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<document id="A9E7A8FC9DA50B369065DA4BC0A81580" ID-DOI="10.5281/zenodo.194252" ID-GBIF-Dataset="d1e5b589-21c5-43a0-b1fc-3c618b0158aa" ID-ISSN="1175-5326" ID-Zenodo-Dep="194252" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460194987408" checkinUser="plazi" docAuthor="Fend, Steven V. &amp; Brinkhurst, Ralph O." docDate="2010" docId="03C5878EFFF0D41DB2CFFF197EC96ECC" docLanguage="en" docName="zt02407p027.pdf" docOrigin="Zootaxa 2407" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Rhynchelmis paraolchonensis Giani &amp; Martinez-Ansemil 1984" docType="treatment" docVersion="9" lastPageNumber="24" masterDocId="FFFCFFF6FFE5D40AB258FF8E7F59683B" masterDocTitle="Contributions towards a review of the genus Rhynchelmis Hoffmeister (Clitellata: Lumbriculidae)" masterLastPageNumber="27" masterPageNumber="1" pageNumber="22" updateTime="1698236526747" updateUser="plazi">
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<mods:title id="A6001632F8B838F31B45A72AE9A51BE8">Contributions towards a review of the genus Rhynchelmis Hoffmeister (Clitellata: Lumbriculidae)</mods:title>
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<mods:namePart id="331342731CE9DCC8118CB1C4786206CE">Fend, Steven V.</mods:namePart>
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<mods:namePart id="A46E28CC5C14E131046F00EBA9B60B13">Brinkhurst, Ralph O.</mods:namePart>
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<mods:date id="A5DCBF6CAE64819CE6532BA608F74BA8">2010</mods:date>
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<mods:identifier id="5DFB6DE131EBE0CF3C8AA730BBA6E69B" type="DOI">10.5281/zenodo.194252</mods:identifier>
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<treatment id="03C5878EFFF0D41DB2CFFF197EC96ECC" ID-DOI="http://doi.org/10.5281/zenodo.6201863" ID-GBIF-Taxon="119383734" ID-Zenodo-Dep="6201863" LSID="urn:lsid:plazi:treatment:03C5878EFFF0D41DB2CFFF197EC96ECC" httpUri="http://treatment.plazi.org/id/03C5878EFFF0D41DB2CFFF197EC96ECC" lastPageId="23" lastPageNumber="24" pageId="21" pageNumber="22">
<subSubSection id="C3766513FFF0D41FB2CFFF197FA468EF" pageId="21" pageNumber="22" type="nomenclature">
<paragraph id="8BD33698FFF0D41FB2CFFF197B5E688A" blockId="21.[151,1031,151,212]" box="[151,1031,151,177]" pageId="21" pageNumber="22">
<heading id="D09B81F4FFF0D41FB2CFFF197B5E688A" bold="true" box="[151,1031,151,177]" fontSize="11" level="1" pageId="21" pageNumber="22" reason="1">
<emphasis id="B918EA8AFFF0D41FB2CFFF197B5E688A" bold="true" box="[151,1031,151,177]" pageId="21" pageNumber="22">
<emphasis id="B918EA8AFFF0D41FB2CFFF197D0A688A" bold="true" box="[151,595,151,177]" italics="true" pageId="21" pageNumber="22">
Pseudorhynchelmis
<taxonomicName id="4C6C4D1BFFF0D41FB3D4FF197D0A688A" ID-CoL="8TXH3" authority="Giani &amp; Martinez-Ansemil, 1984" authorityName="Giani &amp; Martinez-Ansemil" authorityYear="1984" box="[396,595,151,177]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="paraolchonensis">paraolchonensis</taxonomicName>
</emphasis>
(
<bibRefCitation id="EFFD4B69FFF0D41FB03AFF197CA6688A" author="Giani" box="[610,1023,151,177]" pageId="21" pageNumber="26" refString="Giani, N. &amp; Martinez Ansemil, E. (1984) Deux nouvelles especes de Lumbriculidae du Sud-Ouest de l'Europe. Annales de Limnologie, 20, 157 - 165." type="journal article" year="1984">Giani &amp; Martinez-Ansemil, 1984</bibRefCitation>
)
</emphasis>
</heading>
</paragraph>
<paragraph id="8BD33698FFF0D41FB2CFFF347FA468EF" blockId="21.[151,1031,151,212]" box="[151,253,186,212]" pageId="21" pageNumber="22">
(
<figureCitation id="13572A1DFFF0D41FB2C7FF347FAD68EF" box="[159,244,186,212]" captionStart="FIGURE 10. P" captionStartId="21.[151,255,1688,1712]" captionTargetBox="[152,1432,607,1649]" captionTargetId="figure@21.[151,1436,602,1667]" captionTargetPageId="21" captionText="FIGURE 10. P. paraolchonensis from the Giani collection. (A) spermathecal pores. (B) spermathecal duct and part of (broken) ampulla. (C – D) two transverse sections of a male pore. (E) atrium with prostates. (F) transverse section of a post-clitellar segment, showing the supra-intestinal and dorsal blood vessels. Scale bars = 100 μm." httpUri="https://zenodo.org/record/194262/files/figure.png" pageId="21" pageNumber="22">Fig. 10</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C3766513FFF0D41CB2CFFE8B7D0268C7" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22" type="description">
<paragraph id="8BD33698FFF0D41FB2CFFE8B7CB26A1F" blockId="21.[151,1437,261,548]" pageId="21" pageNumber="22">
<emphasis id="B918EA8AFFF0D41FB2CFFE8B7EAB6924" bold="true" box="[151,498,261,287]" pageId="21" pageNumber="22">Supplementary description.</emphasis>
Dorsal pad of pharynx only moderately developed. Longitudinal muscles in weak bands that are not curled at margins; oblique dorsolateral muscles not strongly developed. A single pair of convoluted commissural vessels in each anterior segment may enter adjacent segment; commissures absent in segments posterior to XI, but a loop enters the sperm sacs from X. Dorsal and ventral vessels prominent, with connections to perivisceral sinus; perivisceral sinus forms a supra-intestinal vessel in posterior segments, without cardiac cells (
<figureCitation id="13572A1DFFF0D41FB367FE4E7EC169E1" box="[319,408,448,474]" captionStart="FIGURE 10. P" captionStartId="21.[151,255,1688,1712]" captionTargetBox="[152,1432,607,1649]" captionTargetId="figure@21.[151,1436,602,1667]" captionTargetPageId="21" captionText="FIGURE 10. P. paraolchonensis from the Giani collection. (A) spermathecal pores. (B) spermathecal duct and part of (broken) ampulla. (C – D) two transverse sections of a male pore. (E) atrium with prostates. (F) transverse section of a post-clitellar segment, showing the supra-intestinal and dorsal blood vessels. Scale bars = 100 μm." httpUri="https://zenodo.org/record/194262/files/figure.png" pageId="21" pageNumber="22">Fig. 10</figureCitation>
F). Nephridia may begin in XIII, but absent in many segments. Nephridia with small postseptal expansion; convoluted tubules extend dorsally, but may also follow ventral blood vessel through several segments; duct only slightly widened at inconspicuous nephropore.
