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<mods:title id="6229615C5FA4F23C56007AB081855592">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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<subSubSection id="C32E21EAFFBEFFDA4DBFFA05FD165ADD" box="[158,747,1447,1472]" pageId="14" pageNumber="1117" type="nomenclature">
<paragraph id="8B8B7261FFBEFFDA4DBFFA05FD165ADD" blockId="14.[158,747,1447,1472]" box="[158,747,1447,1472]" pageId="14" pageNumber="1117">
<heading id="D0C3C50DFFBEFFDA4DBFFA05FD165ADD" box="[158,747,1447,1472]" centered="true" fontSize="9" level="2" pageId="14" pageNumber="1117" reason="2">
<emphasis id="B940AE73FFBEFFDA4DBFFA05FF5F5AA2" box="[158,162,1448,1471]" italics="true" pageId="14" pageNumber="1117"></emphasis>
<taxonomicName id="4C3409E2FFBEFFDA4D83FA05FD165ADD" ID-CoL="72Q69" authority="BENHAM, 1905" authorityName="Benham" authorityYear="1905" box="[162,747,1447,1472]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" isUncertain="true" kingdom="Animalia" order="Enchytraeida" pageId="14" pageNumber="1108" phylum="Annelida" rank="species" species="macquariensis">
<emphasis id="B940AE73FFBEFFDA4D83FA05FEAE5AA2" box="[162,339,1448,1471]" italics="true" pageId="14" pageNumber="1117">LUMBRICILLUS</emphasis>
CF.
<emphasis id="B940AE73FFBEFFDA4CA2FA06FDC65AA2" box="[387,571,1451,1471]" italics="true" pageId="14" pageNumber="1117">MACQUARIENSIS</emphasis>
<bibRefCitation id="EFA50F90FFBEFFDA4F63FA0AFD165ADD" author="Benham WB" box="[578,747,1447,1472]" pageId="14" pageNumber="1117" pagination="285 - 297" refId="ref23131" refString="Benham WB. 1905. On the Oligochaeta from the southern islands of the New Zealand region. Transactions and Proceedings of the New Zealand Institute 37: 285 - 297." type="journal article" year="1905">BENHAM, 1905</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
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<paragraph id="8B8B7261FFBEFFDA4CBBFA74FE125AEF" blockId="14.[410,495,1497,1522]" box="[410,495,1497,1522]" pageId="14" pageNumber="1117">
(
<figureCitation id="130F6EE4FFBEFFDA4C82FA74FE1A5AEF" box="[419,487,1497,1522]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="14" pageNumber="1117">FIG. 8</figureCitation>
)
</paragraph>
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<paragraph id="8B8B7261FFBEFFDA4DB0F9BCFD045979" blockId="14.[145,768,1553,1758]" pageId="14" pageNumber="1117">
<treatmentCitationGroup id="AB24554FFFBEFFDA4DB0F9BCFD04593A" box="[145,761,1553,1575]" pageId="14" pageNumber="1117">
?
<treatmentCitation id="0A955470FFBEFFDA4DBDF9BCFD08593A" author="Benham WB" box="[156,757,1553,1575]" page="297" pageId="14" pageNumber="1117" year="1905">
<taxonomicName id="4C3409E2FFBEFFDA4DBDF9BCFD08593A" ID-CoL="72Q69" authority="Benham, 1905: 295 - 297" authorityName="Benham" authorityPageNumber="295 - 297" authorityYear="1905" box="[156,757,1553,1575]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="14" pageNumber="1117" phylum="Annelida" rank="species" species="macquariensis">
<emphasis id="B940AE73FFBEFFDA4DBDF9BCFE26593A" box="[156,475,1553,1575]" italics="true" pageId="14" pageNumber="1117">Lumbricillus macquariensis</emphasis>
<bibRefCitation id="EFA50F90FFBEFFDA4CC2F9BCFD08593A" author="Benham WB" box="[483,757,1553,1575]" pageId="14" pageNumber="1117" pagination="285 - 297" refId="ref23131" refString="Benham WB. 1905. On the Oligochaeta from the southern islands of the New Zealand region. Transactions and Proceedings of the New Zealand Institute 37: 285 - 297." type="journal article" year="1905">Benham, 1905: 295297</bibRefCitation>
</taxonomicName>
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,
</treatmentCitationGroup>
pl. XIV, figs 8, 1113;
<bibRefCitation id="EFA50F90FFBEFFDA4CEBF99DFD075958" author="Benham WB" box="[458,762,1584,1606]" pageId="14" pageNumber="1117" pagination="189 - 191" refId="ref23191" refString="Benham WB. 1915. On Lumbricillus macquariensis Benham. Transactions and Proceedings of the New Zealand Institute, Wellington 47: 189 - 191." type="journal article" year="1915">Benham, 1915: 189191</bibRefCitation>
;
<bibRefCitation id="EFA50F90FFBEFFDA4D92F9E3FE8F5979" author="Benham WB" box="[179,370,1614,1636]" pageId="14" pageNumber="1117" pagination="1 - 38" refId="ref23217" refString="Benham WB. 1922. Oligochaeta of Macquarie Island. Australasian Antarctic expedition 1911 - 14. Scientific Reports C 6: 1 - 38." type="journal article" year="1922">Benham, 1922: 6</bibRefCitation>
;
<bibRefCitation id="EFA50F90FFBEFFDA4C5DF9E3FD4F5979" author="Stephenson J" box="[380,690,1614,1636]" pageId="14" pageNumber="1117" pagination="233 - 264" refId="ref25867" refString="Stephenson J. 1932. Oligochaeta. Part. I. Microdrili. Discovery Reports 4: 233 - 264." type="journal article" year="1932">Stephenson, 1932: 254255</bibRefCitation>
, fig. 7.
