284 lines
54 KiB
XML
284 lines
54 KiB
XML
<document id="DBCED26B6AD9C3598CCEF197AB54CCAF" ID-CLB-Dataset="29233" ID-DOI="10.11646/zootaxa.4497.1.3" ID-GBIF-Dataset="0f383140-efb8-454c-9188-a8fe795437bc" ID-ISSN="1175-5326" ID-Zenodo-Dep="1451092" ID-ZooBank="11725C60-E463-4EB3-A96A-34CEF56923B8" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1538973537549" checkinUser="plazi" docAuthor="Rodriguez, Pilar & Fend, Steven V." docDate="2018" docId="5D62156BFFD8C8421AECD1F3FAAA6812" docLanguage="en" docName="zootaxa.4497.1.3.pdf" docOrigin="Zootaxa 4497 (1)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Paranadrilus descolei Gavrilov 1955" docType="treatment" docVersion="7" lastPageNumber="50" masterDocId="A15B6D13FFDFC84B1A7BD164FF926976" masterDocTitle="On spermatophore-producing aquatic microdrile oligochaetes (Annelida: Clitellata)" masterLastPageNumber="60" masterPageNumber="41" pageNumber="48" updateTime="1698568037511" updateUser="ExternalLinkService">
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<mods:title id="3F1C20715D671F6A1D77E7D01E35DD45">On spermatophore-producing aquatic microdrile oligochaetes (Annelida: Clitellata)</mods:title>
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<mods:namePart id="3AAA56D4236ABAB8B2394BA0C7EC99AE">Rodriguez, Pilar</mods:namePart>
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<mods:namePart id="A53A609F38BDA11431C8390440A7D3A0">Fend, Steven V.</mods:namePart>
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<mods:title id="3917DCA639D4DD3CBE0471E96899399F">Zootaxa</mods:title>
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<mods:date id="FD9EF456A0EEEBE31D33318DCD2D3790">2018</mods:date>
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<treatment id="5D62156BFFD8C8421AECD1F3FAAA6812" ID-DOI="http://doi.org/10.5281/zenodo.5951344" ID-GBIF-Taxon="148651457" ID-Zenodo-Dep="5951344" LSID="urn:lsid:plazi:treatment:5D62156BFFD8C8421AECD1F3FAAA6812" httpUri="http://treatment.plazi.org/id/5D62156BFFD8C8421AECD1F3FAAA6812" lastPageId="9" lastPageNumber="50" pageId="7" pageNumber="48">
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<subSubSection id="9DD1F7F6FFD8C84C1AECD1F3FE9569A5" pageId="7" pageNumber="48" type="nomenclature">
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<paragraph id="D574A47DFFD8C84C1AECD1F3FE0C69C7" blockId="7.[151,414,151,211]" box="[151,414,151,177]" pageId="7" pageNumber="48">
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<heading id="8E3C1311FFD8C84C1AECD1F3FE0C69C7" bold="true" box="[151,414,151,177]" fontSize="11" level="1" pageId="7" pageNumber="48" reason="1">
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<taxonomicName id="12CBDFFEFFD8C84C1AECD1F3FE0C69C7" authorityName="Gavrilov" authorityYear="1955" box="[151,414,151,177]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="48" phylum="Annelida" rank="species" species="descolei">
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<emphasis id="E7BF786FFFD8C84C1AECD1F3FE0C69C7" bold="true" box="[151,414,151,177]" italics="true" pageId="7" pageNumber="48">Paranadrilus descolei</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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<paragraph id="D574A47DFFD8C84C1AECD1DEFE9569A5" blockId="7.[151,414,151,211]" box="[151,263,186,211]" pageId="7" pageNumber="48">
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(
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<figureCitation id="4DF0B8F8FFD8C84C1AE5D1DEFF6C69A5" box="[158,254,186,211]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Figure 3</figureCitation>
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)
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</paragraph>
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</subSubSection>
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<subSubSection id="9DD1F7F6FFD8C8421AECD199FAAA6812" lastPageId="9" lastPageNumber="50" pageId="7" pageNumber="48" type="description">
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<paragraph id="D574A47DFFD8C84C1AECD199FEFE684F" blockId="7.[151,1437,253,2029]" pageId="7" pageNumber="48">
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<emphasis id="E7BF786FFFD8C84C1AECD199FE866860" bold="true" box="[151,276,253,278]" pageId="7" pageNumber="48">Lectotype.</emphasis>
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CH-O-FML 2349, ParVial No.13 (Series
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<date id="A17582BDFFD8C84C197CD19AFCC76860" box="[775,853,254,278]" bridgedPair="-" pageId="7" pageNumber="48" value="2349-01">2349-I</date>
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), longitudinal histological sections (10 µm), on 2 slides. (
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<figureCitation id="4DF0B8F8FFD8C84C1A96D045FECD684F" box="[237,351,289,314]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3K,L</figureCitation>
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).
