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<document id="A44A4A1CC1312FFDD2012D8D1580ECB0" ID-DOI="10.5281/zenodo.193114" ID-GBIF-Dataset="1c228c59-84ad-4a68-812f-2cffafd47c1f" ID-ISSN="1175-5326" ID-Zenodo-Dep="193114" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460097873350" checkinUser="plazi" docAuthor="Rodriguez, Pilar &amp; Achurra, Ainara" docDate="2010" docId="C8764570FFA6AA0A71C8BED3FE93FB4F" docLanguage="en" docName="zt02332p039.pdf" docOrigin="Zootaxa 2332" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Gianius navarroi Rodriguez &amp; Achurra, 2010, n. sp." docType="treatment" docVersion="10" lastPageNumber="31" masterDocId="344F3D08FFA7AA00715FBB4EFFDCFFA3" masterDocTitle="New species of aquatic oligochaetes (Annelida: Clitellata) from groundwaters in karstic areas of northern Spain, with taxonomic remarks on Lophochaeta ignota Štolc, 1886" masterLastPageNumber="39" masterPageNumber="21" pageNumber="22" updateTime="1698234791604" updateUser="plazi">
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<mods:title id="8C7130CB9DB68334192CDA9659E4DF5E">New species of aquatic oligochaetes (Annelida: Clitellata) from groundwaters in karstic areas of northern Spain, with taxonomic remarks on Lophochaeta ignota Štolc, 1886</mods:title>
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<mods:name id="3684EDDFEEDCC0ECC38959A4B3A08B32" type="personal">
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<mods:namePart id="4E951F6B44DDF711B8EE015A889F2E9B">Rodriguez, Pilar</mods:namePart>
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<mods:namePart id="2C98EE7D2A171915577AA30800C1E1FC">Achurra, Ainara</mods:namePart>
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<mods:date id="C1EB0A99BD32F04A641B43FF8D84272C">2010</mods:date>
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<treatment id="C8764570FFA6AA0A71C8BED3FE93FB4F" ID-DOI="http://doi.org/10.5281/zenodo.6211155" ID-GBIF-Taxon="124955401" ID-Zenodo-Dep="6211155" LSID="urn:lsid:plazi:treatment:C8764570FFA6AA0A71C8BED3FE93FB4F" httpUri="http://treatment.plazi.org/id/C8764570FFA6AA0A71C8BED3FE93FB4F" lastPageId="10" lastPageNumber="31" pageId="1" pageNumber="22">
<subSubSection id="08C5A7EDFFA6AA0171C8BED3FEE9FA79" pageId="1" pageNumber="22" type="nomenclature">
<paragraph id="4060F466FFA6AA0171C8BED3FE6CFA14" blockId="1.[151,432,1437,1498]" box="[151,432,1437,1463]" pageId="1" pageNumber="22">
<heading id="1B28430AFFA6AA0171C8BED3FE6CFA14" bold="true" box="[151,432,1437,1463]" fontSize="11" level="1" pageId="1" pageNumber="22" reason="1">
<emphasis id="72AB2874FFA6AA0171C8BED3FE6CFA14" bold="true" box="[151,432,1437,1463]" pageId="1" pageNumber="22">
<taxonomicName id="87DF8FE5FFA6AA0171C8BED3FEBAFA14" box="[151,358,1437,1463]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="1" pageNumber="22" phylum="Annelida" rank="species" species="navarroi" status="sp. nov.">
<emphasis id="72AB2874FFA6AA0171C8BED3FEBAFA14" bold="true" box="[151,358,1437,1463]" italics="true" pageId="1" pageNumber="22">Gianius navarroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA6AA017031BED3FE6CFA14" box="[366,432,1437,1463]" pageId="1" pageNumber="22" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="4060F466FFA6AA0171C8BE8EFEE9FA79" blockId="1.[151,432,1437,1498]" box="[151,309,1472,1498]" pageId="1" pageNumber="22">(Figures 12)</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA6AA0171C8BD43FDA7F987" box="[151,635,1548,1572]" pageId="1" pageNumber="22" type="description">
<paragraph id="4060F466FFA6AA0171C8BD43FDA7F987" blockId="1.[151,635,1548,1572]" box="[151,635,1548,1572]" pageId="1" pageNumber="22">
<taxonomicName id="87DF8FE5FFA6AA0171C8BD43FEEFF987" box="[151,307,1549,1572]" class="Clitellata" family="Tubificidae" kingdom="Animalia" order="Haplotaxida" pageId="1" pageNumber="22" phylum="Annelida" rank="subFamily" subFamily="Phallodrilinae">
<emphasis id="72AB2874FFA6AA0171C8BD43FEEFF987" box="[151,307,1549,1572]" italics="true" pageId="1" pageNumber="22">Phallodrilinae</emphasis>
</taxonomicName>
<emphasis id="72AB2874FFA6AA017066BD42FEA9F987" bold="true" box="[313,373,1548,1572]" pageId="1" pageNumber="22">n. sp.</emphasis>
in
<bibRefCitation id="244E8997FFA6AA0170C8BD43FDABF987" author="Camacho" box="[407,631,1549,1572]" pageId="1" pageNumber="38" refString="Camacho, A. I., Valdecasas, A. G., Rodriguez, J., Cuezva, S., Lario, J. &amp; Sanchez-Moral, S. (2006) Habitat constraints in epikarstic waters of an Iberian Peninsula cave system. Annales de Limnologie - International Journal of Limnology, 42, 127 - 140." type="journal article" year="2006">
Camacho
<emphasis id="72AB2874FFA6AA01735DBD43FDE4F987" box="[514,568,1549,1572]" italics="true" pageId="1" pageNumber="22">et al.</emphasis>
2006
</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA6AA0171C8BD1DFCD1F8DF" pageId="1" pageNumber="22" type="materials_examined">
<paragraph id="4060F466FFA6AA0171C8BD1DFDF2F937" blockId="1.[151,1437,1619,2032]" pageId="1" pageNumber="22">
<emphasis id="72AB2874FFA6AA0171C8BD1DFECAF9CE" bold="true" box="[151,278,1619,1645]" pageId="1" pageNumber="22">
<typeStatus id="9F644AC4FFA6AA0171C8BD1DFECDF9CE" box="[151,273,1619,1645]" pageId="1" pageNumber="22" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="26CE6CA3FFA6AA017040BD1AFEA6F9CD" LSID="urn:lsid:biocol.org:col:35275" box="[287,378,1620,1646]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="1" pageNumber="22">MNCN</collectionCode>
16.03/3051. An incomplete mature specimen, stained in Ehrlichs Haematoxylin and whole-mounted in
<collectingCountry id="38C8B4F6FFA6AA01702BBD34FE10F937" box="[372,460,1658,1684]" name="Canada" pageId="1" pageNumber="22">Canada</collectingCountry>
balsam.
</paragraph>
<paragraph id="4060F466FFA6AA01719ABDEEFC18F941" blockId="1.[151,1437,1619,2032]" pageId="1" pageNumber="22">
<emphasis id="72AB2874FFA6AA01719ABDEEFEB0F919" bold="true" box="[197,364,1696,1722]" pageId="1" pageNumber="22">
<typeStatus id="9F644AC4FFA6AA01719ABDEEFEDFF919" box="[197,259,1696,1722]" pageId="1" pageNumber="22">Type</typeStatus>
locality
</emphasis>
. Ojo Guareña cave (site: “Granja Ortiz”,
<date id="3461D2A6FFA6AA017229BDEFFBCDF918" box="[886,1041,1697,1723]" pageId="1" pageNumber="22" value="2002-06-01">1 June 2002</date>
), northern Burgos,
<collectingCountry id="38C8B4F6FFA6AA01745DBDEFFA90F918" box="[1282,1356,1697,1723]" name="Spain" pageId="1" pageNumber="22">Spain</collectingCountry>
. UTM coordinates for the Palomera entrance: X:446595, Y:4764790, Z:724.
</paragraph>
<paragraph id="4060F466FFA6AA01719ABDA0FCD1F8DF" blockId="1.[151,1437,1619,2032]" pageId="1" pageNumber="22">
<emphasis id="72AB2874FFA6AA01719ABDA0FE93F8AB" bold="true" box="[197,335,1774,1800]" pageId="1" pageNumber="22">
<typeStatus id="9F644AC4FFA6AA01719ABDA0FE96F8AB" box="[197,330,1774,1800]" pageId="1" pageNumber="22" type="paratype">Paratypes</typeStatus>
.
</emphasis>
<collectionCode id="26CE6CA3FFA6AA017007BDA0FE6FF8AB" LSID="urn:lsid:biocol.org:col:35275" box="[344,435,1774,1800]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="1" pageNumber="22">MNCN</collectionCode>
16.03/3052. A complete mature specimen (unmated) from the
<typeStatus id="9F644AC4FFA6AA01759FBDA0FB29F8AB" box="[1216,1269,1774,1800]" pageId="1" pageNumber="22">type</typeStatus>
locality, site: “Gour de las Hojas” (
<date id="3461D2A6FFA6AA0170C5BC5BFD9AF88C" box="[410,582,1813,1839]" pageId="1" pageNumber="22" value="2003-03-08">8 March 2003</date>
). A complete mature specimen in the collection at the University of the Basque Country (UPV/EHU), from the
<typeStatus id="9F644AC4FFA6AA01732DBC72FD79F8F5" box="[626,677,1852,1878]" pageId="1" pageNumber="22">type</typeStatus>
locality,
<date id="3461D2A6FFA6AA01724BBC72FC6BF8F5" box="[788,951,1852,1878]" pageId="1" pageNumber="22" value="2007-06-11">11 June 2007</date>
. Both specimens are stained in Ehrlichs Haematoxylin and whole-mounted in
<collectingCountry id="38C8B4F6FFA6AA01730CBC2CFD77F8DF" box="[595,683,1890,1916]" name="Canada" pageId="1" pageNumber="22">Canada</collectingCountry>
balsam.
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA6AA01719ABCC6FEEDF853" pageId="1" pageNumber="22" type="etymology">
<paragraph id="4060F466FFA6AA01719ABCC6FEEDF853" blockId="1.[151,1437,1619,2032]" pageId="1" pageNumber="22">
<emphasis id="72AB2874FFA6AA01719ABCC6FE96F801" bold="true" box="[197,330,1928,1954]" pageId="1" pageNumber="22">Etymology</emphasis>
. Named after the physiologist Dr. Enrique Navarro who has helped P. Rodriguez in sampling numerous sites for the study of aquatic oligochaetes, and has collaborated in applied research with oligochaetes.
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA5AA03719ABBD9FADEF9D2" lastPageId="3" lastPageNumber="24" pageId="2" pageNumber="23" type="description">
<paragraph id="4060F466FFA5AA02719ABBD9FCEBFD97" blockId="2.[151,1436,151,564]" pageId="2" pageNumber="23">
<emphasis id="72AB2874FFA5AA02719ABBD9FE83FF12" bold="true" box="[197,351,151,177]" pageId="2" pageNumber="23">Description.</emphasis>
Length of preserved worms 1.7 mm; number of segments (in two complete specimens) 23 and 26; maximum diameter 150 µm in segment XI. Prostomium rounded, with glandular epidermis (Fig. 1A). Pygidium short and wide (Fig. 1B). Body wall consisting of a fine cuticule (
<emphasis id="72AB2874FFA5AA027554BBABFBFBFF5D" box="[1035,1063,229,254]" italics="true" pageId="2" pageNumber="23">ca</emphasis>
1 µm thick), an epidermal layer (5 µm thick) and a relatively poorly developed musculature (23 µm thick). Clitellum formed by glands disposed in one layer, from the last third of segment X back to the beginning of segment XII. There is no secondary annulation. One pair of simple spermathecal pores in the most anterior part of segment X, opening in line with ventral chaetae. One pair of male pores in segment XI on porophores, in line with ventral chaetae. All chaetae bifid, 2427 µm long, with distal tooth shorter than proximal. Anterior segments with 25 chaetae per bundle,
<quantity id="87275983FFA5AA02707EBA83FE91FE44" box="[289,333,461,487]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="2" pageNumber="23" unit="in" value="2.0">2 in</quantity>
postclitellar segments. Chaetal bundles are in the posterior third of the segment. A bundle of 5 penial chaetae forms an arc, towards the sagittal plane of the body (
<figureCitation id="D8E4E8E3FFA5AA0272E9BABAFC22FDAD" box="[950,1022,500,526]" captionStart="FIGURE 2" captionStartId="3.[151,256,1406,1430]" captionTargetBox="[303,1266,811,1405]" captionTargetId="figure@3.[303,1283,793,1411]" captionTargetPageId="3" captionText="FIGURE 2. Gianius navarroi n. sp. A: general diagram of the reproductive organs of the species, X and XI: segment number; B: free-hand drawing of somatic chaeta. Abbreviations: a: atrium, mp: male pore, o: ovary, pc: penial chaetae, pgI and pgII: prostate glands I and II, sf: sperm funnel, sa: spermathecal ampulla, sd: spemathecal duct, ss: sperm sacs, t: testis, vd: vas deferens." httpUri="https://zenodo.org/record/193115/files/figure.png" pageId="2" pageNumber="23">Fig. 2</figureCitation>
). Penial chaetae straight, with bent distal end, bifid, and of similar length to somatic chaetae.
</paragraph>
<paragraph id="4060F466FFA5AA02707BBC77FDD7F81B" blockId="2.[151,1437,1849,1976]" pageId="2" pageNumber="23">
<taxonomicName id="87DF8FE5FFA5AA02707BBC77FE3DF8F3" box="[292,481,1849,1872]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="2" pageNumber="23" phylum="Annelida" rank="species" species="navarroi">
<emphasis id="72AB2874FFA5AA02707BBC77FE3DF8F3" box="[292,481,1849,1872]" italics="true" pageId="2" pageNumber="23">Gianius navarroi</emphasis>
</taxonomicName>
A: anterior part of the body in dorso-ventral view, IVVII: segment number; B: anterior and posterior ends of the body in lateral view; C: segments VII and VIII.
