVII XIIIXV backward backward
</paragraph>
<paragraph id="9D59610B4B58FFD8FF84908FBE44CE07" blockId="11.[125,1891,551,815]" box="[125,1854,765,788]" pageId="11" pageNumber="12">
<emphasis id="AF92BD194B58FFD8FF84908FB869CE00" bold="true" box="[125,275,766,788]" pageId="11" pageNumber="12">Penial chaetae</emphasis>
57 simple-pointed or 3(45), simple-pointed, 24 bifid, hook-shaped 2 simple or bifid(?) 2 bifid 12 simple- pointed
</paragraph>
<paragraph id="9D59610B4B58FFD8FEA29168BBD7CE3B" blockId="11.[125,1891,551,815]" box="[347,685,793,815]" pageId="11" pageNumber="12">minutely bifid, fanwise fanwise</paragraph>
</subSubSection>
</treatment>
</document>