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</paragraph>
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<paragraph id="D574A47DFFD8C84C1ABCD022FB5C6B2C" blockId="7.[151,1437,253,2029]" pageId="7" pageNumber="48">
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A review of Gavrilov's
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<typeStatus id="0A701ADFFFD8C84C1B91D022FD886828" box="[490,538,326,350]" pageId="7" pageNumber="48">type</typeStatus>
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collection in Miguel Lillo Museum of Natural Sciences (MMLCN) by P. Rodriguez confirmed
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<bibRefCitation id="B15AD98CFFD8C84C1BEFD00DFDCE68F4" box="[404,604,361,386]" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">Gavrilov's (1958)</bibRefCitation>
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description of the reproductive organs in his very detailed monograph. Gavrilov's
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<typeStatus id="0A701ADFFFD8C84C1B6AD0EAFED368D0" box="[273,321,398,422]" pageId="7" pageNumber="48">type</typeStatus>
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collection is large, composed of almost 1000 individuals from a laboratory culture, initiated with 17 field worms (4 sexually mature and 24 immature) sampled at the
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<typeStatus id="0A701ADFFFD8C84C19F0D0D6FC2968BC" box="[907,955,434,458]" pageId="7" pageNumber="48">type</typeStatus>
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locality in Las Tunas stream (
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<collectingRegion id="170F6A9FFFD8C84C1F61D0D5FA0A68BF" box="[1306,1432,433,457]" country="Argentina" name="Entre Rios" pageId="7" pageNumber="48">Entre Ríos</collectingRegion>
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,
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<collectingCountry id="ADDCE4EDFFD8C84C1AECD0B1FE9B6898" box="[151,265,469,494]" name="Argentina" pageId="7" pageNumber="48">Argentina</collectingCountry>
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) (
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<bibRefCitation id="B15AD98CFFD8C84C1B59D0B1FE486898" box="[290,474,469,494]" pageId="7" pageNumber="48" refId="ref10986" refString="Gavrilov, K. (1955 a) Eine neuer spermathekenloser Vertreter der Tubificiden. Zoological Anzeiger, 155, 294 - 302." type="journal article">Gavrilov 1955a</bibRefCitation>
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,
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<bibRefCitation id="B15AD98CFFD8C84C1B92D0B1FDB76898" box="[489,549,469,494]" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">1958</bibRefCitation>
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). The
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<typeStatus id="0A701ADFFFD8C84C1808D0B2FD316898" box="[627,675,470,494]" pageId="7" pageNumber="48">type</typeStatus>
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collection is at the MMLCN with the reference FML 2349, and consists of
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<typeStatus id="0A701ADFFFD8C84C1B5ED09EFE156B64" box="[293,391,506,530]" pageId="7" pageNumber="48" type="syntype">syntypes</typeStatus>
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(=Kotypen,
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<bibRefCitation id="B15AD98CFFD8C84C185DD09DFD736B64" box="[550,737,505,530]" pageId="7" pageNumber="48" refId="ref10986" refString="Gavrilov, K. (1955 a) Eine neuer spermathekenloser Vertreter der Tubificiden. Zoological Anzeiger, 155, 294 - 302." type="journal article">Gavrilov 1955a</bibRefCitation>
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), thus no specimen was designated as the
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<typeStatus id="0A701ADFFFD8C84C1E82D09DFACD6B64" box="[1273,1375,505,530]" pageId="7" pageNumber="48" type="holotype">holotype</typeStatus>
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. The histological sections most probably used by the author for his drawings and the description of the reproductive system were identified, and among them, we have selected one specimen to be named
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<typeStatus id="0A701ADFFFD8C84C1E1BD325FB586B2C" box="[1120,1226,577,602]" pageId="7" pageNumber="48" type="lectotype">lectotype</typeStatus>
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.