<emphasis id="72AB2874FFA5AA02750BBC15FADDF8D0" bold="true" box="[1108,1281,1883,1907]" pageId="2" pageNumber="23">Abbreviations</emphasis>
: b: brain, cc: chloragogen cells, i: intestine, od: oesophageal diverticula, ph: pharynx, phg: pharyngeal glands, py: pygidium.
<emphasis id="72AB2874FFA5AA02743FBC33FF15F81B" bold="true" pageId="2" pageNumber="23">Scale bars</emphasis>
: A and B: 100 µm, C: 50 µm.
</paragraph>
<paragraph id="4060F466FFA4AA03719ABBD6FAA2FE39" blockId="3.[151,1437,152,758]" pageId="3" pageNumber="24">
Brain with two histologically distinct sections: anterior and posterior parts having large nuclei intensely stained with haematoxylin, and a granulated median part that does not stain well. No coelomocytes observed within the coelomic cavity. Pharyngeal glands in segments IIIV, dorsal in III, and both lateral and ventral in segments IV and V (in a
<typeStatus id="9F644AC4FFA4AA0370E6BA42FDFDFE85" box="[441,545,268,294]" pageId="3" pageNumber="24" type="paratype">paratype</typeStatus>
, glands also ventrally in VI). Chloragogen cells form a layer on the gut from segment VI backwards. In segment VII, a pair of dorsal digestive diverticula, can occupy half or the entire length of the segment, and appear to be attached to the oesophagus at the posterior end of the segment (Figs. 1A, C). A dilatation of the gut in segment VIII is interpreted as the beginning of the intestine (Figs. 1B, C).
</paragraph>
<paragraph id="4060F466FFA4AA03719ABAE8FEA9FD55" blockId="3.[151,1437,152,758]" pageId="3" pageNumber="24">
One pair testes in segment X and one pair ovaries in segment XI. Sperm sacs in segment XI (in one specimen spermatogonia and spermatocytes are free in the coelom of segment X). Male ducts paired, with two prostate glands: one ental, close to the junction of the vas deferens with atrium; and other ectal, close to the male pore, much smaller than the ental one. A ciliated vas deferens (6 µm diameter) joins the atrium apically. Atria tubular (77 µm long, up to 15 µm diameter) and curved; thicker in the middle in lateral view (with homogeneous diameter in dorsal view) and gradually narrowing to both ental and ectal ends. The atrial musculature is thin (23 µm thick) and the lumen is ciliated or filled with granules (as in
<figureCitation id="D8E4E8E3FFA4AA03758FB9C0FAC5FD0B" box="[1232,1305,654,680]" captionStart="FIGURE 2" captionStartId="3.[151,256,1406,1430]" captionTargetBox="[303,1266,811,1405]" captionTargetId="figure@3.[303,1283,793,1411]" captionTargetPageId="3" captionText="FIGURE 2. Gianius navarroi n. sp. A: general diagram of the reproductive organs of the species, X and XI: segment number; B: free-hand drawing of somatic chaeta. Abbreviations: a: atrium, mp: male pore, o: ovary, pc: penial chaetae, pgI and pgII: prostate glands I and II, sf: sperm funnel, sa: spermathecal ampulla, sd: spemathecal duct, ss: sperm sacs, t: testis, vd: vas deferens." httpUri="https://zenodo.org/record/193115/files/figure.png" pageId="3" pageNumber="24">Fig. 2</figureCitation>
). The atria open into salient round porophores (
<emphasis id="72AB2874FFA4AA03730FB9FBFDB1FD6D" box="[592,621,693,718]" italics="true" pageId="3" pageNumber="24">ca</emphasis>
10 µm long), where the penial chaetae are located. Female funnels were not observed.
</paragraph>
<caption id="14A0A4EEFFA4AA0371C8BE30FE4CFA5E" httpUri="https://zenodo.org/record/193115/files/figure.png" pageId="3" pageNumber="24" targetBox="[303,1266,811,1405]" targetPageId="3">
<paragraph id="4060F466FFA4AA0371C8BE30FE4CFA5E" blockId="3.[151,1436,1406,1533]" pageId="3" pageNumber="24">
<emphasis id="72AB2874FFA4AA0371C8BE30FEC7FA35" bold="true" box="[151,283,1406,1430]" pageId="3" pageNumber="24">FIGURE 2.</emphasis>
<taxonomicName id="87DF8FE5FFA4AA03707CBE31FE02FA35" box="[291,478,1407,1430]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="3" pageNumber="24" phylum="Annelida" rank="species" species="navarroi" status="sp. nov.">
<emphasis id="72AB2874FFA4AA03707CBE31FE02FA35" box="[291,478,1407,1430]" italics="true" pageId="3" pageNumber="24">Gianius navarroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA4AA0370BABE30FDFFFA35" box="[485,547,1406,1430]" pageId="3" pageNumber="24" rank="species">
<emphasis id="72AB2874FFA4AA0370BABE30FDFFFA35" bold="true" box="[485,547,1406,1430]" pageId="3" pageNumber="24">n. sp.</emphasis>
</taxonomicNameLabel>
A: general diagram of the reproductive organs of the species, X and XI: segment number; B: free-hand drawing of somatic chaeta.
<emphasis id="72AB2874FFA4AA0373F3BEEFFC91FA1A" bold="true" box="[684,845,1441,1465]" pageId="3" pageNumber="24">Abbreviations</emphasis>
: a: atrium, mp: male pore, o: ovary, pc: penial chaetae, pgI and pgII: prostate glands I and II, sf: sperm funnel, sa: spermathecal ampulla, sd: spemathecal duct, ss: sperm sacs, t: testis, vd: vas deferens.
</paragraph>
</caption>
<paragraph id="4060F466FFA4AA03719ABD7EFADEF9D2" blockId="3.[151,1437,1584,2035]" pageId="3" pageNumber="24">One pair of spermathecae, with a stout duct (20 µm long, 1316 µm wide) and an elongated globular ampulla (4050 µm long, 1617 µm maximum diameter), which contains sperm forming a loop.</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA4AA04719ABD33FF28FCC9" lastPageId="4" lastPageNumber="25" pageId="3" pageNumber="24" type="distribution">
<paragraph id="4060F466FFA4AA03719ABD33FF37F8FA" blockId="3.[151,1437,1584,2035]" pageId="3" pageNumber="24">
<emphasis id="72AB2874FFA4AA03719ABD33FE20F934" bold="true" box="[197,508,1661,1687]" pageId="3" pageNumber="24">Distribution and habitat.</emphasis>
<taxonomicName id="87DF8FE5FFA4AA03735CBD30FD17F934" box="[515,715,1662,1687]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="3" pageNumber="24" phylum="Annelida" rank="species" species="navarroi" status="sp. nov.">
<emphasis id="72AB2874FFA4AA03735CBD30FD17F934" box="[515,715,1662,1687]" italics="true" pageId="3" pageNumber="24">Gianius navarroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA4AA03738CBD33FCC9F934" box="[723,789,1661,1687]" pageId="3" pageNumber="24" rank="species">
<emphasis id="72AB2874FFA4AA03738CBD33FCC9F934" bold="true" box="[723,789,1661,1687]" pageId="3" pageNumber="24">n. sp.</emphasis>
</taxonomicNameLabel>
is known only from Ojo Guareña cave system, and it is likely to be a stygobiont. The species was collected from two different epikarstic sites within the cave, but was rare, at most one specimen per sample. Habitats sampled were puddles and cave streams, having temporary or permanent water, at depths up to
<quantity id="87275983FFA4AA03731ABDBFFD44F8A8" box="[581,664,1777,1803]" metricMagnitude="-1" metricUnit="m" metricValue="4.0" pageId="3" pageNumber="24" unit="cm" value="40.0">40 cm</quantity>
; sediment was composed of mud, sand and/or stones. Water temperature was 8.49.9ºC, pH 7.88.0, dissolved oxygen 74.378.4 % saturation and conductivity 389465 µS/cm.
</paragraph>
<paragraph id="4060F466FFA4AA04719ABC2BFF28FCC9" blockId="3.[151,1437,1584,2035]" lastBlockId="4.[151,1437,152,990]" lastPageId="4" lastPageNumber="25" pageId="3" pageNumber="24">
<emphasis id="72AB2874FFA4AA03719ABC2BFE11F8DC" bold="true" box="[197,461,1893,1919]" pageId="3" pageNumber="24">Taxonomic remarks.</emphasis>
<taxonomicName id="87DF8FE5FFA4AA03708ABC28FD79F8DC" box="[469,677,1894,1919]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="3" pageNumber="24" phylum="Annelida" rank="species" species="navarroi" status="sp. nov.">
<emphasis id="72AB2874FFA4AA03708ABC28FD79F8DC" box="[469,677,1894,1919]" italics="true" pageId="3" pageNumber="24">Gianius navarroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA4AA0373F1BC2BFD2FF8DC" box="[686,755,1893,1919]" pageId="3" pageNumber="24" rank="species">
<emphasis id="72AB2874FFA4AA0373F1BC2BFD2FF8DC" bold="true" box="[686,755,1893,1919]" pageId="3" pageNumber="24">n. sp.</emphasis>
</taxonomicNameLabel>
has spermathecal pores in line with the ventral chaetae, spermathecae with short ducts, vas deferens entering the apical end of an elongated and curved atrium, and two prostate glands per atrium attached to proximal and distal ends, respectively. All of these characteristics are diagnostic for the phallodriline genus
<taxonomicName id="87DF8FE5FFA4AA0373C4BC97FC7BF850" authority="Erseus, 1992" authorityName="Erseus" authorityYear="1992" box="[667,935,2009,2035]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="3" pageNumber="24" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFA4AA0373C4BC97FD20F851" box="[667,764,2009,2034]" italics="true" pageId="3" pageNumber="24">Gianius</emphasis>
<bibRefCitation id="244E8997FFA4AA03725BBC97FC7BF850" author="Erseus" box="[772,935,2009,2035]" pageId="3" pageNumber="38" refString="Erseus, C. (1992) A generic revision of the Phallodrilinae (Oligochaeta, Tubificidae). Zoologica Scripta, 21, 5 - 48." type="journal article" year="1992">Erséus, 1992</bibRefCitation>
</taxonomicName>
. The presence of digestive diverticula in segment VII was previously known only in the species
<taxonomicName id="87DF8FE5FFA3AA047273BBD6FA4BFF11" authority="Rodriguez and Giani, 1989" authorityName="Rodriguez and Giani" authorityYear="1989" box="[812,1431,152,178]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="labouichensis">
<emphasis id="72AB2874FFA3AA047273BBD6FBE9FF12" box="[812,1077,152,177]" italics="true" pageId="4" pageNumber="25">Gianius labouichensis</emphasis>
(
<bibRefCitation id="244E8997FFA3AA04751ABBD6FA53FF11" author="Rodriguez" box="[1093,1423,152,178]" pageId="4" pageNumber="39" refString="Rodriguez, P. &amp; Giani, N. (1989) New species of Phallodrilus (Oligochaeta, Tubificidae) from caves of Northern Spain and Southwestern France. Hydrobiologia, 180, 57 - 63." type="journal article" year="1989">Rodriguez and Giani, 1989</bibRefCitation>
)
</taxonomicName>
; although the connection of the diverticula with the gut appears to be in the posterior end of the segment for the new species, whereas for
<taxonomicName id="87DF8FE5FFA3AA04709ABBABFDA2FF5D" box="[453,638,229,254]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="labouchensis">
<emphasis id="72AB2874FFA3AA04709ABBABFDA2FF5D" box="[453,638,229,254]" italics="true" pageId="4" pageNumber="25">G. labouchensis</emphasis>
</taxonomicName>
the single diverticulum connects anteriorly. However, the latter is a distinctive species, well distinguished from other congeners by the lack of penial chaetae, a uniformly tubular atrium with a protrusible pseudopenis, and elongated narrow spermathecal ampulla. The general structure of the male duct in
<taxonomicName id="87DF8FE5FFA3AA047000BA17FE3AFED1" box="[351,486,345,370]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="navarroi" status="sp. nov.">
<emphasis id="72AB2874FFA3AA047000BA17FE3AFED1" box="[351,486,345,370]" italics="true" pageId="4" pageNumber="25">G. navarroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA3AA0470B0BA16FDEFFED1" box="[495,563,344,370]" pageId="4" pageNumber="25" rank="species">
<emphasis id="72AB2874FFA3AA0470B0BA16FDEFFED1" bold="true" box="[495,563,344,370]" pageId="4" pageNumber="25">n. sp.</emphasis>
</taxonomicNameLabel>
is more similar to the remaining species of the genus, being closest to
<taxonomicName id="87DF8FE5FFA3AA0474DCBA17FE08FE39" authority="Hrabe, 1960" authorityName="Hrabe" authorityYear="1960" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="aquaedulcis">
<emphasis id="72AB2874FFA3AA0474DCBA17FEFAFE3A" italics="true" pageId="4" pageNumber="25">G. aquaedulcis</emphasis>
(
<bibRefCitation id="244E8997FFA3AA047069BA30FE10FE39" author="Hrabe" box="[310,460,382,410]" pageId="4" pageNumber="39" refString="Hrabe, S. (1960) Oligochaeta limicola from the collection of Dr. Husmann. Publications de la Faculte des Sciences de l'Universite J. E. Purkyn e, Brno, 415, 245 - 277." type="journal article" year="1960">Hrabë, 1960</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="87DF8FE5FFA3AA0470BBBACEFD71FE39" authority="Juget" authorityName="Juget" box="[484,685,384,410]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="cavealis">
<emphasis id="72AB2874FFA3AA0470BBBACEFDBBFE3A" box="[484,615,384,409]" italics="true" pageId="4" pageNumber="25">G. cavealis</emphasis>
Juget
</taxonomicName>
and des
<bibRefCitation id="244E8997FFA3AA047247BACEFC3FFE39" author="Chatelliers" box="[792,995,384,410]" pageId="4" pageNumber="39" refString="Juget, J. &amp; des Chatelliers, M. (2001) Taxonomical survey of some stygobiont oligochaetes from the eastern part of France, including description of a new species. Hydrobiologia, 463, 23 - 28." type="journal article" year="2001">Chatelliers, 2001</bibRefCitation>
and
<taxonomicName id="87DF8FE5FFA3AA047540BACEFEECFE63" authority="Arslan, Timm and Erseus, 2007" authorityName="Arslan, Timm and Erseus" authorityYear="2007" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="anatolicus">
<emphasis id="72AB2874FFA3AA047540BACEFB60FE3A" box="[1055,1212,384,409]" italics="true" pageId="4" pageNumber="25">G. anatolicus</emphasis>
<bibRefCitation id="244E8997FFA3AA04759CBACEFEECFE63" author="Arslan" pageId="4" pageNumber="38" refString="Arslan, N., Timm, T. &amp; Erseus, C. (2007) Aquatic Oligochaeta (Annelida) of Balikdami wetland (Turkey), with description of two new species of Phallodrilinae. Biologia, 62, 323 - 334." type="journal article" year="2007">Arslan, Timm and Erséus, 2007</bibRefCitation>
</taxonomicName>
based on the thin atrial musculature and the moderate (45) number of penial chaetae.
