treatments-xml/data/03/9D/C3/039DC377FFA8FFCD4C13FD05FED05BD2.xml

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<mods:title>New insights into the systematics of Lumbricillus and Marionina (Clitellata: Enchytraeidae) inferred from Southern Hemisphere samples, including three new species</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Klinth, Mårten J.</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Rota, Emilia</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Martinsson, Svante</mods:namePart>
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<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Prantoni, Alessandro L.</mods:namePart>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Erséus, Christer</mods:namePart>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:date>2022</mods:date>
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<mods:number>2021-09-24</mods:number>
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<mods:number>194</mods:number>
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<mods:number>4</mods:number>
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<mods:identifier type="DOI">10.1093/zoolinnean/zlab073</mods:identifier>
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<taxonomicName box="[306,478,679,704]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="undetermined">
<emphasis box="[306,439,680,703]" italics="true" pageId="24" pageNumber="1127">MARIONINA</emphasis>
SP.
</taxonomicName>
‘SNOW IS.’
</heading>
</paragraph>
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<subSubSection pageId="24" pageNumber="1127" type="description">
<paragraph blockId="24.[377,528,729,754]" box="[377,528,729,754]" pageId="24" pageNumber="1127">
(
<figureCitation box="[386,478,729,754]" captionStart="Figure 9" captionStartId="19.[828,907,850,872]" captionTargetBox="[830,1438,198,807]" captionTargetId="figure-668@19.[827,1441,195,810]" captionTargetPageId="19" captionText="Figure 9. A, Marionina sp. ‘Snow Is.’ (CE34647), anterior body (lateral view). B, Marionina aestuum, sectioned specimen BMNH 1933.2.23.895, cephalic region (lateral view). C, M. aestuum (CE12477), cephalic region (ventral view). D, Marionina georgiana (paralectotype 13), cephalic region (ventral view). Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461109" httpUri="https://zenodo.org/record/6461109/files/figure.png" pageId="24" pageNumber="1127">FIGS 9A</figureCitation>
, 
<figureCitation box="[492,520,729,754]" captionStart="Figure 14" captionStartId="24.[145,224,1817,1839]" captionTargetBox="[148,757,1616,1774]" captionTargetId="figure-808@24.[145,760,1613,1777]" captionTargetPageId="24" captionText="Figure 14. Marionina sp. ‘Snow Is.’ (CE34647). A, chaetal bundle. B, anterior part of body. Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461125" httpUri="https://zenodo.org/record/6461125/files/figure.png" pageId="24" pageNumber="1127">14</figureCitation>
)
</paragraph>
<paragraph blockId="24.[145,761,801,976]" pageId="24" pageNumber="1127">
<emphasis box="[145,379,801,822]" italics="true" pageId="24" pageNumber="1127">Material examined:</emphasis>
SMNH198155 (CE34647) and SMNH198156 (CE34648), two semi-mature specimens from Snow Island (South Shetland Islands), 
<collectingCountry box="[634,757,862,884]" name="Antarctica" pageId="24" pageNumber="1127">Antarctica</collectingCountry>
. For information on collection localities and GenBank accession numbers for 
<emphasis box="[410,458,924,945]" italics="true" pageId="24" pageNumber="1127">COI</emphasis>
barcodes, see 
<tableCitation box="[624,710,924,946]" captionStart="Table 1" captionStartId="3.[144,208,419,441]" captionTargetPageId="3" captionText="Table 1. Specimens from the Southern Hemisphere used in this study, with specimen identification number, region of the collection site, GenBank accession numbers for the seven genetic markers, and voucher numbers (with museum acronym)" httpUri="http://table.plazi.org/id/DF4B22E9FFB3FFD74DB1FE0EFBE15ECA" pageId="24" pageNumber="1127" tableUuid="DF4B22E9FFB3FFD74DB1FE0EFBE15ECA">Table 1</tableCitation>
and the Supporting Information (
<tableCitation box="[477,576,954,976]" captionStart="Table 1" captionStartId="3.[144,208,419,441]" captionTargetPageId="3" captionText="Table 1. Specimens from the Southern Hemisphere used in this study, with specimen identification number, region of the collection site, GenBank accession numbers for the seven genetic markers, and voucher numbers (with museum acronym)" httpUri="http://table.plazi.org/id/DF4B22E9FFB3FFD74DB1FE0EFBE15ECA" pageId="24" pageNumber="1127" tableUuid="DF4B22E9FFB3FFD74DB1FE0EFBE15ECA">Table S1</tableCitation>
).