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</paragraph>
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<paragraph id="D574A47DFFD8C84C1ABCD301FEFF6D24" blockId="7.[151,1437,253,2029]" pageId="7" pageNumber="48">
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Spermathecae were absent in all mature specimens examined of
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<taxonomicName id="12CBDFFEFFD8C84C19DBD301FB0B6B08" authorityName="Gavrilov" authorityYear="1955" box="[928,1177,613,638]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="48" phylum="Annelida" rank="species" species="descolei">
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<emphasis id="E7BF786FFFD8C84C19DBD301FB0B6B08" box="[928,1177,613,638]" italics="true" pageId="7" pageNumber="48">Paranadrilus descolei</emphasis>
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</taxonomicName>
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. Spermatophores were not observed in the review of the
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<typeStatus id="0A701ADFFFD8C84C1861D3EEFDD86BD4" box="[538,586,650,674]" pageId="7" pageNumber="48">type</typeStatus>
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collection, although they were described in a few specimens in Gavrilov's monograph (
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<bibRefCitation id="B15AD98CFFD8C84C1B53D3C9FDC96BB0" box="[296,603,685,710]" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">
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Gavrilov 1958:
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<figureCitation id="4DF0B8F8FFD8C84C1B9AD3C9FDC96BB0" box="[481,603,685,710]" captionStart="FIGURE 5" captionStartId="10.[151,250,1714,1736]" captionTargetBox="[266,1323,994,1681]" captionTargetId="figure@10.[254,1332,983,1691]" captionTargetPageId="10" captionText="FIGURE 5. Uktena riparia Fend, Rodriguez & Lenat, 2015. A. Ventral side showing 2 stained spermatophores attached to the base of a protruded copulatory organ. B. Detail of the spermatophore protruding from male pore, and genital chaetae. C. Copulatory organ and associated gland within the spermathecal bursa, with spermatophores attached to its base. D. Male bursa with showing one of the several glandular organs. E. Detail of the atrium showing folded epithelium and long prostatic glands. F. Spermatophore forming in the atrial duct. G. Sperm lined up along the epithelium of the spermathecal ampulla. A,C,E from Fend's coll.; D,F,G from MNCN collection; all from the type locality. Abbreviations: a atrium, co copulatory organ, gc genital chaetae, gl gland, mb male bursa, mp male pore, pr prostatic cells, sb spermathecal bursa, sph spermatophore, ssg secretory surface of the gland." httpUri="https://zenodo.org/record/1451104/files/figure.png" pageId="7" pageNumber="48">Fig. 5A–F</figureCitation>
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</bibRefCitation>
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), where spermatophores were attached individually to the body wall. In contrast, all mature specimens in Marchese's field collection had specific attachment sites for the spermatophores, within small lateral pouches not described by
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<bibRefCitation id="B15AD98CFFD8C84C18DDD391FCF16A78" box="[678,867,757,782]" pageId="7" pageNumber="48" refId="ref10986" refString="Gavrilov, K. (1955 a) Eine neuer spermathekenloser Vertreter der Tubificiden. Zoological Anzeiger, 155, 294 - 302." type="journal article">Gavrilov (1955a</bibRefCitation>
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,
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<bibRefCitation id="B15AD98CFFD8C84C1913D391FCE56A78" box="[872,887,757,782]" pageId="7" pageNumber="48" refId="ref11010" refString="Gavrilov, K. (1955 b) Uber die uniparentale Vermehrung von Paranadrilus. Zoological Anzeiger, 155, 302 - 306." type="journal article">b</bibRefCitation>
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,
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<bibRefCitation id="B15AD98CFFD8C84C19FFD391FC526A78" box="[900,960,757,782]" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">1958</bibRefCitation>
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). Lateral pouches were never observed in the
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<typeStatus id="0A701ADFFFD8C84C1ABBD27EFF626A44" box="[192,240,794,818]" pageId="7" pageNumber="48">type</typeStatus>
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collection. In Marchese's collection, these were latero-ventral invaginations of the body wall, containing 1 to 3 spermatophores; the narrow end of each spermatophore was apparently attached with some substance to the outer part of the pouch (
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<figureCitation id="4DF0B8F8FFD8C84C1BD0D205FD8D6A0C" box="[427,543,865,890]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3C,D</figureCitation>
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). These spermatophore pouches are formed in the anterior part of segment XII (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1AE4D2E1FF6A6AE8" box="[159,248,901,926]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3G</figureCitation>
|
||
), with the entrance directed upwards, as described by
|
||
<bibRefCitation id="B15AD98CFFD8C84C192CD2E1FA976AE8" box="[855,1285,901,926]" pageId="7" pageNumber="48" refId="ref9764" refString="Brinkhurst, R. O. & Marchese, M. (1987) A contribution to the taxonomy of the aquatic Oligochaeta (Haplotaxidae, Phreodrilidae, Tubificidae) of South America. Canadian Journal of Zoology, 65, 3154 - 3165. https: // doi. org / 10.1139 / z 87 - 474" type="journal article">Brinkhurst & Marchese (1987: Fig. 7)</bibRefCitation>
|
||
. The walls of the spermatophore pouch are surrounded by musculature (
|
||
<figureCitation id="4DF0B8F8FFD8C84C195BD2CDFCE46AB4" box="[800,886,937,962]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3E</figureCitation>
|
||
), suggesting that as eggs pass through the female pore, the pouch would contract, and sperm from the spermatophores could easily be discharged into the cocoons to fertilize the eggs. The examined spermatophores are conical, teardrop-shaped, 68–75 µm long and 23–34 µm maximum diameter, whereas they were longer (average 93 µm) and thinner (average 16 µm width) in
|
||
<bibRefCitation id="B15AD98CFFD8C84C1F52D571FF736D24" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">Gavrilov's (1958)</bibRefCitation>
|
||
description.