<taxonomicName id="87DF8FE5FFA3AA047460BAE9FF20FE45" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="cavealis">
<emphasis id="72AB2874FFA3AA047460BAE9FF20FE45" italics="true" pageId="4" pageNumber="25">Gianius cavealis</emphasis>
</taxonomicName>
is well distinguished from the new species by its distal atrial duct well separated from an atrial ampulla, and
<taxonomicName id="87DF8FE5FFA3AA047065BABAFE0BFDAE" box="[314,471,500,525]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="anatolicus">
<emphasis id="72AB2874FFA3AA047065BABAFE0BFDAE" box="[314,471,500,525]" italics="true" pageId="4" pageNumber="25">G. anatolicus</emphasis>
</taxonomicName>
by the presence of a long, simple spermathecal duct and three different
<typeStatus id="9F644AC4FFA3AA047463BABAFAA7FDAD" box="[1340,1403,500,526]" pageId="4" pageNumber="25">types</typeStatus>
of somatic chaetae. The general structure of male duct and spermatheca are similar to
<taxonomicName id="87DF8FE5FFA3AA0475F7B954FABCFD90" box="[1192,1376,538,563]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="aquaedulcis">
<emphasis id="72AB2874FFA3AA0475F7B954FABCFD90" box="[1192,1376,538,563]" italics="true" pageId="4" pageNumber="25">G. aquaedulcis</emphasis>
</taxonomicName>
, and differences are mainly related to the relative size of the prostate glands (both glands large and of about equal size, in
<taxonomicName id="87DF8FE5FFA3AA0471AFB926FE42FD22" box="[240,414,616,641]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="aquaedulcis">
<emphasis id="72AB2874FFA3AA0471AFB926FE42FD22" box="[240,414,616,641]" italics="true" pageId="4" pageNumber="25">G. aquaedulcis</emphasis>
</taxonomicName>
but of visibly different size in the new species), as well as the size of chaetae (41 µm in
<taxonomicName id="87DF8FE5FFA3AA0471C8B9C1FE98FD0B" box="[151,324,655,680]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="aquaedulcis">
<emphasis id="72AB2874FFA3AA0471C8B9C1FE98FD0B" box="[151,324,655,680]" italics="true" pageId="4" pageNumber="25">G. aquaedulcis</emphasis>
</taxonomicName>
) and the presence of oesophageal digestive diverticula. The
<taxonomicName id="87DF8FE5FFA3AA047551B9C1FB60FD0B" box="[1038,1212,655,680]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="aquaedulcis">
<emphasis id="72AB2874FFA3AA047551B9C1FB60FD0B" box="[1038,1212,655,680]" italics="true" pageId="4" pageNumber="25">G. aquaedulcis</emphasis>
</taxonomicName>
material described by
<bibRefCitation id="244E8997FFA3AA0471E3B9FBFE3AFD6C" author="Farara" box="[188,486,693,719]" pageId="4" pageNumber="38" refString="Farara, D. G. &amp; Erseus, C. (1991) Phallodrilus aquaedulcis Hrabe, 1960, a meiobenthic fresh-water oligochaete (Tubificidae) previously known only from Europe, recorded from the Niagara River, North America. Canadian Journal of Zoology - Revue Canadienne de Zoologie, 69, 291 - 294." type="journal article" year="1991">Farara and Erséus (1991)</bibRefCitation>
in North
<collectingCountry id="38C8B4F6FFA3AA047306B9FBFD62FD6C" box="[601,702,693,719]" name="United States of America" pageId="4" pageNumber="25">America</collectingCountry>
has a larger body size and wider atria (3338 µm width); it is distinguished from the new species by the absence of noticeable porophores at the male pores, (described as “simple male pore on inconspicuous papilla”), and by the blunt ental end of the atria. The shape of the pygidium in
<emphasis id="72AB2874FFA3AA047073B867FE71FCE1" box="[300,429,809,834]" italics="true" pageId="4" pageNumber="25">G. n a v a r ro i</emphasis>
is unusual, but very similar to that described for
<taxonomicName id="87DF8FE5FFA3AA0472B0B867FB13FCE1" box="[1007,1231,809,834]" class="Clitellata" family="Tubificidae" genus="Mexidrilus" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="obtusus">
<emphasis id="72AB2874FFA3AA0472B0B867FB13FCE1" box="[1007,1231,809,834]" italics="true" pageId="4" pageNumber="25">Mexidrilus obtusus</emphasis>
</taxonomicName>
by
<bibRefCitation id="244E8997FFA3AA0475A5B867FF33FCC9" author="Erseus" pageId="4" pageNumber="38" refString="Erseus, C. (1992) A generic revision of the Phallodrilinae (Oligochaeta, Tubificidae). Zoologica Scripta, 21, 5 - 48." type="journal article" year="1992">Erséus (1992: Fig.13)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA3AA04719AB839FD7CFB1C" pageId="4" pageNumber="25" type="description">
<paragraph id="4060F466FFA3AA04719AB839FD6BFC7D" blockId="4.[151,1437,152,990]" pageId="4" pageNumber="25">
<taxonomicName id="87DF8FE5FFA3AA04719AB839FE4EFC33" box="[197,402,887,912]" class="Clitellata" family="Tubificidae" genus="Gianius" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="navarroi" status="sp. nov.">
<emphasis id="72AB2874FFA3AA04719AB839FE4EFC33" box="[197,402,887,912]" italics="true" pageId="4" pageNumber="25">Gianius navarroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA3AA0470C5B838FE02FC33" box="[410,478,886,912]" pageId="4" pageNumber="25" rank="species">
<emphasis id="72AB2874FFA3AA0470C5B838FE02FC33" bold="true" box="[410,478,886,912]" pageId="4" pageNumber="25">n. sp.</emphasis>
</taxonomicNameLabel>
is separated from the previously known species of the genus by the presence of oesophageal diverticula, salient round porophores with the penial chaetae, and marked difference in size of prostate glands (ental gland larger than distal).
</paragraph>
<paragraph id="4060F466FFA3AA0471C1BF79FEEAFBD7" blockId="4.[151,459,1079,1140]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA0471C1BF79FF7DFBD7" bold="true" pageId="4" pageNumber="25">
<taxonomicName id="87DF8FE5FFA3AA0471C1BF79FE5DFBF2" box="[158,385,1079,1105]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="25" phylum="Annelida" rank="species" species="gianii" status="sp. nov.">
<emphasis id="72AB2874FFA3AA0471C1BF79FE5DFBF2" bold="true" box="[158,385,1079,1105]" italics="true" pageId="4" pageNumber="25">Isochaetides gianii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA3AA0470D7BF79FE17FBF2" box="[392,459,1079,1105]" pageId="4" pageNumber="25" rank="species">n. sp.</taxonomicNameLabel>
(
</emphasis>
<figureCitation id="D8E4E8E3FFA3AA0471FEBF14FEF1FBD7" box="[161,301,1114,1140]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStart-2="FIGURE 5" captionStartId-0="5.[151,255,1602,1626]" captionStartId-1="7.[151,255,1802,1826]" captionStartId-2="8.[151,260,957,981]" captionTargetBox-0="[160,1429,820,1537]" captionTargetBox-1="[264,1345,182,1790]" captionTargetBox-2="[329,1255,217,909]" captionTargetId-0="figure@5.[151,1436,813,1578]" captionTargetId-1="figure@7.[250,1352,170,1795]" captionTargetId-2="figure@8.[319,1267,194,934]" captionTargetPageId-0="5" captionTargetPageId-1="7" captionTargetPageId-2="8" captionText-0="FIGURE 3. Isochaetides gianii n. sp. Somatic chaetae (A D: free-hand drawings at the same scale). A: anterior ventral; B: posterior ventral; C: anterior pectinate dorsal; D: posterior dorsal; E: anterior bundle of pectinate and hair chaetae; F: spermathecal chaeta. Scale bars: E and F: 25 µm." captionText-1="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." captionText-2="FIGURE 5. Isochaetides gianii n. sp. General diagram of the reproductive organs. X and XI: segment number. Abbreviations: a: atrium, ag: accessory gland, cps: cuticular penial sheath, o: ovary, pg: prostate gland, ps: penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sg: chaetal gland, t: testis, vd: vas deferens, vf: ventral fold of the body wall." httpUri-0="https://zenodo.org/record/193116/files/figure.png" httpUri-1="https://zenodo.org/record/193117/files/figure.png" httpUri-2="https://zenodo.org/record/193118/files/figure.png" pageId="4" pageNumber="25">Figures 35</figureCitation>
)
</paragraph>
<paragraph id="4060F466FFA3AA0471C8BFE6FD7CFB1C" blockId="4.[151,672,1192,1215]" box="[151,672,1192,1215]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA0471C8BFE6FECEFB1C" box="[151,274,1192,1215]" italics="true" pageId="4" pageNumber="25">Lamadrilus</emphasis>
sp. in
<bibRefCitation id="244E8997FFA3AA047006BFE6FD7CFB1C" author="Achurra" box="[345,672,1192,1215]" pageId="4" pageNumber="38" refString="Achurra, A. &amp; Rodriguez, P. (2008) Biodiversity of groundwater oligochaetes from a karst unit in northern Iberian Peninsula: ranking subterranean sites for conservation management. Hydrobiologia, 605, 159 - 171." type="journal article" year="2008">Achurra and Rodriguez (2008)</bibRefCitation>
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA3AA0471C8BFA0FA5FF885" pageId="4" pageNumber="25" type="materials_examined">
<paragraph id="4060F466FFA3AA0471C8BFA0FF2EFA8C" blockId="4.[151,1437,1262,2023]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA0471C8BFA0FECCFAAB" bold="true" box="[151,272,1262,1288]" pageId="4" pageNumber="25">
<typeStatus id="9F644AC4FFA3AA0471C8BFA0FED7FAAB" box="[151,267,1262,1288]" pageId="4" pageNumber="25" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="26CE6CA3FFA3AA047047BFA0FEACFAAB" LSID="urn:lsid:biocol.org:col:35275" box="[280,368,1262,1288]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="4" pageNumber="25">MNCN</collectionCode>
16.03/3053. A dissected worm, stained in Ehrlichs Haematoxylin and mounted in
<collectingCountry id="38C8B4F6FFA3AA04741BBFA0FA40FAAB" box="[1348,1436,1262,1288]" name="Canada" pageId="4" pageNumber="25">Canada</collectingCountry>
balsam.
</paragraph>
<paragraph id="4060F466FFA3AA04719ABE75FCC5FADF" blockId="4.[151,1437,1262,2023]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA04719ABE75FEAEFAF6" bold="true" box="[197,370,1339,1365]" pageId="4" pageNumber="25">
<typeStatus id="9F644AC4FFA3AA04719ABE75FED8FAF6" box="[197,260,1339,1365]" pageId="4" pageNumber="25">Type</typeStatus>
locality.
</emphasis>
Apraiz karstic spring (
<date id="3461D2A6FFA3AA0473D2BE72FCADFAF5" box="[653,881,1340,1366]" pageId="4" pageNumber="25" value="2005-09-06">6 September 2005</date>
), Santa EufemiaEreñozar karst unit, Biscay,
<collectingCountry id="38C8B4F6FFA3AA0471C8BE2CFF02FADF" box="[151,222,1378,1404]" name="Spain" pageId="4" pageNumber="25">Spain</collectingCountry>
. UTM coordinates X:524225, Y:4801350, Z: 22.
</paragraph>
<paragraph id="4060F466FFA3AA04719ABEC6FD9CF99D" blockId="4.[151,1437,1262,2023]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA04719ABEC6FE90FA01" bold="true" box="[197,332,1416,1442]" pageId="4" pageNumber="25">
<typeStatus id="9F644AC4FFA3AA04719ABEC6FE9BFA01" box="[197,327,1416,1442]" pageId="4" pageNumber="25" type="paratype">Paratypes</typeStatus>
.
</emphasis>
<collectionCode id="26CE6CA3FFA3AA04700BBEC7FE76FA00" LSID="urn:lsid:biocol.org:col:34871" box="[340,426,1417,1443]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="4" pageNumber="25">USNM</collectionCode>
2052660. A dissected specimen from the
<typeStatus id="9F644AC4FFA3AA0472F8BEC7FC06FA00" box="[935,986,1417,1443]" pageId="4" pageNumber="25">type</typeStatus>
locality (
<date id="3461D2A6FFA3AA047511BEC7FAF3FA00" box="[1102,1327,1417,1443]" pageId="4" pageNumber="25" value="2005-09-06">6 September 2005</date>
).