</paragraph>
<caption id="DF136610FFF0D41FB2CFF9167BE86ECA" httpUri="https://zenodo.org/record/194262/files/figure.png" pageId="21" pageNumber="22" targetBox="[152,1432,607,1649]" targetPageId="21">
<paragraph id="8BD33698FFF0D41FB2CFF9167BE86ECA" blockId="21.[151,1436,1688,1777]" pageId="21" pageNumber="22">
<emphasis id="B918EA8AFFF0D41FB2CFF9167E716E8B" bold="true" box="[151,296,1688,1712]" pageId="21" pageNumber="22">FIGURE 10.</emphasis>
<emphasis id="B918EA8AFFF0D41FB368F9177EA46E8B" box="[304,509,1689,1712]" italics="true" pageId="21" pageNumber="22">
P.
<taxonomicName id="4C6C4D1BFFF0D41FB313F9177EA46E8B" box="[331,509,1689,1712]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="21" pageNumber="22" phylum="Annelida" rank="species" species="paraolchonensis">paraolchonensis</taxonomicName>
</emphasis>
from the Giani collection. (A) spermathecal pores. (B) spermathecal duct and part of (broken) ampulla. (C–D) two transverse sections of a male pore. (E) atrium with prostates. (F) transverse section of a post-clitellar segment, showing the supra-intestinal and dorsal blood vessels. Scale bars = 100 μm.
</paragraph>
</caption>
<paragraph id="8BD33698FFF0D41FB29CF8907B7D6FB8" blockId="21.[151,1437,1822,2038]" pageId="21" pageNumber="22">
Spermathecae with irregular, sacciform ampulla that may enter XI, but does not connect to gut; ampulla contains loose, unordered sperm. Spermathecal duct a long, narrow cylinder, length more than 5 times width; duct slightly widened before pore, but otherwise not modified (
<figureCitation id="13572A1DFFF0D41FB124F8E77C826FB8" box="[892,987,1897,1923]" captionStart="FIGURE 10. P" captionStartId="21.[151,255,1688,1712]" captionTargetBox="[152,1432,607,1649]" captionTargetId="figure@21.[151,1436,602,1667]" captionTargetPageId="21" captionText="FIGURE 10. P. paraolchonensis from the Giani collection. (A) spermathecal pores. (B) spermathecal duct and part of (broken) ampulla. (C – D) two transverse sections of a male pore. (E) atrium with prostates. (F) transverse section of a post-clitellar segment, showing the supra-intestinal and dorsal blood vessels. Scale bars = 100 μm." httpUri="https://zenodo.org/record/194262/files/figure.png" pageId="21" pageNumber="22">Figs. 10</figureCitation>
A–B).
</paragraph>
<paragraph id="8BD33698FFF0D41CB29CF8007D0268C7" blockId="21.[151,1437,1822,2038]" lastBlockId="22.[151,1436,152,252]" lastPageId="22" lastPageNumber="23" pageId="21" pageNumber="22">
Atrial muscle layer to nearly 10 Μm thick, lining cells moderately thick, columnar. Prostates in large, irregular bundles; bundles are loosely packed (
<figureCitation id="13572A1DFFF0D41FB0BDF83D7C156FF6" box="[741,844,1971,1997]" captionStart="FIGURE 10. P" captionStartId="21.[151,255,1688,1712]" captionTargetBox="[152,1432,607,1649]" captionTargetId="figure@21.[151,1436,602,1667]" captionTargetPageId="21" captionText="FIGURE 10. P. paraolchonensis from the Giani collection. (A) spermathecal pores. (B) spermathecal duct and part of (broken) ampulla. (C – D) two transverse sections of a male pore. (E) atrium with prostates. (F) transverse section of a post-clitellar segment, showing the supra-intestinal and dorsal blood vessels. Scale bars = 100 μm." httpUri="https://zenodo.org/record/194262/files/figure.png" pageId="21" pageNumber="22">Figs. 10</figureCitation>
D–E). Both pairs of vasa deferentia functional; diameter about 20 Μm (20–27) near funnels, narrowing very slightly to about 18 Μm (15–22); lining cells cuboidal, not glandular. Both vasa join atrium near ental end, travelling at most a short distance under muscles. Male pore on a small papilla within a shallow depression; epidermis slightly thickened, but muscles not strongly developed (
<figureCitation id="13572A1DFFF3D41CB3EBFF6C7D4B68C7" box="[435,530,226,252]" captionStart="FIGURE 10. P" captionStartId="21.[151,255,1688,1712]" captionTargetBox="[152,1432,607,1649]" captionTargetId="figure@21.[151,1436,602,1667]" captionTargetPageId="21" captionText="FIGURE 10. P. paraolchonensis from the Giani collection. (A) spermathecal pores. (B) spermathecal duct and part of (broken) ampulla. (C – D) two transverse sections of a male pore. (E) atrium with prostates. (F) transverse section of a post-clitellar segment, showing the supra-intestinal and dorsal blood vessels. Scale bars = 100 μm." httpUri="https://zenodo.org/record/194262/files/figure.png" pageId="22" pageNumber="23">Figs. 10</figureCitation>
C–D).