</paragraph>
<paragraph blockId="24.[145,762,1017,1560]" pageId="24" pageNumber="1127">
<emphasis box="[145,289,1018,1039]" italics="true" pageId="24" pageNumber="1127">Description:</emphasis>
Live colour unknown; and specimens apparently without pigmentation. Length of first 16–19 segments 
<quantity box="[366,517,1079,1100]" metricMagnitude="-3" metricUnit="m" metricValue="2.65" metricValueMax="2.9" metricValueMin="2.4" pageId="24" pageNumber="1127" unit="mm" value="2.65" valueMax="2.9" valueMin="2.4">2.4–2.9 mm</quantity>
(fixed, amputated specimens); first 15 segments 
<quantity box="[491,625,1109,1131]" metricMagnitude="-3" metricUnit="m" metricValue="2.25" metricValueMax="2.3" metricValueMin="2.2" pageId="24" pageNumber="1127" unit="mm" value="2.25" valueMax="2.3" valueMin="2.2">2.2–2.3 mm</quantity>
long; width at XII 
<quantity box="[223,327,1140,1161]" metricMagnitude="-4" metricUnit="m" metricValue="2.7" pageId="24" pageNumber="1127" unit="mm" value="0.27">0.27 mm</quantity>
. Chaetae slightly sigmoid (
<figureCitation box="[648,746,1140,1162]" captionStart="Figure 14" captionStartId="24.[145,224,1817,1839]" captionTargetBox="[148,757,1616,1774]" captionTargetId="figure-808@24.[145,760,1613,1777]" captionTargetPageId="24" captionText="Figure 14. Marionina sp. ‘Snow Is.’ (CE34647). A, chaetal bundle. B, anterior part of body. Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461125" httpUri="https://zenodo.org/record/6461125/files/figure.png" pageId="24" pageNumber="1127">Fig. 14A</figureCitation>
). Upper bundles dorsolateral (closer to lateral line than ventral bundles), with three to four chaetae anterior to clitellum, (three) four in postclitellar segments, at least to XIX. Ventral bundles with four chaetae throughout to XIX. The longest measured chaetae of each worm 55 µm long, ~3 µm wide. Epidermis densely covered with rows of pale gland cells. Clitellum not developed. Head pore at 0/I.
</paragraph>
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Coelomocytes numerous, 10–15 µm long; round, oval or spindle shaped; granulated, with distinct nucleus. Paired pharyngeal glands (
<figureCitation box="[638,750,1477,1499]" captionStart-0="Figure 1" captionStart-1="Figure 2" captionStart-2="Figure 3" captionStart-3="Figure 4" captionStartId-0="6.[145,225,1653,1675]" captionStartId-1="7.[161,239,1469,1491]" captionStartId-2="9.[163,246,967,989]" captionStartId-3="9.[829,915,1077,1099]" captionTargetBox-0="[152,1417,198,1611]" captionTargetBox-1="[170,1434,198,1426]" captionTargetBox-2="[168,774,196,926]" captionTargetBox-3="[830,1438,198,1034]" captionTargetId-0="figure-119@6.[149,1420,195,1614]" captionTargetId-1="figure-156@7.[167,1437,195,1429]" captionTargetId-2="figure-428@9.[167,775,195,927]" captionTargetId-3="figure-376@9.[827,1441,195,1037]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-2="9" captionTargetPageId-3="9" captionText-0="Figure 1. Majority-rule consensus tree for COI estimated with Bayesian inference. Specimens shaded in grey are from the Southern Hemisphere. Specimens in bold represent new species. Support values are posterior probabilities. Scale bar represents the estimated number of substitutions per site." captionText-1="Figure 2. Species tree estimated using the multispecies coalescent model in *BEAST. Specimens shaded in grey are from the Southern Hemisphere. Specimens in bold represent new species. Two support values are presented: first, the posterior probabilities from *BEAST; second, the posterior probabilities from the concatenated MRBAYES tree (see Supporting Information, Fig. S3), where ‘–’ indicates the absence of the node in question. Scale bar shows the expected number of changes per site scaled in accordance with the substitution rate of COI." captionText-2="Figure 3. Lumbricillus antarcticus. A, chaetal bundle (CE12480). B, anterior part of body (CE12479). C, nephridium (CE12480). D, genitalia (CE12479). E, F, spermathecae (CE12479 and CE34641). Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." captionText-3="Figure 4. Lumbricillus antarcticus and Marionina colpites. A–C, L. antarcticus, sectioned specimen BMNH 1933.2.23.899. A, B, spermatheca. C, genitalia. D–F, M. colpites. D, male pore and associated glands (BMNH 1933.2.23.888). E, F, spermatheca (BMNH 1933.2.23.886 and BMNH 1933.2.23.888). Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." figureDoi-0="http://doi.org/10.5281/zenodo.6461087" figureDoi-1="http://doi.org/10.5281/zenodo.6461089" figureDoi-2="http://doi.org/10.5281/zenodo.6461093" figureDoi-3="http://doi.org/10.5281/zenodo.6461095" httpUri-0="https://zenodo.org/record/6461087/files/figure.png" httpUri-1="https://zenodo.org/record/6461089/files/figure.png" httpUri-2="https://zenodo.org/record/6461093/files/figure.png" httpUri-3="https://zenodo.org/record/6461095/files/figure.png" pageId="24" pageNumber="1127">Fig. 14B</figureCitation>
) present in IV, V and VI; only third pair with ventral lobes and dorsally converging but connection not discernible. Origin of dorsal vessel indiscernible, but with peristomial bifurcation. Nephridia observed in 7/8–8/9, anteseptale with funnel only, postseptale elongated oval with terminal efferent duct. Brain with posterior incision, with clusters of perikarya on prostomial nerves and at ventral ends of circumoesophageal connectives (
<figureCitation box="[1227,1312,381,403]" captionStart="Figure 9" captionStartId="19.[828,907,850,872]" captionTargetBox="[830,1438,198,807]" captionTargetId="figure-668@19.[827,1441,195,810]" captionTargetPageId="19" captionText="Figure 9. A, Marionina sp. ‘Snow Is.’ (CE34647), anterior body (lateral view). B, Marionina aestuum, sectioned specimen BMNH 1933.2.23.895, cephalic region (lateral view). C, M. aestuum (CE12477), cephalic region (ventral view). D, Marionina georgiana (paralectotype 13), cephalic region (ventral view). Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461109" httpUri="https://zenodo.org/record/6461109/files/figure.png" pageId="24" pageNumber="1127">Fig. 9A</figureCitation>
). Ventral nerve cord with small subbuccal bulb (
<figureCitation box="[1277,1363,412,434]" captionStart="Figure 9" captionStartId="19.[828,907,850,872]" captionTargetBox="[830,1438,198,807]" captionTargetId="figure-668@19.[827,1441,195,810]" captionTargetPageId="19" captionText="Figure 9. A, Marionina sp. ‘Snow Is.’ (CE34647), anterior body (lateral view). B, Marionina aestuum, sectioned specimen BMNH 1933.2.23.895, cephalic region (lateral view). C, M. aestuum (CE12477), cephalic region (ventral view). D, Marionina georgiana (paralectotype 13), cephalic region (ventral view). Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461109" httpUri="https://zenodo.org/record/6461109/files/figure.png" pageId="24" pageNumber="1127">Fig. 9A</figureCitation>
); see 
<taxonomicName authorityName="Michaelsen" authorityYear="1890" box="[809,934,443,464]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="genus">
<emphasis box="[809,934,443,464]" italics="true" pageId="24" pageNumber="1127">Marionina</emphasis>
</taxonomicName>
discussion above.