<collectionCode id="26CE6CA3FFA3AA04741CBEC7FA40FA00" LSID="urn:lsid:biocol.org:col:35275" box="[1347,1436,1417,1443]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:35275" name="Museo Nacional de Ciencias Naturales" pageId="4" pageNumber="25">MNCN</collectionCode>
16.03/3054: two whole-mounted specimens from Argatxa spring (
<date id="3461D2A6FFA3AA0472C4BEFEFB59FA69" box="[923,1157,1456,1482]" pageId="4" pageNumber="25" value="2005-09-30">30 September 2005</date>
). In the collection at the University of the Basque Country (UPV/EHU): a sectioned and a dissected specimen from the
<typeStatus id="9F644AC4FFA3AA0475B3BE98FAC3FA53" box="[1260,1311,1494,1520]" pageId="4" pageNumber="25">type</typeStatus>
locality (
<date id="3461D2A6FFA3AA0474D2BE98FEBCF9B4" pageId="4" pageNumber="25" value="2005-09-06">6 September 2005</date>
); a dissected specimen from Oxiña spring (
<date id="3461D2A6FFA3AA04722FBEB3FBE7F9B4" box="[880,1083,1533,1559]" pageId="4" pageNumber="25" value="1985-02-01">1 February 1985</date>
). All sites in Santa Eufemia Ereñozar karstic unit, Biscay,
<collectingCountry id="38C8B4F6FFA3AA0470A9BD6AFDE0F99D" box="[502,572,1572,1598]" name="Spain" pageId="4" pageNumber="25">Spain</collectingCountry>
.
</paragraph>
<paragraph id="4060F466FFA3AA04719ABD04FAE3F97B" blockId="4.[151,1437,1262,2023]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA04719ABD04FE57F9C7" bold="true" box="[197,395,1610,1636]" pageId="4" pageNumber="25">Other material.</emphasis>
Three specimens preserved in 70% alcohol from the
<typeStatus id="9F644AC4FFA3AA047554BD04FBE2F9C7" box="[1035,1086,1610,1636]" pageId="4" pageNumber="25">type</typeStatus>
locality (
<date id="3461D2A6FFA3AA0475EFBD04FA53F9C7" box="[1200,1423,1610,1636]" pageId="4" pageNumber="25" value="2005-09-06">6 September 2005</date>
). Two dissected and three specimens in 70% alcohol, from Argatxa spring (
<date id="3461D2A6FFA3AA047514BD3FFA97F928" box="[1099,1355,1649,1675]" pageId="4" pageNumber="25" value="2005-09-30">30 September 2005</date>
). Four sectioned, seven dissected, two whole-mounted and
<specimenCount id="56D93FEFFFA3AA047254BDD6FC71F911" box="[779,941,1688,1714]" pageId="4" pageNumber="25" type="generic">17 specimens</specimenCount>
preserved in 70% alcohol, from Aulestia spring (
<date id="3461D2A6FFA3AA0471AFBDF0FE1AF97B" box="[240,454,1726,1752]" pageId="4" pageNumber="25" value="1985-02-23">23 February 1985</date>
). All in the collection at the University of the Basque Country (UPV/EHU).
</paragraph>
<paragraph id="4060F466FFA3AA04719ABDABFA5FF885" blockId="4.[151,1437,1262,2023]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA04719ABDABFE50F95C" bold="true" box="[197,396,1765,1791]" pageId="4" pageNumber="25">Other localities.</emphasis>
Argatxa spring: UTM coordinates X:527790, Y:4800925, Z:2; Oxiña spring: X:529825, Y:4801300, Z:38; Aulestia spring: X: 536494, Y: 4794430, Z:66. All in Santa EufemiaEreñozar karst unit.
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA3AA04719ABC7CFDB6F839" pageId="4" pageNumber="25" type="etymology">
<paragraph id="4060F466FFA3AA04719ABC7CFDB6F839" blockId="4.[151,1437,1262,2023]" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA04719ABC7CFE8FF8EF" bold="true" box="[197,339,1842,1868]" pageId="4" pageNumber="25">Etymology.</emphasis>
Named after Dr. Narcisse Giani for his important contribution to the taxonomy and general biology of aquatic oligochaetes and also for his important role in the formation of many oligochaete taxonomists (including the first author).
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA3AA06719ABCE8FCD4F935" lastPageId="6" lastPageNumber="27" pageId="4" pageNumber="25" type="description">
<paragraph id="4060F466FFA3AA05719ABCE8FCC7FE63" blockId="4.[151,1437,1262,2023]" lastBlockId="5.[151,1436,152,758]" lastPageId="5" lastPageNumber="26" pageId="4" pageNumber="25">
<emphasis id="72AB2874FFA3AA04719ABCE8FE82F863" bold="true" box="[197,350,1958,1984]" pageId="4" pageNumber="25">Description.</emphasis>
Body length
<quantity id="87275983FFA3AA04735FBCE8FDA9F863" box="[512,629,1958,1984]" metricMagnitude="-3" metricUnit="m" metricValue="9.0" metricValueMax="10.0" metricValueMin="8.0" pageId="4" pageNumber="25" unit="mm" value="9.0" valueMax="10.0" valueMin="8.0">810 mm</quantity>
; segment number 5983; mean body diameter 260 µm (range: 180 335 µm) in segment V, and 324 µm (225390 µm) in segment VIII. Prostomium rounded or conical, 75125 µm long, width about the same or wider at its base (all measurements in fixed individuals). Body wall consisting of a thin cuticle (&lt;1 µm), smooth epidermis (510 µm thick, but up to 22 µm in the clitellum) and relatively poorly developed musculature (57 µm thick). There is no secondary annulation. Clitellum formed by a single layer of glandular cells, extended from anterior part of XI, back to the line of chaetae in segment XII; no glands in the region between the male pores. One pair of deep folds hiding spermathecal apertures (irregular in shape) in segment X, in line with ventral chaetae. One pair of male pores in segment XI, close to and slightly lateral to the ventral chaetae. In some specimens, the ventral part of segment XI appears deeply retracted by the action of dorso-ventral muscle strands.
</paragraph>
<paragraph id="4060F466FFA2AA05719ABA83FA9DFD55" blockId="5.[151,1436,152,758]" pageId="5" pageNumber="26">
Ventral chaetae 46 per bundle, reduced to two in posterior part of the body, 5075 μm long, with distal tooth thinner and shorter than proximal (
<figureCitation id="D8E4E8E3FFA2AA057331BABAFD6EFDAD" box="[622,690,500,526]" captionStart="FIGURE 3" captionStartId="5.[151,255,1602,1626]" captionTargetBox="[160,1429,820,1537]" captionTargetId="figure@5.[151,1436,813,1578]" captionTargetPageId="5" captionText="FIGURE 3. Isochaetides gianii n. sp. Somatic chaetae (A D: free-hand drawings at the same scale). A: anterior ventral; B: posterior ventral; C: anterior pectinate dorsal; D: posterior dorsal; E: anterior bundle of pectinate and hair chaetae; F: spermathecal chaeta. Scale bars: E and F: 25 µm." httpUri="https://zenodo.org/record/193116/files/figure.png" pageId="5" pageNumber="26">Fig. 3</figureCitation>
A, B). Hair and pectinate chaetae in dorsal bundles (
<figureCitation id="D8E4E8E3FFA2AA05744DBABAFA89FDAD" box="[1298,1365,500,526]" captionStart="FIGURE 3" captionStartId="5.[151,255,1602,1626]" captionTargetBox="[160,1429,820,1537]" captionTargetId="figure@5.[151,1436,813,1578]" captionTargetPageId="5" captionText="FIGURE 3. Isochaetides gianii n. sp. Somatic chaetae (A D: free-hand drawings at the same scale). A: anterior ventral; B: posterior ventral; C: anterior pectinate dorsal; D: posterior dorsal; E: anterior bundle of pectinate and hair chaetae; F: spermathecal chaeta. Scale bars: E and F: 25 µm." httpUri="https://zenodo.org/record/193116/files/figure.png" pageId="5" pageNumber="26">Fig. 3</figureCitation>
CE). Hair chaetae in preclitellar region 13 (seldom 4) per bundle, 162262 µm long; in postclitellar region 12 per bundle, 75125 µm long, gradually becoming shorter backwards. Pectinate chaetae in preclitellar segments 2 4 per bundle, 4790 µm long and with distal nodulus (at about ⅓ to ¼ from the distal end); 23 somewhat shorter chaetae, posteriorly. Teeth of pectinate chaetae about equally long in anterior segments, but distal tooth shorter and thinner than proximal in posterior segments. The number of intermediate teeth in pectinate chaetae 1 or (usually)
<quantity id="87275983FFA2AA0570C4B992FE14FD55" box="[411,456,732,758]" metricMagnitude="-2" metricUnit="m" metricValue="5.08" pageId="5" pageNumber="26" unit="in" value="2.0">2 in</quantity>
anterior dorsal chaetae; intermediate teeth are absent in posterior segments.
</paragraph>
<caption id="14A0A4EEFFA2AA0571C8BD0CFD77F93C" httpUri="https://zenodo.org/record/193116/files/figure.png" pageId="5" pageNumber="26" targetBox="[160,1429,820,1537]" targetPageId="5">
<paragraph id="4060F466FFA2AA0571C8BD0CFD77F93C" blockId="5.[151,1436,1602,1695]" pageId="5" pageNumber="26">
<emphasis id="72AB2874FFA2AA0571C8BD0CFEC5F9F9" bold="true" box="[151,281,1602,1626]" pageId="5" pageNumber="26">FIGURE 3.</emphasis>
<taxonomicName id="87DF8FE5FFA2AA057040BD0CFE35F9FA" box="[287,489,1602,1625]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="5" pageNumber="26" phylum="Annelida" rank="species" species="gianii" status="sp. nov.">
<emphasis id="72AB2874FFA2AA057040BD0CFE35F9FA" box="[287,489,1602,1625]" italics="true" pageId="5" pageNumber="26">Isochaetides gianii</emphasis>
</taxonomicName>
<emphasis id="72AB2874FFA2AA0570AFBD0CFDF0F9F9" bold="true" box="[496,556,1602,1626]" pageId="5" pageNumber="26">
<taxonomicNameLabel id="6998950FFFA2AA0570AFBD0CFDF0F9F9" box="[496,556,1602,1626]" pageId="5" pageNumber="26" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
Somatic chaetae (AD: free-hand drawings at the same scale). A: anterior ventral; B: posterior ventral; C: anterior pectinate dorsal; D: posterior dorsal; E: anterior bundle of pectinate and hair chaetae; F: spermathecal chaeta.
<emphasis id="72AB2874FFA2AA057023BDC9FE33F93C" bold="true" box="[380,495,1671,1695]" pageId="5" pageNumber="26">Scale bars</emphasis>
: E and F: 25 µm.
</paragraph>
</caption>
<paragraph id="4060F466FFA2AA05719ABD9FFC4AF89B" blockId="5.[151,1437,1745,2003]" pageId="5" pageNumber="26">
Modified, distally grooved spermathecal chaetae in segment X, one per bundle, 102145 µm long and 2 3 µm thick (
<figureCitation id="D8E4E8E3FFA2AA05707BBDB6FEB6F8B1" box="[292,362,1784,1810]" captionStart="FIGURE 3" captionStartId="5.[151,255,1602,1626]" captionTargetBox="[160,1429,820,1537]" captionTargetId="figure@5.[151,1436,813,1578]" captionTargetPageId="5" captionText="FIGURE 3. Isochaetides gianii n. sp. Somatic chaetae (A D: free-hand drawings at the same scale). A: anterior ventral; B: posterior ventral; C: anterior pectinate dorsal; D: posterior dorsal; E: anterior bundle of pectinate and hair chaetae; F: spermathecal chaeta. Scale bars: E and F: 25 µm." httpUri="https://zenodo.org/record/193116/files/figure.png" pageId="5" pageNumber="26">Fig. 3</figureCitation>
F). There are no modified penial chaetae; 12 somatic ventral chaetae per bundle, 5052 µm long, are found ventrally in segment XI, lateral to the male pores.
</paragraph>
<paragraph id="4060F466FFA2AA06719ABC0BFCC5FE85" blockId="5.[151,1437,1745,2003]" lastBlockId="6.[151,1437,152,1995]" lastPageId="6" lastPageNumber="27" pageId="5" pageNumber="26">
Brain extends back to 1/2. Coelomocytes absent or, if present, they are scarce, small, oval (5 µm maximum diameter), nucleated and granular. Commissural blood vessels connecting dorsal and ventral vessels are visible in most examined specimens; they form several loops in anterior segments, particularly in V (
<figureCitation id="D8E4E8E3FFA2AA0571E3BCF7FED8F870" box="[188,260,1977,2003]" captionStart="FIGURE 4" captionStartId="7.[151,255,1802,1826]" captionTargetBox="[264,1345,182,1790]" captionTargetId="figure@7.[250,1352,170,1795]" captionTargetPageId="7" captionText="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="5" pageNumber="26">Fig. 4</figureCitation>
A). Typical oligochaete dorsal, muscular pharyngeal pad not developed. In segments IIIII, pharynx with well-developed epithelium both dorsal and ventrally, and a large mass of glands dorsally (
<figureCitation id="D8E4E8E3FFA1AA06744EBBD6FA87FF11" box="[1297,1371,152,178]" captionStart="FIGURE 4" captionStartId="7.[151,255,1802,1826]" captionTargetBox="[264,1345,182,1790]" captionTargetId="figure@7.[250,1352,170,1795]" captionTargetPageId="7" captionText="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="6" pageNumber="27">Fig. 4</figureCitation>
B). A few clusters of pharingeal glands are located dorso-laterally to the digestive tract in segment IV and laterally in V. A layer of chloragogen cells on the gut extends from segment VI backwards. In segment VIII, oesophagus opens abruptly into the intestine (
<figureCitation id="D8E4E8E3FFA1AA0673F1BA42FD29FE85" box="[686,757,268,294]" captionStart="FIGURE 4" captionStartId="7.[151,255,1802,1826]" captionTargetBox="[264,1345,182,1790]" captionTargetId="figure@7.[250,1352,170,1795]" captionTargetPageId="7" captionText="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="6" pageNumber="27">Fig. 4</figureCitation>
C).