|
||
</paragraph>
|
||
<paragraph id="D574A47DFFD8C84C1ABCD539FD626E9A" blockId="7.[151,1437,253,2029]" pageId="7" pageNumber="48">
|
||
The male duct in
|
||
<taxonomicName id="12CBDFFEFFD8C84C1BF7D53AFE6D6D00" authorityName="Gavrilov" authorityYear="1955" box="[396,511,1117,1142]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="48" phylum="Annelida" rank="species" species="descolei">
|
||
<emphasis id="E7BF786FFFD8C84C1BF7D53AFE6D6D00" box="[396,511,1117,1142]" italics="true" pageId="7" pageNumber="48">P. descolei</emphasis>
|
||
</taxonomicName>
|
||
is composed of a small sperm funnel, placed ventrally on septum 10/11 (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1F42D539FA1C6D00" box="[1337,1422,1117,1142]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3F</figureCitation>
|
||
); a moderately long vas deferens; a tubular atrium, densely covered with diffuse prostatic cells; and a narrow ejaculatory duct which opens to a large male copulatory bursa [referred to as "massive penial sac" by
|
||
<bibRefCitation id="B15AD98CFFD8C84C1F5FD5C0FE1C6D96" pageId="7" pageNumber="48" refId="ref9764" refString="Brinkhurst, R. O. & Marchese, M. (1987) A contribution to the taxonomy of the aquatic Oligochaeta (Haplotaxidae, Phreodrilidae, Tubificidae) of South America. Canadian Journal of Zoology, 65, 3154 - 3165. https: // doi. org / 10.1139 / z 87 - 474" type="journal article">Brinkhurst and Marchese (1987)</bibRefCitation>
|
||
]. This bursa is composed of an ental male sac [= bulbous ental portion of the penial sac in
|
||
<bibRefCitation id="B15AD98CFFD8C84C1AECD58FFEC36C72" box="[151,337,1259,1284]" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">Gavrilov (1958)</bibRefCitation>
|
||
], and an ectal section presumably serving as a copulatory chamber. The ental, bulbous section has a glandular inner epithelium, and contains a pendant, copulatory organ, originally described as a penis. The copulatory organ is long and covered or not by a chinitous sheath in Gavrilov's
|
||
<typeStatus id="0A701ADFFFD8C84C1E52D457FBCB6C3D" box="[1065,1113,1331,1355]" pageId="7" pageNumber="48">type</typeStatus>
|
||
collection (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1E9AD456FAE26C3C" box="[1249,1392,1330,1355]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3H,K,L</figureCitation>
|
||
); in contrast, it is short and glandular in Marchese's collection (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1933D431FC566C18" box="[840,964,1365,1390]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3G,I,J</figureCitation>
|
||
). In all instances, the ectal section has a folded epithelium, and terminates in a longitudinal slit-like male pore in the ventral side of segment XI (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1F66D41DFAE36CE7" box="[1309,1393,1401,1426]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3F</figureCitation>
|
||
). In their description of new material from other localities in
|
||
<collectingCountry id="ADDCE4EDFFD8C84C1945D4F8FC3F6CC3" box="[830,941,1436,1461]" name="Argentina" pageId="7" pageNumber="48">Argentina</collectingCountry>
|
||
and
|
||
<collectingCountry id="ADDCE4EDFFD8C84C1996D4F9FBB16CC3" box="[1005,1059,1437,1461]" name="Peru" pageId="7" pageNumber="48">Perú</collectingCountry>
|
||
,
|
||
<bibRefCitation id="B15AD98CFFD8C84C1E48D4F8FA0E6CC3" box="[1075,1436,1436,1461]" pageId="7" pageNumber="48" refId="ref9764" refString="Brinkhurst, R. O. & Marchese, M. (1987) A contribution to the taxonomy of the aquatic Oligochaeta (Haplotaxidae, Phreodrilidae, Tubificidae) of South America. Canadian Journal of Zoology, 65, 3154 - 3165. https: // doi. org / 10.1139 / z 87 - 474" type="journal article">Brinkhurst & Marchese (1987)</bibRefCitation>
|
||
concluded that the organ described as a penis by Gavrilov was not a true penis, but rather a simple fold in the wall of the sac, not connected to the male duct. However, in both the
|
||
<typeStatus id="0A701ADFFFD8C84C1905D480FC3C6C8A" box="[894,942,1508,1532]" pageId="7" pageNumber="48">type</typeStatus>
|
||
collection and in the Marchese's collection examined here, the glandular organ is formed in the bulbous male sac at the junction of the atrial ejaculatory duct [=efferent duct in
|
||