</paragraph>
</subSubSection>
<subSubSection id="C3766513FFF3D41DB2CFFEA97EC96ECC" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23" type="discussion">
<paragraph id="8BD33698FFF3D41CB2CFFEA97EA6697A" blockId="22.[151,511,295,321]" box="[151,511,295,321]" pageId="22" pageNumber="23">
<emphasis id="B918EA8AFFF3D41CB2CFFEA97E4B697A" bold="true" box="[151,274,295,321]" pageId="22" pageNumber="23">Remarks:</emphasis>
<emphasis id="B918EA8AFFF3D41CB342FEA67EA6697A" box="[282,511,296,321]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
</paragraph>
<paragraph id="8BD33698FFF3D41CB2CFFEE37B056B7C" blockId="22.[151,1438,365,2034]" pageId="22" pageNumber="23">
<bibRefCitation id="EFFD4B69FFF3D41CB2CFFEE37D6869BC" author="Kaygorodova" box="[151,561,365,391]" pageId="22" pageNumber="27" refString="Kaygorodova, I. A. &amp; Liventseva, V. G. (2007) A test of monophyly of the genus Rhynchelmis Hoffmeister, 1843 (Oligochaeta, Lumbriculidae): morphology based phylogeny. Investigations in Russia, 139 e, 1517 - 1527. http: // zhurnal. ape. relarn. ru / articles / 2007 / 139 e. pdf" type="journal article" year="2007">Kaygorodova &amp; Liventseva (2007)</bibRefCitation>
expanded this genus to include ten taxa, including an undescribed species, but some of these species are in need of revision, and at least one is probably invalid (
<bibRefCitation id="EFFD4B69FFF3D41CB6DDFE1C7F8E69E9" author="Martin" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova 2008</bibRefCitation>
). There is little morphological evidence to link these species with
<taxonomicName id="4C6C4D1BFFF3D41CB1B7FE367BDF69EA" box="[1007,1158,440,465]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB1B7FE367BDF69EA" box="[1007,1158,440,465]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
, other than the relative positions of the spermathecae and atria. The same configuration occurs in other accepted genera (
<taxonomicName id="4C6C4D1BFFF3D41CB76DFE537FB96A21" authority="Hrabe" authorityName="Hrabe" class="Clitellata" family="Lumbriculidae" genus="Tatriella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB76DFE537AC269CD" box="[1333,1435,477,502]" italics="true" pageId="22" pageNumber="23">Tatriella</emphasis>
Hrabĕ
</taxonomicName>
and
<taxonomicName id="4C6C4D1BFFF3D41CB378FD8C7DF86A27" authority="Fend &amp; Gustafson" authorityName="Fend &amp; Gustafson" box="[288,673,514,540]" class="Clitellata" family="Lumbriculidae" genus="Secubelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB378FD8C7EF56A20" box="[288,428,514,539]" italics="true" pageId="22" pageNumber="23">Secubelmis</emphasis>
Fend &amp; Gustafson
</taxonomicName>
), and in at least three undescribed Nearctic species that may represent additional lineages (S. Fend, unpublished). Other configurations occur in multiple genera; for example, the spermathecae are in the first preatrial segment in the semiprosoporous genera
<taxonomicName id="4C6C4D1BFFF3D41CB74FFDC37E496AB7" authority="Claparede" authorityName="Claparede" class="Clitellata" family="Lumbriculidae" genus="Stylodrilus" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB74FFDC37AC46A5D" box="[1303,1437,589,614]" italics="true" pageId="22" pageNumber="23">Stylodrilus</emphasis>
Claparède
</taxonomicName>
,
<taxonomicName id="4C6C4D1BFFF3D41CB344FDFC7EB46AB7" authority="Eisen" authorityName="Eisen" box="[284,493,626,652]" class="Clitellata" family="Lumbriculidae" genus="Eclipidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB344FDFC7EFC6AB0" box="[284,421,626,651]" italics="true" pageId="22" pageNumber="23">Eclipidrilus</emphasis>
Eisen
</taxonomicName>
and
<taxonomicName id="4C6C4D1BFFF3D41CB07EFDFC7B196AB7" authority="Eckroth &amp; Brinkhurst. Inclusion" authorityName="Eckroth &amp; Brinkhurst. Inclusion" box="[550,1088,626,652]" class="Clitellata" family="Lumbriculidae" genus="Tenagodrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB07EFDFC7DE76AB0" box="[550,702,626,651]" italics="true" pageId="22" pageNumber="23">Tenagodrilus</emphasis>
Eckroth &amp; Brinkhurst. Inclusion
</taxonomicName>
in
<taxonomicName id="4C6C4D1BFFF3D41CB63EFDFC7BA36AB0" box="[1126,1274,626,651]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB63EFDFC7BA36AB0" box="[1126,1274,626,651]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
, based on the position of the first spermathecae in VIII, would almost certainly result in a polyphyletic genus, and would also require the equally unsupported inclusion of
<taxonomicName id="4C6C4D1BFFF3D41CB0C5FD337DA66AED" box="[669,767,701,726]" class="Clitellata" family="Lumbriculidae" genus="Tatriella" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB0C5FD337DA66AED" box="[669,767,701,726]" italics="true" pageId="22" pageNumber="23">Tatriella</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6C4D1BFFF3D41CB160FD337CE06AED" box="[824,953,701,726]" class="Clitellata" family="Lumbriculidae" genus="Secubelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB160FD337CE06AED" box="[824,953,701,726]" italics="true" pageId="22" pageNumber="23">Secubelmis</emphasis>
</taxonomicName>
. The recent decisions by
<bibRefCitation id="EFFD4B69FFF3D41CB6B9FD337E236AC7" author="Kaygorodova" pageId="22" pageNumber="27" refString="Kaygorodova, I. A. &amp; Liventseva, V. G. (2007) A test of monophyly of the genus Rhynchelmis Hoffmeister, 1843 (Oligochaeta, Lumbriculidae): morphology based phylogeny. Investigations in Russia, 139 e, 1517 - 1527. http: // zhurnal. ape. relarn. ru / articles / 2007 / 139 e. pdf" type="journal article" year="2007">Kaygorodova &amp; Liventseva (2007)</bibRefCitation>
and
<bibRefCitation id="EFFD4B69FFF3D41CB3EFFD6C7C6E6AC7" author="Martin" box="[439,823,738,764]" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
to separate these taxa from
<taxonomicName id="4C6C4D1BFFF3D41CB6C0FD6C7A6C6AC0" box="[1176,1333,738,763]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB6C0FD6C7A6C6AC0" box="[1176,1333,738,763]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
on both morphological and molecular grounds are reasonable from a phylogenetic view, and retention of
<emphasis id="B918EA8AFFF3D41CB2CFFCA37E2F6B7D" box="[151,374,813,838]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
is the best solution for conserving the present taxonomic system.
</paragraph>
<paragraph id="8BD33698FFF3D41CB29CFCDC7B776D3C" blockId="22.[151,1438,365,2034]" pageId="22" pageNumber="23">
While the species assigned to
<emphasis id="B918EA8AFFF3D41CB06EFCDC7C7A6B50" box="[566,803,850,875]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
appear to form a (phenetic) group based on overall similarity, there are few, if any, conclusive synapomorphies distinguishing them from other lumbriculid genera. This problem is evident in the recent numerical analyses by
<bibRefCitation id="EFFD4B69FFF3D41CB19EFC137A356B8C" author="Kaygorodova" box="[966,1388,925,951]" pageId="22" pageNumber="27" refString="Kaygorodova, I. A. &amp; Liventseva, V. G. (2007) A test of monophyly of the genus Rhynchelmis Hoffmeister, 1843 (Oligochaeta, Lumbriculidae): morphology based phylogeny. Investigations in Russia, 139 e, 1517 - 1527. http: // zhurnal. ape. relarn. ru / articles / 2007 / 139 e. pdf" type="journal article" year="2007">Kaygorodova &amp; Liventseva (2007)</bibRefCitation>
: the species do form a cluster in the neighbor-joining tree, but were not resolved as a monophyly in a maximum parsimony tree.