</paragraph>
<paragraph id="8B8B7261FFBEFFDA4DB0F9C0FEC259BC" blockId="14.[145,768,1553,1758]" pageId="14" pageNumber="1117">
?
<taxonomicName id="4C3409E2FFBEFFDA4DBDF9C0FD2E599E" ID-CoL="85594" authority="Benham, 1909: 261 - 262" authorityName="Benham" authorityPageNumber="261 - 262" authorityYear="1909" box="[156,723,1645,1667]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="14" pageNumber="1117" phylum="Annelida" rank="species" species="intermedius">
<emphasis id="B940AE73FFBEFFDA4DBDF9C0FE40599F" box="[156,445,1645,1666]" italics="true" pageId="14" pageNumber="1117">Lumbricillus intermedius</emphasis>
<bibRefCitation id="EFA50F90FFBEFFDA4CE2F9C0FD2E599E" author="Benham WB" box="[451,723,1645,1667]" pageId="14" pageNumber="1117" pagination="251 - 294" refId="ref23163" refString="Benham WB. 1909. Report on Oligochaeta of the subantarctic islands of New Zealand. The Subantarctic Islands of New Zealand 1: 251 - 294." type="journal article" year="1909">Benham, 1909: 261262</bibRefCitation>
</taxonomicName>
, pl. X, figs 811.
</paragraph>
<paragraph id="8B8B7261FFBEFFDA4DB0F906FEE159C3" blockId="14.[145,768,1553,1758]" pageId="14" pageNumber="1117">
?
<taxonomicName id="4C3409E2FFBEFFDA4DBFF907FE1E59A2" authority="(Benham, 1905)" box="[158,483,1706,1727]" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" kingdom="Animalia" order="Enchytraeida" pageId="14" pageNumber="1117" phylum="Annelida" rank="species" species="intermedius">
<emphasis id="B940AE73FFBEFFDA4DBFF907FE1E59A2" box="[158,483,1706,1727]" italics="true" pageId="14" pageNumber="1117">Pachydrilus intermedius</emphasis>
</taxonomicName>
<bibRefCitation id="EFA50F90FFBEFFDA4F2AF907FD0759DD" author="Michaelsen W" box="[523,762,1706,1728]" pageId="14" pageNumber="1117" pagination="197 - 240" refId="ref24596" refString="Michaelsen W. 1924. Papers from Dr. Th. Mortensen's Pacific Expedition 1914 - 16. XVII. Oligochaten von Neuseeland und den Auckland-Campbell-Inseln, Nebst einigen anderen Pacifischen Formen. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i KObenhavn 75: 197 - 240." type="journal article" year="1924">Michaelsen, 1924:</bibRefCitation>
197199.
</paragraph>
<paragraph id="8B8B7261FFBEFFDA4DB0F8B5FBBA5B35" blockId="14.[145,761,1816,1899]" lastBlockId="14.[809,1425,951,1065]" pageId="14" pageNumber="1117">
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<emphasis id="B940AE73FFBEFFDA4DB0F8B5FEC75830" box="[145,314,1816,1837]" italics="true" pageId="14" pageNumber="1117">
<typeStatus id="548FCCC3FFBEFFDA4DB0F8B5FF3A5830" box="[145,199,1816,1837]" pageId="14" pageNumber="1117">Type</typeStatus>
material:
</emphasis>
<location id="8EEB24BAFFBEFFDA4C71F8B5FE8C5830" LSID="urn:lsid:plazi:treatment:039DC377FFBEFFC44DBFFA05FEA95D56:8EEB24BAFFBEFFDA4C71F8B5FE8C5830" box="[336,369,1816,1837]" country="New Zealand" county="Our" municipality="Professor Parker" name="No" pageId="14" pageNumber="1117" stateProvince="Otago">No</location>
information (
<bibRefCitation id="EFA50F90FFBEFFDA4F34F8B5FF345851" author="Reynolds JW &amp; Wetzel MJ" pageId="14" pageNumber="1117" refId="ref25010" refString="Reynolds JW, Wetzel MJ. 2019. Nomenclatura Oligochaetologica - A catalogue of names, descriptions and type specimens. Editio Secunda. Available at: https: // www. inhs. illinois. edu / people / mjwetzel / nomenoligo" type="url" year="2019">Reynolds &amp; Wetzel, 2019</bibRefCitation>
).
