treatments-xml/data/03/BC/86/03BC8649FF8AFF88FE2AFC16DC65B3AE.xml

<document id="A520E4324DF09A239960225DC9E58463" ID-CLB-Dataset="21986" ID-DOI="10.5852/ejt.2017.361" ID-GBIF-Dataset="db3fd15a-2d90-4f3f-87f8-10b8953e26a5" ID-ISSN="2118-9773" ID-Zenodo-Dep="3836508" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1589998651298" checkinUser="carolina" docAuthor="Fend, Steven V., Rodriguez, Pilar, Achurra, Ainara &amp; Erséus, Christer" docDate="2017" docId="03BC8649FF8AFF88FE2AFC16DC65B3AE" docLanguage="en" docName="ejt-2017-361.pdf.imf" docOrigin="European Journal of Taxonomy 361" docStyle="DocumentStyle:EF2B578F1D15862ADE45B0C07C620911.14:EJT.2018-.journal_article.type1" docStyleId="EF2B578F1D15862ADE45B0C07C620911" docStyleName="EJT.2018-.journal_article.type1" docStyleVersion="14" docTitle="Guestphalinus exilis Fend &amp; Rodriguez &amp; Achurra &amp; Erséus 2017, sp. nov." docType="treatment" docUuid="37A5A3E0-3DC2-4D24-84AC-62E21131E70E" docUuidSource="ZooBank" docVersion="9" lastPageNumber="28" masterDocId="FF85FE31FF9FFF93FFD7FFD9DE4DB573" masterDocTitle="On Kincaidiana Altman, 1936 and Guestphalinus Michaelsen, 1933 (Annelida, Clitellata, Lumbriculidae), with the descriptions of three new species" masterLastPageNumber="46" masterPageNumber="1" pageNumber="22" updateTime="1698840559482" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-3.0" zenodo-license-figures="CC-BY-3.0" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="8BAB88CCBD039A169574A0D6E6A52321">On Kincaidiana Altman, 1936 and Guestphalinus Michaelsen, 1933 (Annelida, Clitellata, Lumbriculidae), with the descriptions of three new species</mods:title>
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<mods:namePart id="17DBF4DC28C7569D116637F710DDCA95">Fend, Steven V.</mods:namePart>
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<mods:affiliation id="5F7011AD463E958B531981187EA26CE7">US Geological Survey, 345 Middlefield Rd., Menlo Park, CA, 94025, USA. &amp; urn: lsid: zoobank. org: author: E 0 C 611 FA- 9619 - 4 DFD- 9701 - 026417181232 &amp; Corresponding author: svfend @ usgs. gov</mods:affiliation>
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<mods:namePart id="BD45350FF6BAACA608E4548FA4CB8AE1">Rodriguez, Pilar</mods:namePart>
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<mods:affiliation id="6022F3698E76C0C5D954B1E2DC9C2BA7">University of the Basque Country, UPV / EHU, Sarriena s / n, Leioa 48940, Spain. &amp; Email: pilar. rodriguez @ ehu. eus &amp; urn: lsid: zoobank. org: author: C 4600 D 2 B- 17 E 2 - 4 CD 0 - AE 75 - B 79 BA 9 DD 35 D 6</mods:affiliation>
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<mods:namePart id="ED2D48032D8BBCE7774844A840E2ABDF">Achurra, Ainara</mods:namePart>
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<mods:affiliation id="475E1310ECD063D57C549F8D14C21862">University of the Basque Country, UPV / EHU, Sarriena s / n, Leioa 48940, Spain. &amp; Email: ainara. achurra @ ehu. eus &amp; urn: lsid: zoobank. org: author: 105572 EB-B 070 - 4 BE 4 - 93 D 2 - 50 FA 50847 C 9 E</mods:affiliation>
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<mods:namePart id="C768B097CC48C283B4A35EB22B94C0B7">Erséus, Christer</mods:namePart>
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<mods:affiliation id="EEAC1E823BF5DA98C74E2CB26F415513">University of Gothenburg, SE- 405 30 Göteborg, Sweden. &amp; Email: christer. erseus @ bioenv. gu. se &amp; urn: lsid: zoobank. org: author: D 98 F 606 A-B 273 - 4 F 50 - 95 F 5 - C 35 F 17 B 12 C 85</mods:affiliation>
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<taxonomicName id="4C154CDCFF8AFF86FE2AFC16DD98B69A" ID-CoL="8Q7LH" authority="Fend &amp; Rodriguez" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[509,981,975,1002]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF8AFF86FE2AFC16DCB9B699" bold="true" box="[509,756,975,1002]" italics="true" pageId="21" pageNumber="22">Guestphalinus exilis</emphasis>
Fend &amp; Rodriguez
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urn:lsid:zoobank.org:act: 
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<figureCitation id="132E2BDAFF8AFF86FD30FBC2DD00B146" box="[743,845,1051,1077]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="23.[189,232,1717,1743]" captionStartId-1="25.[189,232,1788,1814]" captionStartId-2="26.[189,232,1645,1671]" captionTargetBox-0="[199,1387,314,1670]" captionTargetBox-1="[204,1385,280,1758]" captionTargetBox-2="[189,1398,301,1602]" captionTargetId-0="figure@23.[189,1398,314,1681]" captionTargetId-1="figure@25.[189,1398,265,1773]" captionTargetId-2="figure@26.[189,1398,300,1604]" captionTargetPageId-0="23" captionTargetPageId-1="25" captionTargetPageId-2="26" captionText-0="Fig. 7. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. A–B. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. C–D. Anterior end, showing orientation of chaetae. E–G. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. H–I. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 Mm." captionText-1="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." captionText-2="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi-0="http://doi.org/10.5281/zenodo.3836522" figureDoi-1="http://doi.org/10.5281/zenodo.3836524" figureDoi-2="http://doi.org/10.5281/zenodo.3836526" httpUri-0="https://zenodo.org/record/3836522/files/figure.png" httpUri-1="https://zenodo.org/record/3836524/files/figure.png" httpUri-2="https://zenodo.org/record/3836526/files/figure.png" pageId="21" pageNumber="22">Figs 7–9</figureCitation>
</paragraph>
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<subSubSection id="C30F64D4FF8AFF86FF6AFB85DB17B1B8" pageId="21" pageNumber="22" type="etymology">
<paragraph id="8BAA375FFF8AFF86FF6AFB85DF09B105" blockId="21.[189,324,1116,1142]" box="[189,324,1116,1142]" pageId="21" pageNumber="22">
<heading id="D0E28033FF8AFF86FF6AFB85DF09B105" bold="true" box="[189,324,1116,1142]" fontSize="11" level="3" pageId="21" pageNumber="22" reason="3">
<emphasis id="B961EB4DFF8AFF86FF6AFB85DF09B105" bold="true" box="[189,324,1116,1142]" pageId="21" pageNumber="22">Etymology</emphasis>
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<paragraph id="8BAA375FFF8AFF86FF6AFB55DB17B1B8" blockId="21.[189,1398,1164,1227]" pageId="21" pageNumber="22">
Derived fom the Latin word ‘ 
<taxonomicName id="4C154CDCFF8AFF86FDCEFB55DC1AB1D5" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[537,599,1164,1190]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="exilis">
<emphasis id="B961EB4DFF8AFF86FDCEFB55DC1AB1D5" box="[537,599,1164,1190]" italics="true" pageId="21" pageNumber="22">exilis</emphasis>
</taxonomicName>
’ (small), with reference to 
<taxonomicName id="4C154CDCFF8AFF86FC4FFB55DA1FB1D5" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[920,1106,1164,1190]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="elephantinus" status="sp. nov.">
<emphasis id="B961EB4DFF8AFF86FC4FFB55DA1FB1D5" box="[920,1106,1164,1190]" italics="true" pageId="21" pageNumber="22">G. elephantinus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF8AFF86FB8DFB55DAF8B1D5" box="[1114,1205,1164,1190]" pageId="21" pageNumber="22" rank="species">sp. nov.</taxonomicNameLabel>
(see below) and to the name of the extinct Californian pygmy mammoth 
<taxonomicName id="4C154CDCFF8AFF86FC84FB69DB1EB1B8" authority="(Stock &amp; Furlong, 1928)" baseAuthorityName="Stock &amp; Furlong" baseAuthorityYear="1928" box="[851,1363,1200,1227]" class="Mammalia" family="Elephantidae" genus="Mammuthus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Proboscidea" pageId="21" pageNumber="22" phylum="Chordata" rank="species" species="exilis">
<emphasis id="B961EB4DFF8AFF86FC84FB69DA67B1B9" box="[851,1066,1200,1226]" italics="true" pageId="21" pageNumber="22">Mammuthus exilis</emphasis>
(Stock &amp; Furlong, 1928)
</taxonomicName>
.