<bibRefCitation id="B15AD98CFFD8C84C1B1ED74EFD8C6F35" box="[357,542,1578,1603]" pageId="7" pageNumber="48" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">Gavrilov (1958)</bibRefCitation>
|
||
] and there are no traces of the musculature expected for an intromittent organ. As
|
||
<bibRefCitation id="B15AD98CFFD8C84C1ABCD729FE076F10" box="[199,405,1613,1638]" pageId="7" pageNumber="48" refId="ref11010" refString="Gavrilov, K. (1955 b) Uber die uniparentale Vermehrung von Paranadrilus. Zoological Anzeiger, 155, 302 - 306." type="journal article">Gavrilov (1955b)</bibRefCitation>
|
||
suggested, this copulatory organ is probably responsible for the formation of the spermatophore capsules and their attachment to the body wall of the mating partner, since in the absence of spermathecae, an intromittent organ seems unnecessary. In
|
||
<taxonomicName id="12CBDFFEFFD8C84C1924D7F0FC666FDB" authorityName="Gavrilov" authorityYear="1955" box="[863,1012,1684,1709]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="48" phylum="Annelida" rank="genus">
|
||
<emphasis id="E7BF786FFFD8C84C1924D7F0FC666FDB" box="[863,1012,1684,1709]" italics="true" pageId="7" pageNumber="48">Paranadrilus</emphasis>
|
||
</taxonomicName>
|
||
, the material required to build the spermatophore capsule seems to be secreted within the male sac, where Gavrilov described a copious glandular secretion (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1B77D7BFFEEC6F82" box="[268,382,1755,1780]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3K,L</figureCitation>
|
||
). As described by Gavrilov, a chitinous sheath covering the dagger-like "penis" (
|
||
<figureCitation id="4DF0B8F8FFD8C84C1F72D7BFFAF26F82" box="[1289,1376,1755,1780]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3H</figureCitation>
|
||
) was observed in six specimens of the
|
||
<typeStatus id="0A701ADFFFD8C84C1868D664FDD16E6E" box="[531,579,1792,1816]" pageId="7" pageNumber="48">type</typeStatus>
|
||
collection, where the male duct was scarcely developed, but the ovary was voluminous and eggs were present. However, it was not observed in the
|
||
<typeStatus id="0A701ADFFFD8C84C19B6D646FBA46E4D" box="[973,1078,1826,1851]" pageId="7" pageNumber="48" type="lectotype">lectotype</typeStatus>
|
||
, where the male duct was more developed, nor was it seen by
|
||
<bibRefCitation id="B15AD98CFFD8C84C1B8ED622FCFB6E29" box="[501,873,1862,1887]" pageId="7" pageNumber="48" refId="ref9764" refString="Brinkhurst, R. O. & Marchese, M. (1987) A contribution to the taxonomy of the aquatic Oligochaeta (Haplotaxidae, Phreodrilidae, Tubificidae) of South America. Canadian Journal of Zoology, 65, 3154 - 3165. https: // doi. org / 10.1139 / z 87 - 474" type="journal article">Brinkhurst and Marchese (1987)</bibRefCitation>
|
||
, or in the present study of Marchese's collection. The chinitous layer in the copulatory organ of Gavrilov's specimens probably represents the material ready for the spermatophore capsule, before it has been fully formed and released. Formation of spermatophores in the bulbous male sac is supported by the finding of one developing spermatophore in the most ental part of the copulatory bursa in one specimen from Marchese's collection (
|
||
<figureCitation id="4DF0B8F8FFD8C84C18E8D6B0FD706E9A" box="[659,738,2004,2029]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="7" pageNumber="48">Fig. 3J</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="81B4F4F5FFD7C8431AECD457FE726F0A" httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="8" pageNumber="49" startId="8.[151,250,1331,1353]" targetBox="[262,1322,204,1300]" targetPageId="8">
|
||
<paragraph id="D574A47DFFD7C8431AECD457FE726F0A" blockId="8.[151,1436,1331,1660]" pageId="8" pageNumber="49">
|
||
<emphasis id="E7BF786FFFD7C8431AECD457FE816C3F" bold="true" box="[151,275,1331,1353]" pageId="8" pageNumber="49">FIGURE 3.</emphasis>
|
||
<taxonomicName id="12CBDFFEFFD7C8431B61D457FD0E6C3F" authority="Gavrilov, 1955" authorityName="Gavrilov" authorityYear="1955" box="[282,668,1331,1353]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="49" phylum="Annelida" rank="species" species="descolei">
|
||
<emphasis id="E7BF786FFFD7C8431B61D457FE686C3F" box="[282,506,1331,1353]" italics="true" pageId="8" pageNumber="49">Paranadrilus descolei</emphasis>
|
||
Gavrilov, 1955
|
||
</taxonomicName>
|
||
.