<bibRefCitation id="EFFD4B69FFF3D41CB302FC667D946C39" author="Martin" box="[346,717,1000,1026]" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
listed three possible synapomorphies, two of which are not shared by all members of the group: (1) vasa deferentia much wider entally than ectally (noted by
<bibRefCitation id="EFFD4B69FFF3D41CB70AFB857F8E6C77" author="Hrabe" pageId="22" pageNumber="27" refString="Hrabe, S. (1982) Contribution to the knowledge of Oligochaeta from the lake Baikal. V e stnik C eskoslovenske spole c nosti zoologicke, 46,174 - 193." type="book" year="1982">Hrabĕ 1982</bibRefCitation>
), (2) atrial lining cells thick and glandular, and (3) pendant penes in penis sacs. Based on our limited sampling of the group, additional potential apomorphies for the Baikal group include the distinct pharyngeal pad, the supra-intestinal blood vessel, and the penes formed by extrusion of atrial lining cells, but these characters are not unusual within the family, and have to be verified on more species. Even allowing for reversals within some characters, defining the genus on the basis of these apparent synapomorphies may require the exclusion of the western European
<emphasis id="B918EA8AFFF3D41CB0E1FB637CCE6D3D" box="[697,919,1261,1286]" italics="true" pageId="22" pageNumber="23">
P.
<taxonomicName id="4C6C4D1BFFF3D41CB08DFB637CCE6D3D" box="[725,919,1261,1286]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="paraolchonensis">paraolchonensis</taxonomicName>
</emphasis>
(see below).
</paragraph>
<paragraph id="8BD33698FFF3D41CB29CFA9C7C646E6C" blockId="22.[151,1438,365,2034]" pageId="22" pageNumber="23">
A loop of the posterior vas deferens into the post-atrial segment is well documented (e.g.,
<bibRefCitation id="EFFD4B69FFF3D41CB76FFA9C7EEF6D69" author="Giani" pageId="22" pageNumber="26" refString="Giani, N. &amp; Martinez Ansemil, E. (1984) Deux nouvelles especes de Lumbriculidae du Sud-Ouest de l'Europe. Annales de Limnologie, 20, 157 - 165." type="journal article" year="1984">Giani &amp; Martinez-Ansemil 1984</bibRefCitation>
,
<bibRefCitation id="EFFD4B69FFF3D41CB39BFAB67DC76D69" author="Martin" box="[451,670,1336,1362]" pageId="22" pageNumber="27" refString="Martin, P., Ferraguti, M. &amp; Kaygorodova, I. (1998) Description of two new Rhynchelmis species (Oligochaeta, Lumbriculidae) from Lake Baikal (Russia), using classical morphological and ultrastructural spermatozoal characters. Annales de Limnologie, 34, 283 - 293." type="journal article" year="1998">
Martin
<emphasis id="B918EA8AFFF3D41CB044FAB67D016D6A" box="[540,600,1336,1361]" italics="true" pageId="22" pageNumber="23">et al.</emphasis>
1998
</bibRefCitation>
,
<bibRefCitation id="EFFD4B69FFF3D41CB0F4FAB67C356D69" author="Semernoy" box="[684,876,1336,1362]" pageId="22" pageNumber="27" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. In Timm, T., Mazepova, G. F. and Timoshkin, O. A. (Eds.) Guides and keys to identification of fauna and flora of Lake Baikal. Russian Academy of Sciences, Siberian Division, Nauka, Novosibirsk (Russia), 528 pp." type="book" year="2004">Semernoy 2004</bibRefCitation>
), and distinguishes
<emphasis id="B918EA8AFFF3D41CB60EFAB67A646D6A" box="[1110,1341,1336,1361]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
species from all
<taxonomicName id="4C6C4D1BFFF3D41CB2A5FAD37ECA6D4D" box="[253,403,1373,1398]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB2A5FAD37ECA6D4D" box="[253,403,1373,1398]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
, but as this is assumed to be a plesiomorphic character state (
<bibRefCitation id="EFFD4B69FFF3D41CB634FAD57BA46D4C" author="Hrabe" box="[1132,1277,1371,1399]" pageId="22" pageNumber="27" refString="Hrabe, S. (1984) Two atavistic characters of some Lumbriculidae and their importance for the classification of Oligochaeta. Hydrobiologia, 115, 15 - 17." type="journal article" year="1984">Hrabĕ 1984</bibRefCitation>
), it cannot be used to define
<emphasis id="B918EA8AFFF3D41CB314FA0D7D616DA7" box="[332,568,1411,1436]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
(
<bibRefCitation id="EFFD4B69FFF3D41CB010FA0C7CED6DA7" author="Martin" box="[584,948,1410,1436]" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova 2008</bibRefCitation>
).
<emphasis id="B918EA8AFFF3D41CB192FA0D7BEF6DA7" box="[970,1206,1411,1436]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
atria are generally shorter than those of
<taxonomicName id="4C6C4D1BFFF3D41CB3D4FA267D7B6DFA" box="[396,546,1448,1473]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB3D4FA267D7B6DFA" box="[396,546,1448,1473]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
, usually extending at most into the ovarian segment, and in some cases they are more an elongate pear-shape than tubular. In the few
<taxonomicName id="4C6C4D1BFFF3D41CB16EFA437C956DDD" box="[822,972,1485,1510]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB16EFA437C956DDD" box="[822,972,1485,1510]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
species having atria commonly limited to X and XI, the atria are convoluted, and would extend through more posterior segments if straight (
<bibRefCitation id="EFFD4B69FFF3D41CB776FA7C7FAB6E09" author="Martin" pageId="22" pageNumber="27" refString="Martin, P., Ferraguti, M. &amp; Kaygorodova, I. (1998) Description of two new Rhynchelmis species (Oligochaeta, Lumbriculidae) from Lake Baikal (Russia), using classical morphological and ultrastructural spermatozoal characters. Annales de Limnologie, 34, 283 - 293." type="journal article" year="1998">
Martin
<emphasis id="B918EA8AFFF3D41CB7DFFA7D7FEC6E0A" italics="true" pageId="22" pageNumber="23">et al.</emphasis>
1998
</bibRefCitation>
,
<bibRefCitation id="EFFD4B69FFF3D41CB2A6F9967D7D6E09" author="Fend" box="[254,548,1560,1586]" pageId="22" pageNumber="26" refString="Fend, S. V. &amp; Brinkhurst, R. O. (2000) New species of Rhynchelmis (Clitellata, Lumbriculidae), with observations on the Nearctic species. Hydrobiologia, 428, 1 - 59." type="journal article" year="2000">Fend &amp; Brinkhurst 2000</bibRefCitation>
). Nevertheless,
<emphasis id="B918EA8AFFF3D41CB086F9967C9A6E0A" box="[734,963,1560,1585]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
atria appear to represent a more general state within the family than the long atria of
<taxonomicName id="4C6C4D1BFFF3D41CB0F8F9B37C6F6E6D" box="[672,822,1597,1622]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB0F8F9B37C6F6E6D" box="[672,822,1597,1622]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BD33698FFF3D41CB29CF9EC7BF46F9C" blockId="22.[151,1438,365,2034]" pageId="22" pageNumber="23">
The suggestion by
<bibRefCitation id="EFFD4B69FFF3D41CB3E8F9EC7C776E47" author="Martin" box="[432,814,1634,1660]" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
that “pendant penes” may be a synapomorphy for
<emphasis id="B918EA8AFFF3D41CB2CFF9067E2F6E9A" box="[151,374,1672,1697]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
may be somewhat refined by the present observations. The presence of elongate cells lining the atrial duct at the male pores of
<emphasis id="B918EA8AFFF3D41CB07CF9237DC56EFD" box="[548,668,1709,1734]" italics="true" pageId="22" pageNumber="23">P. alyonae</emphasis>
and
<emphasis id="B918EA8AFFF3D41CB08BF9237CD86EFD" box="[723,897,1709,1734]" italics="true" pageId="22" pageNumber="23">
P.