<location id="8EEB24BAFFBEFFDA4DFEF89BFEF15851" LSID="urn:lsid:plazi:treatment:039DC377FFBEFFC44DBFFA05FEA95D56:8EEB24BAFFBEFFDA4DFEF89BFEF15851" box="[223,268,1846,1868]" country="New Zealand" county="Our" municipality="Professor Parker" name="The" pageId="14" pageNumber="1117" stateProvince="Otago">The</location>
material is probably located in the
<collectingRegion id="49F0BC83FFBEFFDA4F92F89AFD055851" box="[691,760,1847,1868]" country="New Zealand" name="Otago" pageId="14" pageNumber="1117">Otago</collectingRegion>
<location id="8EEB24BAFFBEFFDA4DB0F8FBFF075876" LSID="urn:lsid:plazi:treatment:039DC377FFBEFFC44DBFFA05FEA95D56:8EEB24BAFFBEFFDA4DB0F8FBFF075876" box="[145,250,1878,1899]" country="New Zealand" county="Our" municipality="Professor Parker" name="Museum" pageId="14" pageNumber="1117" stateProvince="Otago">Museum</location>
,
<collectingCountry id="F32332F1FFBEFFDA4C29F8FBFE555876" box="[264,424,1877,1899]" name="New Zealand" pageId="14" pageNumber="1117">New Zealand</collectingCountry>
, because
<location id="8EEB24BAFFBEFFDA4F3FF8F8FD7F5876" LSID="urn:lsid:plazi:treatment:039DC377FFBEFFC44DBFFA05FEA95D56:8EEB24BAFFBEFFDA4F3FF8F8FD7F5876" box="[542,642,1877,1899]" country="New Zealand" county="Our" municipality="Professor Parker" name="Benham" pageId="14" pageNumber="1117" stateProvince="Otago">Benham</location>
mentions material being deposited and registered in the
<location id="8EEB24BAFFBEFFDA4E08FC7BFC715CF6" LSID="urn:lsid:plazi:treatment:039DC377FFBEFFC44DBFFA05FEA95D56:8EEB24BAFFBEFFDA4E08FC7BFC715CF6" box="[809,908,982,1003]" country="New Zealand" county="Our" municipality="Professor Parker" name="Museum" pageId="14" pageNumber="1117" stateProvince="Otago">Museum</location>
by a
<collectingMunicipality id="6BEFE81BFFBEFFDA4EE6FC78FB7B5CF6" box="[967,1158,981,1003]" pageId="14" pageNumber="1117">Professor Parker</collectingMunicipality>
, who was curator of the
<collectingRegion id="49F0BC83FFBEFFDA4E08FC59FC905B14" box="[809,877,1012,1033]" country="New Zealand" name="Otago" pageId="14" pageNumber="1117">Otago</collectingRegion>
museum collections at the time.
