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<document id="5DFA92D0042D62AFA62A9CC4780D791A" 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="03BC8649FF8CFF86FDF5FED3DDBCB6FC" 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 wiardi" docType="treatment" docVersion="7" lastPageNumber="22" 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="20" 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="8012F3E8CEBBF79F488FF41BAC68D992">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="B41CAF895CCA39D462BA36ABB2A4ABAF">Fend, Steven V.</mods:namePart>
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<mods:affiliation id="CB98BB39A6F5AAF39562667FA38C1EB5">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="EB29CEF57B2F29167DBA78CF56826E93">Rodriguez, Pilar</mods:namePart>
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<mods:affiliation id="B39B97D280E492F7F9F1B85E2B898959">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="84C607F4AF6F06B291F6FD6A14B775E6">Achurra, Ainara</mods:namePart>
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<mods:affiliation id="EA811E06E8A685FAAA040138728AE9B0">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>
<mods:nameIdentifier id="5F1CB8D46DF1F417F5011BDE2404CACB" type="email">ainara.achurra@ehu.eus</mods:nameIdentifier>
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<mods:namePart id="C6318B77ED91D70270B0575CAA6B85E0">Erséus, Christer</mods:namePart>
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<mods:affiliation id="277709F0D8EC385816DD07A11A20DD97">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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<mods:date id="BFE6B116591B8A4DB57DFD2D0CC06128">2017</mods:date>
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<heading id="D0E28033FF8CFF80FDF5FED3DA5CB456" box="[546,1041,266,293]" centered="true" fontSize="11" level="2" pageId="19" pageNumber="20" reason="2">
<taxonomicName id="4C154CDCFF8CFF80FDF5FED3DA5CB456" ID-CoL="6L5FY" authority="(Michaelsen, 1933)" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[546,1041,266,293]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF8CFF80FDF5FED3DD6BB456" bold="true" box="[546,806,266,293]" italics="true" pageId="19" pageNumber="20">Guestphalinus wiardi</emphasis>
(
<bibRefCitation id="EF844AAEFF8CFF80FCE2FED2DA44B456" author="Michaelsen W." box="[821,1033,267,293]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen, 1933</bibRefCitation>
)
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</heading>
</paragraph>
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<paragraph id="8BAA375FFF8CFF80FD20FEE8DD71B438" blockId="19.[546,1041,266,331]" box="[759,828,305,331]" pageId="19" pageNumber="20">
<figureCitation id="132E2BDAFF8CFF80FD20FEE8DD71B438" box="[759,828,305,331]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6</figureCitation>
</paragraph>
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<paragraph id="8BAA375FFF8CFF80FF6AFEA8DDF3B4F8" blockId="19.[189,958,369,395]" box="[189,958,369,395]" pageId="19" pageNumber="20">
<treatmentCitationGroup id="AB051071FF8CFF80FF6AFEA8DDF3B4F8" box="[189,958,369,395]" pageId="19" pageNumber="20">
<treatmentCitation id="0AB4114EFF8CFF80FF6AFEA8DD00B4F8" author="Michaelsen W." box="[189,845,369,395]" page="7" pageId="19" pageNumber="20" year="1933">
<taxonomicName id="4C154CDCFF8CFF80FF6AFEA8DD00B4F8" authority="Michaelsen, 1933: 7" authorityName="Michaelsen" authorityPageNumber="7" authorityYear="1933" box="[189,845,369,395]" class="Clitellata" family="Dorydrilidae" genus="Dorydrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="wiardi" subGenus="Guestphalinus">
<emphasis id="B961EB4DFF8CFF80FF6AFEA8DF73B4F8" box="[189,318,369,395]" italics="true" pageId="19" pageNumber="20">Dorydrilus</emphasis>
(
<emphasis id="B961EB4DFF8CFF80FE98FEA8DFB5B4F8" box="[335,504,369,395]" italics="true" pageId="19" pageNumber="20">Guestphalinus</emphasis>
)
<emphasis id="B961EB4DFF8CFF80FDDEFEA8DC18B4F8" box="[521,597,369,395]" italics="true" pageId="19" pageNumber="20">wiardi</emphasis>
<bibRefCitation id="EF844AAEFF8CFF80FD8BFEA8DD00B4F8" author="Michaelsen W." box="[604,845,369,395]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen, 1933: 7</bibRefCitation>
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, figs 12.