|
||
<emphasis id="E7BF786FFFD7C84318DDD457FD446C3F" bold="true" box="[678,726,1331,1353]" pageId="8" pageNumber="49">A–G</emphasis>
|
||
,
|
||
<emphasis id="E7BF786FFFD7C84318A7D457FD746C3F" bold="true" box="[732,742,1331,1353]" pageId="8" pageNumber="49">I</emphasis>
|
||
,
|
||
<emphasis id="E7BF786FFFD7C8431897D457FD6B6C3F" bold="true" box="[748,761,1331,1353]" pageId="8" pageNumber="49">J</emphasis>
|
||
from Marchese's collection,
|
||
<emphasis id="E7BF786FFFD7C8431E5BD457FBA66C3F" bold="true" box="[1056,1076,1331,1353]" pageId="8" pageNumber="49">H</emphasis>
|
||
,
|
||
<emphasis id="E7BF786FFFD7C8431E41D457FBDC6C3F" bold="true" box="[1082,1102,1331,1353]" pageId="8" pageNumber="49">K</emphasis>
|
||
,
|
||
<emphasis id="E7BF786FFFD7C8431E2FD457FBF76C3F" bold="true" box="[1108,1125,1331,1353]" pageId="8" pageNumber="49">L</emphasis>
|
||
from Gavrilov's collection.
|
||
<emphasis id="E7BF786FFFD7C8431FF8D457FA0E6C3F" bold="true" box="[1411,1436,1331,1353]" pageId="8" pageNumber="49">A.</emphasis>
|
||
Anterior part of the body.
|
||
<emphasis id="E7BF786FFFD7C8431BD9D436FE2B6C1E" bold="true" box="[418,441,1362,1384]" pageId="8" pageNumber="49">B.</emphasis>
|
||
Ventral gland in the posterior part of segment V.
|
||
<emphasis id="E7BF786FFFD7C84319C8D436FC596C1E" bold="true" box="[947,971,1362,1384]" pageId="8" pageNumber="49">C.</emphasis>
|
||
Spermatophores within a lateral pouch in the anterior part of segment XII.
|
||
<emphasis id="E7BF786FFFD7C8431BB9D414FE496CF0" bold="true" box="[450,475,1392,1414]" pageId="8" pageNumber="49">D.</emphasis>
|
||
Detail of the attachment of the spermatophores within the pouch.
|
||
<emphasis id="E7BF786FFFD7C8431E04D414FB056CF0" bold="true" box="[1151,1175,1392,1414]" pageId="8" pageNumber="49">E.</emphasis>
|
||
Transverse section of the empty spermatophore pouch.
|
||
<emphasis id="E7BF786FFFD7C8431BB1D4EBFE496CD3" bold="true" box="[458,475,1423,1445]" pageId="8" pageNumber="49">F.</emphasis>
|
||
Sperm funnel and male pore slit.
|
||
<emphasis id="E7BF786FFFD7C8431939D4EBFCC56CD3" bold="true" box="[834,855,1423,1445]" pageId="8" pageNumber="49">G.</emphasis>
|
||
Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12.
|
||
<emphasis id="E7BF786FFFD7C84318E7D4CAFD246CB2" bold="true" box="[668,694,1454,1476]" pageId="8" pageNumber="49">H.</emphasis>
|
||
Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection.
|
||
<emphasis id="E7BF786FFFD7C84319DED4A8FC276C94" bold="true" box="[933,949,1484,1506]" pageId="8" pageNumber="49">I.</emphasis>
|
||
Male sac with copulatory organ and copulatory bursa.
|
||
<emphasis id="E7BF786FFFD7C8431AA1D48FFF7F6F77" bold="true" box="[218,237,1515,1537]" pageId="8" pageNumber="49">J.</emphasis>
|
||
Spermatophore in the ental part of the copulatory bursa.