</paragraph>
<paragraph id="4060F466FFA1AA06719ABA7CFCDBFCE0" blockId="6.[151,1437,152,1995]" pageId="6" pageNumber="27">
One pair of spermathecae in segment X. A short spermathecal duct (3050 µm long, 3050 µm wide), with a thick epithelium, opens ventrally in segment X at the inner end of a deep epidermal fold (
<figureCitation id="D8E4E8E3FFA1AA067452BA17FA89FED0" box="[1293,1365,345,371]" captionStart="FIGURE 4" captionStartId="7.[151,255,1802,1826]" captionTargetBox="[264,1345,182,1790]" captionTargetId="figure@7.[250,1352,170,1795]" captionTargetPageId="7" captionText="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="6" pageNumber="27">Fig. 4</figureCitation>
D, E). Spherical to globular spermathecal ampulla (175220 µm maximum diameter), with a thin epithelium (37 µm high, although up to 20 µm close to the connection with the spermathecal duct). The ampulla usually contains 45 drop-like, elongate spermatozeugmata (125202 µm long, 3582 µm maximum diameter), which consist of an external clear matrix 715 µm thick around an axial cylinder of longitudinally arranged fertilizing sperm (
<figureCitation id="D8E4E8E3FFA1AA067025B954FE1BFD97" box="[378,455,538,564]" captionStart="FIGURE 4" captionStartId="7.[151,255,1802,1826]" captionTargetBox="[264,1345,182,1790]" captionTargetId="figure@7.[250,1352,170,1795]" captionTargetPageId="7" captionText="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="6" pageNumber="27">Fig. 4</figureCitation>
D). The spermathecal chaetae are orientated with their distal ends towards the spermathecal fold. The proximal end of the chaeta is bent, and associated with a chaetal gland (follicular cells) with several muscular strands; the distal end (
<emphasis id="72AB2874FFA1AA0673A0B926FCC7FD22" box="[767,795,616,641]" italics="true" pageId="6" pageNumber="27">ca</emphasis>
one third of its total length) is grooved and contained within a chaetophore (3250 µm diameter), which is internally glandular, radially arranged and externally surrounded by a muscular wall about 23 µm thick (
<figureCitation id="D8E4E8E3FFA1AA067274B9FBFCA5FD6C" box="[811,889,693,719]" captionStart="FIGURE 5" captionStartId="8.[151,260,957,981]" captionTargetBox="[329,1255,217,909]" captionTargetId="figure@8.[319,1267,194,934]" captionTargetPageId="8" captionText="FIGURE 5. Isochaetides gianii n. sp. General diagram of the reproductive organs. X and XI: segment number. Abbreviations: a: atrium, ag: accessory gland, cps: cuticular penial sheath, o: ovary, pg: prostate gland, ps: penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sg: chaetal gland, t: testis, vd: vas deferens, vf: ventral fold of the body wall." httpUri="https://zenodo.org/record/193118/files/figure.png" pageId="6" pageNumber="27">Fig. 5</figureCitation>
). Close to the chaetophore, and most likely functionally associated with it, 13 accessory glands can be observed, with histological structure similar to that of the prostate gland. The position of the chaetophore is usually anterior to the spermathecal fold, although in some cases it can be posterior to the fold.
</paragraph>
<paragraph id="4060F466FFA1AA06719AB81EFCC8F9EB" blockId="6.[151,1437,152,1995]" pageId="6" pageNumber="27">
One pair testes in X and one pair ovaries in XI. Sperm sacs may extend to IX anteriorly and to XII posteriorly. Egg sac can reach segment XIII. Sperm funnel 80127 µm high in lateral view (about one third the diameter of the body), attached to septum 10/11. Vas deferens ciliated throughout, two to three times longer than the atrium and penis, and restricted to segment XI, where it forms a few loops. It is up to 30 µm in diameter, but thinner in its proximal end (down to 13 µm diameter), and in some specimens also thinner just before its junction with the atrium (15 µm diameter). Atria in XI, tubular (250300 µm long), bent and expanded near proximal end (4260 µm, measured at the level of junction with the prostate), thinner in their distal part (3040 µm), and swelling before entering the penial sac (
<figureCitation id="D8E4E8E3FFA1AA06729CBF10FBD0FBDB" box="[963,1036,1118,1144]" captionStart="FIGURE 5" captionStartId="8.[151,260,957,981]" captionTargetBox="[329,1255,217,909]" captionTargetId="figure@8.[319,1267,194,934]" captionTargetPageId="8" captionText="FIGURE 5. Isochaetides gianii n. sp. General diagram of the reproductive organs. X and XI: segment number. Abbreviations: a: atrium, ag: accessory gland, cps: cuticular penial sheath, o: ovary, pg: prostate gland, ps: penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sg: chaetal gland, t: testis, vd: vas deferens, vf: ventral fold of the body wall." httpUri="https://zenodo.org/record/193118/files/figure.png" pageId="6" pageNumber="27">Fig. 5</figureCitation>
). Distal section of atrium may be bent, forming a loop before entering the penial sac. Two different
<typeStatus id="9F644AC4FFA1AA0672ECBFCBFC2EFB3C" box="[947,1010,1157,1183]" pageId="6" pageNumber="27">types</typeStatus>
of atrial cells, one forming a dense layer (up to 37 µm high) at the concave side of the atrium, and the other
<typeStatus id="9F644AC4FFA1AA067565BFE2FBACFB65" box="[1082,1136,1196,1222]" pageId="6" pageNumber="27">type</typeStatus>
with a clear granulated appearance forming a layer of cells (up of 20 µm high) along the convex side and the proximal part of the atrium. Atrial musculature very thin (&lt;2 µm thick) and atrial lumen narrow, 25 µm. A medium-size, compact prostate gland connects by a short stalk to the concave side, at the proximal expanded end of the atrium. Vas deferens enters subapically on the convex side of the atrium. Penial sac well developed, oval, with maximum diameter parallel to the longitudinal axis of the body (115145 µm), having a ring fold at its inner end. Penis cylindrical (65115 µm long and 3550 µm diameter) and formed by two epithelial layers (
<figureCitation id="D8E4E8E3FFA1AA0675B0BEDAFAE4FA0D" box="[1263,1336,1428,1454]" captionStart="FIGURE 4" captionStartId="7.[151,255,1802,1826]" captionTargetBox="[264,1345,182,1790]" captionTargetId="figure@7.[250,1352,170,1795]" captionTargetPageId="7" captionText="FIGURE 4. Isochaetides gianii n. sp. A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac. Abbreviations: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca. Scale bars: A D 100 µm, E H: 50 µm." httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="6" pageNumber="27">Fig. 4</figureCitation>
FH). A mushroom-shaped cuticular layer, consisting of an inner oval part (24 µm thick) lining both the penial sac and the proximal ring fold; and a central stalk (12 µm thick) forming a cuticular sheath around the penis. This penial sheath is slightly conical and encloses most of the penis, which is free inside it and can project partially outwards. Female funnels were not observed.
</paragraph>
<paragraph id="4060F466FFA1AA06719ABD1BFCD4F935" blockId="6.[151,1437,152,1995]" pageId="6" pageNumber="27">
<emphasis id="72AB2874FFA1AA06719ABD1BFE8DF9CC" bold="true" box="[197,337,1621,1647]" pageId="6" pageNumber="27">Teratology.</emphasis>
One specimen with two male ducts plus one spermatheca on one side, and one male duct, one spermatheca plus two chaetophores on the other side.
</paragraph>
</subSubSection>
<subSubSection id="08C5A7EDFFA1AA0A719ABDECFE93FB4F" lastPageId="10" lastPageNumber="31" pageId="6" pageNumber="27" type="distribution">
<paragraph id="4060F466FFA1AA06719ABDECFF29F893" blockId="6.[151,1437,152,1995]" pageId="6" pageNumber="27">
<emphasis id="72AB2874FFA1AA06719ABDECFE20F91F" bold="true" box="[197,508,1698,1724]" pageId="6" pageNumber="27">Distribution and habitat.</emphasis>
The new species was found in four karstic springs located in the karst of Santa Eufemia-Ereñozar, in sandy sediments. Water characteristics were for Argatxa spring: 13.7ºC, pH 7.2, 446 µS/ cm and 91 % dissolved oxygen; and for Apraiz spring: 14.8ºC, pH 7.0, 469 µS/cm and 70 % dissolved oxygen.
</paragraph>
<paragraph id="4060F466FFA1AA08719ABC73FC6FF9AB" blockId="6.[151,1437,152,1995]" lastBlockId="8.[151,1437,1131,2008]" lastPageId="8" lastPageNumber="29" pageId="6" pageNumber="27">
<emphasis id="72AB2874FFA1AA06719ABC73FE15F8F4" bold="true" box="[197,457,1853,1879]" pageId="6" pageNumber="27">Taxonomic remarks.</emphasis>
The position and general structure of the gonads, male ducts and spermathecae in the new species are characteristic of the oligochaetes of the subfamily
<taxonomicName id="87DF8FE5FFA1AA06728DBC2AFB80F8DD" box="[978,1116,1892,1918]" class="Clitellata" family="Tubificidae" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="27" phylum="Annelida" rank="subFamily" subFamily="Tubificinae">Tubificinae</taxonomicName>
. The new species is related to the tubificine genera
<taxonomicName id="87DF8FE5FFA1AA0670EABCC5FD3AF807" authority="Hrabe, 1966" authorityName="Hrabe" authorityYear="1966" box="[437,742,1928,1956]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="27" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFA1AA0670EABCC5FD96F807" box="[437,586,1931,1956]" italics="true" pageId="6" pageNumber="27">Isochaetides</emphasis>
<bibRefCitation id="244E8997FFA1AA06730DBCC6FD3AF807" author="Hrabe" box="[594,742,1928,1956]" pageId="6" pageNumber="39" refString="Hrabe, S. (1966) New or insufficiently known species of the family Tubificidae. Publications de la Faculte des Sciences de l'Universite J. E. Purkyn e, Brno, 470, 57 - 77." type="journal article" year="1966">Hrabë, 1966</bibRefCitation>
</taxonomicName>
and
<emphasis id="72AB2874FFA1AA06727DBCC5FC77F807" box="[802,939,1931,1956]" italics="true" pageId="6" pageNumber="27">Lamadrilus</emphasis>
<bibRefCitation id="244E8997FFA1AA0672ECBCC4FB9AF807" author="Timm" box="[947,1094,1930,1956]" pageId="6" pageNumber="39" refString="Timm, T. (1998) Tubificidae (Clitellata, Oligochaeta) from Lake Taimyr, North Siberia, with a description of two new species and Lamadrilus gen. nov. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Biologie, 68, 23 - 42." type="journal article" year="1998">Timm, 1998</bibRefCitation>
by the presence of a tubular atrium, a discrete prostate gland attached to atrium by a narrow stalk, a vas deferens longer than the atrium and ciliated throughout, a penis within a penial sac, and modified spermathecal chaetae. Other close genera with similar characteristics of the male duct are
<taxonomicName id="87DF8FE5FFAFAA08738DBFDCFCABFB08" box="[722,887,1170,1195]" class="Clitellata" family="Tubificidae" genus="Tasserkidrilus" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA08738DBFDCFCABFB08" box="[722,887,1170,1195]" italics="true" pageId="8" pageNumber="29">Tasserkidrilus</emphasis>
</taxonomicName>
,
<taxonomicName id="87DF8FE5FFAFAA0872D9BFDCFBD7FB08" box="[902,1035,1170,1195]" class="Clitellata" family="Randiellidae" genus="Arctodrilus" kingdom="Animalia" order="Enchytraeida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA0872D9BFDCFBD7FB08" box="[902,1035,1170,1195]" italics="true" pageId="8" pageNumber="29">Arctodrilus</emphasis>
</taxonomicName>
,
<taxonomicName id="87DF8FE5FFAFAA087544BFDCFB71FB08" box="[1051,1197,1170,1195]" class="Clitellata" family="Tubificidae" genus="Troglodrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA087544BFDCFB71FB08" box="[1051,1197,1170,1195]" italics="true" pageId="8" pageNumber="29">Troglodrilus</emphasis>
</taxonomicName>
and
<taxonomicName id="87DF8FE5FFAFAA0875B8BFDCFAEDFB08" box="[1255,1329,1170,1195]" class="Clitellata" family="Randiellidae" genus="Haber" kingdom="Animalia" order="Enchytraeida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA0875B8BFDCFAEDFB08" box="[1255,1329,1170,1195]" italics="true" pageId="8" pageNumber="29">Haber</emphasis>
</taxonomicName>
, the first three lacking modified genital setae, and the latter with both spermathecal and penial chaetae modified. The new species has modified spermathecal setae in X and no penial chaetae, as well as a general structure of the male duct which can be present in either
<taxonomicName id="87DF8FE5FFAFAA0873DFBE48FCC8FABC" box="[640,788,1286,1311]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA0873DFBE48FCC8FABC" box="[640,788,1286,1311]" italics="true" pageId="8" pageNumber="29">Isochaetides</emphasis>
</taxonomicName>
or
<emphasis id="72AB2874FFAFAA087263BE48FC18FABC" box="[828,964,1286,1311]" italics="true" pageId="8" pageNumber="29">Lamadrilus</emphasis>
. These include the vas deferens joining the atrium subapically, and the expanded proximal end of the atrium, which has two layers of glandular cells: a dark, densely granulated layer in the concave side and a light layer in the convex side. The new species is well distinguished from other species of
<taxonomicName id="87DF8FE5FFAFAA0873D0BE34FCF5FA30" box="[655,809,1402,1427]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA0873D0BE34FCF5FA30" box="[655,809,1402,1427]" italics="true" pageId="8" pageNumber="29">Isochaetides</emphasis>
</taxonomicName>
or
<emphasis id="72AB2874FFAFAA08720CBE34FC3CFA30" box="[851,992,1402,1427]" italics="true" pageId="8" pageNumber="29">Lamadrilus</emphasis>
by a combination of characters that include: hair chaetae and pectinates in dorsal bundles, grooved spermathecal chaetae associated with chaetophores and glands, a well-developed penial sac lined by a cuticle layer with a proximal ring fold, and a cuticular penial sheath (not asymmetrical or in the form of a spade).