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</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFE6ADD35B4BE" blockId="19.[189,1254,435,497]" box="[189,888,435,462]" pageId="19" pageNumber="20">
<treatmentCitationGroup id="AB051071FF8CFF80FF6AFE6ADD35B4BE" box="[189,888,435,462]" pageId="19" pageNumber="20">
<taxonomicName id="4C154CDCFF8CFF80FF6AFE6ADFF7B4BE" authorityName="Michaelsen" authorityYear="1933" box="[189,442,435,461]" class="Clitellata" family="Dorydrilidae" genus="Dorydrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF8CFF80FF6AFE6ADFF7B4BE" box="[189,442,435,461]" italics="true" pageId="19" pageNumber="20">Questphalinus wiardi</emphasis>
</taxonomicName>
<treatmentCitation id="0AB4114EFF8CFF80FE00FE6ADCDDB4BE" author="Hrabe S." box="[471,656,435,461]" page="10" pageId="19" pageNumber="20" year="1936">
<bibRefCitation id="EF844AAEFF8CFF80FE00FE6ADCDDB4BE" author="Hrabe S." box="[471,656,435,461]" pageId="19" pageNumber="20" pagination="3 - 18" refId="ref24011" refString="Hrabe S. 1936. Uber Dorydrilus (Piguetia) mirabilis n. subgen., n. sp. aus einem Sodbrunnen in der Umgebung von Basel sowie uber Dorydrilus (Dorydrilus) michaelseni Pig. und Bichaeta sanguinea Bret. Spisy vydavane prirodovedeckou fakultou Masarykovy University [Aus dem Zoologischen Institut der Masaryk-Universitat in Brno] 227: 3 - 18." type="journal article" year="1936">Hrabě 1936: 10</bibRefCitation>
</treatmentCitation>
;
<treatmentCitation id="0AB4114EFF8CFF80FD4AFE6ADD4AB4BD" author="Hrabe S." box="[669,775,435,462]" page="45" pageId="19" pageNumber="20" year="1973">
<bibRefCitation id="EF844AAEFF8CFF80FD4AFE6ADD4AB4BD" author="Hrabe S." box="[669,775,435,462]" pageId="19" pageNumber="20" pagination="39 - 50" refId="ref24075" refString="Hrabe S. 1973. On a collection of Oligochaeta from various parts of Yugoslavia. Biloski vestnik Ljubljana 21: 39 - 50." type="journal article" year="1973">1973: 45</bibRefCitation>
</treatmentCitation>
, figs 46.
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</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFE0EDAABB482" blockId="19.[189,1254,435,497]" box="[189,1254,471,497]" pageId="19" pageNumber="20">
<treatmentCitationGroup id="AB051071FF8CFF80FF6AFE0EDAABB482" box="[189,1254,471,497]" pageId="19" pageNumber="20">
<taxonomicName id="4C154CDCFF8CFF80FF6AFE0EDFF7B482" ID-CoL="6L5FY" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[189,442,471,497]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="19" pageNumber="20" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF8CFF80FF6AFE0EDFF7B482" box="[189,442,471,497]" italics="true" pageId="19" pageNumber="20">Guestphalinus wiardi</emphasis>
</taxonomicName>
<treatmentCitation id="0AB4114EFF8CFF80FE00FE0EDCDAB482" author="Cook D. G." box="[471,663,471,497]" page="281" pageId="19" pageNumber="20" year="1968">
<bibRefCitation id="EF844AAEFF8CFF80FE00FE0EDCDAB482" author="Cook D. G." box="[471,663,471,497]" pageId="19" pageNumber="20" pagination="273 - 289" refId="ref22931" refString="Cook D. G. 1968. The genera of the family Lumbriculidae and the genus Dorydrilus (Annelida, Oligochaeta). Journal of Zoology 156: 273 - 289. https: // doi. org / 10.1111 / j. 1469 - 7998.1968. tb 04352. x" type="journal article" year="1968">Cook 1968: 281</bibRefCitation>
</treatmentCitation>
, fig. 2e; 1971: 237. —
<treatmentCitation id="0AB4114EFF8CFF80FC72FE0EDA25B482" box="[933,1128,471,497]" pageId="19" pageNumber="20">