</paragraph>
<paragraph blockId="24.[809,1425,197,679]" pageId="24" pageNumber="1127">Male genitalia not fully developed. Developing male cells free floating in XI. Developing sperm funnels in XI, 60–65 µm long, 40 µm wide. Thin line of cells extending backwards from sperm funnels to what will become the male opening. Developing spermathecae in V, shape uncertain. No midventral subneural glands observed.</paragraph>
<paragraph blockId="24.[809,1424,720,772]" pageId="24" pageNumber="1127">
<emphasis box="[809,1128,720,741]" italics="true" pageId="24" pageNumber="1127">Geographical distribution:</emphasis>
<materialsCitation location="Snow Island" pageId="24" pageNumber="1127" specimenCount="1" stateProvince="Shetland Islands">
Known only from 
<location LSID="urn:lsid:plazi:treatment:039DC377FFA8FFCD4C13FD05FED05BD2:8EEB24BAFFA8FFCC4871FD7DFC8C5C19" name="Snow Island" pageId="24" pageNumber="1127" stateProvince="Shetland Islands">Snow Island</location>
(South 
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)
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.
</paragraph>
<paragraph blockId="24.[809,1425,813,1326]" pageId="24" pageNumber="1127">
<emphasis box="[809,920,813,834]" italics="true" pageId="24" pageNumber="1127">Remarks:</emphasis>
The two small specimens of this species were first believed to be juvenile 
<taxonomicName authorityName="Orsted" authorityYear="1844" box="[1214,1369,844,865]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="genus">
<emphasis box="[1214,1369,844,865]" italics="true" pageId="24" pageNumber="1127">Lumbricillus</emphasis>
</taxonomicName>
, but they were placed genetically within a separate clade together with the two larger, darkly pigmented species, 
<taxonomicName authorityName="Stephenson" authorityYear="1932" box="[809,947,936,957]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="aestuum">
<emphasis box="[809,947,936,957]" italics="true" pageId="24" pageNumber="1127">M. aestuum</emphasis>
</taxonomicName>
and 
<taxonomicName box="[1008,1108,936,957]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="fusca">
<emphasis box="[1008,1108,936,957]" italics="true" pageId="24" pageNumber="1127">M. fusca</emphasis>
</taxonomicName>
. Upon closer examination, despite their submature condition, we found our specimens to have free-floating developing male cells, more akin to 
<emphasis box="[961,1122,1028,1049]" italics="true" pageId="24" pageNumber="1127">
<taxonomicName authorityName="Michaelsen" authorityYear="1890" box="[961,1082,1028,1049]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="genus" sensu="stricto">Marionina</taxonomicName>
<taxonomicNameLabel box="[1089,1122,1028,1049]" pageId="24" pageNumber="1127" sensu="stricto">s.s.</taxonomicNameLabel>
</emphasis>
than to 
<taxonomicName authorityName="Orsted" authorityYear="1844" box="[1219,1367,1028,1049]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="genus">
<emphasis box="[1219,1367,1028,1049]" italics="true" pageId="24" pageNumber="1127">Lumbricillus</emphasis>
</taxonomicName>
. Our 
<taxonomicName box="[809,963,1059,1080]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="undetermined">
<emphasis box="[809,929,1059,1080]" italics="true" pageId="24" pageNumber="1127">Marionina</emphasis>
sp.