</paragraph>
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<paragraph id="8BAA375FFF8AFF86FF6AFB2ADFE7B07E" blockId="21.[189,426,1267,1293]" box="[189,426,1267,1293]" pageId="21" pageNumber="22">
<heading id="D0E28033FF8AFF86FF6AFB2ADFE7B07E" bold="true" box="[189,426,1267,1293]" fontSize="11" level="3" pageId="21" pageNumber="22" reason="3">
<emphasis id="B961EB4DFF8AFF86FF6AFB2ADFE7B07E" bold="true" box="[189,426,1267,1293]" pageId="21" pageNumber="22">Material examined</emphasis>
</heading>
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<materialsCitation id="3B7D3D02FF8AFF86FF3DFAFDDC35B0DF" ID-GBIF-Occurrence="2621122304" collectingDate="2012-05-06" collectionCode="USNM" collectorName="S. Fend" country="United States of America" county="Shasta County" location="Squaw Creek" municipality="Canada" pageId="21" pageNumber="22" specimenCode="USNM 1422283" specimenCount="1" stateProvince="California" typeStatus="holotype">
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<collectingCountry id="F30277CFFF8AFF86FF6AFA90DC1AB011" box="[189,599,1352,1379]" name="United States of America" pageId="21" pageNumber="22">UNITED STATES OF AMERICA</collectingCountry>
: a dissected specimen, slide-mounted in 
<collectingMunicipality id="6BCEAD25FF8AFF86FBE1FA91DAC3B011" box="[1078,1166,1352,1378]" pageId="21" pageNumber="22">Canada</collectingMunicipality>
balsam, 
<collectingRegion id="49D1F9BDFF8AFF86FB2FFA91DB3FB011" box="[1272,1394,1352,1378]" country="United States of America" name="California" pageId="21" pageNumber="22">California</collectingRegion>
, 
<collectingCounty id="62CB4FD3FF8AFF86FF6AFAB4DF24B0F4" box="[189,361,1389,1415]" pageId="21" pageNumber="22">Shasta County</collectingCounty>
, 
<location id="8ECA6184FF8AFF86FEA4FAB4DC5DB0F4" LSID="urn:lsid:plazi:treatment:03BC8649FF8AFF88FE2AFC16DC65B3AE:8ECA6184FF8AFF86FEA4FAB4DC5DB0F4" box="[371,528,1389,1415]" country="United States of America" county="Shasta County" municipality="Canada" name="Squaw Creek" pageId="21" pageNumber="22" stateProvince="California">Squaw Creek</location>
at 
<location id="8ECA6184FF8AFF86FDE3FAB4DD5BB0F4" LSID="urn:lsid:plazi:treatment:03BC8649FF8AFF88FE2AFC16DC65B3AE:8ECA6184FF8AFF86FDE3FAB4DD5BB0F4" box="[564,790,1389,1415]" country="United States of America" county="Shasta County" municipality="Canada" name="Chirpchatter Camp" pageId="21" pageNumber="22" stateProvince="California">Chirpchatter Camp</location>
, deep riffle, gravel patches in cobble bottom, 
<date id="FFAB119FFF8AFF86FAFCFAB4DEB6B0DF" pageId="21" pageNumber="22" value="2012-05-06">
<collectingDate id="EFEFE877FF8AFF86FAFCFAB4DEB6B0DF" pageId="21" pageNumber="22" value="2012-05-06">6 May 2012</collectingDate>
</date>
, 
<collectorName id="26E05289FF8AFF86FED1FA4BDF2DB0DF" box="[262,352,1426,1452]" pageId="21" pageNumber="22">S. Fend</collectorName>
leg. (
<specimenCode id="DBB39F24FF8AFF86FE75FA4BDC21B0DF" box="[418,620,1426,1452]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="21" pageNumber="22" type="Museum">USNM 1422283</specimenCode>
).
</paragraph>
</materialsCitation>
<paragraph id="8BAA375FFF8AFF86FF3DFA0CDD35B083" blockId="21.[189,1399,1493,1703]" box="[234,888,1493,1520]" pageId="21" pageNumber="22">
<emphasis id="B961EB4DFF8AFF86FF3DFA0CDF2AB09C" bold="true" box="[234,359,1493,1519]" pageId="21" pageNumber="22">
<typeStatus id="54AE89FDFF8AFF86FF3DFA0CDF2AB09C" box="[234,359,1493,1519]" pageId="21" pageNumber="22" type="paratype">Paratypes</typeStatus>
</emphasis>
(all collected by S. Fend at the type locality)
</paragraph>
<paragraph id="8BAA375FFF8AFF86FF6AFA23DC1DB3D4" blockId="21.[189,1399,1493,1703]" pageId="21" pageNumber="22">
<collectingCountry id="F30277CFFF8AFF86FF6AFA23DC1EB367" box="[189,595,1530,1556]" name="United States of America" pageId="21" pageNumber="22">UNITED STATES OF AMERICA</collectingCountry>
: 1 dissected on 2 slides, 
<date id="FFAB119FFF8AFF86FCB9FA23DA49B367" box="[878,1028,1530,1556]" pageId="21" pageNumber="22" value="2010-04-19">19 Apr. 2010</date>
(
<collectionCode id="ED04AF9AFF8AFF86FBC5FA23DA24B367" box="[1042,1129,1530,1556]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="21" pageNumber="22" type="Museum">USNM</collectionCode>
1422284); 1 dissected, 
<date id="FFAB119FFF8AFF86FF6AF9C7DF10B34B" box="[189,349,1566,1593]" pageId="21" pageNumber="22" value="2010-05-17">17 May 2010</date>
(
<collectionCode id="ED04AF9AFF8AFF86FEB9F9C7DF88B34B" box="[366,453,1566,1592]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="21" pageNumber="22" type="Museum">USNM</collectionCode>
1422285); 1 dissected, 
<date id="FFAB119FFF8AFF86FD35F9C7DD39B34B" box="[738,884,1566,1593]" pageId="21" pageNumber="22" value="2012-05-06">6 May 2012</date>
(
<collectionCode id="ED04AF9AFF8AFF86FC53F9C7DD96B34B" box="[900,987,1566,1592]" pageId="21" pageNumber="22">CASIZ</collectionCode>
220935); 1 whole mount, 
<date id="FFAB119FFF8AFF86FACDF9C7DEB5B32E" pageId="21" pageNumber="22" value="2010-05-17">17 May 2010</date>
(
<collectionCode id="ED04AF9AFF8AFF86FED1F99ADF10B32E" box="[262,349,1603,1629]" pageId="21" pageNumber="22">CASIZ</collectionCode>
220936); 1 sagittally sectioned, 
<date id="FFAB119FFF8AFF86FD01F99ADD3CB32E" box="[726,881,1603,1630]" pageId="21" pageNumber="22" value="2010-05-17">17 May 2010</date>
(
<collectionCode id="ED04AF9AFF8AFF86FCA8F99ADD9BB32E" box="[895,982,1603,1629]" pageId="21" pageNumber="22">CASIZ</collectionCode>
220938); a whole mount, immature but with gonads (DNA voucher), 
<date id="FFAB119FFF8AFF86FD98F9B1DCA1B3F1" box="[591,748,1640,1666]" pageId="21" pageNumber="22" value="2010-04-19">19 Apr. 2010</date>
(
<collectionCode id="ED04AF9AFF8AFF86FD2AF9B1DD19B3F1" box="[765,852,1640,1666]" pageId="21" pageNumber="22">CASIZ</collectionCode>
220937); 2 dissected on slides, 
<date id="FFAB119FFF8AFF86FB01F9B1DB3AB3F1" box="[1238,1399,1640,1666]" pageId="21" pageNumber="22" value="2010-05-17">17 May 2010</date>
(
<collectionCode id="ED04AF9AFF8AFF86FF12F954DF52B3D4" box="[197,287,1677,1703]" country="Spain" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" lsid="urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="21" pageNumber="22" type="Museum">MNCN</collectionCode>
16.03/3100–16.03/3101).
</paragraph>
<paragraph id="8BAA375FFF8AFF86FF3DF916DFACB39A" blockId="21.[189,1399,1743,2027]" box="[234,481,1743,1769]" pageId="21" pageNumber="22">
<emphasis id="B961EB4DFF8AFF86FF3DF916DFACB39A" bold="true" box="[234,481,1743,1769]" pageId="21" pageNumber="22">Additional material</emphasis>
</paragraph>
<paragraph id="8BAA375FFF8AFF85FF6AF92CDD6FB43A" blockId="21.[189,1399,1743,2027]" lastBlockId="22.[189,1398,267,330]" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22">
<collectingCountry id="F30277CFFF8AFF86FF6AF92CDC2DB27D" box="[189,608,1780,1807]" name="United States of America" pageId="21" pageNumber="22">UNITED STATES OF AMERICA</collectingCountry>
: 
<collectingRegion id="49D1F9BDFF8AFF86FDB8F92DDD68B27D" box="[623,805,1780,1806]" country="United States of America" name="California" pageId="21" pageNumber="22">CALIFORNIA</collectingRegion>
: 2 sagittal sections, 3 dissected, 4 whole mounts (all immature or partially mature), Shasta County, Squaw Creek at Chirpchatter Camp, 
<date id="FFAB119FFF8AFF86FB03F8C0DB3CB240" box="[1236,1393,1817,1843]" pageId="21" pageNumber="22" value="1996-06-17">17 Jun. 1996</date>
, S. Fend leg.; 1 dissected (partially mature), same locality as preceding, 
<date id="FFAB119FFF8AFF86FBFDF8E7DA81B22B" box="[1066,1228,1854,1880]" pageId="21" pageNumber="22" value="2010-04-19">19 Apr. 2010</date>
, S. Fend leg.; 4 dissected (1 mature), 8 whole mounts (immature), several in alcohol, same locality as preceding, 
<date id="FFAB119FFF8AFF86FA8EF8BADF74B2D2" pageId="21" pageNumber="22" value="2010-05-17">17 May 2010</date>
, S. Fend leg.; 1 dissected (mature), several immature in alcohol, same locality as preceding, 
<date id="FFAB119FFF8AFF86FF6AF875DF02B2B5" box="[189,335,1964,1990]" pageId="21" pageNumber="22" value="2012-05-06">6 May 2012</date>
, S. Fend leg.; 2 whole mounts (partially mature), Humboldt County, South Fork Eel River at Elk Creek, 
<date id="FFAB119FFF8AFF86FE88F808DFB4B298" box="[351,505,2001,2027]" pageId="21" pageNumber="22" value="2001-06-24">24 Jun. 2001</date>
, S. Fend leg.; 1 whole mount (partially mature), Mendocino County, Garcia River, 
<date id="FFAB119FFF89FF85FED8FED2DFF9B456" box="[271,436,267,294]" pageId="22" pageNumber="23" value="1996-03-23">23 Mar. 1996</date>
, W. Fields leg.; 1 whole mount (immature, with small gonads), Colusa County, Stony Creek at Stonyford, 
<date id="FFAB119FFF89FF85FE20FEF6DCC7B43A" box="[503,650,303,330]" pageId="22" pageNumber="23" value="2002-11-04">4 Nov. 2002</date>
, S. Fend leg.