<bibRefCitation id="EF844AAEFF8CFF80FC72FE0EDA25B482" author="Dembitsky E. B." box="[933,1128,471,497]" pageId="19" pageNumber="20" pagination="12 - 16" refId="ref23145" refString="Dembitsky E. B. 1987. Novyj rod vod nyh maloscetinkovyh cervej v faune Sovetskogo Sojuza. In: Kacalova O. L. &amp; Parele E. A. (eds) Vodnye Maloscetinkovye Cervi. Materialy Sestogo Vsesojuznogo Simpoziuma, Salaspils 27 - 30 Aprelja 1987, Riga: 12 - 16. [In Russian: A new genus of oligochaetes in fauna of the Soviet Union. In: Kacalova O. L. &amp; Parele E. A. (eds) Aquatic Oligochaeta, Materials of the 6 th Symposium, Salaspils, 27 - 30 April 1987]." type="book chapter" year="1987">Dembitsky 1987</bibRefCitation>
</treatmentCitation>
: figs IIII.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C30F64D4FF8CFF80FF6AFDC0DB77B79F" pageId="19" pageNumber="20" type="materials_examined">
<paragraph id="8BAA375FFF8CFF80FF6AFDC0DFE7B740" blockId="19.[189,426,537,563]" box="[189,426,537,563]" pageId="19" pageNumber="20">
<heading id="D0E28033FF8CFF80FF6AFDC0DFE7B740" bold="true" box="[189,426,537,563]" fontSize="11" level="3" pageId="19" pageNumber="20" reason="3">
<emphasis id="B961EB4DFF8CFF80FF6AFDC0DFE7B740" bold="true" box="[189,426,537,563]" pageId="19" pageNumber="20">Material examined</emphasis>
</heading>
</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFD91DCFAB7F5" blockId="19.[189,1399,584,647]" pageId="19" pageNumber="20">
<collectingCountry id="F30277CFFF8CFF80FF6AFD91DF19B711" box="[189,340,584,610]" name="Slovenia" pageId="19" pageNumber="20">SLOVENIA</collectingCountry>
: a single specimen, sagittally sectioned on 3 slides, near Dornberk (Czech National Museum (Prague), Hrabě collection,
<date id="FFAB119FFF8CFF80FDD5FDB5DC36B7F5" box="[514,635,620,647]" pageId="19" pageNumber="20" value="1970-01-25">197025 I</date>
13).
</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFD77DB77B79F" blockId="19.[189,1398,686,748]" pageId="19" pageNumber="20">
<collectingCountry id="F30277CFFF8CFF80FF6AFD77DF5FB7BB" box="[189,274,686,712]" name="Italy" pageId="19" pageNumber="20">ITALY</collectingCountry>
: a sagittally dissected anterior end, Grotta Sulfurea, Frassasi Cave system, Genga/Ancona,
<geoCoordinate id="EE215198FF8CFF80FF6AFD0BDF74B79F" box="[189,313,722,748]" degrees="43.401" direction="north" orientation="latitude" pageId="19" pageNumber="20" precision="55" value="43.401">43.401° N</geoCoordinate>
,
<geoCoordinate id="EE215198FF8CFF80FE92FD0BDFF1B79F" box="[325,444,722,748]" degrees="12.966" direction="east" orientation="longitude" pageId="19" pageNumber="20" precision="55" value="12.966">12.966° E</geoCoordinate>
,
<collectingRegion id="49D1F9BDFF8CFF80FE1EFD0BDC6BB79F" box="[457,550,722,748]" country="Italy" name="Marche" pageId="19" pageNumber="20">Marche</collectingRegion>
, Sharmishtha Dattagupta and Jennifer L. Macalady leg.,
<date id="FFAB119FFF8CFF80FB17FD0BDB7BB79F" box="[1216,1334,722,748]" pageId="19" pageNumber="20" value="2007-06">Jun. 2007</date>
.
</paragraph>
</subSubSection>
<subSubSection id="C30F64D4FF8CFF86FF6AFCCDDDBCB6FC" lastPageId="21" lastPageNumber="22" pageId="19" pageNumber="20" type="description">
<paragraph id="8BAA375FFF8CFF80FF6AFCCDDC8DB65D" blockId="19.[189,704,788,814]" box="[189,704,788,814]" pageId="19" pageNumber="20">
<heading id="D0E28033FF8CFF80FF6AFCCDDC8DB65D" bold="true" box="[189,704,788,814]" fontSize="11" level="3" pageId="19" pageNumber="20" reason="3">
<emphasis id="B961EB4DFF8CFF80FF6AFCCDDC8DB65D" bold="true" box="[189,704,788,814]" pageId="19" pageNumber="20">Description of new material and remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFC9ADFA4B12B" blockId="19.[189,1399,835,1112]" pageId="19" pageNumber="20">
<bibRefCitation id="EF844AAEFF8CFF80FF6AFC9ADF11B62E" author="Hrabe S." box="[189,348,835,861]" pageId="19" pageNumber="20" pagination="39 - 50" refId="ref24075" refString="Hrabe S. 1973. On a collection of Oligochaeta from various parts of Yugoslavia. Biloski vestnik Ljubljana 21: 39 - 50." type="journal article" year="1973">Hrabě (1973)</bibRefCitation>