</taxonomicName>
‘SnowIs.’ isclose to 
<taxonomicName baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[1171,1314,1059,1080]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="georgiana">
<emphasis box="[1171,1314,1059,1080]" italics="true" pageId="24" pageNumber="1127">M.georgiana</emphasis>
</taxonomicName>
inlacking pigmentation, having a densely glandular epidermis, about four chaetae per bundle, similar pharyngeal glands, presence of perikarya on the prostomial nerves and at ventral ends of circumoesophageal connectives, and in the presence of the subbuccal bulb. However, fully mature specimens from Snow Island and/or DNA from 
<taxonomicName baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[869,1017,1274,1295]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="georgiana">
<emphasis box="[869,1017,1274,1295]" italics="true" pageId="24" pageNumber="1127">M. georgiana</emphasis>
</taxonomicName>
will be needed to establish whether the two are the same or separate species.
</paragraph>
<paragraph blockId="24.[842,1392,1402,1458]" pageId="24" pageNumber="1127">
<taxonomicName class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[842,1184,1402,1426]" italics="true" pageId="24" pageNumber="1127">CHRISTENSENIDRILUS BLOCKI</emphasis>
DÓZSA- FARKAS &amp; CONVEY, 1998
</taxonomicName>
</paragraph>
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<paragraph blockId="24.[809,1426,1473,1587]" pageId="24" pageNumber="1127">
<treatmentCitationGroup pageId="24" pageNumber="1127">
<treatmentCitation author="Dozsa-Farkas K &amp; Convey P" page="486" pageId="24" pageNumber="1127" year="1997">
<taxonomicName authority="Dózsa-Farkas &amp; Convey, 1997" box="[809,1419,1473,1495]" class="Clitellata" family="Enchytraeidae" genus="Christensenia " higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[809,1052,1473,1494]" italics="true" pageId="24" pageNumber="1127">Christensenia blocki</emphasis>
<bibRefCitation author="Dozsa-Farkas K &amp; Convey P" pageId="24" pageNumber="1127" pagination="482 - 486" refId="ref23684" refString="Dozsa-Farkas K, Convey P. 1997. Christensenia, a new terrestrial enchytraeid genus from Antarctica. Polar Biology 17: 482 - 486." type="journal article" year="1997">Dózsa-Farkas &amp; Convey, 1997: 483–486</bibRefCitation>
</taxonomicName>
</treatmentCitation>
, figs 1–8.
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</paragraph>
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<treatmentCitationGroup pageId="24" pageNumber="1127">
<taxonomicName authority="(Dózsa-Farkas &amp; Convey, 1997)" box="[833,1119,1534,1555]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[833,1119,1534,1555]" italics="true" pageId="24" pageNumber="1127">Christensenidrilus blocki</emphasis>
</taxonomicName>
– 
<treatmentCitation author="Dozsa-Farkas K &amp; Convey P" page="292" pageId="24" pageNumber="1127" year="1998">
<bibRefCitation author="Dozsa-Farkas K &amp; Convey P" pageId="24" pageNumber="1127" pagination="292" refId="ref23710" refString="Dozsa-Farkas K, Convey P. 1998. Erratum. Christensenia, a new terrestrial enchytraeid genus from Antarctica. Polar Biology 20: 292." type="journal article" year="1998">Dózsa-Farkas &amp; Convey, 1998: 292</bibRefCitation>
</treatmentCitation>
.