<taxonomicName id="4C6C4D1BFFF3D41CB0B7F9237CD86EFD" box="[751,897,1709,1734]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="species" species="shamanensis">shamanensis</taxonomicName>
</emphasis>
(protruding in the latter) suggest penes formed by extrusion of atrial lining cells (
<typeStatus id="54D7883AFFF3D41CB045F95C7D096ED7" box="[541,592,1746,1772]" pageId="22" pageNumber="23">type</typeStatus>
2 penes, sensu
<bibRefCitation id="EFFD4B69FFF3D41CB15DF95C7CD36ED7" author="Cook" box="[773,906,1746,1772]" pageId="22" pageNumber="26" refString="Cook, D. G. (1967) Studies on the Lumbriculidae (Oligochaeta) in Britain. Journal of Zoology (London), 153, 353 - 368." type="journal article" year="1967">Cook 1967</bibRefCitation>
).
<figureCitation id="13572A1DFFF3D41CB1C6F95C7CA66ED7" box="[926,1023,1746,1772]" captionStart="FIGURE 2. R" captionStartId="5.[151,255,1776,1800]" captionTargetBox="[155,1434,204,1742]" captionTargetId="figure@5.[151,1436,194,1755]" captionTargetPageId="5" captionText="FIGURE 2. R. tetratheca. (A) curled longitudinal muscles, oblique muscles. (B) ventral glands in an anterior segment. (C) septal sac on 9 / 10. (D) nephridial duct following a lateral blood vessel. (E) accessory organ in IX. (F) detail of accessory organ. (G, H) vas deferens and atrium within sperm sac. (I) vasa deferentia joining atrium entally. (J) ectal end of atrium, male funnel on 10 / 11, and vas deferens. (K) male pore. (L) spermathecal duct entering ampulla (right) and diverticulum (left). (A, B, D, G, K transverse sections, C, E, F, H – J, L sagittal sections). Scale bars: A – E, H – L = 200 μm; F – G = 100 μm." httpUri="https://zenodo.org/record/194254/files/figure.png" pageId="22" pageNumber="23">Figure 2</figureCitation>
B in
<bibRefCitation id="EFFD4B69FFF3D41CB66FF95C7AC56ED7" author="Martin" box="[1079,1436,1746,1772]" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
suggests that similar penes also occur in
<emphasis id="B918EA8AFFF3D41CB034F9767CCE6F2A" box="[620,919,1784,1809]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis parva</emphasis>
(Michaelsen), and illustrations by
<bibRefCitation id="EFFD4B69FFF3D41CB770F9767D5E6F0C" author="Semernoy" pageId="22" pageNumber="27" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. In Timm, T., Mazepova, G. F. and Timoshkin, O. A. (Eds.) Guides and keys to identification of fauna and flora of Lake Baikal. Russian Academy of Sciences, Siberian Division, Nauka, Novosibirsk (Russia), 528 pp." type="book" year="2004">Semernoy (2004, Figs. 225, 227, and 234)</bibRefCitation>
may show the same penial structure in other
<emphasis id="B918EA8AFFF3D41CB643F8937BA76F0D" box="[1051,1278,1821,1846]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
species. This character may be difficult to observe in most preparations, and needs confirmation in well-fixed specimens of other species. Although various
<typeStatus id="54D7883AFFF3D41CB043F8E67D036FB9" box="[539,602,1896,1922]" pageId="22" pageNumber="23">types</typeStatus>
of penes occur in
<taxonomicName id="4C6C4D1BFFF3D41CB161F8E67C886FBA" box="[825,977,1896,1921]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB161F8E67C886FBA" box="[825,977,1896,1921]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
(e.g.,
<bibRefCitation id="EFFD4B69FFF3D41CB646F8E67A1C6FB9" author="Fend" box="[1054,1349,1896,1922]" pageId="22" pageNumber="26" refString="Fend, S. V. &amp; Brinkhurst, R. O. (2000) New species of Rhynchelmis (Clitellata, Lumbriculidae), with observations on the Nearctic species. Hydrobiologia, 428, 1 - 59." type="journal article" year="2000">Fend &amp; Brinkhurst 2000</bibRefCitation>
for the Nearctic species), their basic structure differs from that of these
<emphasis id="B918EA8AFFF3D41CB136F8037B146F9D" box="[878,1101,1933,1958]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
species.
</paragraph>
<paragraph id="8BD33698FFF3D41DB29CF83C7D326919" blockId="22.[151,1438,365,2034]" lastBlockId="23.[151,1438,152,1783]" lastPageId="23" lastPageNumber="24" pageId="22" pageNumber="23">
The atrial histology of at least some
<emphasis id="B918EA8AFFF3D41CB037F83D7C0D6FF7" box="[623,852,1971,1996]" italics="true" pageId="22" pageNumber="23">Pseudorhynchelmis</emphasis>
differs from that of
<taxonomicName id="4C6C4D1BFFF3D41CB619F83D7B8E6FF7" box="[1089,1239,1971,1996]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="22" pageNumber="23" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF3D41CB619F83D7B8E6FF7" box="[1089,1239,1971,1996]" italics="true" pageId="22" pageNumber="23">Rhynchelmis</emphasis>
</taxonomicName>
. The lining cells of the observed species, plus those observed by
<bibRefCitation id="EFFD4B69FFF3D41CB08DF8567B106FC9" author="Martin" box="[725,1097,2008,2034]" pageId="22" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
, are thick and glandular, but based on their descriptions,
<emphasis id="B918EA8AFFF2D41DB3BCFF167DE6688A" box="[484,703,152,177]" italics="true" pageId="23" pageNumber="24">P. spermatochaeta</emphasis>
and
<emphasis id="B918EA8AFFF2D41DB0A1FF167CD7688A" box="[761,910,152,177]" italics="true" pageId="23" pageNumber="24">P. minimaris</emphasis>
may be exceptions (
<bibRefCitation id="EFFD4B69FFF2D41DB6DCFF167A1A6889" author="Semernoy" box="[1156,1347,152,178]" pageId="23" pageNumber="27" refString="Semernoy, V. P. (1982) Novye vidy oligokhet iz ozera Baikal [New oligochaete species from Lake Baikal]. In: Galazii, G. I. (ed.), Novoe o faune Baikala, Izdatel'stvo &quot; Nauka &quot;, Sibirskoe otdelenie, Novosibirsk, USSR, pp. 58 - 85. [In Russian; English version Canadian Translation of Fisheries and Aquatic Sciences No 5023]" type="journal article" year="1982">Semernoy 1982</bibRefCitation>
,
<bibRefCitation id="EFFD4B69FFF2D41DB708FF167AC96889" author="Semernoy" box="[1360,1424,152,178]" pageId="23" pageNumber="27" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. In Timm, T., Mazepova, G. F. and Timoshkin, O. A. (Eds.) Guides and keys to identification of fauna and flora of Lake Baikal. Russian Academy of Sciences, Siberian Division, Nauka, Novosibirsk (Russia), 528 pp." type="book" year="2004">2004</bibRefCitation>
). The prostate bundles in the material observed here are relatively larger, and more irregular in size, shape, and placement than those of
<taxonomicName id="4C6C4D1BFFF2D41DB3E0FF6C7D1468C0" box="[440,589,226,251]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB3E0FF6C7D1468C0" box="[440,589,226,251]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
, in accordance with the observations by
<bibRefCitation id="EFFD4B69FFF2D41DB677FF6C7AC268C7" author="Martin" box="[1071,1435,226,252]" pageId="23" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
and
<bibRefCitation id="EFFD4B69FFF2D41DB290FE867D3E6919" author="Kaygorodova" box="[200,615,264,290]" pageId="23" pageNumber="27" refString="Kaygorodova, I. A. &amp; Liventseva, V. G. (2007) A test of monophyly of the genus Rhynchelmis Hoffmeister, 1843 (Oligochaeta, Lumbriculidae): morphology based phylogeny. Investigations in Russia, 139 e, 1517 - 1527. http: // zhurnal. ape. relarn. ru / articles / 2007 / 139 e. pdf" type="journal article" year="2007">Kaygorodova &amp; Liventseva (2007)</bibRefCitation>
.