<collectingCounty id="62EA0AEDFFBEFFDA49C1FC59FAF05B14" box="[1248,1293,1012,1033]" pageId="14" pageNumber="1117">Our</collectingCounty>
inquiry did not receive any response
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B7261FFBEFFDA4E08FBFFFB3D5B9B" blockId="14.[809,1425,1105,1158]" pageId="14" pageNumber="1117">
<materialsCitation id="3B5C783CFFBEFFDA4E08FBFFFB415B9B" ID-GBIF-Occurrence="3751562307" country="Australia" location="Macquarie Island" pageId="14" pageNumber="1117" specimenCount="1" typeStatus="holotype">
<emphasis id="B940AE73FFBEFFDA4E08FBFFFC3D5B7B" box="[809,960,1105,1127]" italics="true" pageId="14" pageNumber="1117">
<typeStatus id="548FCCC3FFBEFFDA4E08FBFFFCA35B7A" box="[809,862,1106,1127]" pageId="14" pageNumber="1117">Type</typeStatus>
locality:
</emphasis>
<collectingCountry id="F32332F1FFBEFFDA4EF5FBFCFB5D5B7A" box="[980,1184,1105,1127]" name="Australia" pageId="14" pageNumber="1117">Macquarie Island</collectingCountry>
, brackish pools, with planarians and
<taxonomicName id="4C3409E2FFBEFFDA4EFCFBDDFBA35B98" ID-CoL="7HND" authorityName="G. B. Sowerby I" authorityYear="1823" box="[989,1118,1136,1157]" class="Chytridiomycetes" family="Chytriomycetaceae" genus="Siphonaria" kingdom="Fungi" order="Chytridiales" pageId="14" pageNumber="1117" phylum="Chytridiomycota" rank="genus">
<emphasis id="B940AE73FFBEFFDA4EFCFBDDFBA35B98" box="[989,1118,1136,1157]" italics="true" pageId="14" pageNumber="1117">Siphonaria</emphasis>
</taxonomicName>
limpets
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B7261FFBEFFDA4E08FB02FBC05A86" blockId="14.[809,1424,1199,1435]" pageId="14" pageNumber="1117">
<emphasis id="B940AE73FFBEFFDA4E08FB02FBD55BD9" box="[809,1064,1199,1220]" italics="true" pageId="14" pageNumber="1117">Material examined:</emphasis>
<materialsCitation id="3B5C783CFFBEFFDA4963FB02FB8F5BFE" ID-GBIF-Occurrence="3751562303" collectionCode="SMNH, CE" pageId="14" pageNumber="1117" specimenCode="SMNH198150, CE12483" specimenCount="1">
<specimenCode id="DB92DA1AFFBEFFDA4963FB02FB085BD8" box="[1090,1269,1199,1221]" collectionCode="SMNH" country="Sweden" httpUri="http://grbio.org/cool/8z6v-f8is" name="Department of Paleozoology, Swedish Museum of Natural History" pageId="14" pageNumber="1117">SMNH198150</specimenCode>
(
<specimenCode id="DB92DA1AFFBEFFDA482BFB02FA7F5BD8" box="[1290,1410,1199,1221]" collectionCode="CE" pageId="14" pageNumber="1117">CE12483</specimenCode>
), one mature specimen, and
</materialsCitation>
<materialsCitation id="3B5C783CFFBEFFDA495DFB60FBC55A86" ID-GBIF-Occurrence="3751562304" collectionCode="SMNH, CE" country="South Georgia and the South Sandwich Islands" county="Supporting Information" location="For" municipality="GenBank" pageId="14" pageNumber="1117" specimenCode="SMNH198151-198153, CE12484" specimenCount="1" specimenCount-immature="1">
<specimenCode id="DB92DA1AFFBEFFDA495DFB60FA725BFE" box="[1148,1423,1229,1251]" collectionCode="SMNH" country="Sweden" httpUri="http://grbio.org/cool/8z6v-f8is" name="Department of Paleozoology, Swedish Museum of Natural History" pageId="14" pageNumber="1117">SMNH198151198153</specimenCode>
(
<specimenCode id="DB92DA1AFFBEFFDA4E10FB41FC555A1F" box="[817,936,1260,1282]" collectionCode="CE" pageId="14" pageNumber="1117">CE12484</specimenCode>
CE12486),
<specimenCount id="9D32B9E8FFBEFFDA496BFB41FAE55A1F" box="[1098,1304,1260,1282]" pageId="14" pageNumber="1117" type="immature">three immature</specimenCount>
or semimature specimens, all collected in 2010 from
<collectingCountry id="F32332F1FFBEFFDA486BFAA6FC6D5A22" name="South Georgia and the South Sandwich Islands" pageId="14" pageNumber="1117">South Georgia</collectingCountry>
.