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</paragraph>
</subSubSection>
<subSubSection lastPageId="25" lastPageNumber="1128" pageId="24" pageNumber="1127" type="materials_examined">
<paragraph blockId="24.[809,1425,1628,1803]" pageId="24" pageNumber="1127">
<emphasis box="[809,1086,1628,1650]" italics="true" pageId="24" pageNumber="1127">Material examined:</emphasis>
<materialsCitation collectionCode="SMNH" pageId="24" pageNumber="1127" specimenCode="SMNH198141 - 198143, CE35374-35376" specimenCount="1">
<specimenCode box="[1119,1422,1628,1649]" collectionCode="SMNH" country="Sweden" httpUri="http://grbio.org/cool/8z6v-f8is" name="Department of Paleozoology, Swedish Museum of Natural History" pageId="24" pageNumber="1127">SMNH198141 – 198143</specimenCode>
(
<specimenCode box="[816,1012,1658,1680]" collectionCode="CE" pageId="24" pageNumber="1127">CE35374–35376</specimenCode>
)
</materialsCitation>
, 
<materialsCitation collectionCode="CE" country="United Kingdom" county="Supporting Information" location="Signy Island" municipality="GenBank" pageId="24" pageNumber="1127" specimenCount="1">
three mature specimens from the 
<typeStatus box="[809,859,1690,1711]" pageId="24" pageNumber="1127">type</typeStatus>
locality on 
<location LSID="urn:lsid:plazi:treatment:039DC377FFA8FFCD4C13FD05FED05BD2:8EEB24BAFFA8FFCC4EC0F934FB8959B2" box="[993,1140,1689,1711]" country="United Kingdom" county="Supporting Information" municipality="GenBank" name="Signy Island" pageId="24" pageNumber="1127">Signy Island</location>
(
<location LSID="urn:lsid:plazi:treatment:039DC377FFA8FFCD4C13FD05FED05BD2:8EEB24BAFFA8FFCC49A2F934FB3559B2" box="[1155,1224,1689,1711]" country="United Kingdom" county="Supporting Information" municipality="GenBank" name="South" pageId="24" pageNumber="1127">South</location>
<collectingCountry box="[1232,1413,1689,1711]" name="United Kingdom" pageId="24" pageNumber="1127">Orkney Islands</collectingCountry>
). 
<location LSID="urn:lsid:plazi:treatment:039DC377FFA8FFCD4C13FD05FED05BD2:8EEB24BAFFA8FFCC4E08F915FCAD59D0" box="[809,848,1720,1741]" country="United Kingdom" county="Supporting Information" municipality="GenBank" name="For" pageId="24" pageNumber="1127">For</location>
information on collection localities and 
<collectingMunicipality box="[1315,1424,1720,1742]" pageId="24" pageNumber="1127">GenBank</collectingMunicipality>
accession numbers for 
<emphasis box="[1075,1123,1751,1772]" italics="true" pageId="24" pageNumber="1127">COI</emphasis>
barcodes, see 
<tableCitation box="[1288,1374,1750,1772]" captionStart="Table 1" captionStartId="3.[144,208,419,441]" captionTargetPageId="3" captionText="Table 1. Specimens from the Southern Hemisphere used in this study, with specimen identification number, region of the collection site, GenBank accession numbers for the seven genetic markers, and voucher numbers (with museum acronym)" httpUri="http://table.plazi.org/id/DF4B22E9FFB3FFD74DB1FE0EFBE15ECA" pageId="24" pageNumber="1127" tableUuid="DF4B22E9FFB3FFD74DB1FE0EFBE15ECA">Table 1</tableCitation>
and the 
<collectingCounty box="[853,1124,1781,1803]" pageId="24" pageNumber="1127">Supporting Information</collectingCounty>
(
<tableCitation box="[1139,1238,1781,1803]" captionStart="Table 1" captionStartId="3.[144,208,419,441]" captionTargetPageId="3" captionText="Table 1. Specimens from the Southern Hemisphere used in this study, with specimen identification number, region of the collection site, GenBank accession numbers for the seven genetic markers, and voucher numbers (with museum acronym)" httpUri="http://table.plazi.org/id/DF4B22E9FFB3FFD74DB1FE0EFBE15ECA" pageId="24" pageNumber="1127" tableUuid="DF4B22E9FFB3FFD74DB1FE0EFBE15ECA">Table S1</tableCitation>
)
</materialsCitation>
.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.6461125" ID-Zenodo-Dep="6461125" httpUri="https://zenodo.org/record/6461125/files/figure.png" pageId="24" pageNumber="1127" startId="24.[145,224,1817,1839]" targetBox="[148,757,1616,1774]" targetPageId="24">
<paragraph blockId="24.[145,761,1817,1898]" pageId="24" pageNumber="1127">
<emphasis bold="true" box="[145,263,1817,1839]" pageId="24" pageNumber="1127">Figure 14.</emphasis>
<taxonomicName box="[274,422,1817,1839]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="24" pageNumber="1127" phylum="Annelida" rank="species" species="undetermined">
<emphasis box="[274,386,1817,1839]" italics="true" pageId="24" pageNumber="1127">Marionina</emphasis>
sp.