</paragraph>
<paragraph id="8BD33698FFF2D41DB29CFEA37EBB6B17" blockId="23.[151,1438,152,1783]" pageId="23" pageNumber="24">
Spermathecae of
<emphasis id="B918EA8AFFF2D41DB3D7FEA37D2D697D" box="[399,628,301,326]" italics="true" pageId="23" pageNumber="24">Pseudorhynchelmis</emphasis>
species do not resemble those of any of the
<taxonomicName id="4C6C4D1BFFF2D41DB621FEA37A56697D" box="[1145,1295,301,326]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB621FEA37A56697D" box="[1145,1295,301,326]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
groups (see also
<bibRefCitation id="EFFD4B69FFF2D41DB28EFEDC7DD56957" author="Giani" box="[214,652,338,364]" pageId="23" pageNumber="26" refString="Giani, N. &amp; Martinez Ansemil, E. (1984) Deux nouvelles especes de Lumbriculidae du Sud-Ouest de l'Europe. Annales de Limnologie, 20, 157 - 165." type="journal article" year="1984">Giani &amp; Martinez-Ansemil 1984</bibRefCitation>
,
<bibRefCitation id="EFFD4B69FFF2D41DB0C3FEDC7C376957" author="Semernoy" box="[667,878,338,364]" pageId="23" pageNumber="27" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. In Timm, T., Mazepova, G. F. and Timoshkin, O. A. (Eds.) Guides and keys to identification of fauna and flora of Lake Baikal. Russian Academy of Sciences, Siberian Division, Nauka, Novosibirsk (Russia), 528 pp." type="book" year="2004">Semernoy 2004</bibRefCitation>
,
<bibRefCitation id="EFFD4B69FFF2D41DB125FEDC7A5F6957" author="Martin" box="[893,1286,338,364]" pageId="23" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova 2008</bibRefCitation>
). Unlike
<emphasis id="B918EA8AFFF2D41DB7D9FEDC7AC46950" box="[1409,1437,338,363]" italics="true" pageId="23" pageNumber="24">R.</emphasis>
(
<taxonomicName id="4C6C4D1BFFF2D41DB2F9FEF67E6169AA" box="[161,312,376,401]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB2F9FEF67E6169AA" box="[161,312,376,401]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
), the
<taxonomicName id="4C6C4D1BFFF2D41DB326FEF67E9569AA" box="[382,460,376,401]" class="Clitellata" family="Lumbriculidae" genus="Sutroa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB326FEF67E9569AA" box="[382,460,376,401]" italics="true" pageId="23" pageNumber="24">Sutroa</emphasis>
</taxonomicName>
group, some of the
<emphasis id="B918EA8AFFF2D41DB0E4FEF67C2769AA" box="[700,894,376,401]" italics="true" pageId="23" pageNumber="24">Rhynchelmoides</emphasis>
group, and
<taxonomicName id="4C6C4D1BFFF2D41DB655FEF67BDE69AA" box="[1037,1159,376,401]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="brooksi">
<emphasis id="B918EA8AFFF2D41DB655FEF67BDE69AA" box="[1037,1159,376,401]" italics="true" pageId="23" pageNumber="24">R. brooksi</emphasis>
</taxonomicName>
/
<taxonomicName id="4C6C4D1BFFF2D41DB6D7FEF67BAA69AA" box="[1167,1267,376,401]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="malevici">
<emphasis id="B918EA8AFFF2D41DB6D7FEF67BAA69AA" box="[1167,1267,376,401]" italics="true" pageId="23" pageNumber="24">malevici</emphasis>
</taxonomicName>
, the ampullae are rather simple sacs, without diverticula or obvious ducts to the gut. In contrast to those
<emphasis id="B918EA8AFFF2D41DB681FE137AC4698D" box="[1241,1437,413,438]" italics="true" pageId="23" pageNumber="24">Rhynchelmoides</emphasis>
group species lacking a spermatheca-gut connection, the ampullae are not divided into histologically distinctive sections, with viable sperm concentrated in the ectal region. The ectal ducts do not resemble those of any of the Nearctic
<taxonomicName id="4C6C4D1BFFF2D41DB3FCFD837D626A1D" box="[420,571,525,550]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB3FCFD837D626A1D" box="[420,571,525,550]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
; and are typically more narrowly tubular than in
<emphasis id="B918EA8AFFF2D41DB6D3FD837BFD6A1D" box="[1163,1188,525,550]" italics="true" pageId="23" pageNumber="24">R.</emphasis>
(
<taxonomicName id="4C6C4D1BFFF2D41DB6EEFD837A146A1D" box="[1206,1357,525,550]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB6EEFD837A146A1D" box="[1206,1357,525,550]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
) (e.g.,
<figureCitation id="13572A1DFFF2D41DB2CFFDBC7FBB6A77" box="[151,226,562,588]" captionStart="FIGURE 9. P" captionStartId="20.[151,256,1776,1800]" captionTargetBox="[153,1434,231,1743]" captionTargetId="figure@20.[151,1436,197,1755]" captionTargetPageId="20" captionText="FIGURE 9. P. shamanensis and P. alyonae. (A – B, I) P. shamanensis from Martin collection, (C – H, J – L) from Hrabė collection as olchonensis. (A) anterior end showing pharyngeal pad. (B – C) mid-body, showing dorsal and supraintestinal blood vessels. (D) transverse section of pharynx (E) transverse section of a post-clitellar segment showing dorsal and supra-intestinal blood vessels. (F) transverse section of male pores. (G – H) sections of penes. (I) external view of extruded penis. (J) sagittal section showing wide ental portion and narrow ectal portion of vas deferens. (K) atrium joined by narrow portion of a vas deferens. (L) detail of atrial wall. (M) P. alyonae spermatheca. (A, B, I, M whole mounts, C, H, K, L sagittal sections, D – G, J transverse). Scale bars: A – C, E, F = 200 μm; D, K, M = 100 μm; G – J, L = 50 μm." httpUri="https://zenodo.org/record/194261/files/figure.png" pageId="23" pageNumber="24">Fig. 9</figureCitation>
M vs.