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AFEA8DF36B4F8" blockId="22.[189,379,369,395]" box="[189,379,369,395]" pageId="22" pageNumber="23">
<heading id="D0E28033FF89FF85FF6AFEA8DF36B4F8" bold="true" box="[189,379,369,395]" fontSize="11" level="3" pageId="22" pageNumber="23" reason="3">
<emphasis id="B961EB4DFF89FF85FF6AFEA8DF36B4F8" bold="true" box="[189,379,369,395]" pageId="22" pageNumber="23">Molecular data</emphasis>
</heading>
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AFE79DA9FB4C8" blockId="22.[189,1234,416,443]" box="[189,1234,416,443]" pageId="22" pageNumber="23">
COI, 28S and 16S sequences correspond to a topotypic 
<typeStatus id="54AE89FDFF89FF85FC9DFE78DDE2B4C8" box="[842,943,417,443]" pageId="22" pageNumber="23" type="paratype">paratype</typeStatus>
(see 
<tableCitation id="C69702E4FF89FF85FC3AFE79DA09B4C9" box="[1005,1092,416,442]" captionStart="Table 1" captionStartId="3.[187,256,597,625]" captionTargetPageId="3" captionText="Table 1 (continued on next page). List of taxa and specimens used in the molecular study, corresponding GenBank accession numbers, voucher data, localities and sampling data. “Specimen” numbers refer to the catalog number in the database of C. Erséus. New sequences are indicated by bold. Both COI sequences for G. elephantinus sp. nov. are identical." httpUri="http://table.plazi.org/id/DF6A67D7FF9CFF90FF6CFD8CDD15B7C4" pageId="22" pageNumber="23" tableUuid="DF6A67D7FF9CFF90FF6CFD8CDD15B7C4">Table 1</tableCitation>
for details).
</paragraph>
</subSubSection>
<subSubSection id="C30F64D4FF89FF8BFF6AFE3BDB39B298" lastPageId="24" lastPageNumber="25" pageId="22" pageNumber="23" type="description">
<paragraph id="8BAA375FFF89FF85FF6AFE3BDF03B48F" blockId="22.[189,334,482,508]" box="[189,334,482,508]" pageId="22" pageNumber="23">
<heading id="D0E28033FF89FF85FF6AFE3BDF03B48F" bold="true" box="[189,334,482,508]" fontSize="11" level="3" pageId="22" pageNumber="23" reason="3">
<emphasis id="B961EB4DFF89FF85FF6AFE3BDF03B48F" bold="true" box="[189,334,482,508]" pageId="22" pageNumber="23">Description</emphasis>
</heading>
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AFDC8DB04B7C9" blockId="22.[189,1399,529,698]" pageId="22" pageNumber="23">
Specimens from the 
<typeStatus id="54AE89FDFF89FF85FE7DFDC8DF91B758" box="[426,476,529,555]" pageId="22" pageNumber="23">type</typeStatus>
locality: body measurements in 
<tableCitation id="C69702E4FF89FF85FC85FDC8DDE4B758" box="[850,937,529,555]" captionStart="Table 3" captionStartId="28.[189,258,268,294]" captionTargetPageId="28" captionText="Table 3. Descriptive measurements (mean, range, and n) for different populations of G. exilis Fend &amp; Rodriguez sp. nov. and G. elephantinus Fend &amp; Rodriguez sp. nov. Measurements of “nodulus” refer to the proportional distance from the tip of the chaeta (i.e., “0.5” means the nodulus is at the midpoint of the chaeta)." pageId="22" pageNumber="23">Table 3</tableCitation>
. Prostomium rounded to nearly conical; filiform proboscis 
<quantity id="4CED9ABAFF89FF85FE46FDECDC7DB73C" box="[401,560,565,591]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" metricValueMax="2.1" metricValueMin="0.9" pageId="22" pageNumber="23" unit="mm" value="1.5" valueMax="2.1" valueMin="0.9">0.9 to 2.1 mm</quantity>
long, diameter at midpoint 
<quantity id="4CED9ABAFF89FF85FCBAFDEDDA45B73C" box="[877,1032,564,591]" metricMagnitude="-4" metricUnit="m" metricValue="1.25" metricValueMax="1.5" metricValueMin="1.0" pageId="22" pageNumber="23" unit="mm" value="0.125" valueMax="0.15" valueMin="0.1">0.1–0.15 mm</quantity>
, appearing ringed with multiple shallow constrictions (
<figureCitation id="132E2BDAFF89FF85FE19FD81DC51B701" box="[462,540,600,626]" captionStart="Fig" captionStartId="23.[189,232,1717,1743]" captionTargetBox="[199,1387,314,1670]" captionTargetId="figure@23.[189,1398,314,1681]" captionTargetPageId="23" captionText="Fig. 7. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. A–B. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. C–D. Anterior end, showing orientation of chaetae. E–G. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. H–I. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836522" httpUri="https://zenodo.org/record/3836522/files/figure.png" pageId="22" pageNumber="23">Figs 7</figureCitation>
A–C, 9A). Body segmentation not strong in external view; secondary annulation a narrow anterior ring in IV–IX, weak in post-clitellar segments. Clitellum from VIII to XII or mid-XIII, absent ventrally in IX in the area surrounding male and spermathecal pores (
<figureCitation id="132E2BDAFF89FF85FB7EFD79DB45B7C9" box="[1193,1288,672,698]" captionStart="Fig" captionStartId="23.[189,232,1717,1743]" captionTargetBox="[199,1387,314,1670]" captionTargetId="figure@23.[189,1398,314,1681]" captionTargetPageId="23" captionText="Fig. 7. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. A–B. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. C–D. Anterior end, showing orientation of chaetae. E–G. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. H–I. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836522" httpUri="https://zenodo.org/record/3836522/files/figure.png" pageId="22" pageNumber="23">Figs 7B</figureCitation>
, 
<figureCitation id="132E2BDAFF89FF85FAC3FD79DB76B7C9" box="[1300,1339,672,698]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="22" pageNumber="23">9A</figureCitation>
).
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AFD3BDFA6B685" blockId="22.[189,1399,738,1014]" pageId="22" pageNumber="23">
Chaetae paired, in 4 bundles in each segment, beginning in II. Chaetal measurements given in 
<tableCitation id="C69702E4FF89FF85FACFFD3BDB3CB78F" box="[1304,1393,738,764]" captionStart="Table 3" captionStartId="28.[189,258,268,294]" captionTargetPageId="28" captionText="Table 3. Descriptive measurements (mean, range, and n) for different populations of G. exilis Fend &amp; Rodriguez sp. nov. and G. elephantinus Fend &amp; Rodriguez sp. nov. Measurements of “nodulus” refer to the proportional distance from the tip of the chaeta (i.e., “0.5” means the nodulus is at the midpoint of the chaeta)." pageId="22" pageNumber="23">Table 3</tableCitation>
. Chaetae in II bluntly simple-pointed to shallowly notched, directed anteriorly (
<figureCitation id="132E2BDAFF89FF85FBBCFCDCDAFAB66C" box="[1131,1207,773,799]" captionStart="Fig" captionStartId="23.[189,232,1717,1743]" captionTargetBox="[199,1387,314,1670]" captionTargetId="figure@23.[189,1398,314,1681]" captionTargetPageId="23" captionText="Fig. 7. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. A–B. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. C–D. Anterior end, showing orientation of chaetae. E–G. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. H–I. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836522" httpUri="https://zenodo.org/record/3836522/files/figure.png" pageId="22" pageNumber="23">Figs 7</figureCitation>
E–G, 9B–C, E); these chaetae slightly sigmoid, with distal nodulus; within each bundle, lateral chaeta slightly longer than the median, with more distal nodulus. Posterior to II, chaetae sigmoid, mostly simple pointed, with nodulus about ⅓ of distance from tip; perpendicular to body axis or posteriorly directed; chaetae in III slightly shorter, but those in more posterior segments similar in length to those in II. Chaetae in posterior segments may have a slight dorsal keel (
<figureCitation id="132E2BDAFF89FF85FCDCFC61DD27B6A1" box="[779,874,952,978]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="22" pageNumber="23">Fig. 9D</figureCitation>
). Ventral chaetae absent in IX in mature and post-reproductive worms.
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AFBC4DA97B026" blockId="22.[189,1399,1053,1365]" pageId="22" pageNumber="23">
Epidermis in anterior segments 
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thick, in clitellum 
<quantity id="4CED9ABAFF89FF85FC4FFBC4DA59B14B" box="[920,1044,1053,1080]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" metricValueMax="5.0" metricValueMin="3.0" pageId="22" pageNumber="23" unit="mm" value="40.0" valueMax="50.0" valueMin="30.0">30–50 µm</quantity>
, posteriorly 
<quantity id="4CED9ABAFF89FF85FB7DFBC4DB54B14B" box="[1194,1305,1053,1080]" metricMagnitude="-3" metricUnit="m" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="22" pageNumber="23" unit="mm" value="7.5" valueMax="10.0" valueMin="5.0">5–10 µm</quantity>
. In preclitellar segments, circular muscle layer of body wall arranged in a series of transverse bands (cf. 