</paragraph>
<caption id="14A0A4EEFFA0AA0771C8BC44FE5EF852" httpUri="https://zenodo.org/record/193117/files/figure.png" pageId="7" pageNumber="28" targetBox="[264,1345,182,1790]" targetPageId="7">
<paragraph id="4060F466FFA0AA0771C8BC44FE5EF852" blockId="7.[151,1437,1802,2033]" pageId="7" pageNumber="28">
<emphasis id="72AB2874FFA0AA0771C8BC44FEC5F881" bold="true" box="[151,281,1802,1826]" pageId="7" pageNumber="28">FIGURE 4.</emphasis>
<taxonomicName id="87DF8FE5FFA0AA07707EBC44FE31F882" box="[289,493,1802,1825]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="28" phylum="Annelida" rank="species" species="gianii" status="sp. nov.">
<emphasis id="72AB2874FFA0AA07707EBC44FE31F882" box="[289,493,1802,1825]" italics="true" pageId="7" pageNumber="28">Isochaetides gianii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFA0AA0770ACBC44FDECF881" box="[499,560,1802,1826]" pageId="7" pageNumber="28" rank="species">
<emphasis id="72AB2874FFA0AA0770ACBC44FDECF881" bold="true" box="[499,560,1802,1826]" pageId="7" pageNumber="28">n. sp.</emphasis>
</taxonomicNameLabel>
A: commissural blood vessels in body segments V and VI; B: anterior end of the body, arrow pointing to the glands on the pharynx, in segment III; C: sudden transition to the intestine, from segments VII to VIII; D: spermatheca; E: detail of spermatheca in a histological section; F: penial sac with penis; G: cross histological section of penial sac; H: longitudinal histological section of penial sac.
<emphasis id="72AB2874FFA0AA07754ABC3CFB6AF829" bold="true" box="[1045,1206,1906,1930]" pageId="7" pageNumber="28">Abbreviations</emphasis>
: ag: accessory gland, cps: cuticular penial sheath, i: intestine, p: penis, rf: ring fold of the penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sz: spermatozeugmata, vf: ventral fold of the spermatheca.
<emphasis id="72AB2874FFA0AA0775B9BCF9FA86F86C" bold="true" box="[1254,1370,1975,1999]" pageId="7" pageNumber="28">Scale bars</emphasis>
: AD 100 µm, EH: 50 µm.
</paragraph>
</caption>
<caption id="14A0A4EEFFAFAA0871C8B8F3FE2FFB9E" httpUri="https://zenodo.org/record/193118/files/figure.png" pageId="8" pageNumber="29" targetBox="[329,1255,217,909]" targetPageId="8">
<paragraph id="4060F466FFAFAA0871C8B8F3FE2FFB9E" blockId="8.[151,1438,957,1085]" pageId="8" pageNumber="29">
<emphasis id="72AB2874FFAFAA0871C8B8F3FEFDFC76" bold="true" box="[151,289,957,981]" pageId="8" pageNumber="29">FIGURE 5.</emphasis>
<taxonomicName id="87DF8FE5FFAFAA087075B8F0FDD8FC76" box="[298,516,958,981]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="species" species="gianii" status="sp. nov.">
<emphasis id="72AB2874FFAFAA087075B8F0FDD8FC76" box="[298,516,958,981]" italics="true" pageId="8" pageNumber="29">Isochaetides gianii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFAFAA087353B8F3FD92FC76" box="[524,590,957,981]" pageId="8" pageNumber="29" rank="species">
<emphasis id="72AB2874FFAFAA087353B8F3FD92FC76" bold="true" box="[524,590,957,981]" pageId="8" pageNumber="29">n. sp.</emphasis>
</taxonomicNameLabel>
General diagram of the reproductive organs. X and XI: segment number.
<emphasis id="72AB2874FFAFAA0871C8B8AEFE9EFC5B" bold="true" box="[151,322,992,1016]" pageId="8" pageNumber="29">Abbreviations:</emphasis>
a: atrium, ag: accessory gland, cps: cuticular penial sheath, o: ovary, pg: prostate gland, ps: penial sac, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, sg: chaetal gland, t: testis, vd: vas deferens, vf: ventral fold of the body wall.
</paragraph>
</caption>
<paragraph id="4060F466FFAFAA09719ABD5AFC4CFEEF" blockId="8.[151,1437,1131,2008]" lastBlockId="9.[151,1437,152,526]" lastPageId="9" lastPageNumber="30" pageId="8" pageNumber="29">
Due to the feeble differences between the genera
<taxonomicName id="87DF8FE5FFAFAA08724FBD5BFC7FF98D" box="[784,931,1557,1582]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA08724FBD5BFC7FF98D" box="[784,931,1557,1582]" italics="true" pageId="8" pageNumber="29">Isochaetides</emphasis>
</taxonomicName>
and
<emphasis id="72AB2874FFAFAA087282BD5BFBB9F98D" box="[989,1125,1557,1582]" italics="true" pageId="8" pageNumber="29">Lamadrilus</emphasis>
, the new species was first identified as
<emphasis id="72AB2874FFAFAA087072BD75FE68F9F7" box="[301,436,1595,1620]" italics="true" pageId="8" pageNumber="29">Lamadrilus</emphasis>
sp. (
<bibRefCitation id="244E8997FFAFAA0870B3BD75FCEFF9F6" author="Achurra" box="[492,819,1595,1621]" pageId="8" pageNumber="38" refString="Achurra, A. &amp; Rodriguez, P. (2008) Biodiversity of groundwater oligochaetes from a karst unit in northern Iberian Peninsula: ranking subterranean sites for conservation management. Hydrobiologia, 605, 159 - 171." type="journal article" year="2008">Achurra &amp; Rodriguez 2008</bibRefCitation>
), mainly based on the presence of hair chaetae. Most species in this genus have only bifid somatic chaetae, except
<taxonomicName id="87DF8FE5FFAFAA08722BBD2CFA9BF9DF" authority="Chekanovskaya, 1975" authorityName="Chekanovskaya" authorityYear="1975" box="[884,1351,1634,1660]" class="Clitellata" family="Naididae" genus="Lophochaeta" higherTaxonomySource="CoL" kingdom="Animalia" order="Tubificida" pageId="8" pageNumber="29" phylum="Annelida" rank="species" species="bazikalovae">
<emphasis id="72AB2874FFAFAA08722BBD2CFBFDF9D8" box="[884,1057,1634,1659]" italics="true" pageId="8" pageNumber="29">L. bazikalovae</emphasis>
(
<bibRefCitation id="244E8997FFAFAA08756DBD2CFAE2F9DF" author="Chekanovskaya" box="[1074,1342,1634,1660]" pageId="8" pageNumber="38" refString="Chekanovskaya, O. V. (1975) New tubificids (Oligochaeta, Tubificidae) from the abyssal zone of Lake Baikal. Trudy Baikal'skoi Limnologeskoi Stantsii, 18, 112 - 130." type="journal article" year="1975">Chekanovskaya, 1975</bibRefCitation>
)
</taxonomicName>
, which presents different rates of reduction in hair chaetae and pectinates (
<bibRefCitation id="244E8997FFAFAA087295BDC7FB82F900" author="Timm" box="[970,1118,1673,1699]" pageId="8" pageNumber="39" refString="Timm, T. (1998) Tubificidae (Clitellata, Oligochaeta) from Lake Taimyr, North Siberia, with a description of two new species and Lamadrilus gen. nov. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Biologie, 68, 23 - 42." type="journal article" year="1998">Timm 1998</bibRefCitation>
). However, we agree with
<bibRefCitation id="244E8997FFAFAA0871C8BDE1FDFDF96A" author="Brinkhurst" box="[151,545,1711,1737]" pageId="8" pageNumber="38" refString="Brinkhurst, R. O. &amp; Kathman, R. D. (1983) Varichaetadrilus, a new name for Va ri cha eta Brinkhurst, 1981, non Speiser, 1903, (Diptera) with a description of a new species V. fulleri. Proceedings of the Biological Society of Washington, 96, 301 - 306." type="journal article" year="1983">Brinkhurst and Kathman (1983)</bibRefCitation>
that the absence of species with hair chaetae in a genus should not be a problem for the inclusion of a new species with hair chaetae; and
<taxonomicName id="87DF8FE5FFAFAA0872EDBD98FBCEF94C" box="[946,1042,1750,1775]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="species" species="gianii" status="sp. nov.">
<emphasis id="72AB2874FFAFAA0872EDBD98FBCEF94C" box="[946,1042,1750,1775]" italics="true" pageId="8" pageNumber="29">I. gianii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFAFAA087544BD98FB83F953" box="[1051,1119,1750,1776]" pageId="8" pageNumber="29" rank="species">
<emphasis id="72AB2874FFAFAA087544BD98FB83F953" bold="true" box="[1051,1119,1750,1776]" pageId="8" pageNumber="29">n. sp.</emphasis>
</taxonomicNameLabel>
has been classified on the basis of the structure of the reproductive organs. The genus
<taxonomicName id="87DF8FE5FFAFAA087234BDB3FB41F8B5" authority="Hrabe, 1966" authorityName="Hrabe" authorityYear="1966" box="[875,1181,1786,1814]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAFAA087234BDB3FBDCF8B5" box="[875,1024,1789,1814]" italics="true" pageId="8" pageNumber="29">Isochaetides</emphasis>
<bibRefCitation id="244E8997FFAFAA087557BDB4FB41F8B5" author="Hrabe" box="[1032,1181,1786,1814]" pageId="8" pageNumber="39" refString="Hrabe, S. (1966) New or insufficiently known species of the family Tubificidae. Publications de la Faculte des Sciences de l'Universite J. E. Purkyn e, Brno, 470, 57 - 77." type="journal article" year="1966">Hrabë, 1966</bibRefCitation>
</taxonomicName>
includes species that possess modified spermathecal chaetae (except
<taxonomicName id="87DF8FE5FFAFAA087384BC6DFBE6F89E" authority="Semernoy, 2001" authorityName="Semernoy" authorityYear="2001" box="[731,1082,1827,1853]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="species" species="distinctus">
<emphasis id="72AB2874FFAFAA087384BC6DFCB5F89F" box="[731,873,1827,1852]" italics="true" pageId="8" pageNumber="29">I. distinctus</emphasis>
<bibRefCitation id="244E8997FFAFAA08722EBC6DFBE6F89E" author="Semernoy" box="[881,1082,1827,1853]" pageId="8" pageNumber="39" refString="Semernoy, V. P. (2001) Annelida: Oligochaeta and Aelosomatidae. In: Timoshkin, O. (Ed.), Index of Animal Species Inhabiting Lake Baikal and Its Catchment Area. Novosibirsk, Nauka, pp. 377 - 427." type="book chapter" year="2001">Semernoy, 2001</bibRefCitation>
</taxonomicName>
). These are characteristically grooved in the distal end (except for
<taxonomicName id="87DF8FE5FFAFAA087306BC04FC0DF8C7" authority="Lastockin, 1937" authorityName="Lastockin" authorityYear="1937" box="[601,977,1864,1892]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="species" species="michaelseni">
<emphasis id="72AB2874FFAFAA087306BC04FCDFF8C0" box="[601,771,1866,1891]" italics="true" pageId="8" pageNumber="29">I. michaelseni</emphasis>
<bibRefCitation id="244E8997FFAFAA087253BC06FC0DF8C7" author="Lastockin" box="[780,977,1864,1892]" pageId="8" pageNumber="39" refString="Lastockin, D. A. (1937) New species of Oligochaeta Limicola in the European part of the USSR. Doklady Akademii Nauk SSSR, 17, 233 - 235." type="journal article" year="1937">Lastočkin, 1937</bibRefCitation>
</taxonomicName>
, which has bifid genital chaetae), and generally appear associated with spermathecal glands. In several species, these glands are poorly described, but their remarkable distal position with respect to the spermathecal chaeta, and their structure (sometimes only seen in drawings), is similar to the chaetophores that we have described in
<taxonomicName id="87DF8FE5FFAFAA087515BCF0FB7BF874" box="[1098,1191,1982,2007]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="29" phylum="Annelida" rank="species" species="gianii" status="sp. nov.">
<emphasis id="72AB2874FFAFAA087515BCF0FB7BF874" box="[1098,1191,1982,2007]" italics="true" pageId="8" pageNumber="29">I. gianii</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6998950FFFAFAA0875EFBCF0FB2FF87B" box="[1200,1267,1982,2008]" pageId="8" pageNumber="29" rank="species">
<emphasis id="72AB2874FFAFAA0875EFBCF0FB2FF87B" bold="true" box="[1200,1267,1982,2008]" pageId="8" pageNumber="29">n. sp.</emphasis>
</taxonomicNameLabel>
Chaetophores associated with spermathecal chaetae are also known in other oligochaete genera [e.g. the rhyacodriline
<taxonomicName id="87DF8FE5FFAEAA0971C8BBF1FCB7FF7B" authority="Giani &amp; Martinez-Ansemil, 1979" authorityName="Giani &amp; Martinez-Ansemil" authorityYear="1979" box="[151,875,190,216]" class="Clitellata" family="Tubificidae" genus="Protuberodrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="tourenqui">
<emphasis id="72AB2874FFAEAA0971C8BBF1FE0DFF7B" box="[151,465,191,216]" italics="true" pageId="9" pageNumber="30">Protuberodrilus tourenqui</emphasis>
<bibRefCitation id="244E8997FFAEAA097085BBF0FCB7FF7B" author="Giani" box="[474,875,190,216]" pageId="9" pageNumber="38" refString="Giani, N. &amp; Martinez-Ansemil, E. (1979) Description d'une nouvelle espece de Tubificidae du Nord-Ouest de l'Espagne: Protuberodrilus tourenqui n. g., n. sp. Annales de Limnologie - International Journal of Limnology, 15, 291 - 297." type="journal article" year="1979">Giani &amp; Martínez-Ansemil, 1979</bibRefCitation>
</taxonomicName>
and the telmatodriline
<taxonomicName id="87DF8FE5FFAEAA0975DBBBF1FE58FF5C" authority="Sokolskaya, 1961" authorityName="Sokolskaya" authorityYear="1961" class="Clitellata" family="Tubificidae" genus="Alexandrovia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="ringulata">
<emphasis id="72AB2874FFAEAA0975DBBBF1FA40FF7B" box="[1156,1436,191,216]" italics="true" pageId="9" pageNumber="30">Alexandrovia ringulata</emphasis>
(
<bibRefCitation id="244E8997FFAEAA0971C0BBABFEA7FF5C" author="Sokolskaya" box="[159,379,229,255]" pageId="9" pageNumber="39" refString="Sokolskaya, N. L. (1961) Materialy po faune presnovodnych maloscetinkovych cervej Kamcatki. Bjulleten' Moskovskogo obscestva ispytatelej prirody, otdel biologii, 66, 54 - 68." type="journal article" year="1961">Sokolskaya, 1961</bibRefCitation>
)
</taxonomicName>
]. On the other hand, only one of the six known species of
<emphasis id="72AB2874FFAEAA09751DBBABFB17FF5D" box="[1090,1227,229,254]" italics="true" pageId="9" pageNumber="30">Lamadrilus</emphasis>
(
<taxonomicName id="87DF8FE5FFAEAA097582BBABFF0FFE85" authority="Timm, 1998" authorityName="Timm" authorityYear="1998" class="Clitellata" family="Naididae" genus="Lophochaeta" higherTaxonomySource="CoL" kingdom="Animalia" order="Tubificida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="sorosi">
<emphasis id="72AB2874FFAEAA097582BBABFA99FF5D" box="[1245,1349,229,254]" italics="true" pageId="9" pageNumber="30">L. sorosi</emphasis>
<bibRefCitation id="244E8997FFAEAA097412BBABFF0FFE85" author="Timm" pageId="9" pageNumber="39" refString="Timm, T. (1998) Tubificidae (Clitellata, Oligochaeta) from Lake Taimyr, North Siberia, with a description of two new species and Lamadrilus gen. nov. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Biologie, 68, 23 - 42." type="journal article" year="1998">Timm, 1998</bibRefCitation>
</taxonomicName>
) has modified spermathecal chaetae of the same
<typeStatus id="9F644AC4FFAEAA097251BA42FC9DFE85" box="[782,833,268,294]" pageId="9" pageNumber="30">type</typeStatus>
as that described in
<taxonomicName id="87DF8FE5FFAEAA097571BA42FB1DFE86" box="[1070,1217,268,293]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAEAA097571BA42FB1DFE86" box="[1070,1217,268,293]" italics="true" pageId="9" pageNumber="30">Isochaetides</emphasis>
</taxonomicName>
species, as well as a chaetophore and accessory glands associated to the chaetal sac.