described a mature, unmated specimen from
<collectingCountry id="F30277CFFF8CFF80FCA9FC9ADDA5B62E" box="[894,1000,835,861]" name="Slovenia" pageId="19" pageNumber="20">Slovenia</collectingCountry>
, with sperm on the male funnels. This specimen from Hraběs collection is reexamined here, but only the anterior body section was available. The figures in
<bibRefCitation id="EF844AAEFF8CFF80FE3DFC53DCCDB6D6" author="Hrabe S." box="[490,640,906,933]" pageId="19" pageNumber="20" pagination="39 - 50" refId="ref24075" refString="Hrabe S. 1973. On a collection of Oligochaeta from various parts of Yugoslavia. Biloski vestnik Ljubljana 21: 39 - 50." type="journal article" year="1973">Hrabě (1973</bibRefCitation>
: figs 67) correspond to photographs 6B and 6C, respectively, in the present paper (note that labels for spermathecal and male pores should be reversed in Hrabě: fig. 6). The Italian specimen is mature, with clitellum from VIIIXIII and sperm in the spermathecae; the anterior 100 segments are represented. Descriptions by
<bibRefCitation id="EF844AAEFF8CFF80FC5EFC2FDA24B162" author="Michaelsen W." box="[905,1129,1014,1041]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen (1933)</bibRefCitation>
and Hrabě were quite detailed, and were reviewed by
<bibRefCitation id="EF844AAEFF8CFF80FD92FBC3DD6CB147" author="Dembitsky E. B." box="[581,801,1050,1076]" pageId="19" pageNumber="20" pagination="12 - 16" refId="ref23145" refString="Dembitsky E. B. 1987. Novyj rod vod nyh maloscetinkovyh cervej v faune Sovetskogo Sojuza. In: Kacalova O. L. &amp; Parele E. A. (eds) Vodnye Maloscetinkovye Cervi. Materialy Sestogo Vsesojuznogo Simpoziuma, Salaspils 27 - 30 Aprelja 1987, Riga: 12 - 16. [In Russian: A new genus of oligochaetes in fauna of the Soviet Union. In: Kacalova O. L. &amp; Parele E. A. (eds) Aquatic Oligochaeta, Materials of the 6 th Symposium, Salaspils, 27 - 30 April 1987]." type="book chapter" year="1987">Dembitsky (1987)</bibRefCitation>
; nevertheless, we can add the following remarks based on both specimens.
</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFB59DFF3B075" blockId="19.[189,1399,1152,1286]" pageId="19" pageNumber="20">
A pair of lateral blood vessels is visible in posterior segments of the Italian specimen, and some of these have a few blind branches (
<figureCitation id="132E2BDAFF8CFF80FDD2FB7DDC14B1CC" box="[517,601,1188,1215]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6I</figureCitation>
). Chaetae in anterior segments are simple-pointed, but tips of most chaetae in posterior segments bear a distinct dorsal groove, which may appear as a small dorsal tooth in lateral view (
<figureCitation id="132E2BDAFF8CFF80FE80FB32DFFDB075" box="[343,432,1259,1286]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6F</figureCitation>
).
</paragraph>
<paragraph id="8BAA375FFF8CFF80FF6AFAF4DD0AB2F6" blockId="19.[189,1399,1325,1925]" pageId="19" pageNumber="20">
The anterior vasa deferentia form a loop in the pre-atrial segment (VIII) before penetrating 8/9 and entering IX to join the atrium (
<figureCitation id="132E2BDAFF8CFF80FDEAFA88DCA5B018" box="[573,744,1361,1388]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6A, E, G</figureCitation>
) in both of our specimens; this unique character was described and illustrated by
<bibRefCitation id="EF844AAEFF8CFF80FDDCFAACDCA7B0E3" author="Michaelsen W." box="[523,746,1397,1424]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen (1933)</bibRefCitation>
, but not mentioned in later descriptions. Posterior vasa deferentia do not penetrate septum 9/10.