<figureCitation id="132E2BDAFF89FF85FF6AFBBCDF61B10C" box="[189,300,1125,1151]" captionStart="Fig" captionStartId="33.[189,232,1609,1635]" captionTargetBox="[189,1398,267,1566]" captionTargetId="figure@33.[189,1398,265,1566]" captionTargetPageId="33" captionText="Fig. 12. Guestphalinus elephantinus Fend &amp; Rodriguez sp. nov.A–B (upper) from Clearwater River, WA; C, H, K from Shale Creek, WA (type locality); D–E from Satsop River, WA; B (lower), F–G, N–Q from Guadalupe Creek, CA; I–J, L–M from Peavine Ridge, OR. A. Anterior end of cleared worm. B. Chaetae in II-III, comparing worms from northern (upper) and southern (lower) populations. C. Dorsal chaetae in II. D. The outer chaeta in II, showing the “secondary nodulus”. E–F. Inner dorsal chaeta in II; E shows folded keel. G. Modified dorsal chaetae in II. H. Posterior ventral chaeta. I. Posterior ventral chaeta with bifid tip. J. Transverse section of anterior segment, showing lobes of pharyngeal glands. K. Body wall in anterior segment. L. Spermathecal pore. M. Male pore. N. Ectal part of spermatheca from a dissected, mated worm. O. Expanded atrial duct (type 2 penis?) at male pore. P. Copulatory gland. Q. Female funnel and pore on intersegment 10/11. Scale bars: A = 1 mm; B–I, K–M = 50 µm; J, N–Q = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836532" httpUri="https://zenodo.org/record/3836532/files/figure.png" pageId="22" pageNumber="23">Fig. 12K</figureCitation>
), 
<quantity id="4CED9ABAFF89FF85FE92FBBCDF8CB10C" box="[325,449,1125,1152]" metricMagnitude="-2" metricUnit="m" metricValue="1.25" metricValueMax="1.5" metricValueMin="1.0" pageId="22" pageNumber="23" unit="mm" value="12.5" valueMax="15.0" valueMin="10.0">10–15 µm</quantity>
thick; posteriorly a simple layer about 5 μm thick. Longitudinal muscle layer 
<quantity id="4CED9ABAFF89FF85FF6AFB51DF7BB1D0" box="[189,310,1160,1187]" metricMagnitude="-2" metricUnit="m" metricValue="5.5" metricValueMax="6.0" metricValueMin="5.0" pageId="22" pageNumber="23" unit="mm" value="55.0" valueMax="60.0" valueMin="50.0">50–60 µm</quantity>
thick. Brain in the peristomium, lateral lobes rounded. Pharynx begins dorsally and laterally in II, ventrally in III, extending through IV. Pharyngeal glands in IV to VI or VII; on each side, three lobes (dorsal, lateral and median) (
<figureCitation id="132E2BDAFF89FF85FD83FB09DCFDB199" box="[596,688,1232,1258]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="22" pageNumber="23">Fig. 9G</figureCitation>
) broadly connected at posterior septum of each segment and extending anteriorly, joining corresponding lobe in previous segment by a thin extension. No abrupt division between esophagus and intestine. Chloragogen cells cover the gut usually beginning in VII; in the most posterior segments many free eleocytes present in the coelomic cavity (
<figureCitation id="132E2BDAFF89FF85FBB8FAE2DA80B026" box="[1135,1229,1339,1365]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="22" pageNumber="23">Fig. 9H</figureCitation>
).
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AFAA4DB38B33A" blockId="22.[189,1399,1405,1610]" pageId="22" pageNumber="23">
First nephridia usually on 6/7, absent in VIII–XI, usually paired on 11/12, occuring irregularly in posterior segments. Each nephridium with small anteseptal funnel; granular postseptal expansion elongate to ovate (length 
<quantity id="4CED9ABAFF89FF85FEBFFA1DDC48B0AC" box="[360,517,1476,1503]" metricMagnitude="-1" metricUnit="m" metricValue="1.9" metricValueMax="2.5" metricValueMin="1.3" pageId="22" pageNumber="23" unit="mm" value="190.0" valueMax="250.0" valueMin="130.0">130–250 µm</quantity>
, diameter 40–80 μm), directed posteriad or somewhat dorsad (
<figureCitation id="132E2BDAFF89FF85FAC7FA1DDB2AB0AD" box="[1296,1383,1476,1502]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="22" pageNumber="23">Fig. 9I</figureCitation>
); convoluted efferent duct may pass through one or more adjacent (anterior or posterior) segments, ventral or ventrolateral to the gut, usually near the ventral blood vessel, terminating in a short ectal branch to a simple nephridiopore anterior to the ventral chaetae; indistinct vesicle at the pore in some specimens.
</paragraph>
<paragraph id="8BAA375FFF89FF85FF6AF9A8DB1BB211" blockId="22.[189,1399,1649,1890]" pageId="22" pageNumber="23">Dorsal blood vessel passes under brain: two forks pass around the pharynx and join in IV, forming the ventral vessel. Dorsal vessel free anteriorly, closely appressed to top of gut posterior to VI or VII. Ventral vessel separate from gut, but 2–3 short vessels join it to the perivisceral sinus in each segment posterior to about VII. One pair of commissural vessels in anterior segments; those in II–VI (or VII) long and sinuous, extending through most of originating segment; those in VIII to XII (or XIV) restricted to posterior part of the segment, but a posterior loop from those in IX and X may enter sperm and egg sacs. No lateral blood vessels observed behind about segment XIV. Perivisceral sinus begins in about VII.</paragraph>
<paragraph id="8BAA375FFF89FF8BFF6AF850DB1CB4C5" blockId="22.[189,1398,1929,2027]" lastBlockId="24.[189,1398,267,438]" lastPageId="24" lastPageNumber="25" pageId="22" pageNumber="23">
Genital field covers ventral side of IX, flattened or slightly concave in preserved, mature worms. Male and spermathecal pores paired in IX. Male pores near posterior intersegmental groove (
<figureCitation id="132E2BDAFF89FF85FB69F874DB4EB2B4" box="[1214,1283,1965,1991]" captionStart="Fig" captionStartId="23.[189,232,1717,1743]" captionTargetBox="[199,1387,314,1670]" captionTargetId="figure@23.[189,1398,314,1681]" captionTargetPageId="23" captionText="Fig. 7. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. A–B. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. C–D. Anterior end, showing orientation of chaetae. E–G. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. H–I. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836522" httpUri="https://zenodo.org/record/3836522/files/figure.png" pageId="22" pageNumber="23">Fig. 7</figureCitation>
A–B) and inside lines of ventral chaetae; spermathecal pores in line with, and in front of male pores, about level with dorsal chaetal bundles (ventral chaetae absent in IX). Female pores paired, on chaetal line at intersegmental groove 10/11. Secretory openings of copulatory glands in IX, small and circular areas, lateral to genital pores, and either in front of or behind them. Paired testes on anterior septa in VIII and IX may be large, extending through anterior half of segment; ovaries in X, reaching to posterior septum, or into XI. Sperm sacs extending back to XIV–XVII in mature worms, egg sacs as far as XV–XVII.
</paragraph>
<caption id="DF6A67D7FF88FF84FF6AF96CDC2BB29A" ID-DOI="http://doi.org/10.5281/zenodo.3836522" ID-Zenodo-Dep="3836522" httpUri="https://zenodo.org/record/3836522/files/figure.png" pageId="23" pageNumber="24" startId="23.[189,232,1717,1743]" targetBox="[199,1387,314,1670]" targetPageId="23">
<paragraph id="8BAA375FFF88FF84FF6AF96CDC2BB29A" blockId="23.[189,1399,1717,2025]" pageId="23" pageNumber="24">
<emphasis id="B961EB4DFF88FF84FF6AF96CDF40B3BC" bold="true" box="[189,269,1717,1743]" pageId="23" pageNumber="24">Fig. 7.</emphasis>
<taxonomicName id="4C154CDCFF88FF84FEC0F96CDCBFB3BC" authority="Fend &amp; Rodriguez" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[279,754,1717,1743]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF88FF84FEC0F96CDC44B3BC" box="[279,521,1717,1743]" italics="true" pageId="23" pageNumber="24">Guestphalinus exilis</emphasis>
Fend &amp; Rodriguez
</taxonomicName>
<taxonomicNameLabel id="A2525636FF88FF84FD2BF96FDD11B3A3" box="[764,860,1718,1744]" pageId="23" pageNumber="24" rank="species">sp. nov.</taxonomicNameLabel>
, external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. 
<emphasis id="B961EB4DFF88FF84FB08F901DB5BB381" bold="true" box="[1247,1302,1752,1778]" pageId="23" pageNumber="24">A–B</emphasis>
. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. 
<emphasis id="B961EB4DFF88FF84FB1DF89BDB4EB22F" bold="true" box="[1226,1283,1858,1884]" pageId="23" pageNumber="24">C–D</emphasis>
. Anterior end, showing orientation of chaetae. 
<emphasis id="B961EB4DFF88FF84FDA3F8BCDCE0B20C" bold="true" box="[628,685,1893,1919]" pageId="23" pageNumber="24">E–G</emphasis>
. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. 
<emphasis id="B961EB4DFF88FF84FD53F851DCF8B2D1" bold="true" box="[644,693,1928,1954]" pageId="23" pageNumber="24">H–I</emphasis>
. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 
<emphasis id="B961EB4DFF88FF84FDE0F815DC05B29B" bold="true" box="[567,584,1996,2024]" pageId="23" pageNumber="24">M</emphasis>
m.