<location id="8EEB24BAFFBEFFDA4E81FA87FC375A22" LSID="urn:lsid:plazi:treatment:039DC377FFBEFFC44DBFFA05FEA95D56:8EEB24BAFFBEFFDA4E81FA87FC375A22" box="[928,970,1322,1343]" country="South Georgia and the South Sandwich Islands" county="Supporting Information" municipality="GenBank" name="For" pageId="14" pageNumber="1117">For</location>
details of collection and
<collectingMunicipality id="6BEFE81BFFBEFFDA483AFA84FA725A22" box="[1307,1423,1321,1343]" pageId="14" pageNumber="1117">GenBank</collectingMunicipality>
accession numbers for
<emphasis id="B940AE73FFBEFFDA4979FAE5FB775A40" box="[1112,1162,1352,1373]" italics="true" pageId="14" pageNumber="1117">COI</emphasis>
barcodes and other gene sequences, see
<tableCitation id="C6B647DAFFBEFFDA490EFACAFB705A61" box="[1071,1165,1383,1405]" 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="14" pageNumber="1117" tableUuid="DF4B22E9FFB3FFD74DB1FE0EFBE15ECA">Table 1</tableCitation>
and the
<collectingCounty id="62EA0AEDFFBEFFDA4825FACAFC4B5A86" pageId="14" pageNumber="1117">Supporting Information</collectingCounty>
(
<tableCitation id="C6B647DAFFBEFFDA4EE7FA28FBD15A86" box="[966,1068,1413,1435]" 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="14" pageNumber="1117" tableUuid="DF4B22E9FFB3FFD74DB1FE0EFBE15ECA">Table S1</tableCitation>
)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8B7261FFBEFFDA4E08FA68FBBA5874" blockId="14.[809,1426,1476,1897]" pageId="14" pageNumber="1117">
<emphasis id="B940AE73FFBEFFDA4E08FA68FC435AC7" box="[809,958,1477,1498]" italics="true" pageId="14" pageNumber="1117">Description:</emphasis>
Length of first 1548 segments
<quantity id="4CCCDF84FFBEFFDA487FFA69FC775AE5" metricMagnitude="-3" metricUnit="m" metricValue="4.8" metricValueMax="7.1" metricValueMin="2.5" pageId="14" pageNumber="1117" unit="mm" value="4.8" valueMax="7.1" valueMin="2.5">2.5 7.1 mm</quantity>
(fixed, amputated specimens); first 15 segments 2.53.0 mm long; width at clitellum
<quantity id="4CCCDF84FFBEFFDA4870F9ACFC6E592B" metricMagnitude="-4" metricUnit="m" metricValue="9.75" metricValueMax="11.0" metricValueMin="8.5" pageId="14" pageNumber="1117" unit="mm" value="0.975" valueMax="1.1" valueMin="0.85">0.85 1.10 mm</quantity>
. Chaetae slightly sigmoid (
<figureCitation id="130F6EE4FFBEFFDA49F8F98DFAD25928" box="[1241,1327,1568,1590]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="14" pageNumber="1117">Fig. 8A</figureCitation>
). Upper bundles dorsolateral (closer to lateral line than the ventral bundles), with four to six chaetae anterior to clitellum, and three to four chaetae in postclitellar segments, at least to segment XLVIII. Ventral bundles with (four) five to seven chaetae anterior to clitellum, and four to six chaetae posteriorly. The longest measured chaetae of each worm 135145 µm long, ~67 µm wide. Epidermis loosely covered with rows of pale gland cells. Clitellum not fully developed. Head pore not observed.
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4D9AFF68FDCD5D56" blockId="15.[163,780,197,1354]" pageId="15" pageNumber="1118">
Coelomocytes numerous, 1530 µm long; round, oval or spindle shaped; granulated with distinct nucleus. Paired pharyngeal glands (
<figureCitation id="130F6EE4FFBFFFDB4F6CFEAFFD5C5E05" box="[589,673,258,280]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="15" pageNumber="1118">Fig. 8B</figureCitation>
) present in IV, V and VI, with third pair extending back into VII; each pair converging dorsally, but connections not discernible; dorsal lobes of about equal size, ventral lobes in IVVI, increasing in size from IV to VI, absent in VII. Dorsal vessel originating in XVIXVII, with peristomial bifurcation. Nephridia (
<figureCitation id="130F6EE4FFBFFFDB4F79FE17FD525ECD" box="[600,687,442,464]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="15" pageNumber="1118">Fig. 8C</figureCitation>
) ~110 190 µm long, observed in 7/89/10 and postclitellar segments. Anteseptale small, consisting of funnel only. Postseptale oval, tapering into posteroventral efferent duct. Brain with posterior incision.
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4D9AFDF9FD535CA6" blockId="15.[163,780,197,1354]" pageId="15" pageNumber="1118">
Male genitalia paired (
<figureCitation id="130F6EE4FFBFFFDB4C9CFDF9FDF05D74" box="[445,525,596,618]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="15" pageNumber="1118">Fig. 8E</figureCitation>
). Testes originating in anterior of XI, with testis sacs forming regular lobes extending forwards into X, but these lobes are fingerlike, thats is, much thinner than those of most species of
<taxonomicName id="4C3409E2FFBFFFDB4D9FFD63FEAF5DFE" ID-CoL="84PT8" authorityName="Orsted" authorityYear="1844" box="[190,338,718,739]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="genus">
<emphasis id="B940AE73FFBFFFDB4D9FFD63FEAF5DFE" box="[190,338,718,739]" italics="true" pageId="15" pageNumber="1118">Lumbricillus</emphasis>
</taxonomicName>
. Sperm funnels in XI, in
<specimenCount id="9D32B9E8FFBFFFDB4F53FD62FCF65DF9" box="[626,779,718,740]" pageId="15" pageNumber="1118" type="generic">one specimen</specimenCount>
extending backwards into XII (see
<figureCitation id="130F6EE4FFBFFFDB4F67FD40FD665C1F" box="[582,667,749,771]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="15" pageNumber="1118">Fig. 8E</figureCitation>
), 425 µm long, 155 µm wide, making them about three times as long as wide; funnels tapering towards vasa deferentia. Most of vasa irregularly coiled in XII, 25 µm wide. Penial bulbs round or pear shaped, 125 µm in diameter, discharging into deep invagination of body wall (
<figureCitation id="130F6EE4FFBFFFDB4FC1FC2BFF3F5CA7" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="15" pageNumber="1118">Fig. 8E</figureCitation>
). Ovaries in XII. No mature eggs observed.