|
||
<emphasis id="E7BF786FFFD7C843194AD48FFCD76F77" bold="true" box="[817,837,1515,1537]" pageId="8" pageNumber="49">K</emphasis>
|
||
,
|
||
<emphasis id="E7BF786FFFD7C8431930D48FFCF06F77" bold="true" box="[843,866,1515,1537]" pageId="8" pageNumber="49">L.</emphasis>
|
||
Segment XI of the lectotype of
|
||
<taxonomicName id="12CBDFFEFFD7C8431ED3D488FA836F77" authorityName="Gavrilov" authorityYear="1955" box="[1192,1297,1515,1537]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="49" phylum="Annelida" rank="species" species="descolei">
|
||
<emphasis id="E7BF786FFFD7C8431ED3D488FA836F77" box="[1192,1297,1515,1537]" italics="true" pageId="8" pageNumber="49">P. descolei</emphasis>
|
||
</taxonomicName>
|
||
, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations:
|
||
<emphasis id="E7BF786FFFD7C84318F0D74CFD326F48" bold="true" box="[651,672,1576,1598]" pageId="8" pageNumber="49">at</emphasis>
|
||
attachment site,
|
||
<emphasis id="E7BF786FFFD7C8431929D74CFCFE6F48" bold="true" box="[850,876,1576,1598]" pageId="8" pageNumber="49">cb</emphasis>
|
||
copulatory bursa,
|
||
<emphasis id="E7BF786FFFD7C8431E56D74CFBD76F48" bold="true" box="[1069,1093,1576,1598]" pageId="8" pageNumber="49">co</emphasis>
|
||
copulatory organ,
|
||
<emphasis id="E7BF786FFFD7C8431F71D74CFAB16F48" bold="true" box="[1290,1315,1576,1598]" pageId="8" pageNumber="49">ed</emphasis>
|
||
ejaculatory duct,
|
||
<emphasis id="E7BF786FFFD7C8431AA9D723FF756F2B" bold="true" box="[210,231,1607,1629]" pageId="8" pageNumber="49">m</emphasis>
|
||
muscular layer,
|
||
<emphasis id="E7BF786FFFD7C8431BE3D723FE296F2B" bold="true" box="[408,443,1607,1629]" pageId="8" pageNumber="49">mp</emphasis>
|
||
male pore,
|
||
<emphasis id="E7BF786FFFD7C8431847D723FDC96F2B" bold="true" box="[572,603,1607,1629]" pageId="8" pageNumber="49">ms</emphasis>
|
||
male sac,
|
||
<emphasis id="E7BF786FFFD7C84318B5D723FD496F2B" bold="true" box="[718,731,1607,1629]" pageId="8" pageNumber="49">o</emphasis>
|
||
ovary,
|
||
<emphasis id="E7BF786FFFD7C8431957D723FCDA6F2B" bold="true" box="[812,840,1607,1629]" pageId="8" pageNumber="49">ph</emphasis>
|
||
pharynx,
|
||
<emphasis id="E7BF786FFFD7C84319CFD723FC546F2B" bold="true" box="[948,966,1607,1629]" pageId="8" pageNumber="49">sf</emphasis>
|
||
sperm funnel,
|
||
<emphasis id="E7BF786FFFD7C8431E1CD723FBED6F2B" bold="true" box="[1127,1151,1607,1629]" pageId="8" pageNumber="49">sp</emphasis>
|
||
spermatophore pouch,
|
||
<emphasis id="E7BF786FFFD7C8431F0DD723FA0E6F2B" bold="true" box="[1398,1436,1607,1629]" pageId="8" pageNumber="49">sph</emphasis>
|
||
spermatophore,
|
||
<emphasis id="E7BF786FFFD7C8431B41D702FEC76F0A" bold="true" box="[314,341,1638,1660]" pageId="8" pageNumber="49">vd</emphasis>
|
||
vas deferens.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="D574A47DFFD7C8421ABCD7A6FF7E6981" blockId="8.[151,1436,1730,2007]" lastBlockId="9.[151,1436,151,357]" lastPageId="9" lastPageNumber="50" pageId="8" pageNumber="49">
|
||
Only 11.2% of the individuals examined from cultures by
|
||
<bibRefCitation id="B15AD98CFFD7C8431903D7A6FBA36FAD" box="[888,1073,1730,1755]" pageId="8" pageNumber="49" refId="ref11034" refString="Gavrilov, K. (1958) Notas adicionales sobre Paranadrilus. Acta Zoologica Lilloana, XVI, 149 - 235." type="book chapter">Gavrilov (1958)</bibRefCitation>
|
||
had spermatophores (only 1–2 spermatophores per individual); however, in the specimens from Marchese's collection we have counted up to 6 spermatophores (
|
||
<quantity id="12330998FFD7C8431B1FD66EFE086E55" box="[356,410,1802,1827]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" pageId="8" pageNumber="49" unit="in" value="3.0">3 in</quantity>
|
||
each ventral pouch). In Gavrilov's studies of laboratory cultures, parthenogenetic reproduction predominated; however, this can be an alteration of the reproductive biology of the species under culture conditions, and should be contrasted with field data. Gavrilov never observed the spermatophore pouches in segment XII, even though he worked with live organisms and performed histological sections. Spermatophore pouches were not seen in the present review of the