</paragraph>
</caption>
<paragraph id="8BAA375FFF87FF8BFF6AFE04DB3AB66B" blockId="24.[189,1399,477,792]" pageId="24" pageNumber="25">
Spermathecae to over 
<quantity id="4CED9ABAFF87FF8BFE1FFE04DC79B48B" box="[456,564,477,504]" metricMagnitude="0" metricUnit="m" metricValue="2.0" pageId="24" pageNumber="25" unit="mm" value="2000.0">2000 µm</quantity>
long in mated worms, nearly tubular, with two, weakly differentiated sections, both containing sperm. Ectal, duct-like section gradually expanding to form ental ampulla; ampulla may be folded within IX (both in unmated or mated specimens) or extending into X–XII. Ectally, spermatheca passes through a short (12 μm), narrow constriction within ring of muscle fibers (
<figureCitation id="132E2BDAFF87FF8BFF12FDB4DF5CB7F4" box="[197,273,621,647]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Figs 8</figureCitation>
A–B, 9J) and terminates in a shallow epidermal infolding, about 70–110 μm deep. Duct-like section (80–100 μm diameter) with irregular, columnar epithelium (up to 40–45 μm high); dense sperm may be lined up along epithelium (
<figureCitation id="132E2BDAFF87FF8BFD88FD6CDCF3B7BC" box="[607,702,693,719]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="24" pageNumber="25">Fig. 9K</figureCitation>
); very thin outer muscle layer about 2 μm thick. Ampullalike section nearly tubular, diameter about 
<quantity id="4CED9ABAFF87FF8BFD63FD00DD42B787" box="[692,783,729,756]" metricMagnitude="-1" metricUnit="m" metricValue="1.0" pageId="24" pageNumber="25" unit="mm" value="100.0">100 µm</quantity>
for much of its length, but may be entally expanded to over 200 μm (
<figureCitation id="132E2BDAFF87FF8BFEA9FD24DF92B664" box="[382,479,765,791]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Figs 8A</figureCitation>
, 
<figureCitation id="132E2BDAFF87FF8BFE3CFD24DC42B664" box="[491,527,765,791]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="24" pageNumber="25">9L</figureCitation>
), with thinner epithelium and wider lumen containing sparse spermatozoa.
</paragraph>
<paragraph id="8BAA375FFF87FF8BFF6AFCE6DB66B69A" blockId="24.[189,1399,831,1002]" pageId="24" pageNumber="25">
Typically, 3–6 stalked copulatory glands are associated with genital pores in mature and nearly-mature worms (
<figureCitation id="132E2BDAFF87FF8BFF12FCBADF45B60E" box="[197,264,867,893]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Fig. 8</figureCitation>
A–C); they are absent at early stages of reproductive development. Glands are a pyriform cluster of granular, petiolate cells 160–300 μm long (
<figureCitation id="132E2BDAFF87FF8BFD49FC5EDC92B6D2" box="[670,735,903,929]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="24" pageNumber="25">Fig. 9</figureCitation>
M–N), sometimes appearing as a group of smaller clusters. Conjoined cell extensions of the gland are constricted by a ring of circular muscle fibers and then surrounded by thin epithelium before opening in a round secretory surface (to 
<quantity id="4CED9ABAFF87FF8BFC72FC16DDA2B699" box="[933,1007,975,1002]" metricMagnitude="-2" metricUnit="m" metricValue="5.0" pageId="24" pageNumber="25" unit="mm" value="50.0">50 µm</quantity>
diameter) on the body wall.
</paragraph>
<paragraph id="8BAA375FFF87FF8BFF6AFBC8DB12B054" blockId="24.[189,1399,1041,1319]" pageId="24" pageNumber="25">
Anterior and posterior male funnels about equal in size, to 240 μm high; anterior pair rather flat on septum 8/9; posterior pair on 9/10, but usually directed back into X; both pairs functional, with sperm when fully mature. Anterior vas deferens extends into VIII (
<figureCitation id="132E2BDAFF87FF8BFC4BFB80DDA5B100" box="[924,1000,1113,1139]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Figs 8</figureCitation>
A–C, 9O), forming a convoluted mass, then penetrates 8/9, running along ventral body wall to near the male pore, then follows atrium to near the ental end, joining the atrium apically. Length of anterior vasa deferentia to 2600 μm, width 36–50 μm. Posterior vas deferens forms a compact, convoluted mass in posterior IX, then follows atrium within the sperm sac, joining it at or near the apical end. Length of posterior vasa over 2000 μm, width 38–46 μm. Vasa deferentia histologically similar throughout, with ciliated, non-glandular epithelium.
</paragraph>
<paragraph id="8BAA375FFF87FF8BFF6AFA96DF29B3FA" blockId="24.[189,1399,1359,1673]" pageId="24" pageNumber="25">
Atria of mature worms usually extend back into X or XI; in nearly-mature worms they may be entirely in IX. Atria petiolate in mature worms (
<figureCitation id="132E2BDAFF87FF8BFD51FAAADC84B0FE" box="[646,713,1395,1421]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Fig. 8</figureCitation>
A–B); a short ectal duct (150 μm long) has thick, columnar epithelium and a thin (to 2 μm) muscle coat; no distinct penis, although duct may be somewhat expanded at male pore, with thickened epithelium (
<figureCitation id="132E2BDAFF87FF8BFD72FA62DD4BB0A6" box="[677,774,1467,1493]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Figs 8B</figureCitation>
, 
<figureCitation id="132E2BDAFF87FF8BFCC4FA62DD76B0A6" box="[787,827,1467,1493]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="24" pageNumber="25">9Q</figureCitation>
). An abrupt transition from the atrial duct to the tubular or sacciform atrial ampulla. Ampulla length to 
<date id="FFAB119FFF87FF8BFC95FA06DD88B089" box="[834,965,1503,1530]" pageId="24" pageNumber="25" value="1060" valueMax="1105">1060–1105</date>
μm, width 130–150 μm, thin-walled (5–8 μm) with cuboidal epithelium and wide lumen (
<figureCitation id="132E2BDAFF87FF8BFCEEF9DADDD1B36E" box="[825,924,1539,1565]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Figs 8A</figureCitation>
, 
<figureCitation id="132E2BDAFF87FF8BFC7CF9DADD80B36E" box="[939,973,1539,1565]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="24" pageNumber="25">9P</figureCitation>
); clusters of prostatic cells, 70–140 μm long, sparsely cover the atrial ampulla. In nearly-mature, unmated worms, atria tubular (
<figureCitation id="132E2BDAFF87FF8BFADDF9FEDB25B332" box="[1290,1384,1575,1601]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Fig. 8C</figureCitation>
); ampulla and duct not greatly differentiated; prostates small but appear more densely packed than in mated worms.
</paragraph>
<paragraph id="8BAA375FFF87FF8BFF6AF968DB39B298" blockId="24.[189,1399,1713,2027]" pageId="24" pageNumber="25">
Eel River, Stony Creek and Garcia River, northern 
<collectingRegion id="49D1F9BDFF87FF8BFCDDF968DDCBB3B8" box="[778,902,1713,1739]" country="United States of America" name="California" pageId="24" pageNumber="25">California</collectingRegion>
: The few, partially-mature specimens from these localities are tentatively assigned to 
<taxonomicName id="4C154CDCFF87FF8BFD65F90CDDEFB39C" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[690,930,1749,1775]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="24" pageNumber="25" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF87FF8BFD65F90CDDEFB39C" box="[690,930,1749,1775]" italics="true" pageId="24" pageNumber="25">Guestphalinus exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF87FF8BFC7DF90FDA4BB383" box="[938,1030,1750,1776]" pageId="24" pageNumber="25" rank="species">sp. nov.</taxonomicNameLabel>
based on the ringed proboscis, the chaetae, and gonads in VIII, IX and X. They differ from the 
<typeStatus id="54AE89FDFF87FF8BFC63F920DDAAB260" box="[948,999,1785,1811]" pageId="24" pageNumber="25">type</typeStatus>
diagnosis in that chaetae in II are smaller than other anterior chaetae, and simple-pointed (but oriented forward, as in the 
<typeStatus id="54AE89FDFF87FF8BFB35F8C4DB58B244" box="[1250,1301,1821,1847]" pageId="24" pageNumber="25">type</typeStatus>
locality population). Ventral chaetae in posterior segments distinctly keeled and larger than corresponding dorsals (
<figureCitation id="132E2BDAFF87FF8BFEF5F8BCDF24B20C" box="[290,361,1893,1919]" captionStart="Fig" captionStartId="23.[189,232,1717,1743]" captionTargetBox="[199,1387,314,1670]" captionTargetId="figure@23.[189,1398,314,1681]" captionTargetPageId="23" captionText="Fig. 7. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., external characters. A–C, E–G from Squaw Creek, CA(the type locality); D, H from South Fork Eel River, CA; I from Stony Creek, CA. A–B. Ventral views of two worms, showing positions of genital pores: A is partially-mature, with 3 glands opening around male and spermathecal pores; B is from a reproductively-mature worm, dashed line encloses the non-clitellar genital field with 5 glands opening around spermathecal and male pores. C–D. Anterior end, showing orientation of chaetae. E–G. Selected chaetae from typical specimens; G shows relative position of dorsal chaetae in II and III. H–I. Selected chaetae from partially-mature worms. Chaetae are numbered by segment (Roman numerals), followed by “d” (dorsal) or “v” (ventral). Scale bars: A–B = 2 mm; C–D = 1 mm; E–I = 100 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836522" httpUri="https://zenodo.org/record/3836522/files/figure.png" pageId="24" pageNumber="25">Fig. 7</figureCitation>
H–I). As in typical 
<taxonomicName id="4C154CDCFF87FF8BFD84F8BCDCFAB20C" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[595,695,1893,1919]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="24" pageNumber="25" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF87FF8BFD84F8BCDCFAB20C" box="[595,695,1893,1919]" italics="true" pageId="24" pageNumber="25">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF87FF8BFD17F8BCDD52B20C" box="[704,799,1893,1919]" pageId="24" pageNumber="25" rank="species">sp. nov.</taxonomicNameLabel>
, specimens from Eel River have semiprosoporous male ducts, with vasa deferentia joining atrium apically and the anterior vas deferens penetrating 8/9 (
<figureCitation id="132E2BDAFF87FF8BFF11F874DF69B2B4" box="[198,292,1965,1991]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="24" pageNumber="25">Fig. 8D</figureCitation>
). Spermathecal and male pores are behind ventral chaetae of IX and 5 large copulatory glands surround the genital pores. One specimen has an additional, developing spermatheca on one side, in X.