</paragraph>
<paragraph id="4060F466FFAEAA09719ABA17FADCFDAD" blockId="9.[151,1437,152,526]" pageId="9" pageNumber="30">
The two different epithelial layers of the atrium described in the new species are probably equivalent to the dark and light layers described in
<taxonomicName id="87DF8FE5FFAEAA09730DBACEFD65FE3A" box="[594,697,384,409]" class="Clitellata" family="Naididae" genus="Lophochaeta" higherTaxonomySource="CoL" kingdom="Animalia" order="Tubificida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="sorosi">
<emphasis id="72AB2874FFAEAA09730DBACEFD65FE3A" box="[594,697,384,409]" italics="true" pageId="9" pageNumber="30">L. sorosi</emphasis>
</taxonomicName>
by
<bibRefCitation id="244E8997FFAEAA0973BABACEFC58FE39" author="Timm" box="[741,900,384,410]" pageId="9" pageNumber="39" refString="Timm, T. (1998) Tubificidae (Clitellata, Oligochaeta) from Lake Taimyr, North Siberia, with a description of two new species and Lamadrilus gen. nov. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Biologie, 68, 23 - 42." type="journal article" year="1998">Timm (1998)</bibRefCitation>
, and are also reminiscent of the layers shown in drawings of some
<taxonomicName id="87DF8FE5FFAEAA0970FABAE9FD98FE63" box="[421,580,423,448]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAEAA0970FABAE9FD98FE63" box="[421,580,423,448]" italics="true" pageId="9" pageNumber="30">Isochaetides</emphasis>
</taxonomicName>
species. This histological characteristic has been confirmed in the examination of
<typeStatus id="9F644AC4FFAEAA097001BA83FE48FE44" box="[350,404,461,487]" pageId="9" pageNumber="30">type</typeStatus>
specimens of
<taxonomicName id="87DF8FE5FFAEAA097314BA83FC48FE44" authority="Brinkhurst, 1965" authorityName="Brinkhurst" authorityYear="1965" box="[587,916,461,487]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="freyi">
<emphasis id="72AB2874FFAEAA097314BA83FD7CFE45" box="[587,672,461,486]" italics="true" pageId="9" pageNumber="30">I. freyi</emphasis>
(
<bibRefCitation id="244E8997FFAEAA0973EDBA83FC50FE44" author="Brinkhurst" box="[690,908,461,487]" pageId="9" pageNumber="38" refString="Brinkhurst, R. O. (1965) Studies on the North American aquatic Oligochaeta II: Tubificidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 117, 117 - 172." type="journal article" year="1965">Brinkhurst, 1965</bibRefCitation>
)
</taxonomicName>
(
<figureCitation id="D8E4E8E3FFAEAA0972F8BA83FC2FFE44" box="[935,1011,461,487]" captionStart="FIGURE 6" captionStartId="9.[151,255,1722,1746]" captionTargetBox="[155,1425,593,1680]" captionTargetId="figure@9.[151,1436,581,1698]" captionTargetPageId="9" captionText="FIGURE 6. Isochaetides freyi (Brinkhurst, 1965). A: histological section of the atrium; B: penial sac with penis lined by cuticle; C: ventral fold of the body wall, where the spermathecal duct opens; D: spermatheca. Abbreviation s: a: atrium, c: chaetophore, cps: cuticular penial sheath, p: penis, pg: prostate gland, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, vd: vas deferens, vf: ventral fold of the body wall. Scale bars: 100 µm." httpUri="https://zenodo.org/record/193119/files/figure.png" pageId="9" pageNumber="30">Fig. 6</figureCitation>
A), although it has probably been overlooked in most descriptions, not only in species of these genera but also in other tubificines.
</paragraph>
<caption id="14A0A4EEFFAEAA0971C8BDF4FB09F89A" httpUri="https://zenodo.org/record/193119/files/figure.png" pageId="9" pageNumber="30" targetBox="[155,1425,593,1680]" targetPageId="9">
<paragraph id="4060F466FFAEAA0971C8BDF4FB09F89A" blockId="9.[151,1437,1722,1850]" pageId="9" pageNumber="30">
<emphasis id="72AB2874FFAEAA0971C8BDF4FEC5F971" bold="true" box="[151,281,1722,1746]" pageId="9" pageNumber="30">FIGURE 6.</emphasis>
<taxonomicName id="87DF8FE5FFAEAA097040BDF4FD76F972" authority="Brinkhurst, 1965" authorityName="Brinkhurst" authorityYear="1965" box="[287,682,1722,1745]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="freyi">
<emphasis id="72AB2874FFAEAA097040BDF4FE07F972" box="[287,475,1722,1745]" italics="true" pageId="9" pageNumber="30">Isochaetides freyi</emphasis>
(Brinkhurst, 1965)
</taxonomicName>
. A: histological section of the atrium; B: penial sac with penis lined by cuticle; C: ventral fold of the body wall, where the spermathecal duct opens; D: spermatheca.
<emphasis id="72AB2874FFAEAA0975D9BD93FAC0F956" bold="true" box="[1158,1308,1757,1781]" pageId="9" pageNumber="30">Abbreviation</emphasis>
s: a: atrium, c: chaetophore, cps: cuticular penial sheath, p: penis, pg: prostate gland, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, vd: vas deferens, vf: ventral fold of the body wall.
<emphasis id="72AB2874FFAEAA0972A6BC6CFBB0F899" bold="true" box="[1017,1132,1826,1850]" pageId="9" pageNumber="30">Scale bars</emphasis>
: 100 µm.
</paragraph>
</caption>
<paragraph id="4060F466FFAEAA0A719ABC22FEA9FDAD" blockId="9.[151,1436,1900,2003]" lastBlockId="10.[151,1437,152,1260]" lastPageId="10" lastPageNumber="31" pageId="9" pageNumber="30">
The presence of a penial sac with an apical ring fold is known in other
<taxonomicName id="87DF8FE5FFAEAA09754FBC22FB7FF826" box="[1040,1187,1900,1925]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFAEAA09754FBC22FB7FF826" box="[1040,1187,1900,1925]" italics="true" pageId="9" pageNumber="30">Isochaetides</emphasis>
</taxonomicName>
species (e.g.
<taxonomicName id="87DF8FE5FFAEAA09741BBC22FA4DF826" box="[1348,1425,1900,1925]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="freyi">
<emphasis id="72AB2874FFAEAA09741BBC22FA4DF826" box="[1348,1425,1900,1925]" italics="true" pageId="9" pageNumber="30">I. freyi</emphasis>
</taxonomicName>
) (
<figureCitation id="D8E4E8E3FFAEAA0971FFBCDCFF34F80F" box="[160,232,1938,1964]" captionStart="FIGURE 6" captionStartId="9.[151,255,1722,1746]" captionTargetBox="[155,1425,593,1680]" captionTargetId="figure@9.[151,1436,581,1698]" captionTargetPageId="9" captionText="FIGURE 6. Isochaetides freyi (Brinkhurst, 1965). A: histological section of the atrium; B: penial sac with penis lined by cuticle; C: ventral fold of the body wall, where the spermathecal duct opens; D: spermatheca. Abbreviation s: a: atrium, c: chaetophore, cps: cuticular penial sheath, p: penis, pg: prostate gland, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, vd: vas deferens, vf: ventral fold of the body wall. Scale bars: 100 µm." httpUri="https://zenodo.org/record/193119/files/figure.png" pageId="9" pageNumber="30">Fig. 6</figureCitation>
B). Several lateral folds are present in the penial sac of some
<emphasis id="72AB2874FFAEAA097296BCDDFB8EF80F" box="[969,1106,1939,1964]" italics="true" pageId="9" pageNumber="30">Lamadrilus</emphasis>
species (e.g.
<taxonomicName id="87DF8FE5FFAEAA0975AABCDDFA81F80F" box="[1269,1373,1939,1964]" class="Clitellata" family="Naididae" genus="Lophochaeta" higherTaxonomySource="CoL" kingdom="Animalia" order="Tubificida" pageId="9" pageNumber="30" phylum="Annelida" rank="species" species="sorosi">
<emphasis id="72AB2874FFAEAA0975AABCDDFA81F80F" box="[1269,1373,1939,1964]" italics="true" pageId="9" pageNumber="30">L. sorosi</emphasis>
</taxonomicName>
), but never in the form of an ental ring fold. The presence of a short, mostly asymmetrical cuticular penial sheath (sometimes with a spade-shaped distal appendage) is a diagnostic character in
<emphasis id="72AB2874FFADAA0A753EBBD6FB31FF12" box="[1121,1261,152,177]" italics="true" pageId="10" pageNumber="31">Lamadrilus</emphasis>
; however, the genus incorporates different
<typeStatus id="9F644AC4FFADAA0A70AEBBF0FDECFF7B" box="[497,560,190,216]" pageId="10" pageNumber="31">types</typeStatus>
of penial sheaths (sometimes variable in shape within the same species, with a more or less marked asymmetry, see
<bibRefCitation id="244E8997FFADAA0A7389BBABFCA5FF5C" author="Timm" box="[726,889,229,255]" pageId="10" pageNumber="39" refString="Timm, T. (1998) Tubificidae (Clitellata, Oligochaeta) from Lake Taimyr, North Siberia, with a description of two new species and Lamadrilus gen. nov. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Biologie, 68, 23 - 42." type="journal article" year="1998">Timm, 1998</bibRefCitation>
: Figs. 4550). The diagnosis of the genus
<taxonomicName id="87DF8FE5FFADAA0A71C8BA42FEF1FE86" box="[151,301,268,293]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFADAA0A71C8BA42FEF1FE86" box="[151,301,268,293]" italics="true" pageId="10" pageNumber="31">Isochaetides</emphasis>
</taxonomicName>
by
<bibRefCitation id="244E8997FFADAA0A7004BA42FD81FE85" author="Snimschikova" box="[347,605,268,294]" pageId="10" pageNumber="39" refString="Snimschikova, L. N. (1998) Revision of the genus Lycodrilus Grube and description of the evolution of Limnodrilus Claparede (Oligochaeta, Tubificidae) species in Lake Baikal. 2. Revision of the species Lycodrilus grubei with discussion of the current taxonomic position of Limnodrilus species from Lake Baikal and their evolution. Zoologicheskii Zhurnal, 77, 639 - 647." type="journal article" year="1998">Snimschikova (1998)</bibRefCitation>
describes the cuticular penial sheath as thin or absent and
<bibRefCitation id="244E8997FFADAA0A747CBA42FF39FEEF" author="Semernoy" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy (2004)</bibRefCitation>
describes it as more or less thin, either of uniform width or conical but without oblique cut on its distal end. In fact, all
<taxonomicName id="87DF8FE5FFADAA0A700EBA17FE38FED1" box="[337,484,345,370]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFADAA0A700EBA17FE38FED1" box="[337,484,345,370]" italics="true" pageId="10" pageNumber="31">Isochaetides</emphasis>
</taxonomicName>
species known to date, except for
<taxonomicName id="87DF8FE5FFADAA0A7223BA17FC26FED1" box="[892,1018,345,370]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="lacustris">
<emphasis id="72AB2874FFADAA0A7223BA17FC26FED1" box="[892,1018,345,370]" italics="true" pageId="10" pageNumber="31">I. lacustris</emphasis>
</taxonomicName>
, have a thin cuticular penial sheath or a thin cuticular covering on the penis (although thick in
<taxonomicName id="87DF8FE5FFADAA0A7212BACEFB7EFE39" authority="Semernoy, 2004" authorityName="Semernoy" authorityYear="2004" box="[845,1186,384,410]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="distinctus">
<emphasis id="72AB2874FFADAA0A7212BACEFC0AFE3A" box="[845,982,384,409]" italics="true" pageId="10" pageNumber="31">I. distinctus</emphasis>
<bibRefCitation id="244E8997FFADAA0A7281BACEFB7EFE39" author="Semernoy" box="[990,1186,384,410]" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy, 2004</bibRefCitation>
</taxonomicName>
). The examination of the
<typeStatus id="9F644AC4FFADAA0A719DBAE8FF29FE63" box="[194,245,422,448]" pageId="10" pageNumber="31">type</typeStatus>
specimens of
<taxonomicName id="87DF8FE5FFADAA0A70C4BAE9FE3BFE63" box="[411,487,423,448]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="freyi">
<emphasis id="72AB2874FFADAA0A70C4BAE9FE3BFE63" box="[411,487,423,448]" italics="true" pageId="10" pageNumber="31">I. freyi</emphasis>
</taxonomicName>
have revealed a conspicuous cuticular layer, covering the inner side of the penial sac, the large ental fold and the penis, and taking the shape of their contour (
<figureCitation id="D8E4E8E3FFADAA0A7544BA83FBBEFE44" box="[1051,1122,461,487]" captionStart="FIGURE 6" captionStartId="9.[151,255,1722,1746]" captionTargetBox="[155,1425,593,1680]" captionTargetId="figure@9.[151,1436,581,1698]" captionTargetPageId="9" captionText="FIGURE 6. Isochaetides freyi (Brinkhurst, 1965). A: histological section of the atrium; B: penial sac with penis lined by cuticle; C: ventral fold of the body wall, where the spermathecal duct opens; D: spermatheca. Abbreviation s: a: atrium, c: chaetophore, cps: cuticular penial sheath, p: penis, pg: prostate gland, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, vd: vas deferens, vf: ventral fold of the body wall. Scale bars: 100 µm." httpUri="https://zenodo.org/record/193119/files/figure.png" pageId="10" pageNumber="31">Fig. 6</figureCitation>
B), in a similar way to that in the new species.