<bibRefCitation id="EF844AAEFF8CFF80FD41FA40DD3FB0C0" author="Michaelsen W." box="[662,882,1433,1459]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen (1933)</bibRefCitation>
described and illustrated a greatly expanded posterior sperm funnel extending well into the posterior sperm sacs. Both posterior and anterior male funnels are small in the Slovenian worm (
<figureCitation id="132E2BDAFF8CFF80FD74FA39DD4DB088" box="[675,768,1504,1531]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6A</figureCitation>
); they are much larger in the Italian specimen, and the posterior extends back into X (
<figureCitation id="132E2BDAFF8CFF80FDFDF9DDDCC4B36C" box="[554,649,1540,1567]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6G</figureCitation>
). Vasa deferentia are very thick (to
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) in the Italian worm, joining the atrium subapically; they join the atrium before the apex in the Slovenian worm, running a short distance under the muscle layer to enter the lumen apically, as described by
<bibRefCitation id="EF844AAEFF8CFF80FBA0F995DB00B314" author="Dembitsky E. B." box="[1143,1357,1612,1639]" pageId="19" pageNumber="20" pagination="12 - 16" refId="ref23145" refString="Dembitsky E. B. 1987. Novyj rod vod nyh maloscetinkovyh cervej v faune Sovetskogo Sojuza. In: Kacalova O. L. &amp; Parele E. A. (eds) Vodnye Maloscetinkovye Cervi. Materialy Sestogo Vsesojuznogo Simpoziuma, Salaspils 27 - 30 Aprelja 1987, Riga: 12 - 16. [In Russian: A new genus of oligochaetes in fauna of the Soviet Union. In: Kacalova O. L. &amp; Parele E. A. (eds) Aquatic Oligochaeta, Materials of the 6 th Symposium, Salaspils, 27 - 30 April 1987]." type="book chapter" year="1987">Dembitsky (1987)</bibRefCitation>
for Crimean specimens. Compared with other descriptions, the atrial ampulla appears rather short and ovate in the Slovenian and Italian worms, although the total atrium length (320 and
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, respectively) and width are similar to those of the Crimean specimens (
<bibRefCitation id="EF844AAEFF8CFF80FC97F961DA48B3A1" author="Dembitsky E. B." box="[832,1029,1720,1746]" pageId="19" pageNumber="20" pagination="12 - 16" refId="ref23145" refString="Dembitsky E. B. 1987. Novyj rod vod nyh maloscetinkovyh cervej v faune Sovetskogo Sojuza. In: Kacalova O. L. &amp; Parele E. A. (eds) Vodnye Maloscetinkovye Cervi. Materialy Sestogo Vsesojuznogo Simpoziuma, Salaspils 27 - 30 Aprelja 1987, Riga: 12 - 16. [In Russian: A new genus of oligochaetes in fauna of the Soviet Union. In: Kacalova O. L. &amp; Parele E. A. (eds) Aquatic Oligochaeta, Materials of the 6 th Symposium, Salaspils, 27 - 30 April 1987]." type="book chapter" year="1987">Dembitsky 1987</bibRefCitation>
). Male pores are on a rounded protrusion (referred to as a porophore by
<bibRefCitation id="EF844AAEFF8CFF80FD6FF902DDC8B385" author="Michaelsen W." box="[696,901,1755,1782]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen 1933</bibRefCitation>
), closely behind a groove containing the spermathecal and copulatory gland pores in the Slovenian worm (
<figureCitation id="132E2BDAFF8CFF80FC19F926DA58B269" box="[974,1045,1791,1818]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6</figureCitation>
AB). Although this structure was also illustrated by
<bibRefCitation id="EF844AAEFF8CFF80FE00F8FADCEBB24E" author="Dembitsky E. B." box="[471,678,1827,1853]" pageId="19" pageNumber="20" pagination="12 - 16" refId="ref23145" refString="Dembitsky E. B. 1987. Novyj rod vod nyh maloscetinkovyh cervej v faune Sovetskogo Sojuza. In: Kacalova O. L. &amp; Parele E. A. (eds) Vodnye Maloscetinkovye Cervi. Materialy Sestogo Vsesojuznogo Simpoziuma, Salaspils 27 - 30 Aprelja 1987, Riga: 12 - 16. [In Russian: A new genus of oligochaetes in fauna of the Soviet Union. In: Kacalova O. L. &amp; Parele E. A. (eds) Aquatic Oligochaeta, Materials of the 6 th Symposium, Salaspils, 27 - 30 April 1987]." type="book chapter" year="1987">Dembitsky (1987</bibRefCitation>
: fig. III) it was not seen in our Italian specimen (
<figureCitation id="132E2BDAFF8CFF80FAD0F8FADB24B24D" box="[1287,1385,1827,1854]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6G</figureCitation>
). Prostate glands are large (to about
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high), petiolate clusters of cells, more similar to fig.
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<bibRefCitation id="EF844AAEFF8CFF80FF6AF8B2DFD7B2F6" author="Michaelsen W." box="[189,410,1899,1925]" pageId="19" pageNumber="20" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen (1933)</bibRefCitation>
than to fig. III in
<bibRefCitation id="EF844AAEFF8CFF80FDBEF8B2DD0FB2F6" author="Dembitsky E. B." box="[617,834,1899,1925]" pageId="19" pageNumber="20" pagination="12 - 16" refId="ref23145" refString="Dembitsky E. B. 1987. Novyj rod vod nyh maloscetinkovyh cervej v faune Sovetskogo Sojuza. In: Kacalova O. L. &amp; Parele E. A. (eds) Vodnye Maloscetinkovye Cervi. Materialy Sestogo Vsesojuznogo Simpoziuma, Salaspils 27 - 30 Aprelja 1987, Riga: 12 - 16. [In Russian: A new genus of oligochaetes in fauna of the Soviet Union. In: Kacalova O. L. &amp; Parele E. A. (eds) Aquatic Oligochaeta, Materials of the 6 th Symposium, Salaspils, 27 - 30 April 1987]." type="book chapter" year="1987">Dembitsky (1987)</bibRefCitation>
.