</paragraph>
</subSubSection>
<caption id="DF6A67D7FF86FF8AFF6AF925DD0BB29A" ID-DOI="http://doi.org/10.5281/zenodo.3836524" ID-Zenodo-Dep="3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="25" pageNumber="26" startId="25.[189,232,1788,1814]" targetBox="[204,1385,280,1758]" targetPageId="25">
<paragraph id="8BAA375FFF86FF8AFF6AF925DD0BB29A" blockId="25.[189,1399,1788,2025]" pageId="25" pageNumber="26">
<emphasis id="B961EB4DFF86FF8AFF6AF925DF45B265" bold="true" box="[189,264,1788,1814]" pageId="25" pageNumber="26">Fig. 8.</emphasis>
<taxonomicName id="4C154CDCFF86FF8AFED9F925DC9BB265" authority="Fend &amp; Rodriguez" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[270,726,1788,1814]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="25" pageNumber="26" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF86FF8AFED9F925DFB6B265" box="[270,507,1788,1814]" italics="true" pageId="25" pageNumber="26">Guestphalinus exilis</emphasis>
Fend &amp; Rodriguez
</taxonomicName>
<taxonomicNameLabel id="A2525636FF86FF8AFD0BF924DD7AB264" box="[732,823,1789,1815]" pageId="25" pageNumber="26" rank="species">sp. nov.</taxonomicNameLabel>
, reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River 
<emphasis id="B961EB4DFF86FF8AFB08F8C6DAB9B24A" bold="true" box="[1247,1268,1823,1849]" pageId="25" pageNumber="26">A</emphasis>
. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. 
<emphasis id="B961EB4DFF86FF8AFF6AF8BFDE9CB2F3" bold="true" box="[189,209,1894,1920]" pageId="25" pageNumber="26">B</emphasis>
. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. 
<emphasis id="B961EB4DFF86FF8AFF6AF850DE9FB2D0" bold="true" box="[189,210,1929,1955]" pageId="25" pageNumber="26">C</emphasis>
. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. 
<emphasis id="B961EB4DFF86FF8AFCC9F874DD7EB2B4" bold="true" box="[798,819,1965,1991]" pageId="25" pageNumber="26">D</emphasis>
. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular.
</paragraph>
</caption>
<caption id="DF6A67D7FF85FF89FF6AF9B4DF3FB29B" ID-DOI="http://doi.org/10.5281/zenodo.3836526" ID-Zenodo-Dep="3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="26" pageNumber="27" startId="26.[189,232,1645,1671]" targetBox="[189,1398,301,1602]" targetPageId="26">
<paragraph id="8BAA375FFF85FF89FF6AF9B4DF3FB29B" blockId="26.[189,1399,1645,2024]" pageId="26" pageNumber="27">
<emphasis id="B961EB4DFF85FF89FF6AF9B4DF46B3F4" bold="true" box="[189,267,1645,1671]" pageId="26" pageNumber="27">Fig. 9.</emphasis>
<taxonomicName id="4C154CDCFF85FF89FEC4F9B4DCAAB3F4" authority="Fend &amp; Rodriguez" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[275,743,1645,1671]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="26" pageNumber="27" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF85FF89FEC4F9B4DC4EB3F4" box="[275,515,1645,1671]" italics="true" pageId="26" pageNumber="27">Guestphalinus exilis</emphasis>
Fend &amp; Rodriguez
</taxonomicName>
<taxonomicNameLabel id="A2525636FF85FF89FD38F9B7DD00B3FB" box="[751,845,1646,1672]" pageId="26" pageNumber="27" rank="species">sp. nov.</taxonomicNameLabel>
, from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. 
<emphasis id="B961EB4DFF85FF89FE2DF949DC42B3D9" bold="true" box="[506,527,1680,1706]" pageId="26" pageNumber="27">A</emphasis>
. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. 
<emphasis id="B961EB4DFF85FF89FE29F96DDC5FB3BD" bold="true" box="[510,530,1716,1742]" pageId="26" pageNumber="27">B</emphasis>
. Tips of chaetae in II dorsal. 
<emphasis id="B961EB4DFF85FF89FCA4F96ADDC5B3BE" bold="true" box="[883,904,1715,1741]" pageId="26" pageNumber="27">C</emphasis>
. Chaetae in II ventral. 
<emphasis id="B961EB4DFF85FF89FB4AF96DDAFFB3BD" bold="true" box="[1181,1202,1716,1742]" pageId="26" pageNumber="27">D</emphasis>
. Tips of chaetae in two posterior segments, showing dorsal keel. 
<emphasis id="B961EB4DFF85FF89FCDDF90EDD53B382" bold="true" box="[778,798,1751,1777]" pageId="26" pageNumber="27">E</emphasis>
. Chaetae in II dorsal. 
<emphasis id="B961EB4DFF85FF89FBE5F90EDA09B382" bold="true" box="[1074,1092,1751,1777]" pageId="26" pageNumber="27">F</emphasis>
. Pharynx. 
<emphasis id="B961EB4DFF85FF89FB1DF90EDAACB382" bold="true" box="[1226,1249,1751,1777]" pageId="26" pageNumber="27">G</emphasis>
. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). 
<emphasis id="B961EB4DFF85FF89FC65F923DD84B267" bold="true" box="[946,969,1786,1812]" pageId="26" pageNumber="27">H</emphasis>
. Eleocytes, in a posterior segment. 
<emphasis id="B961EB4DFF85FF89FF6AF8C4DE85B244" bold="true" box="[189,200,1821,1847]" pageId="26" pageNumber="27">I</emphasis>
. Nephridial funnel (left) and postseptal expansion. 
<emphasis id="B961EB4DFF85FF89FCE9F8C4DD00B244" bold="true" box="[830,845,1821,1847]" pageId="26" pageNumber="27">J</emphasis>
. Spermathecal pore and constricted ectal end of duct. 
<emphasis id="B961EB4DFF85FF89FEFEF898DF0DB228" bold="true" box="[297,320,1857,1883]" pageId="26" pageNumber="27">K</emphasis>
. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. 
<emphasis id="B961EB4DFF85FF89FB78F898DA8EB228" bold="true" box="[1199,1219,1857,1883]" pageId="26" pageNumber="27">L</emphasis>
. Spermathecal ampulla, ental part with weakly-staining sperm. 
<emphasis id="B961EB4DFF85FF89FCD0F8BDDD6EB20D" bold="true" box="[775,803,1892,1918]" pageId="26" pageNumber="27">M</emphasis>
. Ventral copulatory gland. 
<emphasis id="B961EB4DFF85FF89FBB8F8BDDAC9B20D" bold="true" box="[1135,1156,1892,1918]" pageId="26" pageNumber="27">N</emphasis>
. Pore of copulatory gland. 
<emphasis id="B961EB4DFF85FF89FEDBF85EDF6EB2D2" bold="true" box="[268,291,1927,1953]" pageId="26" pageNumber="27">O</emphasis>
. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. 
<emphasis id="B961EB4DFF85FF89FB48F85EDAFCB2D2" bold="true" box="[1183,1201,1927,1953]" pageId="26" pageNumber="27">P</emphasis>
. Expanded atrial ampulla with prostates. 
<emphasis id="B961EB4DFF85FF89FE0BF873DFBEB2B7" bold="true" box="[476,499,1962,1988]" pageId="26" pageNumber="27">Q</emphasis>
. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm.