</taxonomicName>
‘Snow Is.’ (CE34647). A, chaetal bundle. B, anterior part of body. Abbreviations are defined under ‘Taxonomy’. Scale bars: 100 μm.
</paragraph>
</caption>
<paragraph blockId="24.[809,1425,1845,1898]" lastBlockId="25.[163,779,197,1231]" lastPageId="25" lastPageNumber="1128" pageId="24" pageNumber="1127">
<emphasis box="[809,920,1845,1866]" italics="true" pageId="24" pageNumber="1127">Remarks:</emphasis>
Our specimens perfectly resemble those originally described by Dózsa-Farkas &amp; Convey; therefore, we decided not to give complete descriptions and illustrations, but instead a few additional comments about the morphology. 
<bibRefCitation author="Schmelz RM &amp; Collado R" pageId="25" pageNumber="1128" pagination="7 - 22" refId="ref25564" refString="Schmelz RM, Collado R. 2008. A type-based redescription of Pachydrilus georgianus Michaelsen, 1888, the type species of Marionina Michaelsen, 1890, with comments on Christensenidrilus Dozsa-Farkas &amp; Convey, 1998 (Enchytraeidae, &quot; Oligochaeta &quot;, Annelida). Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 44: 7 - 22." type="journal article" year="2008">Schmelz &amp; Collado (2008)</bibRefCitation>
, in their comparison between 
<taxonomicName box="[612,725,289,310]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[612,725,289,310]" italics="true" pageId="25" pageNumber="1128">Ch.blocki</emphasis>
</taxonomicName>
and 
<taxonomicName baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[163,315,320,341]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="georgiana">
<emphasis box="[163,315,320,341]" italics="true" pageId="25" pageNumber="1128">M. georgiana</emphasis>
</taxonomicName>
, were left with a few questions that we will try to answer. First, we did not observe neuron perikarya in the prostomium of 
<taxonomicName box="[556,677,381,402]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[556,677,381,402]" italics="true" pageId="25" pageNumber="1128">Ch. blocki</emphasis>
</taxonomicName>
, similar to those described by 
<bibRefCitation author="Schmelz RM &amp; Collado R" box="[430,745,412,434]" pageId="25" pageNumber="1128" pagination="7 - 22" refId="ref25564" refString="Schmelz RM, Collado R. 2008. A type-based redescription of Pachydrilus georgianus Michaelsen, 1888, the type species of Marionina Michaelsen, 1890, with comments on Christensenidrilus Dozsa-Farkas &amp; Convey, 1998 (Enchytraeidae, &quot; Oligochaeta &quot;, Annelida). Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 44: 7 - 22." type="journal article" year="2008">Schmelz &amp; Collado (2008)</bibRefCitation>
in 
<taxonomicName baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[163,315,443,464]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="georgiana">
<emphasis box="[163,315,443,464]" italics="true" pageId="25" pageNumber="1128">M. georgiana</emphasis>
</taxonomicName>
and observed by us also in 
<taxonomicName authorityName="Stephenson" authorityYear="1932" box="[642,779,443,464]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="aestuum">
<emphasis box="[642,779,443,464]" italics="true" pageId="25" pageNumber="1128">M. aestuum</emphasis>
</taxonomicName>
and 
<emphasis box="[214,246,474,495]" italics="true" pageId="25" pageNumber="1128">M.</emphasis>
sp. ‘Snow Is.’. Second, the dorsal blood vessel bifurcates in both species under the brain in a lumbricilline (peristomial) pattern. Third, the penial bulbs of 
<taxonomicName box="[262,377,565,586]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[262,377,565,586]" italics="true" pageId="25" pageNumber="1128">Ch. blocki</emphasis>
</taxonomicName>
, if they can be considered as such, which were described originally as compact and medium sized, were hard to distinguish, composed of a small aggregation of cells (&lt;50 µm in diameter) surrounding the superficial (non-invaginate) male pores, at least apparently similar to the penial bodies of 