</paragraph>
<paragraph id="8BAA375FFF8CFF86FF6AF874DF8FB4AE" blockId="19.[189,1399,1965,2027]" lastBlockId="21.[189,1399,267,478]" lastPageId="21" lastPageNumber="22" pageId="19" pageNumber="20">
The large copulatory glands (
<figureCitation id="132E2BDAFF8CFF80FDE7F874DCF0B2B4" box="[560,701,1965,1991]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="19" pageNumber="20">Fig. 6C, H</figureCitation>
) are quite similar in structure to the copulatory glands described below in the new Nearctic species, although a distinct muscle layer was not seen at the duct. The illustration in
<bibRefCitation id="EF844AAEFF8AFF86FE4CFED2DC7DB456" author="Hrabe S." box="[411,560,267,293]" pageId="21" pageNumber="22" pagination="39 - 50" refId="ref24075" refString="Hrabe S. 1973. On a collection of Oligochaeta from various parts of Yugoslavia. Biloski vestnik Ljubljana 21: 39 - 50." type="journal article" year="1973">Hrabě (1973</bibRefCitation>
: fig. 7) is similar to ours. The slight difference in placement (adjacent to the spermathecae in both of our specimens), compared with Michaelsens material (at the ventral chaetae), suggests that the position of these glands may be variable, as in
<taxonomicName id="4C154CDCFF8AFF86FBF2FE8CDF7AB4E0" authority="Fend &amp; Rodriguez" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" 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="B961EB4DFF8AFF86FBF2FE8CDB58B41C" box="[1061,1301,341,367]" italics="true" pageId="21" pageNumber="22">Guestphalinus exilis</emphasis>
Fend &amp; Rodriguez
</taxonomicName>
<taxonomicNameLabel id="A2525636FF8AFF86FEEAFEA3DFDBB4E7" box="[317,406,378,404]" pageId="21" pageNumber="22" rank="species">sp. nov.</taxonomicNameLabel>
(see below). Their occurrence may also vary with stage of reproductive development, as Dembitsky (1967) was unable to find these glands in the Crimean specimens, attributing their absence to possible resorption.
</paragraph>
<caption id="DF6A67D7FF8BFF87FF6AF903DD0EB29F" ID-DOI="http://doi.org/10.5281/zenodo.3836520" ID-Zenodo-Dep="3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="20" pageNumber="21" startId="20.[189,232,1754,1780]" targetBox="[209,1378,277,1716]" targetPageId="20">
<paragraph id="8BAA375FFF8BFF87FF6AF903DD0EB29F" blockId="20.[189,1399,1754,2028]" pageId="20" pageNumber="21">
<emphasis id="B961EB4DFF8BFF87FF6AF903DF5CB387" bold="true" box="[189,273,1754,1780]" pageId="20" pageNumber="21">Fig. 6.</emphasis>
<taxonomicName id="4C154CDCFF8BFF87FEC9F902DD51B386" authority="(Michaelsen, 1933)" baseAuthorityName="Michaelsen" baseAuthorityYear="1933" box="[286,796,1755,1781]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="20" pageNumber="21" phylum="Annelida" rank="species" species="wiardi">
<emphasis id="B961EB4DFF8BFF87FEC9F902DC6CB386" box="[286,545,1755,1781]" italics="true" pageId="20" pageNumber="21">Guestphalinus wiardi</emphasis>
(
<bibRefCitation id="EF844AAEFF8BFF87FDE1F902DD5EB386" author="Michaelsen W." box="[566,787,1755,1781]" pageId="20" pageNumber="21" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen, 1933</bibRefCitation>
)
</taxonomicName>
; AE a sectioned worm from Slovenia, Hraběs collection; FI, a dissected Italian specimen.
<emphasis id="B961EB4DFF8BFF87FD01F927DCA6B26B" bold="true" box="[726,747,1790,1816]" pageId="20" pageNumber="21">A</emphasis>
. Male duct and spermatheca in IX, reconstructed from sections; pore from a copulatory gland (not shown) opens next to spermathecal pore.