</paragraph>
</caption>
<subSubSection id="C30F64D4FF84FF88FF6AFED2DD47B1F7" pageId="27" pageNumber="28" type="discussion">
<paragraph id="8BAA375FFF84FF88FF6AFED2DF62B456" blockId="27.[189,303,267,293]" box="[189,303,267,293]" pageId="27" pageNumber="28">
<heading id="D0E28033FF84FF88FF6AFED2DF62B456" bold="true" box="[189,303,267,293]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="3">
<emphasis id="B961EB4DFF84FF88FF6AFED2DF62B456" bold="true" box="[189,303,267,293]" pageId="27" pageNumber="28">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BAA375FFF84FF88FF6AFEE2DF03B7F3" blockId="27.[189,1399,315,640]" pageId="27" pageNumber="28">
<taxonomicName id="4C154CDCFF84FF88FF6AFEE2DFFEB426" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[189,435,315,341]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FF6AFEE2DFFEB426" box="[189,435,315,341]" italics="true" pageId="27" pageNumber="28">Guestphalinus exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FE16FEE5DC6FB425" box="[449,546,316,342]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
from western North America resembles the Palearctic 
<taxonomicName id="4C154CDCFF84FF88FB0CFEE5DB1FB426" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[1243,1362,315,342]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF84FF88FB0CFEE5DB1FB426" box="[1243,1362,315,342]" italics="true" pageId="27" pageNumber="28">G. wiardi</emphasis>
</taxonomicName>
in characters considered diagnostic for the genus (see above). Compared with 
<taxonomicName id="4C154CDCFF84FF88FBE2FEB8DAE8B408" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[1077,1189,353,379]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF84FF88FBE2FEB8DAE8B408" box="[1077,1189,353,379]" italics="true" pageId="27" pageNumber="28">G. wiardi</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C154CDCFF84FF88FB64FEB8DB58B408" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[1203,1301,353,379]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FB64FEB8DB58B408" box="[1203,1301,353,379]" italics="true" pageId="27" pageNumber="28">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FACBFEB8DB3AB408" box="[1308,1399,353,379]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
has modified chaetae in II, a much larger, sacciform atrium, and more copulatory glands; the male pore was never strongly protruding (
<figureCitation id="132E2BDAFF84FF88FDEFFE72DCD8B4B6" box="[568,661,427,453]" captionStart="Fig" captionStartId="20.[189,232,1754,1780]" captionTargetBox="[209,1378,277,1716]" captionTargetId="figure@20.[202,1385,263,1727]" captionTargetPageId="20" captionText="Fig. 6. Guestphalinus wiardi (Michaelsen, 1933); A–E a sectioned worm from Slovenia, Hrabě’s collection; F–I, a dissected Italian specimen. A. Male duct and spermatheca in IX, reconstructed from sections; pore from a copulatory gland (not shown) opens next to spermathecal pore. B. Atrium and male porophore. C. Copulatory gland. D. Spermathecal pore and duct. E. Anterior male funnel with vas deferens entering pre-atrial segment. F. Posterior ventral chaeta, showing grooved dorsal tooth. G. Reproductive organs in IX-XI, figure rotated to face to right. H. Detail of G, showing position of copulatory gland, from median view. I. Branched lateral blood vessel in posterior segment. Scale bars: A, G–H = 200 Mm; B, E, I = 100 Mm; C–D, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="27" pageNumber="28">Fig. 6B</figureCitation>
vs 
<figureCitation id="132E2BDAFF84FF88FD15FE72DD45B4B5" box="[706,776,427,454]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="27" pageNumber="28">Fig. 8</figureCitation>
A–B). The reproductive organs of 
<taxonomicName id="4C154CDCFF84FF88FB7CFE75DB5DB4B6" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[1195,1296,427,454]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FB7CFE75DB5DB4B6" box="[1195,1296,427,454]" italics="true" pageId="27" pageNumber="28">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FACDFE75DB3AB4B5" box="[1306,1399,428,454]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
are morphologically similar to those of 
<taxonomicName id="4C154CDCFF84FF88FD44FE08DD03B498" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[659,846,465,491]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="elephantinus" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FD44FE08DD03B498" box="[659,846,465,491]" italics="true" pageId="27" pageNumber="28">G. elephantinus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FC80FE08DDFFB498" box="[855,946,465,491]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
(see below); both lack a distinct male porophore or penis, although the atrial duct may be somewhat expanded at the male pore, with thickened epithelium, suggesting that lining cells may be protrusible (
<figureCitation id="132E2BDAFF84FF88FC7FFDC2DA42B745" box="[936,1039,539,566]" captionStart="Fig" captionStartId="26.[189,232,1645,1671]" captionTargetBox="[189,1398,301,1602]" captionTargetId="figure@26.[189,1398,300,1604]" captionTargetPageId="26" captionText="Fig. 9. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., from Squaw Creek (the type locality); F, I–N, P–Q from sagittal sections. A. A complete worm, anterior end, lightly stained with hematoxylin, showing clitellum and genital field. B. Tips of chaetae in II dorsal. C. Chaetae in II ventral. D. Tips of chaetae in two posterior segments, showing dorsal keel. E. Chaetae in II dorsal. F. Pharynx. G. Pharyngeal glands from a whole mount, 3 lobes facing anteriad (to left). H. Eleocytes, in a posterior segment. I. Nephridial funnel (left) and postseptal expansion. J. Spermathecal pore and constricted ectal end of duct. K. Spermathecal ampulla, ectal part, with sperm penetrating epithelium. L. Spermathecal ampulla, ental part with weakly-staining sperm. M. Ventral copulatory gland. N. Pore of copulatory gland. O. Anterior vas deferens protruding into VII (to left), with male funnel on 8/9. P. Expanded atrial ampulla with prostates. Q. Male pore. Scale bars: A = 2 mm; B–E, H–J, N, Q = 50 µm; F–G, K–M, O–P = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836526" httpUri="https://zenodo.org/record/3836526/files/figure.png" pageId="27" pageNumber="28">Figs 9Q</figureCitation>
, 
<figureCitation id="132E2BDAFF84FF88FBF5FDC5DAC8B745" box="[1058,1157,540,566]" captionStart="Fig" captionStartId="33.[189,232,1609,1635]" captionTargetBox="[189,1398,267,1566]" captionTargetId="figure@33.[189,1398,265,1566]" captionTargetPageId="33" captionText="Fig. 12. Guestphalinus elephantinus Fend &amp; Rodriguez sp. nov.A–B (upper) from Clearwater River, WA; C, H, K from Shale Creek, WA (type locality); D–E from Satsop River, WA; B (lower), F–G, N–Q from Guadalupe Creek, CA; I–J, L–M from Peavine Ridge, OR. A. Anterior end of cleared worm. B. Chaetae in II-III, comparing worms from northern (upper) and southern (lower) populations. C. Dorsal chaetae in II. D. The outer chaeta in II, showing the “secondary nodulus”. E–F. Inner dorsal chaeta in II; E shows folded keel. G. Modified dorsal chaetae in II. H. Posterior ventral chaeta. I. Posterior ventral chaeta with bifid tip. J. Transverse section of anterior segment, showing lobes of pharyngeal glands. K. Body wall in anterior segment. L. Spermathecal pore. M. Male pore. N. Ectal part of spermatheca from a dissected, mated worm. O. Expanded atrial duct (type 2 penis?) at male pore. P. Copulatory gland. Q. Female funnel and pore on intersegment 10/11. Scale bars: A = 1 mm; B–I, K–M = 50 µm; J, N–Q = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836532" httpUri="https://zenodo.org/record/3836532/files/figure.png" pageId="27" pageNumber="28">12M, O</figureCitation>
) as “type-2 penes” (temporary structures formed by extruded lining cells of the male duct, see 
<bibRefCitation id="EF844AAEFF84FF88FB91FD98DB22B728" author="Rodriguez P. &amp; Giani N." box="[1094,1391,577,603]" pageId="27" pageNumber="28" pagination="35 - 51" refId="ref24981" refString="Rodriguez P. &amp; Giani N. 1994. A preliminary review of the taxonomic characters used for the systematics of the genus Trichodrilus Claparede (Oligochaeta, Lumbriculidae). Hydrobiologia 278: 35 - 51. https: // doi. org / 10.1007 / BF 00142310" type="journal article" year="1994">Rodriguez &amp; Giani 1994</bibRefCitation>
: fig. 17D–F).
</paragraph>
<paragraph id="8BAA375FFF84FF88FF6AFD73DD47B1F7" blockId="27.[189,1399,682,1156]" pageId="27" pageNumber="28">
Molecular results strongly suggest more than one Nearctic species (see below). Both new 
<taxonomicName id="4C154CDCFF84FF88FB1AFD73DB3BB7B7" authorityName="Michaelsen" authorityYear="1933" box="[1229,1398,682,708]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF84FF88FB1AFD73DB3BB7B7" box="[1229,1398,682,708]" italics="true" pageId="27" pageNumber="28">Guestphalinus</emphasis>
</taxonomicName>
species present distinctive chaetal morphotypes, and these chaetal characters are therefore used here as main diagnostic characters. Nevertheless, there is some variation among populations assigned to both basic morphotypes. In segment II chaetae of typical (from Squaw Creek) 
<taxonomicName id="4C154CDCFF84FF88FBABFCC3DAACB647" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[1148,1249,794,820]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FBABFCC3DAACB647" box="[1148,1249,794,820]" italics="true" pageId="27" pageNumber="28">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FB3CFCC3DB04B647" box="[1259,1353,794,820]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
are slightly modified relative to those in other segments. In most Squaw Creek specimens the chaetae appear slightly notched, slightly larger than those in III, and a little shorter than those in middle segments (
<tableCitation id="C69702E4FF84FF88FF12FC50DF6CB6D7" box="[197,289,905,932]" captionStart="Table 3" captionStartId="28.[189,258,268,294]" captionTargetPageId="28" captionText="Table 3. Descriptive measurements (mean, range, and n) for different populations of G. exilis Fend &amp; Rodriguez sp. nov. and G. elephantinus Fend &amp; Rodriguez sp. nov. Measurements of “nodulus” refer to the proportional distance from the tip of the chaeta (i.e., “0.5” means the nodulus is at the midpoint of the chaeta)." pageId="27" pageNumber="28">Table 3</tableCitation>
). In specimens from two other 