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4D9AFC69FE2A5A54" blockId="15.[163,780,197,1354]" pageId="15" pageNumber="1118">
Spermathecae (
<figureCitation id="130F6EE4FFBFFFDB4C59FC69FE325CC4" box="[376,463,964,986]" captionStart="Figure 8" captionStartId="14.[809,886,730,752]" captionTargetBox="[812,1421,198,688]" captionTargetId="figure-484@14.[809,1424,195,691]" captionTargetPageId="14" captionText="Figure 8. Lumbricillus cf.macquariensis. A, chaetal bundle (CE12486).B, anterior part of body (CE12485).C, nephridium (CE12484). D, spermatheca (CE12483). E, genitalia (CE12483); the testis sacs are illustrated from another specimen (CE12486) but with the same scale. Abbreviations are defined under Taxonomy. Scale bars: 100 μm." figureDoi="http://doi.org/10.5281/zenodo.6461107" httpUri="https://zenodo.org/record/6461107/files/figure.png" pageId="15" pageNumber="1118">Fig. 8D</figureCitation>
) in V, pouch shaped, with short ectal duct gradually widening into ampulla. Ampulla with thicker epithelium in the ectal parts, which transitions to thinner epithelium in the more ental parts. Ental part seemingly connected with oesophagus. No sperm observed. Spermathecae 340 µm long, 55 µm wide at the ectal duct, 120 µm wide at widest part of ampulla. Gland cells surrounding ectal duct, forming compact mass, glandular body 125 µm in diameter at its widest part. In the mature specimen, we observed only one midventral subneural gland, 190 µm long, in XIV, but said specimen was amputated and ended in this segment.
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4D82FA20FE265985" blockId="15.[163,780,1421,1688]" pageId="15" pageNumber="1118">
<emphasis id="B940AE73FFBFFFDB4D82FA20FE0B5ABF" box="[163,502,1421,1442]" italics="true" pageId="15" pageNumber="1118">Geographical distribution:</emphasis>
Our specimens were collected from
<collectingCountry id="F32332F1FFBFFFDB4C57FA01FD5F5ADF" box="[374,674,1452,1474]" name="South Georgia Island" pageId="15" pageNumber="1118">South Georgia Island</collectingCountry>
, where
<taxonomicName id="4C3409E2FFBFFFDB4D82FA66FE955AFD" ID-CoL="72Q69" authorityName="Benham" authorityYear="1905" box="[163,360,1483,1504]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="macquariensis">
<emphasis id="B940AE73FFBFFFDB4D82FA66FE955AFD" box="[163,360,1483,1504]" italics="true" pageId="15" pageNumber="1118">L. macquariensis</emphasis>
</taxonomicName>
was recorded by
<bibRefCitation id="EFA50F90FFBFFFDB4F0EFA67FCF85AFD" author="Stephenson J" box="[559,773,1482,1504]" pageId="15" pageNumber="1118" pagination="233 - 264" refId="ref25867" refString="Stephenson J. 1932. Oligochaeta. Part. I. Microdrili. Discovery Reports 4: 233 - 264." type="journal article" year="1932">Stephenson (1932)</bibRefCitation>
. However, the species was originally described from specimens that came from
<collectingCountry id="F32332F1FFBFFFDB4CEEF9A5FD6A5903" box="[463,663,1544,1566]" name="Australia" pageId="15" pageNumber="1118">Macquarie Island</collectingCountry>
(
<bibRefCitation id="EFA50F90FFBFFFDB4F85F9A5FF275921" author="Benham WB" pageId="15" pageNumber="1118" pagination="285 - 297" refId="ref23131" refString="Benham WB. 1905. On the Oligochaeta from the southern islands of the New Zealand region. Transactions and Proceedings of the New Zealand Institute 37: 285 - 297." type="journal article" year="1905">Benham, 1905</bibRefCitation>
) and since then has also been reported from the Campbell and Auckland Islands (
<bibRefCitation id="EFA50F90FFBFFFDB4F69F9E8FD075947" author="Benham WB" box="[584,762,1605,1627]" pageId="15" pageNumber="1118" pagination="1 - 38" refId="ref23217" refString="Benham WB. 1922. Oligochaeta of Macquarie Island. Australasian Antarctic expedition 1911 - 14. Scientific Reports C 6: 1 - 38." type="journal article" year="1922">Benham, 1922</bibRefCitation>