<emphasis id="B961EB4DFF8BFF87FB1DF8F8DA93B248" bold="true" box="[1226,1246,1825,1851]" pageId="20" pageNumber="21">B</emphasis>
. Atrium and male porophore.
<emphasis id="B961EB4DFF8BFF87FE5DF89DDFD2B22D" bold="true" box="[394,415,1860,1886]" pageId="20" pageNumber="21">C</emphasis>
. Copulatory gland.
<emphasis id="B961EB4DFF8BFF87FD45F89DDCEAB22D" bold="true" box="[658,679,1860,1886]" pageId="20" pageNumber="21">D</emphasis>
. Spermathecal pore and duct.
<emphasis id="B961EB4DFF8BFF87FBCEF89DDA60B22D" bold="true" box="[1049,1069,1860,1886]" pageId="20" pageNumber="21">E</emphasis>
. Anterior male funnel with vas deferens entering pre-atrial segment.
<emphasis id="B961EB4DFF8BFF87FD64F8B1DC88B2F1" bold="true" box="[691,709,1896,1922]" pageId="20" pageNumber="21">F</emphasis>
. Posterior ventral chaeta, showing grooved dorsal tooth.
<emphasis id="B961EB4DFF8BFF87FF6AF853DE99B2D7" bold="true" box="[189,212,1930,1956]" pageId="20" pageNumber="21">G</emphasis>
. Reproductive organs in IX-XI, figure rotated to face to right.
<emphasis id="B961EB4DFF8BFF87FC1FF852DD92B2D6" bold="true" box="[968,991,1931,1957]" pageId="20" pageNumber="21">H</emphasis>
. Detail of G, showing position of copulatory gland, from median view.
<emphasis id="B961EB4DFF8BFF87FDAFF877DCCEB2BB" bold="true" box="[632,643,1966,1992]" pageId="20" pageNumber="21">I</emphasis>
. Branched lateral blood vessel in posterior segment. Scale bars: A, GH = 200
<emphasis id="B961EB4DFF8BFF87FEBBF817DF30B299" bold="true" box="[364,381,1998,2026]" pageId="20" pageNumber="21">M</emphasis>
m; B, E, I = 100
<emphasis id="B961EB4DFF8BFF87FD94F817DC19B299" bold="true" box="[579,596,1998,2026]" pageId="20" pageNumber="21">M</emphasis>
m; CD, F = 50
<emphasis id="B961EB4DFF8BFF87FCC3F817DD68B299" bold="true" box="[788,805,1998,2026]" pageId="20" pageNumber="21">M</emphasis>
m.
</paragraph>
</caption>
<paragraph id="8BAA375FFF8AFF86FF6AFDDFDDBCB6FC" blockId="21.[189,1399,518,912]" pageId="21" pageNumber="22">
The spermatheca is not clearly differentiated into duct vs ampullar sections, although there is a sphincterlike constriction, with thickened circular muscles near the pore of the Slovenian worm (
<figureCitation id="132E2BDAFF8AFF86FAD2FDF2DB24B735" box="[1285,1385,555,582]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="21" pageNumber="22">Fig. 6D</figureCitation>
). The female funnel is clearly visible only in the Italian worm (
<figureCitation id="132E2BDAFF8AFF86FC71FD96DA4BB719" box="[934,1030,591,618]" 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); AE a sectioned worm from Slovenia, Hraběs collection; FI, 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, GH = 200 Mm; B, E, I = 100 Mm; CD, F = 50 Mm." figureDoi="http://doi.org/10.5281/zenodo.3836520" httpUri="https://zenodo.org/record/3836520/files/figure.png" pageId="21" pageNumber="22">Fig. 6G</figureCitation>
), where it appears to penetrate the posterior septum 10/11 (the plesiopore condition), as in the illustration in
<bibRefCitation id="EF844AAEFF8AFF86FB5AFDADDB3CB7FC" author="Michaelsen W." box="[1165,1393,628,655]" pageId="21" pageNumber="22" pagination="1 - 19" refId="ref24546" refString="Michaelsen W. 1933. Uber Hohlen Oligochaten. Mitteilungen uber Hohlen- und Karstforschung, Zeitschrift des Hauptverbandes Deutscher Hohlenforscher 1: 1 - 19." type="journal article" year="1933">Michaelsen (1933)</bibRefCitation>
.