<collectingRegion id="49D1F9BDFF84FF88FD48FC50DD5BB6D0" box="[671,790,905,931]" country="United States of America" name="California" pageId="27" pageNumber="28">California</collectingRegion>
sites, chaetae in II are simple-pointed and smaller than those in III – and thus similar to most other lumbriculids. These are provisionally assigned to 
<taxonomicName id="4C154CDCFF84FF88FF6AFC0CDF52B69D" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[189,287,980,1007]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FF6AFC0CDF52B69D" box="[189,287,980,1007]" italics="true" pageId="27" pageNumber="28">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FEF1FC0CDFCEB69C" box="[294,387,981,1007]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
, based on the limited and only partially-mature available material. Partially developed reproductive organs (
<figureCitation id="132E2BDAFF84FF88FE60FC20DC5FB167" box="[439,530,1017,1044]" captionStart="Fig" captionStartId="25.[189,232,1788,1814]" captionTargetBox="[204,1385,280,1758]" captionTargetId="figure@25.[189,1398,265,1773]" captionTargetPageId="25" captionText="Fig. 8. Guestphalinus exilis Fend &amp; Rodriguez sp. nov., reproductive structures, all oriented with anterior end to left. A–C from Squaw Creek (the type locality), D from the South Fork Eel River A. Segments VIII–XI of a mature worm, showing reproductive organs; anterior vas deferens broken at spermatheca. B. Segments IX–X of a recently-mated worm; spermathecal ampulla not yet developed; atrium expanded. C. Segments IX–X of a partially-mature worm; atrium and spermathecae tubular, but copulatory glands well-developed; ental end of spermatheca missing. D. Segment IX of a partially-mature worm; glands well developed; atrium and spermatheca small, tubular." figureDoi="http://doi.org/10.5281/zenodo.3836524" httpUri="https://zenodo.org/record/3836524/files/figure.png" pageId="27" pageNumber="28">Fig. 8D</figureCitation>
) clearly indicate that the Eel River specimens belong to 
<taxonomicName id="4C154CDCFF84FF88FB7AFC23DB1BB167" authorityName="Michaelsen" authorityYear="1933" box="[1197,1366,1018,1044]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF84FF88FB7AFC23DB1BB167" box="[1197,1366,1018,1044]" italics="true" pageId="27" pageNumber="28">Guestphalinus</emphasis>
</taxonomicName>
as defined here. The single specimens from Stony Creek and the Garcia River, although immature, have gonads in VIII–X, distinguishing them from other proboscis-bearing lumbriculid genera in the region (e.g., 
<taxonomicName id="4C154CDCFF84FF88FED7FBB3DFDCB1F7" authorityName="Altman" authorityYear="1936" box="[256,401,1130,1156]" class="Clitellata" family="Lumbriculidae" genus="Kincaidiana" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF84FF88FED7FBB3DFDCB1F7" box="[256,401,1130,1156]" italics="true" pageId="27" pageNumber="28">Kincaidiana</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C154CDCFF84FF88FE48FBB3DC78B1F7" authorityName="Hoffmeister" authorityYear="1843" box="[415,565,1130,1156]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF84FF88FE48FBB3DC78B1F7" box="[415,565,1130,1156]" italics="true" pageId="27" pageNumber="28">Rhynchelmis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C154CDCFF84FF88FDB9FBB3DCB4B1F7" authorityName="Fend &amp; Rodriguez" authorityYear="2003" box="[622,761,1130,1156]" class="Clitellata" family="Lumbriculidae" genus="Eremidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF84FF88FDB9FBB3DCB4B1F7" box="[622,761,1130,1156]" italics="true" pageId="27" pageNumber="28">Eremidrilus</emphasis>
</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="C30F64D4FF84FF88FF6AFB74DC65B3AE" pageId="27" pageNumber="28" type="biology_ecology">
<paragraph id="8BAA375FFF84FF88FF6AFB74DF50B1B4" blockId="27.[189,285,1197,1223]" box="[189,285,1197,1223]" pageId="27" pageNumber="28">
<heading id="D0E28033FF84FF88FF6AFB74DF50B1B4" bold="true" box="[189,285,1197,1223]" fontSize="11" level="3" pageId="27" pageNumber="28" reason="3">
<emphasis id="B961EB4DFF84FF88FF6AFB74DF50B1B4" bold="true" box="[189,285,1197,1223]" pageId="27" pageNumber="28">Habitat</emphasis>
</heading>
</paragraph>
<paragraph id="8BAA375FFF84FF88FF6AFB07DC65B3AE" blockId="27.[189,1399,1246,1757]" pageId="27" pageNumber="28">
The type locality is Squaw Creek, a tributary to the Sacramento River, in northern 
<collectingRegion id="49D1F9BDFF84FF88FB10FB07DB0FB18B" box="[1223,1346,1246,1272]" country="United States of America" name="California" pageId="27" pageNumber="28">California</collectingRegion>
; the collection site is a 5 
<superScript id="7C609A17FF84FF88FE7BFAD8DFF4B07C" attach="left" box="[428,441,1281,1295]" fontSize="6" pageId="27" pageNumber="28">th</superScript>
order riffle-pool stream with gravel-cobble sediment (
<collectingCountry id="F30277CFFF84FF88FB94FADADA3BB06E" box="[1091,1142,1283,1309]" name="United States of America" pageId="27" pageNumber="28">U.S.</collectingCountry>
D.A. 1999). The 
<typeStatus id="54AE89FDFF84FF88FA93FADDDB3AB06D" box="[1348,1399,1284,1310]" pageId="27" pageNumber="28">type</typeStatus>
series was found only at the lower end of a single riffle, and no additional material has been found at other sites within the stream. Other oligochaetes were collected throughout the stream; dominant species were 
<taxonomicName id="4C154CDCFF84FF88FED7FAAADCF0B0FE" authority="Eisen, 1904" authorityName="Eisen" authorityYear="1904" box="[256,701,1395,1421]" class="Clitellata" family="Enchytraeidae" genus="Mesenchytraeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Tubificida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="pedatus">
<emphasis id="B961EB4DFF84FF88FED7FAAADC69B0FE" box="[256,548,1395,1421]" italics="true" pageId="27" pageNumber="28">Mesenchytraeus pedatus</emphasis>
Eisen, 1904
</taxonomicName>
and 
<taxonomicName id="4C154CDCFF84FF88FD2CFAAADB48B0FE" authority="Rodriguez &amp; Fend, 2013" authorityName="Rodriguez &amp; Fend" authorityYear="2013" box="[763,1285,1395,1421]" class="Clitellata" family="Naididae" genus="Rhyacodrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="clio">
<emphasis id="B961EB4DFF84FF88FD2CFAAADD83B0FE" box="[763,974,1395,1421]" italics="true" pageId="27" pageNumber="28">Rhyacodrilus clio</emphasis>
<bibRefCitation id="EF844AAEFF84FF88FC0FFAAADB48B0FE" author="Rodriguez P. &amp; Fend S. V." box="[984,1285,1395,1421]" pageId="27" pageNumber="28" pagination="1 - 44" refId="ref24921" refString="Rodriguez P. &amp; Fend S. V. 2013. New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879). Zootaxa 3664 (1): 1 - 44. https: // doi. org / 10.11646 / zootaxa. 3664.1.1" type="journal article" year="2013">Rodriguez &amp; Fend, 2013</bibRefCitation>
</taxonomicName>
(also the 
<typeStatus id="54AE89FDFF84FF88FF6AFA40DEBDB0C0" box="[189,240,1433,1459]" pageId="27" pageNumber="28">type</typeStatus>
locality for 
<taxonomicName id="4C154CDCFF84FF88FE52FA40DF9EB0C1" authorityName="Rodriguez &amp; Fend" authorityYear="2013" box="[389,467,1432,1458]" class="Clitellata" family="Naididae" genus="Rhyacodrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="clio">
<emphasis id="B961EB4DFF84FF88FE52FA40DF9EB0C1" box="[389,467,1432,1458]" italics="true" pageId="27" pageNumber="28">R. clio</emphasis>
</taxonomicName>
). The four streams where 
<taxonomicName id="4C154CDCFF84FF88FCC6FA40DD38B0C1" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[785,885,1432,1459]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FCC6FA40DD38B0C1" box="[785,885,1432,1459]" italics="true" pageId="27" pageNumber="28">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FCA9FA40DD97B0C0" box="[894,986,1433,1459]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
was collected were similar in size and sediment characteristics, and are coldwater streams, supporting salmonid fishes. The species was usually found in patches of finer gravel within cobble-boulder riffles. Specific conductance at the Squaw Creek and Eel River sites (
<date id="FFAB119FFF84FF88FE21F9D1DC37B351" box="[502,634,1544,1570]" pageId="27" pageNumber="28" value="2014-04">April 2014</date>
) was 265 and 250 µS 
<superScript id="7C609A17FF84FF88FC57F9D0DDFFB366" attach="left" box="[896,946,1543,1571]" fontSize="6" pageId="27" pageNumber="28">cm-1</superScript>
, respectively. Earlier measurements at a downstream site on Squaw Creek (
<bibRefCitation id="EF844AAEFF84FF88FDB9F9F7DDDCB33B" author="Rettig S. A. &amp; Bortleson G. C." box="[622,913,1582,1608]" pageId="27" pageNumber="28" pagination="82 - 4081" refId="ref24808" refString="Rettig S. A. &amp; Bortleson G. C. 1983. Limnological Study of Shasta Lake, Shasta County, California, with Emphasis on the Effests of the 1977 Drought. U. S. Geological Survey Water-Resources Investigations 82 - 4081. Available from https: // pubs. er. usgs. gov / publication / wri 824081 [accessed 19 Sep. 2017]." type="book chapter" year="1983">Rettig &amp; Bortleson 1983</bibRefCitation>
) gave temperatures from 13.5 to 22.5 °C and a specific conductance of 109–261 µS 
<superScript id="7C609A17FF84FF88FD6AF98ADCA2B32C" attach="left" box="[701,751,1617,1645]" fontSize="6" pageId="27" pageNumber="28">cm-1</superScript>
. Visits to the 
<typeStatus id="54AE89FDFF84FF88FC42F98ADD85B31E" box="[917,968,1619,1645]" pageId="27" pageNumber="28">type</typeStatus>
locality from July to September did not produce any specimens of 
<taxonomicName id="4C154CDCFF84FF88FDC8F9A0DCCDB3E1" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[543,640,1656,1683]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="exilis" status="sp. nov.">
<emphasis id="B961EB4DFF84FF88FDC8F9A0DCCDB3E1" box="[543,640,1656,1683]" italics="true" pageId="27" pageNumber="28">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF84FF88FD51F9A0DCADB3E0" box="[646,736,1657,1683]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
The sporadic occurrence and patchy distribution suggest that, like 
<taxonomicName id="4C154CDCFF84FF88FEFDF947DFD7B3CB" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[298,410,1694,1720]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="27" pageNumber="28" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF84FF88FEFDF947DFD7B3CB" box="[298,410,1694,1720]" italics="true" pageId="27" pageNumber="28">G. wiardi</emphasis>
</taxonomicName>
, they may also be primarily groundwater or hyporheic worms that only occasionally show up in surface collections.
</paragraph>
</subSubSection>
</treatment>
</document>