),
<collectingCountry id="F32332F1FFBFFFDB4D82F9C9FE045967" box="[163,505,1636,1658]" name="Heard Island and McDonald Islands" pageId="15" pageNumber="1118">Heard and McDonald Islands</collectingCountry>
(
<bibRefCitation id="EFA50F90FFBFFFDB4F28F9C9FD855967" author="Lee KE" box="[521,632,1636,1658]" pageId="15" pageNumber="1118" pagination="149 - 165" refId="ref24151" refString="Lee KE. 1968. Oligochaeta from subantarctic islands. British, Australian and New Zealand Antarctic Research Expedition 1929 - 1931 Reports Series B (Zoology and Botany) 8: 149 - 165." type="journal article" year="1968">Lee, 1968</bibRefCitation>
) and Bishop Island (
<bibRefCitation id="EFA50F90FFBFFFDB4DDAF92FFE36598A" author="Davies KF &amp; Greenslade P &amp; Melbourne BA" box="[251,459,1666,1688]" pageId="15" pageNumber="1118" pagination="455 - 458" refId="ref23552" refString="Davies KF, Greenslade P, Melbourne BA. 1997. The invertebrates of sub-antarctic Bishop Island. Polar Biology 17: 455 - 458." type="journal article" year="1997">
Davies
<emphasis id="B940AE73FFBFFFDB4C6EF92EFE75598A" box="[335,392,1666,1688]" italics="true" pageId="15" pageNumber="1118">et al.</emphasis>
, 1997
</bibRefCitation>
).
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4D82F96CFC1F5B6F" blockId="15.[163,780,1729,1904]" lastBlockId="15.[827,1444,197,1905]" pageId="15" pageNumber="1118">
<emphasis id="B940AE73FFBFFFDB4D82F96CFEE759CB" box="[163,282,1729,1750]" italics="true" pageId="15" pageNumber="1118">Remarks:</emphasis>
Our specimens are similar to those identified as
<taxonomicName id="4C3409E2FFBFFFDB4C1CF94DFDF859E8" ID-CoL="72Q69" authorityName="Benham" authorityYear="1905" box="[317,517,1760,1781]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="macquariensis">
<emphasis id="B940AE73FFBFFFDB4C1CF94DFDF859E8" box="[317,517,1760,1781]" italics="true" pageId="15" pageNumber="1118">L. macquariensis</emphasis>
</taxonomicName>
by
<bibRefCitation id="EFA50F90FFBFFFDB4F10F94DFCF659EB" author="Stephenson J" box="[561,779,1760,1782]" pageId="15" pageNumber="1118" pagination="233 - 264" refId="ref25867" refString="Stephenson J. 1932. Oligochaeta. Part. I. Microdrili. Discovery Reports 4: 233 - 264." type="journal article" year="1932">Stephenson (1932)</bibRefCitation>
from
<collectingCountry id="F32332F1FFBFFFDB4DFEF953FE2B5809" box="[223,470,1790,1812]" name="South Georgia Island" pageId="15" pageNumber="1118">South Georgia Island</collectingCountry>
. In the original description from
<collectingCountry id="F32332F1FFBFFFDB4DC0F8B0FE50582E" box="[225,429,1821,1843]" name="Australia" pageId="15" pageNumber="1118">Macquarie Island</collectingCountry>
(which lies south of
<collectingCountry id="F32332F1FFBFFFDB4FBCF8B0FCF7582E" box="[669,778,1821,1843]" name="Australia" pageId="15" pageNumber="1118">Australia</collectingCountry>
and
<collectingCountry id="F32332F1FFBFFFDB4DFDF891FE7B584F" box="[220,390,1852,1874]" name="New Zealand" pageId="15" pageNumber="1118">New Zealand</collectingCountry>
),
<bibRefCitation id="EFA50F90FFBFFFDB4CBEF891FD9D584C" author="Benham WB" box="[415,608,1851,1874]" pageId="15" pageNumber="1118" pagination="285 - 297" refId="ref23131" refString="Benham WB. 1905. On the Oligochaeta from the southern islands of the New Zealand region. Transactions and Proceedings of the New Zealand Institute 37: 285 - 297." type="journal article" year="1905">Benham (1905)</bibRefCitation>
illustrated a spermatheca with a narrow ental duct connecting the sac-like ampulla with the oesophagus.