|
||
<typeStatus id="0A701ADFFFD7C84318A2D6FFFC9B6EC5" box="[729,777,1947,1971]" pageId="8" pageNumber="49">type</typeStatus>
|
||
collection. This suggests that the lateral pouches were not formed under culture conditions, possibly due to a high proportion of some
|
||
<typeStatus id="0A701ADFFFD7C8431982D6DBFBBB6EA1" box="[1017,1065,1983,2007]" pageId="8" pageNumber="49">type</typeStatus>
|
||
of parthenogenetic reproduction [e.g. merospermy or pseudogamy, suggested by
|
||
<bibRefCitation id="B15AD98CFFD6C84218C3D1F3FC1069C6" box="[696,898,151,176]" pageId="9" pageNumber="50" refId="ref11010" refString="Gavrilov, K. (1955 b) Uber die uniparentale Vermehrung von Paranadrilus. Zoological Anzeiger, 155, 302 - 306." type="journal article">Gavrilov (1955b)</bibRefCitation>
|
||
]. Nevertheless, it is also possible that the genus
|
||
<taxonomicName id="12CBDFFEFFD6C8421AECD1D8FEBE69A3" authorityName="Gavrilov" authorityYear="1955" box="[151,300,188,213]" class="Clitellata" family="Naididae" genus="Paranadrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="50" phylum="Annelida" rank="genus">
|
||
<emphasis id="E7BF786FFFD6C8421AECD1D8FEBE69A3" box="[151,300,188,213]" italics="true" pageId="9" pageNumber="50">Paranadrilus</emphasis>
|
||
</taxonomicName>
|
||
includes additional species in the Neotropical region, a question that future genetic analyses can help answer.
|
||
</paragraph>
|
||
<paragraph id="D574A47DFFD6C8421ABCD060FAAA6812" blockId="9.[151,1436,151,357]" pageId="9" pageNumber="50">
|
||
Although not related to spermatophore production, some organs in Marchese's collection are worth mentioning because they can be of interest in future studies of the genus: the pharynx is voluminous and dorsally folded (
|
||
<figureCitation id="4DF0B8F8FFD6C8421F14D043FF2E6812" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="9" pageNumber="50">Fig. 3A</figureCitation>
|
||
); and small (single?) mid-ventral glands are shown in the posterior part of segments IV–VII (
|
||
<figureCitation id="4DF0B8F8FFD6C8421EA8D028FAB86812" box="[1235,1322,332,357]" captionStart="FIGURE 3" captionStartId="8.[151,250,1331,1353]" captionTargetBox="[262,1322,204,1300]" captionTargetId="figure@8.[254,1332,193,1310]" captionTargetPageId="8" captionText="FIGURE 3. Paranadrilus descolei Gavrilov, 1955. A–G,I,J from Marchese's collection, H,K,L from Gavrilov's collection. A. Anterior part of the body. B. Ventral gland in the posterior part of segment V. C. Spermatophores within a lateral pouch in the anterior part of segment XII. D. Detail of the attachment of the spermatophores within the pouch. E. Transverse section of the empty spermatophore pouch. F. Sperm funnel and male pore slit. G. Bulbous male sac containing the copulatory organ, and empty spermatophore pouch behind septum 11/12. H. Scarcely developed male sac containing a long copulatory organ covered by a chitinous layer in a dissected specimen from Gavrilov's type collection. I. Male sac with copulatory organ and copulatory bursa. J. Spermatophore in the ental part of the copulatory bursa. K,L. Segment XI of the lectotype of P. descolei, showing the large ovary, the bulbous male sac filled by glandular secretions and spermatozoids, and the copulatory bursa with folded walls. A,E,G,I–L, histological sections. Abbreviations: at attachment site, cb copulatory bursa, co copulatory organ, ed ejaculatory duct, m muscular layer, mp male pore, ms male sac, o ovary, ph pharynx, sf sperm funnel, sp spermatophore pouch, sph spermatophore, vd vas deferens." httpUri="https://zenodo.org/record/1451100/files/figure.png" pageId="9" pageNumber="50">Fig. 3B</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |