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<document id="7854DA2C3660FB7DC6C23A4E2B1DC0CA" ID-CLB-Dataset="29912" ID-DOI="10.11646/zootaxa.4433.3.10" ID-GBIF-Dataset="b57a5098-dc9f-4615-a3df-c9f4f5aee313" ID-ISSN="1175-5326" ID-Zenodo-Dep="1290333" ID-ZooBank="4E1F06CF-00B8-47E0-BD40-FA52B9C59F03" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1529055623786" checkinUser="plazi" docAuthor="Kvist, Sebastian &amp; Erséus, Christer" docDate="2018" docId="03DA87C59D57FFD164826381FA60FD76" docLanguage="en" docName="zootaxa.4433.3.10.pdf" docOrigin="Zootaxa 4433 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Tubificoides mackiei Kvist &amp; Erséus 2018, n. sp." docType="treatment" docVersion="7" lastPageNumber="569" masterDocId="FFE3FFBD9D53FFD964156575FFE8FFF2" masterDocTitle="Two new European species of the marine genus Tubificoides (Annelida: Clitellata: Naididae) with notes on the morphology of T. pseudogaster (Dahl, 1960)" masterLastPageNumber="573" masterPageNumber="561" pageNumber="565" updateTime="1698548329325" updateUser="ExternalLinkService">
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<mods:title id="F6D66E71B4DB5D9C8389D0EA7B95A632">Two new European species of the marine genus Tubificoides (Annelida: Clitellata: Naididae) with notes on the morphology of T. pseudogaster (Dahl, 1960)</mods:title>
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<mods:namePart id="73DC930F45BE88D8F4F4C8AAAB9F38BD">Kvist, Sebastian</mods:namePart>
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<mods:namePart id="9A17AFF8AB091A5913075F8E64AC7C52">Erséus, Christer</mods:namePart>
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<mods:date id="6948C37B4B02A35ACEA877931E7326B4">2018</mods:date>
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<treatment id="03DA87C59D57FFD164826381FA60FD76" ID-DOI="http://doi.org/10.5281/zenodo.5998486" ID-GBIF-Taxon="145952759" ID-Zenodo-Dep="5998486" LSID="urn:lsid:plazi:treatment:03DA87C59D57FFD164826381FA60FD76" httpUri="http://treatment.plazi.org/id/03DA87C59D57FFD164826381FA60FD76" lastPageId="8" lastPageNumber="569" pageId="4" pageNumber="565">
<subSubSection id="C36965589D57FFDD64826381FEF4F8DD" pageId="4" pageNumber="565" type="nomenclature">
<paragraph id="8BCC36D39D57FFDD64826381FE33F8FC" blockId="4.[151,475,1780,1839]" box="[151,475,1780,1806]" pageId="4" pageNumber="565">
<heading id="D08481BF9D57FFDD64826381FE33F8FC" bold="true" box="[151,475,1780,1806]" fontSize="11" level="1" pageId="4" pageNumber="565" reason="1">
<emphasis id="B907EAC19D57FFDD64826381FE33F8FC" bold="true" box="[151,475,1780,1806]" pageId="4" pageNumber="565">
<taxonomicName id="4C734D509D57FFDD64826381FE79F8FC" authority="Kvist &amp; Erséus, 2018" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[151,401,1780,1806]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="565" phylum="Annelida" rank="species" species="mackiei" status="sp. nov.">
<emphasis id="B907EAC19D57FFDD64826381FE79F8FC" bold="true" box="[151,401,1780,1806]" italics="true" pageId="4" pageNumber="565">Tubificoides mackiei</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23457BA9D57FFDD658D6381FE33F8FC" box="[408,475,1780,1806]" pageId="4" pageNumber="565" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BCC36D39D57FFDD64826263FEF4F8DD" blockId="4.[151,475,1780,1839]" box="[151,284,1814,1839]" pageId="4" pageNumber="565">
<figureCitation id="13482A569D57FFDD64826263FEF4F8DD" box="[151,284,1814,1839]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStartId-0="6.[151,250,1377,1399]" captionStartId-1="7.[151,250,1839,1861]" captionTargetBox-0="[151,1436,193,1356]" captionTargetBox-1="[219,1263,592,1814]" captionTargetId-0="figure@6.[151,1436,193,1356]" captionTargetId-1="figure@7.[202,1413,556,1824]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionText-0="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." captionText-1="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri-0="https://zenodo.org/record/1290339/files/figure.png" httpUri-1="https://zenodo.org/record/1290341/files/figure.png" pageId="4" pageNumber="565">Figures 34</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="C36965589D57FFDD6482622BFD92F840" pageId="4" pageNumber="565" type="description">
<paragraph id="8BCC36D39D57FFDD6482622BFCE1F886" blockId="4.[151,777,1886,1970]" box="[151,777,1886,1908]" pageId="4" pageNumber="565">
<taxonomicName id="4C734D509D57FFDD6482622BFE48F886" box="[151,416,1886,1908]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="565" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D57FFDD6482622BFE48F886" box="[151,416,1886,1908]" italics="true" pageId="4" pageNumber="565">Tubificoides pseudogaster</emphasis>
</taxonomicName>
;
<bibRefCitation id="EFE24B229D57FFDD65B8622BFD59F886" box="[429,689,1886,1908]" pageId="4" pageNumber="565" refId="ref7065" refString="Brinkhurst, R. O. &amp; Baker, H. R. (1979) A review of the marine Tubificidae (Oligochaeta) of North America. Canadian Journal of Zoology, 57 (8), 1553 - 1569. https: // doi. org / 10.1139 / z 79 - 203" type="journal article">Brinkhurst &amp; Baker 1979</bibRefCitation>
, partim?
</paragraph>
<paragraph id="8BCC36D39D57FFDD64826208FD44F861" blockId="4.[151,777,1886,1970]" box="[151,684,1917,1939]" pageId="4" pageNumber="565">
<taxonomicName id="4C734D509D57FFDD64826208FE48F861" box="[151,416,1917,1939]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="565" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D57FFDD64826208FE48F861" box="[151,416,1917,1939]" italics="true" pageId="4" pageNumber="565">Tubificoides pseudogaster</emphasis>
</taxonomicName>
;
<bibRefCitation id="EFE24B229D57FFDD65B86208FDBCF861" box="[429,596,1917,1939]" pageId="4" pageNumber="565" refId="ref7012" refString="Brinkhurst, R. O. (1986) Taxonomy of the genus Tubificoides Lastockin (Oligochaeta, Tubificidae): species with bifid setae. Canadian Journal of Zoology, 64 (6), 1270 - 1279. https: // doi. org / 10.1139 / z 86 - 189" type="journal article">Brinkhurst 1986</bibRefCitation>
, partim?
</paragraph>
<paragraph id="8BCC36D39D57FFDD648262E9FD92F840" blockId="4.[151,777,1886,1970]" box="[151,634,1948,1970]" pageId="4" pageNumber="565">
<taxonomicName id="4C734D509D57FFDD648262E9FEFBF840" authorityName="Lastockin" authorityYear="1937" box="[151,275,1948,1970]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="565" phylum="Annelida" rank="genus">
<emphasis id="B907EAC19D57FFDD648262E9FEFBF840" box="[151,275,1948,1970]" italics="true" pageId="4" pageNumber="565">Tubificoides</emphasis>
</taxonomicName>
&quot;
<taxonomicName id="4C734D509D57FFDD653662E9FE42F840" box="[291,426,1948,1970]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="4" pageNumber="565" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D57FFDD653662E9FE42F840" box="[291,426,1948,1970]" italics="true" pageId="4" pageNumber="565">pseudogaster</emphasis>
</taxonomicName>
I&quot;;
<bibRefCitation id="EFE24B229D57FFDD65C462E9FD92F840" box="[465,634,1948,1970]" pageId="4" pageNumber="565" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">
Kvist
<emphasis id="B907EAC19D57FFDD661B62E8FDA9F840" box="[526,577,1948,1970]" italics="true" pageId="4" pageNumber="565">et al.</emphasis>
2010
</bibRefCitation>
</paragraph>
</subSubSection>
<subSubSection id="C36965589D57FFDC648262ACFACEFE96" lastPageId="5" lastPageNumber="566" pageId="4" pageNumber="565" type="diagnosis">
<paragraph id="8BCC36D39D57FFDC648262ACFACEFE96" blockId="4.[151,1436,2009,2034]" lastBlockId="5.[151,1437,151,2038]" lastPageId="5" lastPageNumber="566" pageId="4" pageNumber="565">
<emphasis id="B907EAC19D57FFDD648262ACFEF9F800" bold="true" box="[151,273,2009,2034]" pageId="4" pageNumber="565">Diagnosis.</emphasis>
This species can be distinguished from congeners on the basis of the lack of hair chaetae and cuticular papillation, and the detailed morphology of the cuticular penis sheaths. In comparison with its closest known relative,
<emphasis id="B907EAC19D56FFDC64EC65C8FE46FF27" box="[249,430,188,213]" italics="true" pageId="5" pageNumber="566">
<taxonomicName id="4C734D509D56FFDC64EC65C8FE42FF27" box="[249,426,188,213]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="5" pageNumber="566" phylum="Annelida" rank="species" species="pseudogaster">T. pseudogaster</taxonomicName>
,
</emphasis>
the new species is distinguished by its wider body at segment XI, its wider ental portions of the atria, its longer penis sheath, its lack of an outward-turned collar at the ental end of the penial tube, and its lower maximal number of preclitellar chaetae. In addition, the average COI distance between
<taxonomicName id="4C734D509D56FFDC60A66470FACEFEEF" authority="Kvist &amp; Erséus, 2018" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[1203,1318,260,285]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="5" pageNumber="566" phylum="Annelida" rank="species" species="mackiei" status="sp. nov.">
<emphasis id="B907EAC19D56FFDC60A66470FACEFEEF" box="[1203,1318,260,285]" italics="true" pageId="5" pageNumber="566">T. mackiei</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23457BA9D56FFDC61386471FA84FEEF" box="[1325,1388,260,285]" pageId="5" pageNumber="566" rank="species">
<emphasis id="B907EAC19D56FFDC61386471FA84FEEF" bold="true" box="[1325,1388,260,285]" pageId="5" pageNumber="566">n. sp.</emphasis>
</taxonomicNameLabel>
and
<taxonomicName id="4C734D509D56FFDC6482645CFEA4FEB2" box="[151,332,295,320]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="5" pageNumber="566" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D56FFDC6482645CFEA4FEB2" box="[151,332,295,320]" italics="true" pageId="5" pageNumber="566">T. pseudogaster</emphasis>
</taxonomicName>
is 22.07% (see
<bibRefCitation id="EFE24B229D56FFDC66066452FD37FEB2" box="[531,735,295,320]" pageId="5" pageNumber="566" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">
Kvist
<emphasis id="B907EAC19D56FFDC6649645CFD72FEB2" box="[604,666,295,320]" italics="true" pageId="5" pageNumber="566">et al.</emphasis>
2010
</bibRefCitation>
) and a total of 127 characteristic attributes (
<emphasis id="B907EAC19D56FFDC60E3645CFAF1FEB2" box="[1270,1305,297,320]" italics="true" pageId="5" pageNumber="566">i.e.</emphasis>
, molecular synapomorphies; see
<bibRefCitation id="EFE24B229D56FFDC659D6439FDB9FE96" box="[392,593,332,357]" pageId="5" pageNumber="566" refId="ref9493" refString="Sarkar, I. N., Planet, P. J., Bael, T. E., Stanley, S. E., Siddall, M., DeSalle, R. &amp; Figurski, D. H. (2002) Characteristic attributes in cancer microarrays. Journal of Biomedical Informatics, 35 (2), 111 - 122. https: // doi. org / 10.1016 / S 1532 - 0464 (02) 00504 - X" type="journal article">
Sarkar
<emphasis id="B907EAC19D56FFDC65CD6438FDF8FE97" box="[472,528,332,357]" italics="true" pageId="5" pageNumber="566">et al.</emphasis>
2002
</bibRefCitation>
) separate the COI sequences of these species (
<bibRefCitation id="EFE24B229D56FFDC604C6439FAF2FE96" box="[1113,1306,332,357]" pageId="5" pageNumber="566" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">
Kvist
<emphasis id="B907EAC19D56FFDC608B6438FB3FFE97" box="[1182,1239,332,357]" italics="true" pageId="5" pageNumber="566">et al.</emphasis>
2010
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C36965589D56FFDC64D2641AFAF3FC72" pageId="5" pageNumber="566" type="materials_examined">
<paragraph id="8BCC36D39D56FFDC64D2641AFE07FE22" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
<emphasis id="B907EAC19D56FFDC64D2641AFED3FE7A" bold="true" box="[199,315,367,392]" pageId="5" pageNumber="566">Holotype.</emphasis>
ZMBN 123768 (specimen ID CE21703), stained, whole-mount on microscope slide with coverslip, sexually mature specimen, collected by C. Erséus and M. Klinth,
<date id="FFCD10139D56FFDC676D64E0FBF0FE5E" box="[888,1048,404,429]" pageId="5" pageNumber="566" value="2014-05-12">May 12, 2014</date>
; holotype COI sequence GenBank accession number
<accessionNumber id="9420AB309D56FFDC657364C2FE03FE22" box="[358,491,439,464]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MG652362" pageId="5" pageNumber="566">MG652362</accessionNumber>
.
</paragraph>
<paragraph id="8BCC36D39D56FFDC64D264A9FBE7FDEA" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
<materialsCitation id="3B1B3C8E9D56FFDC64D264A9FBE7FDEA" ID-GBIF-Occurrence="1849038037" country="Norway" elevation="4" latitude="58.86472" location="Gunnarsholmen Island" longLatPrecision="17" longitude="9.411666" municipality="Kragero" pageId="5" pageNumber="566" specimenCount="1" stateProvince="Telemark" typeStatus="holotype">
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<typeStatus id="54C888719D56FFDC64D264A9FEEAFE07" box="[199,258,476,501]" pageId="5" pageNumber="566">Type</typeStatus>
locality.
</emphasis>
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(inside of island, close to marina),
<collectingMunicipality id="6BA8ACA99D56FFDC605A64A8FB45FE07" box="[1103,1197,477,501]" pageId="5" pageNumber="566">Kragerö</collectingMunicipality>
,
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,
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(
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,
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),
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depth, anaerobic, coarse sand and gravel.
</materialsCitation>
</paragraph>
<paragraph id="8BCC36D39D56FFDC64D26751FB57FD5A" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
<emphasis id="B907EAC19D56FFDC64D26751FEAEFDCF" bold="true" box="[199,326,548,573]" pageId="5" pageNumber="566">Paratypes.</emphasis>
ZMBN 123769-123771 (specimen IDs CE21701, CE21702, CE21704); SMNH Type coll. 8955 8957 (specimen IDs CE21705CE21707), stained, whole-mounts on microscope slide with coverslip, all specimens fully mature (except CE21707), collected by C. Erséus and M. Klinth,
<date id="FFCD10139D56FFDC60336718FB2CFD76" box="[1062,1220,620,645]" pageId="5" pageNumber="566" value="2014-05-12">May 12, 2014</date>
, from type locality. Paratype COI sequence GenBank accession numbers
<accessionNumber id="9420AB309D56FFDC66FA67FAFC9BFD5A" box="[751,883,655,680]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MG652360" pageId="5" pageNumber="566">MG652360</accessionNumber>
,
<accessionNumber id="9420AB309D56FFDC676B67FAFC17FD5A" box="[894,1023,655,680]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MG652361" pageId="5" pageNumber="566">MG652361</accessionNumber>
and
<accessionNumber id="9420AB309D56FFDC602367FAFB53FD5A" box="[1078,1211,655,680]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/MG652363" pageId="5" pageNumber="566">MG652363</accessionNumber>
.
</paragraph>
<paragraph id="8BCC36D39D56FFDC64D267C1FAF3FC72" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
<emphasis id="B907EAC19D56FFDC64D267C1FDEBFD3F" bold="true" box="[199,515,692,717]" pageId="5" pageNumber="566">Other material examined.</emphasis>
<collectionCode id="ED62AE169D56FFDC661B67C1FDB7FD3E" box="[526,607,692,716]" country="Canada" httpUri="http://grbio.org/cool/8z6v-f8is" name="Saskatchewan Museum of Natural History" pageId="5" pageNumber="566">SMNH</collectionCode>
108985,
<collectionCode id="ED62AE169D56FFDC66DA67C1FCC8FD3E" box="[719,800,692,716]" country="Canada" httpUri="http://grbio.org/cool/8z6v-f8is" name="Saskatchewan Museum of Natural History" pageId="5" pageNumber="566">SMNH</collectionCode>
108986, and
<collectionCode id="ED62AE169D56FFDC67D167C1FBFDFD3E" box="[964,1045,692,716]" country="Canada" httpUri="http://grbio.org/cool/8z6v-f8is" name="Saskatchewan Museum of Natural History" pageId="5" pageNumber="566">SMNH</collectionCode>
108988 (specimen IDs CE199-3, CE2077, CE
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<bibRefCitation id="EFE24B229D56FFDC658567A2FDB4FD02" box="[400,604,727,752]" pageId="5" pageNumber="566" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">Kvist et al. 2010</bibRefCitation>
), stained, whole-mounts on microscope slide with coverslip, one fully mature, one half-mature and one immature specimen, collected at Tjärnö Marine Biological Laboratory, Bohuslän,
<collectingCountry id="F36476439D56FFDC6482666AFF18FCCA" box="[151,240,799,824]" name="Sweden" pageId="5" pageNumber="566">Sweden</collectingCountry>
(
<geoCoordinate id="EE4750149D56FFDC64EA666AFE67FCCA" box="[255,399,799,824]" direction="north" orientation="latitude" pageId="5" pageNumber="566" precision="15" value="58.87472">58°52' 29&quot;N</geoCoordinate>
,
<geoCoordinate id="EE4750149D56FFDC658E6655FDCBFCCA" box="[411,547,800,824]" direction="east" orientation="longitude" pageId="5" pageNumber="566" precision="15" value="11.1455555">11°08' 44&quot;E</geoCoordinate>
) by C. Erséus,
<date id="FFCD10139D56FFDC66C76655FCAEFCCA" box="[722,838,800,824]" pageId="5" pageNumber="566" value="2000-09">Sep. 2000</date>
,
<date id="FFCD10139D56FFDC67446655FC2BFCCA" box="[849,963,799,824]" pageId="5" pageNumber="566" value="2006-10">Oct. 2006</date>
,
<date id="FFCD10139D56FFDC67DA6655FBABFCCA" box="[975,1091,800,824]" pageId="5" pageNumber="566" value="2007-09">Sep. 2007</date>
;
<collectionCode id="ED62AE169D56FFDC605A6655FB48FCCA" box="[1103,1184,800,824]" country="Canada" httpUri="http://grbio.org/cool/8z6v-f8is" name="Saskatchewan Museum of Natural History" pageId="5" pageNumber="566">SMNH</collectionCode>
108987 (specimen ID CE
<quantity id="4C8B9B369D56FFDC64AE6631FEFAFCAF" box="[187,274,836,861]" metricMagnitude="1" metricUnit="m" metricValue="7.89178" pageId="5" pageNumber="566" unit="in" value="3107.0">3107 in</quantity>
<bibRefCitation id="EFE24B229D56FFDC650F6631FE35FCAE" box="[282,477,836,861]" pageId="5" pageNumber="566" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">Kvist et al. 2010</bibRefCitation>
), half mature specimen, collected at Hanhalsholme,
<collectingRegion id="49B7F8319D56FFDC60206631FB79FCAF" box="[1077,1169,836,861]" country="Sweden" name="Halland" pageId="5" pageNumber="566">Halland</collectingRegion>
,
<collectingCountry id="F36476439D56FFDC60886631FB1EFCAF" box="[1181,1270,836,861]" name="Sweden" pageId="5" pageNumber="566">Sweden</collectingCountry>
(
<geoCoordinate id="EE4750149D56FFDC61126631FA70FCAF" box="[1287,1432,836,861]" direction="north" orientation="latitude" pageId="5" pageNumber="566" precision="15" value="57.449444">57°26' 58&quot;N</geoCoordinate>
,
<geoCoordinate id="EE4750149D56FFDC6482661DFEC8FC72" box="[151,288,872,896]" direction="east" orientation="longitude" pageId="5" pageNumber="566" precision="15" value="12.067223">12°04' 02&quot;E</geoCoordinate>
) by S. Kvist and M. Lindström,
<date id="FFCD10139D56FFDC6699661DFD17FC72" box="[652,767,872,896]" pageId="5" pageNumber="566" value="2007-09">Sep. 2007</date>
; for genetic information, see
<bibRefCitation id="EFE24B229D56FFDC605D6612FAFEFC72" box="[1096,1302,871,896]" pageId="5" pageNumber="566" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">Kvist et al. (2010)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C36965589D56FFDC64D266F9FE0EFBE2" pageId="5" pageNumber="566" type="etymology">
<paragraph id="8BCC36D39D56FFDC64D266F9FE0EFBE2" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
<emphasis id="B907EAC19D56FFDC64D266F9FEA6FC57" bold="true" box="[199,334,908,933]" pageId="5" pageNumber="566">Etymology.</emphasis>
Named after the esteemed polychaete systematist, Andrew (Andy) S.Y. Mackie, for his contributions to annelid systematics, and for his hospitality, generosity and aid during the first authors collection of numerous naidids (from various localities in
<collectingCountry id="F36476439D56FFDC66BD66A1FD04FC1F" box="[680,748,980,1005]" name="United Kingdom" pageId="5" pageNumber="566">Wales</collectingCountry>
in 2007), including
<emphasis id="B907EAC19D56FFDC67DB66A0FB6BFC1F" box="[974,1155,980,1005]" italics="true" pageId="5" pageNumber="566">
<taxonomicName id="4C734D509D56FFDC67DB66A0FB96FC1F" box="[974,1150,980,1005]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="5" pageNumber="566" phylum="Annelida" rank="species" species="pseudogaster">T. pseudogaster</taxonomicName>
,
</emphasis>
which partly enabled the separation of the new species.
</paragraph>
</subSubSection>
<subSubSection id="C36965589D56FFDC64D26169FC0DF807" pageId="5" pageNumber="566" type="description">
<paragraph id="8BCC36D39D56FFDC64D26169FAADFB1A" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
<emphasis id="B907EAC19D56FFDC64D26169FEB1FBC7" bold="true" box="[199,345,1052,1077]" pageId="5" pageNumber="566">Description.</emphasis>
All specimens incomplete, posterior ends used for DNA extraction.
<typeStatus id="54C888719D56FFDC60466169FB53FBC7" box="[1107,1211,1052,1077]" pageId="5" pageNumber="566" type="holotype">Holotype</typeStatus>
<quantity id="4C8B9B369D56FFDC60D46169FACDFBC6" box="[1217,1317,1052,1077]" metricMagnitude="-3" metricUnit="m" metricValue="4.52" pageId="5" pageNumber="566" unit="mm" value="4.52">4.52 mm</quantity>
long, with anterior 20 segments (
<figureCitation id="13482A569D56FFDC6587614AFE05FBAA" box="[402,493,1087,1112]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="5" pageNumber="566">Fig. 3A</figureCitation>
),
<typeStatus id="54C888719D56FFDC66176135FD85FBAA" box="[514,621,1088,1112]" pageId="5" pageNumber="566" type="paratype">paratypes</typeStatus>
<quantity id="4C8B9B369D56FFDC66606135FD15FBA5" box="[629,765,1088,1112]" metricMagnitude="-3" metricUnit="m" metricValue="2.9" metricValueMax="4.1" metricValueMin="1.7" pageId="5" pageNumber="566" unit="mm" value="2.9" valueMax="4.1" valueMin="1.7">1.74.1 mm</quantity>
long, with anterior 1123 segments. Width at XI,
<quantity id="4C8B9B369D56FFDC612D6135FA75FBA5" box="[1336,1437,1088,1112]" metricMagnitude="-4" metricUnit="m" metricValue="8.8" pageId="5" pageNumber="566" unit="mm" value="0.88">0.88 mm</quantity>
in
<typeStatus id="54C888719D56FFDC64AF6111FEF7FB8F" box="[186,287,1124,1149]" pageId="5" pageNumber="566" type="holotype">holotype</typeStatus>
,
<quantity id="4C8B9B369D56FFDC65256111FE31FB8E" box="[304,473,1124,1148]" metricMagnitude="-4" metricUnit="m" metricValue="5.6499999999999995" metricValueMax="7.4" metricValueMin="3.9" pageId="5" pageNumber="566" unit="mm" value="0.565" valueMax="0.74" valueMin="0.39">0.390.74 mm</quantity>
in
<typeStatus id="54C888719D56FFDC661D6110FD9FFB8F" box="[520,631,1125,1149]" pageId="5" pageNumber="566" type="paratype">paratypes</typeStatus>
; all specimens mounted and compressed.
<typeStatus id="54C888719D56FFDC607A6111FB3FFB8F" box="[1135,1239,1124,1149]" pageId="5" pageNumber="566" type="holotype">Holotype</typeStatus>
and all but one
<typeStatus id="54C888719D56FFDC648261FCFF1FFB53" box="[151,247,1161,1185]" pageId="5" pageNumber="566" type="paratype">paratype</typeStatus>
sexually mature, other specimens in various stages of maturity. Prostomium rounded in most specimens, slightly more triangular in some, shorter than basally wide (
<figureCitation id="13482A569D56FFDC675661D9FC77FB36" box="[835,927,1196,1221]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="5" pageNumber="566">Fig. 3A</figureCitation>
). Cuticle smooth, cuticular papillae nowhere present. Clitellum present in XXI in
<typeStatus id="54C888719D56FFDC662A61BAFD49FB1A" box="[575,673,1231,1256]" pageId="5" pageNumber="566" type="holotype">holotype</typeStatus>
and most
<typeStatus id="54C888719D56FFDC670061A4FC6BFB1B" box="[789,899,1233,1257]" pageId="5" pageNumber="566" type="paratype">paratypes</typeStatus>
, not developed in remaining specimens.
</paragraph>
<paragraph id="8BCC36D39D56FFDC64D26180FE5CF9B9" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
In preclitellar segments, dorsal chaetal bundles with (2)35 bifid crotchets (
<figureCitation id="13482A569D56FFDC60326181FB6CFAFE" box="[1063,1156,1268,1293]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4A</figureCitation>
), 4070 µm long. Bifids with teeth of about same length; upper tooth slightly thinner than lower. Postclitellar, dorsal bundles with 12(3) bifid crotchets (
<figureCitation id="13482A569D56FFDC6559604EFE4EFAA1" box="[332,422,1339,1364]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4B</figureCitation>
), 3555 µm long; upper tooth longer and slightly thinner than lower. Preclitellar ventral bundles (
<figureCitation id="13482A569D56FFDC64E8602BFEBBFA85" box="[253,339,1374,1399]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4C</figureCitation>
) with (2)34 bifid crotchets (3565 µm long); upper tooth thinner but about same length as lower. Postclitellar ventral bundles (
<figureCitation id="13482A569D56FFDC65F760F4FDD3FA68" box="[482,571,1409,1434]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4D</figureCitation>
) with 12 bifid crotchets (3540 µm long), upper teeth slightly thinner than lower; teeth of about same length, and rather diverging. No ventral chaetae in segment XI in mature specimens. Dorsal bifid chaetae weakly sigmoidal, some ventral bundles with heavily bent, sigmoidal chaetae (sometimes broken); nodulus inconspicuous in most specimens, somewhat more developed in
<typeStatus id="54C888719D56FFDC605E6099FB59F9F7" box="[1099,1201,1516,1541]" pageId="5" pageNumber="566" type="holotype">holotype</typeStatus>
. Spermathecal pores paired, located slightly above ventral chaetae in X. Male pores paired, in line with assumed ventral chaetae about halfway into segment XI.
</paragraph>
<paragraph id="8BCC36D39D56FFDC64D26323FC0DF807" blockId="5.[151,1437,151,2038]" pageId="5" pageNumber="566">
Pharyngeal glands present in segments IVV. Vas deferens (
<figureCitation id="13482A569D56FFDC67636323FC27F99D" box="[886,975,1622,1647]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4E</figureCitation>
; vd) about 35 µm wide in
<typeStatus id="54C888719D56FFDC61166323FA80F99D" box="[1283,1384,1622,1647]" pageId="5" pageNumber="566" type="holotype">holotype</typeStatus>
, full length not measurable in any specimen, but longer than 200 µm in one
<typeStatus id="54C888719D56FFDC67DD630EFBC4F961" box="[968,1068,1659,1683]" pageId="5" pageNumber="566" type="paratype">paratype</typeStatus>
; densely ciliated towards atrium insertion point. Prostate (
<figureCitation id="13482A569D56FFDC65BA63E8FDEDF944" box="[431,517,1693,1718]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4E</figureCitation>
; pr) rather large, but possibly compressed. Atrium (
<figureCitation id="13482A569D56FFDC605663E8FB4DF944" box="[1091,1189,1693,1718]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="5" pageNumber="566">Figs. 3B</figureCitation>
,
<figureCitation id="13482A569D56FFDC60A563EBFB3AF944" box="[1200,1234,1694,1718]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">4E</figureCitation>
; at) about 430 µm long, about 100 µm wide at widest point; tripartite with coarse cellular matrix in ental cap-like portion, finer cellular structure in mid-portion epithelium, and denser epithelium in muscular ectal portion. Atrium terminating in cuticular penis sheath with slight inward turn at ectal end, but still rather large terminal opening (
<figureCitation id="13482A569D56FFDC60CE6272FAD5F8D2" box="[1243,1341,1799,1824]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="5" pageNumber="566">Figs. 3B</figureCitation>
,
<figureCitation id="13482A569D56FFDC615C627DFA70F8D2" box="[1353,1432,1800,1824]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">4E, 4F</figureCitation>
; cu), about 90 µm long, 3035 µm wide at base. Soft tissue of penis protruding through terminal opening of penis sheath in
<typeStatus id="54C888719D56FFDC651F623BFE84F895" box="[266,364,1870,1895]" pageId="5" pageNumber="566" type="holotype">holotype</typeStatus>
and most
<typeStatus id="54C888719D56FFDC65F96225FDB3F89A" box="[492,603,1872,1896]" pageId="5" pageNumber="566" type="paratype">paratypes</typeStatus>
. Spermathecae (
<figureCitation id="13482A569D56FFDC670C623BFC97F895" box="[793,895,1870,1895]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="5" pageNumber="566">Figs. 3C</figureCitation>
,
<figureCitation id="13482A569D56FFDC679A623AFC5CF895" box="[911,948,1871,1895]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">4G</figureCitation>
; sp) extending from rather large but nonswollen pore, with narrower ectal duct (
<figureCitation id="13482A569D56FFDC66686207FD35F879" box="[637,733,1906,1931]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4G</figureCitation>
; sd) opening up into wider ampulla entally (
<figureCitation id="13482A569D56FFDC61176207FA8AF879" box="[1282,1378,1906,1931]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">Fig. 4G</figureCitation>
; sa), spermathecae much compressed in all specimens and often pressed into surrounding segments; duct about 80 µm long, 40 µm wide at widest point, ampullae about 400 µm long, 150 µm wide at widest part. Spermatozeugmata rather stout and often coiled (170 µm long, 40 µm wide) (
<figureCitation id="13482A569D56FFDC670B62A8FC68F807" box="[798,896,2013,2038]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="5" pageNumber="566">Figs. 3C</figureCitation>
,
<figureCitation id="13482A569D56FFDC679962A8FC59F807" box="[908,945,2013,2037]" captionStart="FIGURE 4" captionStartId="7.[151,250,1839,1861]" captionTargetBox="[219,1263,592,1814]" captionTargetId="figure@7.[202,1413,556,1824]" captionTargetPageId="7" captionText="FIGURE 4. Tubificoides mackiei n. sp., holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown)." httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="5" pageNumber="566">4G</figureCitation>
; sz).
</paragraph>
</subSubSection>
<caption id="DF0C665B9D55FFDF64826014FC7EFA46" httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="6" pageNumber="567" startId="6.[151,250,1377,1399]" targetBox="[151,1436,193,1356]" targetPageId="6">
<paragraph id="8BCC36D39D55FFDF64826014FC7EFA46" blockId="6.[151,1437,1377,1460]" pageId="6" pageNumber="567">
<emphasis id="B907EAC19D55FFDF64826014FEFBFA84" bold="true" box="[151,275,1377,1399]" pageId="6" pageNumber="567">FIGURE 3.</emphasis>
<taxonomicName id="4C734D509D55FFDF650F6014FE03FA85" authority="Kvist &amp; Erséus, 2018" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[282,491,1377,1399]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="mackiei" status="sp. nov.">
<emphasis id="B907EAC19D55FFDF650F6014FE03FA85" box="[282,491,1377,1399]" italics="true" pageId="6" pageNumber="567">Tubificoides mackiei</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23457BA9D55FFDF65E66014FDC4FA84" box="[499,556,1377,1398]" pageId="6" pageNumber="567" rank="species">
<emphasis id="B907EAC19D55FFDF65E66014FDC4FA84" bold="true" box="[499,556,1377,1398]" pageId="6" pageNumber="567">n. sp.</emphasis>
</taxonomicNameLabel>
, holotype (AC) and
<taxonomicName id="4C734D509D55FFDF67196017FC44FA85" box="[780,940,1377,1399]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF67196017FC44FA85" box="[780,940,1377,1399]" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
lectotype (D).
<emphasis id="B907EAC19D55FFDF605D6014FB88FA84" bold="true" box="[1096,1120,1377,1398]" pageId="6" pageNumber="567">A.</emphasis>
Anterior end of body.
<emphasis id="B907EAC19D55FFDF615F6014FA89FA84" bold="true" box="[1354,1377,1377,1398]" pageId="6" pageNumber="567">B.</emphasis>
Male genital apparatus.
<emphasis id="B907EAC19D55FFDF654460F5FE82FA64" bold="true" box="[337,362,1408,1430]" pageId="6" pageNumber="567">C.</emphasis>
Spermatheca.
<emphasis id="B907EAC19D55FFDF661460F5FDF2FA67" bold="true" box="[513,538,1408,1429]" pageId="6" pageNumber="567">D.</emphasis>
Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma.
</paragraph>
</caption>
<subSubSection id="C36965589D55FFDF64D260A9FC39F9EA" pageId="6" pageNumber="567" type="distribution">
<paragraph id="8BCC36D39D55FFDF64D260A9FC39F9EA" blockId="6.[151,1437,1500,2029]" pageId="6" pageNumber="567">
<emphasis id="B907EAC19D55FFDF64D260A9FE19FA07" bold="true" box="[199,497,1500,1525]" pageId="6" pageNumber="567">Distribution and habitat.</emphasis>
Known from the southeast coast of
<collectingCountry id="F36476439D55FFDF679260A8FC09FA07" box="[903,993,1501,1525]" name="Norway" pageId="6" pageNumber="567">Norway</collectingCountry>
and west coast of
<collectingCountry id="F36476439D55FFDF60A760A9FAE6FA07" box="[1202,1294,1500,1525]" name="Sweden" pageId="6" pageNumber="567">Sweden</collectingCountry>
, in fine sand and silt, as well as coarse sand and gravel; intertidal and shallow subtidal.
</paragraph>
</subSubSection>
<subSubSection id="C36965589D55FFD164D26351FA60FD76" lastPageId="8" lastPageNumber="569" pageId="6" pageNumber="567" type="discussion">
<paragraph id="8BCC36D39D55FFDE64D26351FF16FF0A" blockId="6.[151,1437,1500,2029]" lastBlockId="7.[151,1436,151,501]" lastPageId="7" lastPageNumber="568" pageId="6" pageNumber="567">
<emphasis id="B907EAC19D55FFDF64D26351FED3F9CF" bold="true" box="[199,315,1572,1597]" pageId="6" pageNumber="567">Remarks.</emphasis>
Molecular data support that
<taxonomicName id="4C734D509D55FFDF668F6351FC6EF9CF" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[666,902,1572,1597]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D55FFDF668F6351FC6EF9CF" box="[666,902,1572,1597]" italics="true" pageId="6" pageNumber="567">Tubificoides mackiei</emphasis>
</taxonomicName>
and the north European
<taxonomicName id="4C734D509D55FFDF60A36350FA84F9CF" box="[1206,1388,1572,1597]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF60A36350FA84F9CF" box="[1206,1388,1572,1597]" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
are closely related (
<bibRefCitation id="EFE24B229D55FFDF655E6332FDF9F992" box="[331,529,1607,1632]" pageId="6" pageNumber="567" refId="ref8712" refString="Kvist, S., Sarkar, I. N. &amp; Erseus, C. (2010) Genetic variation and phylogeny of the cosmopolitan marine genus Tubificoides (Annelida: Clitellata: Naididae: Tubificinae). Molecular Phylogenetics and Evolution, 57 (2), 687 - 702. https: // doi. org / 10.1016 / j. ympev. 2010.08.018" type="journal article">
Kvist
<emphasis id="B907EAC19D55FFDF6587633CFE24F992" box="[402,460,1607,1632]" italics="true" pageId="6" pageNumber="567">et al.</emphasis>
2010
</bibRefCitation>
). The two taxa strongly resemble each other, both in general appearance and in some details of the chaetal morphology. In particular,
<taxonomicName id="4C734D509D55FFDF67006319FBE8F977" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[789,1024,1644,1669]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D55FFDF67006319FBE8F977" box="[789,1024,1644,1669]" italics="true" pageId="6" pageNumber="567">Tubificoides mackiei</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D509D55FFDF602A6318FB1CF977" box="[1087,1268,1644,1669]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF602A6318FB1CF977" box="[1087,1268,1644,1669]" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
are similar in general body length from prostomium to segment XI, the lack of hair chaetae and cuticular papillation, and in the detailed morphology of dorsal chaetae (
<bibRefCitation id="EFE24B229D55FFDF664263C1FD3AF93F" box="[599,722,1716,1741]" pageId="6" pageNumber="567" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl 1960</bibRefCitation>
;
<bibRefCitation id="EFE24B229D55FFDF66CA63C1FC8EF93E" box="[735,870,1716,1741]" pageId="6" pageNumber="567" refId="ref6620" refString="Baker, H. R. (1980) A redescription of Tubificoides pseudogaster (Dahl) (Oligochaeta: Tubificidae). Transactions of the American Microscopical Society, 99 (3), 337 - 342. https: // doi. org / 10.2307 / 3226011" type="journal article">Baker 1980</bibRefCitation>
;
<bibRefCitation id="EFE24B229D55FFDF676463C1FB57F93F" box="[881,1215,1716,1741]" pageId="6" pageNumber="567" refId="ref9713" refString="Takashima, Y. &amp; Mawatari, S. F. (1996) Marine Tubificidae (Oligochaeta) from Hokkaido, northern Japan, with descriptions of two new species. Species Diversity, 1, 55 - 70. https: // doi. org / 10.12782 / specdiv. 1.55" type="journal article">Takashima &amp; Mawatari 1996</bibRefCitation>
). However, the two species can be distinguished on the basis of body width,
<emphasis id="B907EAC19D55FFDF670163ACFCD4F902" box="[788,828,1753,1776]" italics="true" pageId="6" pageNumber="567">e.g.</emphasis>
using the width at segment X (
<taxonomicName id="4C734D509D55FFDF608B63ACFAF9F902" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[1182,1297,1751,1776]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D55FFDF608B63ACFAF9F902" box="[1182,1297,1751,1776]" italics="true" pageId="6" pageNumber="567">T. mackiei</emphasis>
</taxonomicName>
seems to be about twice as wide as
<taxonomicName id="4C734D509D55FFDF65B36388FDB1F8E7" box="[422,601,1788,1813]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF65B36388FDB1F8E7" box="[422,601,1788,1813]" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
in X [see
<bibRefCitation id="EFE24B229D55FFDF66C36389FCBBF8E7" box="[726,851,1788,1813]" pageId="6" pageNumber="567" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl 1960</bibRefCitation>
], although such a difference may depend on, or be exaggerated by, coverslip compression on a microscope slide), the shape of the atrium (wider ental portion in
<taxonomicName id="4C734D509D55FFDF61926254FF06F8AF" authorityName="Kvist &amp; Erséus" authorityYear="2018" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D55FFDF61926254FF06F8AF" italics="true" pageId="6" pageNumber="567">T. mackiei</emphasis>
</taxonomicName>
), and the maximal number of preclitellar, dorsal chaetae (the reported [up to] six chaetae per bundle in
<taxonomicName id="4C734D509D55FFDF61926230FEC4F872" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF61926230FEC4F872" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
[
<bibRefCitation id="EFE24B229D55FFDF652C6212FE5CF872" box="[313,436,1895,1920]" pageId="6" pageNumber="567" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl 1960</bibRefCitation>
] are nowhere found in
<taxonomicName id="4C734D509D55FFDF66A0621CFCC0F872" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[693,808,1895,1920]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D55FFDF66A0621CFCC0F872" box="[693,808,1895,1920]" italics="true" pageId="6" pageNumber="567">T. mackiei</emphasis>
</taxonomicName>
). Also, the shape and size of the penis sheath (shorter in
<taxonomicName id="4C734D509D55FFDF648262F8FEA5F857" box="[151,333,1932,1957]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF648262F8FEA5F857" box="[151,333,1932,1957]" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
), and the details of the penial tube (with an outward-turned collar at the ental portion in
<taxonomicName id="4C734D509D55FFDF619262F8FEC4F83A" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="6" pageNumber="567" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D55FFDF619262F8FEC4F83A" italics="true" pageId="6" pageNumber="567">T. pseudogaster</emphasis>
</taxonomicName>
) differs between the taxa. It should be noted that, with regard to the detailed morphology of the chaetae, in particular the length of the upper teeth in the dorsal bundles, Dahls (1960) original description and Bakers (1980) re-description of
<taxonomicName id="4C734D509D54FFDE661F65ECFD53FF42" box="[522,699,151,176]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="568" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D54FFDE661F65ECFD53FF42" box="[522,699,151,176]" italics="true" pageId="7" pageNumber="568">T. pseudogaster</emphasis>
</taxonomicName>
report slightly different information.
<bibRefCitation id="EFE24B229D54FFDE607165E2FB04FF42" box="[1124,1260,151,176]" pageId="7" pageNumber="568" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl (1960)</bibRefCitation>
mentioned that the upper teeth in dorsal bundles are sometimes shorter than the lower teeth, whereas
<bibRefCitation id="EFE24B229D54FFDE607365C9FB13FF26" box="[1126,1275,188,213]" pageId="7" pageNumber="568" refId="ref6620" refString="Baker, H. R. (1980) A redescription of Tubificoides pseudogaster (Dahl) (Oligochaeta: Tubificidae). Transactions of the American Microscopical Society, 99 (3), 337 - 342. https: // doi. org / 10.2307 / 3226011" type="journal article">Baker (1980)</bibRefCitation>
suggested the opposite.
</paragraph>
<paragraph id="8BCC36D39D54FFDE64D26471FAEFFE06" blockId="7.[151,1436,151,501]" pageId="7" pageNumber="568">
<bibRefCitation id="EFE24B229D54FFDE64D26471FEBCFEEF" box="[199,340,260,285]" pageId="7" pageNumber="568" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl (1960)</bibRefCitation>
reported the following for
<emphasis id="B907EAC19D54FFDE66896470FCB3FEEF" box="[668,859,260,285]" italics="true" pageId="7" pageNumber="568">
<taxonomicName id="4C734D509D54FFDE66896470FCBEFEEF" box="[668,854,260,285]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="568" phylum="Annelida" rank="species" species="pseudogaster">T. pseudogaster</taxonomicName>
:
</emphasis>
&quot;In segment 9 the intestinal walls are thickened dorsally and laterally with a glandular structure&quot;. These structures are indicated in the scientific name for the species (Gr. pseudēs meaning false, and Gr. gastēr meaning stomach). All of our slide-mounted, non-juvenile specimens of both
<taxonomicName id="4C734D509D54FFDE657C6404FDF1FE7A" box="[361,537,367,392]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="568" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D54FFDE657C6404FDF1FE7A" box="[361,537,367,392]" italics="true" pageId="7" pageNumber="568">T. pseudogaster</emphasis>
</taxonomicName>
and
<taxonomicName id="4C734D509D54FFDE665A6404FD2AFE7A" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[591,706,367,392]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="568" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D54FFDE665A6404FD2AFE7A" box="[591,706,367,392]" italics="true" pageId="7" pageNumber="568">T. mackiei</emphasis>
</taxonomicName>
display a feature very similar to that detailed by
<bibRefCitation id="EFE24B229D54FFDE60F1641AFA86FE7A" box="[1252,1390,367,392]" pageId="7" pageNumber="568" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl (1960)</bibRefCitation>
, yet it is difficult to discern if the structures are of digestive epithelial origin or chloragogenic cells, and whether they are directly related to the intestinal tract or free in the coelomic cavity. Regardless, because both species seem to have this structure, it may be a synapomorphy for them (and perhaps additional, closely related taxa).
</paragraph>
<caption id="DF0C665B9D54FFDE6482625AFCBAF82C" httpUri="https://zenodo.org/record/1290341/files/figure.png" pageId="7" pageNumber="568" startId="7.[151,250,1839,1861]" targetBox="[219,1263,592,1814]" targetPageId="7">
<paragraph id="8BCC36D39D54FFDE6482625AFCBAF82C" blockId="7.[151,1437,1839,2014]" pageId="7" pageNumber="568">
<emphasis id="B907EAC19D54FFDE6482625AFEFCF8B6" bold="true" box="[151,276,1839,1861]" pageId="7" pageNumber="568">FIGURE 4.</emphasis>
<taxonomicName id="4C734D509D54FFDE6508625AFE18F8B7" authority="Kvist &amp; Erséus, 2018" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[285,496,1839,1861]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="7" pageNumber="568" phylum="Annelida" rank="species" species="mackiei" status="sp. nov.">
<emphasis id="B907EAC19D54FFDE6508625AFE18F8B7" box="[285,496,1839,1861]" italics="true" pageId="7" pageNumber="568">Tubificoides mackiei</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A23457BA9D54FFDE65ED625AFDDBF8B6" box="[504,563,1839,1860]" pageId="7" pageNumber="568" rank="species">
<emphasis id="B907EAC19D54FFDE65ED625AFDDBF8B6" bold="true" box="[504,563,1839,1860]" pageId="7" pageNumber="568">n. sp.</emphasis>
</taxonomicNameLabel>
, holotype. A. Preclitellar dorsal chaeta. B. Postclitellar dorsal chaeta. C. Preclitellar ventral chaeta. D. Postclitellar ventral chaeta. AD distal ends of chaetae only. E. Male genital apparatus (for clarity, penial sac is not drawn in figure) and ovary (note that the ovary is not connected to any male structure, but abuts the atrium). F. Lateral contours of penis sheaths (broken lines indicate protruding soft parts). G. Spermatheca with spermatozeugmata. Abbreviations: at, atrium; cu, cuticular penis sheath; ov, ovary; pr, prostate gland; sa, spermathecal ampulla; sd, spermathecal duct; sz, spermatozeugma; vd, vas deferens (only a short part of vd is shown).
</paragraph>
</caption>
<paragraph id="8BCC36D39D5BFFD164D265E2FA60FD76" blockId="8.[151,1437,151,645]" pageId="8" pageNumber="569">
<emphasis id="B907EAC19D5BFFD164D265E2FD37FF42" bold="true" box="[199,735,151,176]" pageId="8" pageNumber="569">
On the penis morphology of
<taxonomicName id="4C734D509D5BFFD1663065EDFD3FFF42" box="[549,727,151,176]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D5BFFD1663065EDFD3FFF42" bold="true" box="[549,727,151,176]" italics="true" pageId="8" pageNumber="569">T. pseudogaster</emphasis>
</taxonomicName>
.
</emphasis>
Historically, delineations of species within
<taxonomicName id="4C734D509D5BFFD160C065E2FAB6FF42" authorityName="Lastockin" authorityYear="1937" box="[1237,1374,151,176]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="genus">
<emphasis id="B907EAC19D5BFFD160C065E2FAB6FF42" box="[1237,1374,151,176]" italics="true" pageId="8" pageNumber="569">Tubificoides</emphasis>
</taxonomicName>
have relied heavily on the shape and size of penis sheaths (
<bibRefCitation id="EFE24B229D5BFFD1671865C9FBD2FF26" box="[781,1082,188,213]" pageId="8" pageNumber="569" refId="ref7065" refString="Brinkhurst, R. O. &amp; Baker, H. R. (1979) A review of the marine Tubificidae (Oligochaeta) of North America. Canadian Journal of Zoology, 57 (8), 1553 - 1569. https: // doi. org / 10.1139 / z 79 - 203" type="journal article">Brinkhurst &amp; Baker 1979</bibRefCitation>
;
<bibRefCitation id="EFE24B229D5BFFD1605D65C9FAECFF26" box="[1096,1284,188,213]" pageId="8" pageNumber="569" refId="ref6919" refString="Brinkhurst, R. O. (1981) A contribution to the taxonomy of the Tubificinae (Oligochaeta: Tubificidae). Proceedings of the Biological Society of Washington, 94 (4), 1048 - 1067." type="journal article">Brinkhurst 1981</bibRefCitation>
,
<bibRefCitation id="EFE24B229D5BFFD1610665C9FAA6FF27" box="[1299,1358,188,213]" pageId="8" pageNumber="569" refId="ref6959" refString="Brinkhurst, R. O. (1985) A further contribution to the taxonomy of the genus Tubificoides Lastockin (Oligochaeta: Tubificidae). Canadian Journal of Zoology, 63 (2), 400 - 410. https: // doi. org / 10.1139 / z 85 - 061" type="journal article">1985</bibRefCitation>
,
<bibRefCitation id="EFE24B229D5BFFD1614965C9FA7FFF27" box="[1372,1431,188,213]" pageId="8" pageNumber="569" refId="ref7012" refString="Brinkhurst, R. O. (1986) Taxonomy of the genus Tubificoides Lastockin (Oligochaeta, Tubificidae): species with bifid setae. Canadian Journal of Zoology, 64 (6), 1270 - 1279. https: // doi. org / 10.1139 / z 86 - 189" type="journal article">1986</bibRefCitation>
;
<bibRefCitation id="EFE24B229D5BFFD1648265AAFEF7FF0A" box="[151,287,223,248]" pageId="8" pageNumber="569" refId="ref6669" refString="Baker, H. R. (1983) New species of Tubificoides Lastockin (Oligochaeta; Tubificidae) from the Pacific Northeast and the Arctic. Canadian Journal of Zoology, 61 (6), 1270 - 1283. https: // doi. org / 10.1139 / z 83 - 171" type="journal article">Baker 1983</bibRefCitation>
; Helgasson &amp;
<bibRefCitation id="EFE24B229D5BFFD165D86595FDB5FF0A" box="[461,605,224,248]" pageId="8" pageNumber="569" refId="ref8221" refString="Helgason, G. V. &amp; Erseus, C. (1987) Three new species of Tubificoides (Oligochaeta, Tubificidae) from the North - west Atlantic and notes on geographic variation in the circumpolar T. kozloffi. Sarsia, 72 (2), 159 - 169. https: // doi. org / 10.1080 / 00364827.1987.10419713" type="journal article">Erséus 1987</bibRefCitation>
); this character also seems to separate
<taxonomicName id="4C734D509D5BFFD160316594FB71FF0A" authorityName="Kvist &amp; Erséus" authorityYear="2018" box="[1060,1177,223,248]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="mackiei">
<emphasis id="B907EAC19D5BFFD160316594FB71FF0A" box="[1060,1177,223,248]" italics="true" pageId="8" pageNumber="569">T. mackiei</emphasis>
</taxonomicName>
from
<emphasis id="B907EAC19D5BFFD160F16594FA75FF0A" box="[1252,1437,223,248]" italics="true" pageId="8" pageNumber="569">
<taxonomicName id="4C734D509D5BFFD160F16594FA70FF0A" box="[1252,1432,223,248]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="pseudogaster">T. pseudogaster</taxonomicName>
.
</emphasis>
Although the resolution of Dahls (1960) figure showing the penis sheath of
<taxonomicName id="4C734D509D5BFFD167ED6470FB41FEEF" box="[1016,1193,260,285]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D5BFFD167ED6470FB41FEEF" box="[1016,1193,260,285]" italics="true" pageId="8" pageNumber="569">T. pseudogaster</emphasis>
</taxonomicName>
is less than ideal, the language in her description is clear.
<bibRefCitation id="EFE24B229D5BFFD1663B6452FD51FEB2" box="[558,697,295,320]" pageId="8" pageNumber="569" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl (1960)</bibRefCitation>
, when describing the penis sheath, states that: &quot;Its length is about twice its breadth&quot; and that &quot;Distally the penial tube has an outward turned collar&quot;. In addition, the figures supporting Bakers (1980) and Takashima &amp; Mawataris (1996) re-descriptions of the species clearly show a &quot;cone-shaped&quot; penis sheath, with an outward turn at the ental end [referred to as &quot;ental flange&quot; in
<bibRefCitation id="EFE24B229D5BFFD1611164E1FEB1FE22" pageId="8" pageNumber="569" refId="ref9713" refString="Takashima, Y. &amp; Mawatari, S. F. (1996) Marine Tubificidae (Oligochaeta) from Hokkaido, northern Japan, with descriptions of two new species. Species Diversity, 1, 55 - 70. https: // doi. org / 10.12782 / specdiv. 1.55" type="journal article">Takashima &amp; Mawatari (1996)</bibRefCitation>
]. In order to aid future morphological identifications of
<taxonomicName id="4C734D509D5BFFD167DD64CCFB90FE22" box="[968,1144,439,464]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D5BFFD167DD64CCFB90FE22" box="[968,1144,439,464]" italics="true" pageId="8" pageNumber="569">T. pseudogaster</emphasis>
</taxonomicName>
(see also above regarding the DNA barcodes for this species) and owing to the low quality of the figures presented by
<bibRefCitation id="EFE24B229D5BFFD160B964A9FADFFE07" box="[1196,1335,476,501]" pageId="8" pageNumber="569" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl (1960)</bibRefCitation>
, we here present a photograph of the penis sheath from the
<typeStatus id="54C888719D5BFFD16714648AFC80FDEA" box="[769,872,511,536]" pageId="8" pageNumber="569" type="lectotype">lectotype</typeStatus>
of
<taxonomicName id="4C734D509D5BFFD1678C6774FBB8FDEA" box="[921,1104,511,536]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D5BFFD1678C6774FBB8FDEA" box="[921,1104,511,536]" italics="true" pageId="8" pageNumber="569">T. pseudogaster</emphasis>
</taxonomicName>
(
<figureCitation id="13482A569D5BFFD16073648AFB2FFDEA" box="[1126,1223,511,536]" captionStart="FIGURE 3" captionStartId="6.[151,250,1377,1399]" captionTargetBox="[151,1436,193,1356]" captionTargetId="figure@6.[151,1436,193,1356]" captionTargetPageId="6" captionText="FIGURE 3. Tubificoides mackiei n. sp., holotype (AC) and T. pseudogaster lectotype (D). A. Anterior end of body. B. Male genital apparatus. C. Spermatheca. D. Penis sheath of lectotype (ZMUC-OLI-000032; see text for discussion). Abbreviations: at, atrium; cu, cuticular penis sheath; sp, spermatheca; sz, spermatozeugma." httpUri="https://zenodo.org/record/1290339/files/figure.png" pageId="8" pageNumber="569">Fig. 3D</figureCitation>
). The structure is morphologically compatible with the detailed description by
<bibRefCitation id="EFE24B229D5BFFD1674B6751FC05FDCF" box="[862,1005,548,573]" pageId="8" pageNumber="569" refId="ref7440" refString="Dahl, I. O. (1960) The oligochaete fauna of 3 Danish brackish water areas (taxonomic and biological observations). Meddelelser fra Danmarks Fiskeri- og HavundersOgelser, New Series, 26, 1 - 20." type="journal article">Dahl (1960)</bibRefCitation>
, as well as both the text and figures presented in Bakers (1980) re-description, and we hope that these new photographs, in combination with the genetic data, will mediate the difficulty of delimiting species that are morphologically similar to
<taxonomicName id="4C734D509D5BFFD160C46718FA69FD77" box="[1233,1409,620,645]" class="Clitellata" family="Tubificidae" genus="Tubificoides" higherTaxonomySource="GBIF" kingdom="Animalia" order="Haplotaxida" pageId="8" pageNumber="569" phylum="Annelida" rank="species" species="pseudogaster">
<emphasis id="B907EAC19D5BFFD160C46718FA69FD77" box="[1233,1409,620,645]" italics="true" pageId="8" pageNumber="569">T. pseudogaster</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>