<taxonomicName id="4C154CDCFF8AFF86FF6AFD40DF2BB7C0" authorityName="Michaelsen" authorityYear="1933" box="[189,358,665,691]" class="Clitellata" family="Lumbriculidae" genus="Guestphalinus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF8AFF86FF6AFD40DF2BB7C0" box="[189,358,665,691]" italics="true" pageId="21" pageNumber="22">Guestphalinus</emphasis>
</taxonomicName>
was the only lumbriculid genus considered by
<bibRefCitation id="EF844AAEFF8AFF86FC49FD40DA24B7C0" author="Brinkhurst R. O." box="[926,1129,665,691]" pageId="21" pageNumber="22" pagination="2731 - 2739" refId="ref22789" refString="Brinkhurst R. O. 1989. A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Canadian Journal of Zoology 67: 2731 - 2739. https: // doi. org / 10.1139 / z 89 - 387" type="journal article" year="1989">Brinkhurst (1989</bibRefCitation>
: character 16) to have plesiopore female ducts, in common with
<taxonomicName id="4C154CDCFF8AFF86FD77FD67DD6CB7AB" box="[672,801,702,728]" class="Clitellata" family="Haplotaxidae" genus="Haplotaxis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="21" pageNumber="22" phylum="Annelida" rank="genus">
<emphasis id="B961EB4DFF8AFF86FD77FD67DD6CB7AB" box="[672,801,702,728]" italics="true" pageId="21" pageNumber="22">Haplotaxis</emphasis>
</taxonomicName>
, one of the outgroup taxa in that analysis. However, this character has not been observed in
<taxonomicName id="4C154CDCFF8AFF86FD58FD3ADD07B78F" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[655,842,738,765]" 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="B961EB4DFF8AFF86FD58FD3ADD07B78F" box="[655,842,738,765]" italics="true" pageId="21" pageNumber="22">G. elephantinus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF8AFF86FC85FD3ADDE0B78E" box="[850,941,739,765]" pageId="21" pageNumber="22" rank="species">sp. nov.</taxonomicNameLabel>
nor in
<taxonomicName id="4C154CDCFF8AFF86FBD5FD3ADA28B78F" authority="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus, 2017" authorityName="Fend &amp; Rodriguez &amp; Achurra &amp; Erséus" authorityYear="2017" box="[1026,1125,738,765]" 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="B961EB4DFF8AFF86FBD5FD3ADA28B78F" box="[1026,1125,738,765]" italics="true" pageId="21" pageNumber="22">G. exilis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2525636FF8AFF86FBBAFD3ADA86B78E" box="[1133,1227,739,765]" pageId="21" pageNumber="22" rank="species">sp. nov.</taxonomicNameLabel>
, where female pores appear to be intersegmental (see below,
<figureCitation id="132E2BDAFF8AFF86FD06FCDEDD71B652" box="[721,828,775,801]" 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.AB (upper) from Clearwater River, WA; C, H, K from Shale Creek, WA (type locality); DE from Satsop River, WA; B (lower), FG, NQ from Guadalupe Creek, CA; IJ, LM 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”. EF. 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; BI, KM = 50 µm; J, NQ = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.3836532" httpUri="https://zenodo.org/record/3836532/files/figure.png" pageId="21" pageNumber="22">Fig. 12Q</figureCitation>
). Plesiopore female ducts appear to be unusual in the
<taxonomicName id="4C154CDCFF8AFF86FF30FCF5DFD9B635" authorityName="Vejdovsky" authorityYear="1884" box="[231,404,812,838]" class="Clitellata" family="Lumbriculidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="family">Lumbriculidae</taxonomicName>
(
<bibRefCitation id="EF844AAEFF8AFF86FE73FCF5DC2EB635" author="Brinkhurst R. O." box="[420,611,812,838]" pageId="21" pageNumber="22" pagination="2731 - 2739" refId="ref22789" refString="Brinkhurst R. O. 1989. A phylogenetic analysis of the Lumbriculidae (Annelida, Oligochaeta). Canadian Journal of Zoology 67: 2731 - 2739. https: // doi. org / 10.1139 / z 89 - 387" type="journal article" year="1989">Brinkhurst 1989</bibRefCitation>
;
<bibRefCitation id="EF844AAEFF8AFF86FDA6FCF5DD2BB635" author="Fend S. V. &amp; Ohtaka A." box="[625,870,812,838]" pageId="21" pageNumber="22" pagination="677 - 683" refId="ref23651" refString="Fend S. V. &amp; Ohtaka A. 2004. Yamaguchia toyensis n. sp., n. gen. (Annelida, Clitellata, Lumbriculidae) from profundal lake habitat in Japan. Zoological Science 21: 677 - 683. https: // doi. org / 10.2108 / zsj. 21.677" type="journal article" year="2004">Fend &amp; Ohtaka 2004</bibRefCitation>
), but they may simply have escaped notice in other species descriptions. The exact position of the septum may be difficult to define, as muscle fibers from septa may join the body wall on either side of the female funnel.
</paragraph>
</subSubSection>
</treatment>
</document>