<taxonomicName baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[191,342,750,771]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="georgiana">
<emphasis box="[191,342,750,771]" italics="true" pageId="25" pageNumber="1128">M. georgiana</emphasis>
</taxonomicName>
, but lacking any prostate glands. The developing male gametes, although numerous and completely occupying segments X–XII, do not show any sign of being contained in lobed testis sacs. Based on these new elements and the presence of anucleate coelomic corpuscles in place of nucleated coelomocytes in 
<taxonomicName box="[196,316,933,954]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[196,316,933,954]" italics="true" pageId="25" pageNumber="1128">Ch. blocki</emphasis>
</taxonomicName>
, we cannot but agree with 
<bibRefCitation author="Rota E &amp; Matamoros L &amp; Erseus C" pageId="25" pageNumber="1128" pagination="417 - 436" refId="ref25417" refString="Rota E, Matamoros L, Erseus C. 2008. In search of Marionina (Clitellata, Enchytraeidae): a taxonomic history of the genus and redescription of the type species Pachydrilus georgianus Michaelsen, 1888. Italian Journal of Zoology 75: 417 - 436." type="journal article" year="2008">
Rota 
<emphasis box="[718,779,933,955]" italics="true" pageId="25" pageNumber="1128">et al.</emphasis>
(2008)
</bibRefCitation>
and 
<bibRefCitation author="Schmelz RM &amp; Collado R" box="[287,576,964,986]" pageId="25" pageNumber="1128" pagination="7 - 22" refId="ref25564" refString="Schmelz RM, Collado R. 2008. A type-based redescription of Pachydrilus georgianus Michaelsen, 1888, the type species of Marionina Michaelsen, 1890, with comments on Christensenidrilus Dozsa-Farkas &amp; Convey, 1998 (Enchytraeidae, &quot; Oligochaeta &quot;, Annelida). Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 44: 7 - 22." type="journal article" year="2008">Schmelz &amp; Collado (2008)</bibRefCitation>
, in that 
<taxonomicName box="[668,779,964,985]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="blocki">
<emphasis box="[668,779,964,985]" italics="true" pageId="25" pageNumber="1128">Ch. blocki</emphasis>
</taxonomicName>
does not share enough morphological characters with 
<taxonomicName baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[163,323,1026,1047]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="species" species="georgiana">
<emphasis box="[163,323,1026,1047]" italics="true" pageId="25" pageNumber="1128">M. georgiana</emphasis>
</taxonomicName>
to assume it to be closely related to 
<emphasis box="[163,327,1056,1077]" italics="true" pageId="25" pageNumber="1128">
<taxonomicName authorityName="Michaelsen" authorityYear="1890" box="[163,286,1056,1077]" class="Clitellata" family="Enchytraeidae" genus="Marionina" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="genus" sensu="stricto">Marionina</taxonomicName>
<taxonomicNameLabel box="[294,327,1056,1077]" pageId="25" pageNumber="1128" sensu="stricto">s.s.</taxonomicNameLabel>
</emphasis>
, which has also been supported by our molecular data. Unfortunately, we found no support for any specific placement of 
<taxonomicName authorityName="Dozsa-Farkas &amp; Convey" authorityYear="1998" box="[487,698,1117,1138]" class="Clitellata" family="Enchytraeidae" genus="Christensenidrilus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="genus">
<emphasis box="[487,698,1117,1138]" italics="true" pageId="25" pageNumber="1128">Christensenidrilus</emphasis>
</taxonomicName>
within the family 
<taxonomicName box="[284,451,1148,1170]" class="Clitellata" family="Enchytraeidae" kingdom="Animalia" order="Enchytraeida" pageId="25" pageNumber="1128" phylum="Annelida" rank="family">Enchytraeidae</taxonomicName>
as a whole, probably because of the lack of suitable outgroups with which to make comparison.
</paragraph>
</subSubSection>
</treatment>
</document>