</paragraph>
<paragraph id="4060F466FFADAA0A719AB954FEECFBA7" blockId="10.[151,1437,152,1260]" pageId="10" pageNumber="31">
The structure of the spermatheca is not commonly used as a diagnostic character in tubificines. However, in
<taxonomicName id="87DF8FE5FFADAA0A71E7B90FFE8FFDF9" box="[184,339,577,602]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFADAA0A71E7B90FFE8FFDF9" box="[184,339,577,602]" italics="true" pageId="10" pageNumber="31">Isochaetides</emphasis>
</taxonomicName>
the short spermathecal ducts, associated with modified spermathecal chaetae located in adjacent glandular sacs (referred to as chaetophores in present work), appear to be a very common feature. The structure of the spermathecal duct is contentious and what is described here as a short spermathecal duct opening at the ental end of a ventral body fold, appears to be interpreted as a short ental sphincter of the spermathecal duct by
<bibRefCitation id="244E8997FFADAA0A70C8B992FDBFFD55" author="Semernoy" box="[407,611,732,758]" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy (2004)</bibRefCitation>
in
<taxonomicName id="87DF8FE5FFADAA0A73D6B992FC34FD55" authority="Semernoy, 2004" authorityName="Semernoy" authorityYear="2004" box="[649,1000,732,758]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="peniacerus">
<emphasis id="72AB2874FFADAA0A73D6B992FCFFFD56" box="[649,803,732,757]" italics="true" pageId="10" pageNumber="31">I. peniacerus</emphasis>
<bibRefCitation id="244E8997FFADAA0A7276B992FC34FD55" author="Semernoy" box="[809,1000,732,758]" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy, 2004</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="87DF8FE5FFADAA0A72A9B992FA8EFD55" authority="Semernoy, 2004" authorityName="Semernoy" authorityYear="2004" box="[1014,1362,732,758]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="compactus">
<emphasis id="72AB2874FFADAA0A72A9B992FB50FD56" box="[1014,1164,732,757]" italics="true" pageId="10" pageNumber="31">I. compactus</emphasis>
<bibRefCitation id="244E8997FFADAA0A75CBB992FA8EFD55" author="Semernoy" box="[1172,1362,732,758]" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy, 2004</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="87DF8FE5FFADAA0A74D4B992FE1DFCBF" authority="Semernoy, 2004" authorityName="Semernoy" authorityYear="2004" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="septatus">
<emphasis id="72AB2874FFADAA0A74D4B992FF2BFCB8" italics="true" pageId="10" pageNumber="31">I. septatus</emphasis>
<bibRefCitation id="244E8997FFADAA0A71A1B84CFE1DFCBF" author="Semernoy" box="[254,449,770,796]" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy, 2004</bibRefCitation>
</taxonomicName>
. The histology of the duct in the new species reveals that the cells are not muscular, and the irregular aperture at the body surface does not have the structure of a pore, typically formed by cells arranged in a circle around the opening. Short spermathecal ducts, opening into ventral folds of the body wall, associated with grooved spermathecal chaetae and chaetophores, are also present in
<taxonomicName id="87DF8FE5FFADAA0A75FCB839FB29FC33" box="[1187,1269,887,912]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="freyi">
<emphasis id="72AB2874FFADAA0A75FCB839FB29FC33" box="[1187,1269,887,912]" italics="true" pageId="10" pageNumber="31">I. freyi</emphasis>
</taxonomicName>
(
<figureCitation id="D8E4E8E3FFADAA0A7458B838FA8DFC33" box="[1287,1361,886,912]" captionStart="FIGURE 6" captionStartId="9.[151,255,1722,1746]" captionTargetBox="[155,1425,593,1680]" captionTargetId="figure@9.[151,1436,581,1698]" captionTargetPageId="9" captionText="FIGURE 6. Isochaetides freyi (Brinkhurst, 1965). A: histological section of the atrium; B: penial sac with penis lined by cuticle; C: ventral fold of the body wall, where the spermathecal duct opens; D: spermatheca. Abbreviation s: a: atrium, c: chaetophore, cps: cuticular penial sheath, p: penis, pg: prostate gland, sa: spermathecal ampulla, sch: spermathecal chaeta, sd: spermathecal duct, vd: vas deferens, vf: ventral fold of the body wall. Scale bars: 100 µm." httpUri="https://zenodo.org/record/193119/files/figure.png" pageId="10" pageNumber="31">Fig. 6</figureCitation>
C, D). These characters are shared by most
<taxonomicName id="87DF8FE5FFADAA0A7308B8D3FD32FC15" box="[599,750,925,950]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFADAA0A7308B8D3FD32FC15" box="[599,750,925,950]" italics="true" pageId="10" pageNumber="31">Isochaetides</emphasis>
</taxonomicName>
species, and we have considered them together with the general structure of the male duct and penis, for the classification of the new species as a member of the genus
<taxonomicName id="87DF8FE5FFADAA0A71C8B8A4FEF6FBA0" box="[151,298,1002,1027]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFADAA0A71C8B8A4FEF6FBA0" box="[151,298,1002,1027]" italics="true" pageId="10" pageNumber="31">Isochaetides</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="4060F466FFADAA0A719ABF5FFE93FB4F" blockId="10.[151,1437,152,1260]" pageId="10" pageNumber="31">
The finding of an
<taxonomicName id="87DF8FE5FFADAA0A70C3BF5FFDF3FB89" box="[412,559,1041,1066]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="genus">
<emphasis id="72AB2874FFADAA0A70C3BF5FFDF3FB89" box="[412,559,1041,1066]" italics="true" pageId="10" pageNumber="31">Isochaetides</emphasis>
</taxonomicName>
species in karstic areas in northern Iberian Peninsula is noteworthy, since the genus is represented mainly in eastern Europe (Timm &amp; Veldhuijzen van
<bibRefCitation id="244E8997FFADAA0A7569BF76FB12FBF1" author="Zanten" box="[1078,1230,1080,1106]" pageId="10" pageNumber="39" refString="Timm, T. &amp; Veldhuijzen van Zanten, H. H. (2002) Freshwater Oligochaeta of North-West Europe. World Biodiversity Database, CD-ROM Series. Expert Center for Taxonomic Identification, University of Amsterdam" type="book" year="2002">Zanten 2002</bibRefCitation>
) and Asia (many species in Lake Baikal, see
<bibRefCitation id="244E8997FFADAA0A7080BF10FD42FBDB" author="Semernoy" box="[479,670,1118,1144]" pageId="10" pageNumber="39" refString="Semernoy, V. P. (2004) Oligochaeta of Lake Baikal. Novosibirsk, Nauka, 528 pp." type="book" year="2004">Semernoy 2004</bibRefCitation>
), although three species have been described in
<collectingCountry id="38C8B4F6FFADAA0A7587BF10FA9EFBDB" box="[1240,1346,1118,1144]" name="United States of America" pageId="10" pageNumber="31">America</collectingCountry>
:
<taxonomicName id="87DF8FE5FFADAA0A7411BF11FEABFB3C" authority="Brinkhurst, 1965" authorityName="Brinkhurst" authorityYear="1965" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="freyi">
<emphasis id="72AB2874FFADAA0A7411BF11FA40FBDB" box="[1358,1436,1119,1144]" italics="true" pageId="10" pageNumber="31">I. freyi</emphasis>
(
<bibRefCitation id="244E8997FFADAA0A71C0BFCBFEB1FB3C" author="Brinkhurst" box="[159,365,1157,1183]" pageId="10" pageNumber="38" refString="Brinkhurst, R. O. (1965) Studies on the North American aquatic Oligochaeta II: Tubificidae. Proceedings of the Academy of Natural Sciences of Philadelphia, 117, 117 - 172." type="journal article" year="1965">Brinkhurst, 1965</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="87DF8FE5FFADAA0A70EDBFCBFC72FB3C" authority="Brinkhurst &amp; Cook, 1966" authorityName="Brinkhurst &amp; Cook" authorityYear="1966" box="[434,942,1157,1183]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="curvisetosus">
<emphasis id="72AB2874FFADAA0A70EDBFCBFD83FB3D" box="[434,607,1157,1182]" italics="true" pageId="10" pageNumber="31">I. curvisetosus</emphasis>
(
<bibRefCitation id="244E8997FFADAA0A7330BFCBFC7AFB3C" author="Brinkhurst" box="[623,934,1157,1183]" pageId="10" pageNumber="38" refString="Brinkhurst, R. O. &amp; Cook, D. G. (1966) Studies on the North American aquatic Oligochaeta III: Lumbriculidae and additional notes and records of other families. Proceedings of the Academy of Natural Sciences of Philadelphia, 118, 1 - 33." type="journal article" year="1966">Brinkhurst &amp; Cook, 1966</bibRefCitation>
)
</taxonomicName>
[considered as a probably synonym of
<taxonomicName id="87DF8FE5FFADAA0A74D4BFCBFE6BFB65" authority="Moore, 1905" authorityName="Moore" authorityYear="1905" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="hamatus">
<emphasis id="72AB2874FFADAA0A74D4BFCBFF21FB66" italics="true" pageId="10" pageNumber="31">I. hamatus</emphasis>
(
<bibRefCitation id="244E8997FFADAA0A7051BFE2FE72FB65" author="Moore" box="[270,430,1196,1222]" pageId="10" pageNumber="39" refString="Moore, J. P. (1905) Some marine Oligochaeta of New England. Proceedings of the Academy of Natural Sciences of Philadelphia, 57, 373 - 399." type="journal article" year="1905">Moore, 1905</bibRefCitation>
)
</taxonomicName>
after
<bibRefCitation id="244E8997FFADAA0A735FBFE2FD0DFB65" author="Brinkhurst" box="[512,721,1196,1222]" pageId="10" pageNumber="38" refString="Brinkhurst, R. O. (1986) Guide to the Freshwater Aquatic Microdrile Oligochaetes of North America. Canadian Special Publication of Fisheries and Aquatic Sciences, Ottawa, 259 pp." type="book" year="1986">Brinkhurst, 1986</bibRefCitation>
] in North
<collectingCountry id="38C8B4F6FFADAA0A720FBFE2FC60FB65" box="[848,956,1196,1222]" name="United States of America" pageId="10" pageNumber="31">America</collectingCountry>
; and
<taxonomicName id="87DF8FE5FFADAA0A72A2BFE2FAA0FB65" authority="Cernosvitov, 1939" authorityName="Cernosvitov" authorityYear="1939" box="[1021,1404,1189,1222]" class="Clitellata" family="Tubificidae" genus="Isochaetides" kingdom="Animalia" order="Haplotaxida" pageId="10" pageNumber="31" phylum="Annelida" rank="species" species="lacustris">
<emphasis id="72AB2874FFADAA0A72A2BFE2FB5DFB66" box="[1021,1153,1196,1221]" italics="true" pageId="10" pageNumber="31">I. lacustris</emphasis>
(Černosvitov, 1939)
</taxonomicName>
in South
<collectingCountry id="38C8B4F6FFADAA0A71BCBF9CFE90FB4F" box="[227,332,1234,1260]" name="United States of America" pageId="10" pageNumber="31">America</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>