<date id="FFD21058FFF2D41DB36EFDBC7E0D6A77" box="[310,340,562,588]" pageId="23" pageNumber="24">3I</date>
). The spermathecal ampulla joins the gut via a short, but distinct ental duct in those
<taxonomicName id="4C6C4D1BFFF2D41DB2CFFDD67E686A4A" box="[151,305,600,625]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB2CFFDD67E686A4A" box="[151,305,600,625]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
species having a spermatheca-gut connection (e.g.,
<figureCitation id="13572A1DFFF2D41DB1F4FDD67CAD6A49" box="[940,1012,600,626]" captionStart="FIGURE 3. R" captionStartId="7.[151,257,1869,1893]" captionTargetBox="[218,1352,281,1826]" captionTargetId="figure@7.[218,1352,271,1843]" captionTargetPageId="7" captionText="FIGURE 3. R. limosella from Hrabė collection. (A) segments IX – XII, showing atrium, male funnels, and accessory organ. (B) male funnels and vasa deferentia at ectal end of atrium. (C) atrium and vas deferens. (D) ental end of atrium, with vasa deferentia embedded under the muscle layer. (E – F) male pore. (G) atrium and vas deferens. (H) accessory organ. (I – J) spermatheca and diverticulum. (K) spermathecal ampullae at gut. (A – B, F – J sagittal sections, C – E, K transverse sections). Scale bars: C, D, F – H = 100 μm; A, B, E, I – K = 200 μm." httpUri="https://zenodo.org/record/194255/files/figure.png" pageId="23" pageNumber="24">Fig. 3</figureCitation>
K). In contrast, Semernoy’s (2004) illustrations of several Baikal
<emphasis id="B918EA8AFFF2D41DB074FDF37C756AAD" box="[556,812,637,662]" italics="true" pageId="23" pageNumber="24">Pseudorhynchelmis</emphasis>
species (not available for study here) show an inconspicuous median junction, which may need to be verified with sectioned material.
<bibRefCitation id="EFFD4B69FFF2D41DB77BFD2C7EC36AD9" author="Martin" pageId="23" pageNumber="27" refString="Martin, P. &amp; Kaygorodova, I. (2008) A new species of Pseudorhynchelmis Hrabe, 1982 (Clitellata: Lumbriculidae) from Lake Baikal, with re-descriptions of P. parva and P. olchonensis. Zootaxa, 1938, 23 - 39." type="journal article" year="2008">Martin &amp; Kaygorodova (2008)</bibRefCitation>
questioned the validity of the spermatheca-gut connection in these taxa; although
<bibRefCitation id="EFFD4B69FFF2D41DB2CFFD637D1A6B3C" author="Kaygorodova" box="[151,579,749,775]" pageId="23" pageNumber="27" refString="Kaygorodova, I. A. &amp; Liventseva, V. G. (2007) A test of monophyly of the genus Rhynchelmis Hoffmeister, 1843 (Oligochaeta, Lumbriculidae): morphology based phylogeny. Investigations in Russia, 139 e, 1517 - 1527. http: // zhurnal. ape. relarn. ru / articles / 2007 / 139 e. pdf" type="journal article" year="2007">Kaygorodova &amp; Liventseva (2007)</bibRefCitation>
assumed that it was correct for purposes of analysis, they defined the genus in part by its absence.
</paragraph>
<paragraph id="8BD33698FFF2D41DB29CFCB67D0C6C37" blockId="23.[151,1438,152,1783]" pageId="23" pageNumber="24">
The structure of the pharynx is usually not described in detail in lumbriculids. The two observed Baikal
<emphasis id="B918EA8AFFF2D41DB2CFFCD37E256B4D" box="[151,380,861,886]" italics="true" pageId="23" pageNumber="24">Pseudorhynchelmis</emphasis>
(but not
<taxonomicName id="4C6C4D1BFFF2D41DB3BCFCD37DFF6B4D" box="[484,678,861,886]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="paraolchonensis">
<emphasis id="B918EA8AFFF2D41DB3BCFCD37DFF6B4D" box="[484,678,861,886]" italics="true" pageId="23" pageNumber="24">paraolchonensis</emphasis>
</taxonomicName>
) had a strongly developed dorsal pad in II–III or IV, whereas this did not occur in any observed
<emphasis id="B918EA8AFFF2D41DB055FC0C7D7E6BA0" box="[525,551,898,923]" italics="true" pageId="23" pageNumber="24">R.</emphasis>
(
<taxonomicName id="4C6C4D1BFFF2D41DB062FC0C7D8C6BA0" box="[570,725,898,923]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB062FC0C7D8C6BA0" box="[570,725,898,923]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
), or in
<taxonomicName id="4C6C4D1BFFF2D41DB16AFC0C7C936BA0" box="[818,970,898,923]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="orientalis">
<emphasis id="B918EA8AFFF2D41DB16AFC0C7C936BA0" box="[818,970,898,923]" italics="true" pageId="23" pageNumber="24">R. orientalis</emphasis>
</taxonomicName>
. Semernoy (1984, Figs. 233 and 234) illustrates pharyngeal pads in two other
<emphasis id="B918EA8AFFF2D41DB037FC267C0D6BFA" box="[623,852,936,961]" italics="true" pageId="23" pageNumber="24">Pseudorhynchelmis</emphasis>
species. The dorsal wall of the pharynx was only slightly developed, and both dorsal and ventral walls were eversible in other
<emphasis id="B918EA8AFFF2D41DB611FC437A496BDD" box="[1097,1296,973,998]" italics="true" pageId="23" pageNumber="24">Rhynchelmoides</emphasis>
and
<taxonomicName id="4C6C4D1BFFF2D41DB714FC437AC56BDD" box="[1356,1436,973,998]" class="Clitellata" family="Lumbriculidae" genus="Sutroa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB714FC437AC56BDD" box="[1356,1436,973,998]" italics="true" pageId="23" pageNumber="24">Sutroa</emphasis>
</taxonomicName>
group taxa (
<bibRefCitation id="EFFD4B69FFF2D41DB37AFC7C7D106C37" author="Fend" box="[290,585,1010,1036]" pageId="23" pageNumber="26" refString="Fend, S. V. &amp; Brinkhurst, R. O. (2000) New species of Rhynchelmis (Clitellata, Lumbriculidae), with observations on the Nearctic species. Hydrobiologia, 428, 1 - 59." type="journal article" year="2000">Fend &amp; Brinkhurst 2000</bibRefCitation>
).
</paragraph>
<paragraph id="8BD33698FFF2D41DB29CFB967D2D6D29" blockId="23.[151,1438,152,1783]" pageId="23" pageNumber="24">
No mention of the blood vessels was made in the original descriptions of
<emphasis id="B918EA8AFFF2D41DB674FB967BFE6C0A" box="[1068,1191,1048,1073]" italics="true" pageId="23" pageNumber="24">P. alyonae</emphasis>
and
<emphasis id="B918EA8AFFF2D41DB6B9FB967ACC6C0A" box="[1249,1429,1048,1073]" italics="true" pageId="23" pageNumber="24">
P.
<taxonomicName id="4C6C4D1BFFF2D41DB6A7FB967ACC6C0A" box="[1279,1429,1048,1073]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="shamanensis">shamanensis</taxonomicName>
</emphasis>
, and the vascular system was not completely visible in the available material. However, our observations suggest that these species differ from most
<taxonomicName id="4C6C4D1BFFF2D41DB0D7FBED7C7D6C47" box="[655,804,1123,1148]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB0D7FBED7C7D6C47" box="[655,804,1123,1148]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
species in the complete lack of posterior lateral blood vessels, the anterior commissures with a strong anterior loop, and the supra-intestinal vessel formed from the perivisceral sinus. Nephridia were difficult to see clearly, as no sagittally sectioned specimens were available. Although taxonomically important, nephridia and blood vessels are difficult to see in most preparations, and are not described for many lumbriculids.
</paragraph>
<paragraph id="8BD33698FFF2D41DB29CFA937EC96ECC" blockId="23.[151,1438,152,1783]" pageId="23" pageNumber="24">
The lack of a proboscis, the lack of a spermathecal-gut connection, and the testes with functional vasa deferentia in XI support the removal of
<emphasis id="B918EA8AFFF2D41DB0CDFACD7CDC6D67" box="[661,901,1347,1372]" italics="true" pageId="23" pageNumber="24">
P.
<taxonomicName id="4C6C4D1BFFF2D41DB0EDFACD7CDC6D67" box="[693,901,1347,1372]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="paraolchonensis">paraolchonensis</taxonomicName>
</emphasis>
from
<taxonomicName id="4C6C4D1BFFF2D41DB18CFACD7B2D6D67" box="[980,1140,1347,1372]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="B918EA8AFFF2D41DB18CFACD7B2D6D67" box="[980,1140,1347,1372]" italics="true" pageId="23" pageNumber="24">Rhynchelmis</emphasis>
</taxonomicName>
, but as these are likely plesiomorphic states, they only indirectly support its placement in
<emphasis id="B918EA8AFFF2D41DB1FCFAE67BD16DBA" box="[932,1160,1384,1409]" italics="true" pageId="23" pageNumber="24">Pseudorhynchelmis</emphasis>
. The blood vessels, the somewhat median placement of genital pores, the posterior vas deferens entering XI, the structure of the spermathecae, and the histology of the atrium also suggest affinity with
<emphasis id="B918EA8AFFF2D41DB194FA3D7B376DF7" box="[972,1134,1459,1484]" italics="true" pageId="23" pageNumber="24">
P.
<taxonomicName id="4C6C4D1BFFF2D41DB1B0FA3D7B376DF7" box="[1000,1134,1459,1484]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="olchonensis">olchonensis</taxonomicName>
</emphasis>
and similar species.
<bibRefCitation id="EFFD4B69FFF2D41DB704FA3C7EA86DC9" author="Giani" pageId="23" pageNumber="26" refString="Giani, N. &amp; Martinez Ansemil, E. (1984) Deux nouvelles especes de Lumbriculidae du Sud-Ouest de l'Europe. Annales de Limnologie, 20, 157 - 165." type="journal article" year="1984">Giani &amp; Martinez-Ansemil (1984)</bibRefCitation>
differentiated
<emphasis id="B918EA8AFFF2D41DB0FFFA567CC96DCA" box="[679,912,1496,1521]" italics="true" pageId="23" pageNumber="24">
P.
<taxonomicName id="4C6C4D1BFFF2D41DB09EFA567CC96DCA" box="[710,912,1496,1521]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="paraolchonensis">paraolchonensis</taxonomicName>
</emphasis>
from
<emphasis id="B918EA8AFFF2D41DB184FA567BD56DCA" box="[988,1164,1496,1521]" italics="true" pageId="23" pageNumber="24">
P.
<taxonomicName id="4C6C4D1BFFF2D41DB1A4FA567BD56DCA" box="[1020,1164,1496,1521]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="olchonensis">olchonensis</taxonomicName>
</emphasis>
by the position of the clitellum, number of segments, and the posterior vas deferens length not greater than that of the anterior.
<bibRefCitation id="EFFD4B69FFF2D41DB76BFA737E516E07" pageId="23" pageNumber="27" refString="Martin, P., Ferraguti, M. &amp; Kaygorodova, I. (1998) Description of two new Rhynchelmis species (Oligochaeta, Lumbriculidae) from Lake Baikal (Russia), using classical morphological and ultrastructural spermatozoal characters. Annales de Limnologie, 34, 283 - 293." type="journal article">
Martin
<emphasis id="B918EA8AFFF2D41DB7DFFA737FEC6E07" italics="true" pageId="23" pageNumber="24">et al.</emphasis>
(1998)
</bibRefCitation>
noted the distinctive chaetae, which were illustrated in the original description as being proportionally much thicker than in other members of the genus, and also noted the short atria and greater overall size.
<emphasis id="B918EA8AFFF2D41DB2CFF9E37D016EBD" box="[151,600,1645,1670]" italics="true" pageId="23" pageNumber="24">
Pseudorhynchelmis
<taxonomicName id="4C6C4D1BFFF2D41DB3D6F9E37D016EBD" box="[398,600,1645,1670]" class="Clitellata" family="Lumbriculidae" genus="Rhynchelmis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lumbriculida" pageId="23" pageNumber="24" phylum="Annelida" rank="species" species="paraolchonensis">paraolchonensis</taxonomicName>
</emphasis>
differs from at least some of the Baikal species in lacking potential synapomorphies for
<emphasis id="B918EA8AFFF2D41DB3DBF91D7D3A6E97" box="[387,611,1683,1708]" italics="true" pageId="23" pageNumber="24">Pseudorhynchelmis</emphasis>
: the ectal end of the atrium forms a slight papilla, but the lining cells do not appear modified; the vasa deferentia are not strongly tapered; and the pharynx does not have a strongly developed dorsal pad.
</paragraph>
</subSubSection>
</treatment>
</document>
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