<bibRefCitation id="EFA50F90FFBFFFDB49D5FF68FA5E5FC6" author="Benham WB" box="[1268,1443,197,219]" pageId="15" pageNumber="1118" pagination="251 - 294" refId="ref23163" refString="Benham WB. 1909. Report on Oligochaeta of the subantarctic islands of New Zealand. The Subantarctic Islands of New Zealand 1: 251 - 294." type="journal article" year="1909">Benham (1909)</bibRefCitation>
described
<taxonomicName id="4C3409E2FFBFFFDB4E8AFF49FBB05FE4" authority="Benham, 1909" box="[939,1101,228,249]" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="intermedius">
<emphasis id="B940AE73FFBFFFDB4E8AFF49FBB05FE4" box="[939,1101,228,249]" italics="true" pageId="15" pageNumber="1118">L. intermedius</emphasis>
</taxonomicName>
(from
<collectingRegion id="49F0BC83FFBFFFDB49B2FF49FB025FE7" box="[1171,1279,228,250]" country="New Zealand" name="Auckland" pageId="15" pageNumber="1118">Auckland</collectingRegion>
Island), which he considered as intermediate between
<taxonomicName id="4C3409E2FFBFFFDB482DFEAFFB705E2B" ID-CoL="72Q66" authority="(Michaelsen, 1888)" baseAuthorityName="Michaelsen" baseAuthorityYear="1888" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="maximus">
<emphasis id="B940AE73FFBFFFDB482DFEAFFC5B5E2A" italics="true" pageId="15" pageNumber="1118">Lumbricillus maximus</emphasis>
(
<bibRefCitation id="EFA50F90FFBFFFDB4E99FE8CFB795E2B" author="Michaelsen W" box="[952,1156,289,311]" pageId="15" pageNumber="1118" pagination="53 - 73" refId="ref24401" refString="Michaelsen W. 1888. Die Oligochaeten von Sud-Georgien nach der Ausbeute der Deutschen Station von 1882 - 1883. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 5: 53 - 73." type="journal article" year="1888">Michaelsen, 1888</bibRefCitation>
)
</taxonomicName>
and
<taxonomicName id="4C3409E2FFBFFFDB49EAFE8FFA9A5E2A" ID-CoL="4BRXM" baseAuthorityName="Claparede" baseAuthorityYear="1861" box="[1227,1383,290,311]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="verrucosus">
<emphasis id="B940AE73FFBFFFDB49EAFE8FFA9A5E2A" box="[1227,1383,290,311]" italics="true" pageId="15" pageNumber="1118">L. verrucosus</emphasis>
</taxonomicName>
, and which has a spermatheca with a small pore connecting the ampulla to the oesophagus without any narrow ental duct. However,
<bibRefCitation id="EFA50F90FFBFFFDB490CFED0FB205E8E" author="Benham WB" box="[1069,1245,381,403]" pageId="15" pageNumber="1118" pagination="189 - 191" refId="ref23191" refString="Benham WB. 1915. On Lumbricillus macquariensis Benham. Transactions and Proceedings of the New Zealand Institute, Wellington 47: 189 - 191." type="journal article" year="1915">Benham (1915)</bibRefCitation>
revisited his two species (
<taxonomicName id="4C3409E2FFBFFFDB4E82FE31FB8F5EAC" ID-CoL="72Q69" authorityName="Benham" authorityYear="1905" box="[931,1138,412,433]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="macquariensis">
<emphasis id="B940AE73FFBFFFDB4E82FE31FB8F5EAC" box="[931,1138,412,433]" italics="true" pageId="15" pageNumber="1118">L. macquariensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C3409E2FFBFFFDB4993FE31FA995EAD" authority="Benham, 1909" box="[1202,1380,411,433]" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="intermedius">
<emphasis id="B940AE73FFBFFFDB4993FE31FA995EAD" box="[1202,1380,411,433]" italics="true" pageId="15" pageNumber="1118">L. intermedius</emphasis>
</taxonomicName>
) and concluded that they were, in fact, the same, making
<taxonomicName id="4C3409E2FFBFFFDB4E1AFE77FC1E5EF3" authority="Benham, 1909" box="[827,995,473,495]" class="Clitellata" family="Enchytraeidae" genus="Pachydrilus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="intermedius">
<emphasis id="B940AE73FFBFFFDB4E1AFE77FC1E5EF3" box="[827,995,473,495]" italics="true" pageId="15" pageNumber="1118">L. intermedius</emphasis>
</taxonomicName>
a junior synonym of
<emphasis id="B940AE73FFBFFFDB49F6FE77FA5E5EF3" box="[1239,1443,473,495]" italics="true" pageId="15" pageNumber="1118">
<taxonomicName id="4C3409E2FFBFFFDB49F6FE77FA625EF3" ID-CoL="72Q69" authorityName="Benham" authorityYear="1905" box="[1239,1439,473,495]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="macquariensis">L. macquariensis</taxonomicName>
.
</emphasis>
He also concluded that in both samples the spermatheca did not have a narrow ental duct, but a pore connecting it directly to the oesophagus. The spermatheca illustrated by
<bibRefCitation id="EFA50F90FFBFFFDB4ED7FDFEFB4A5D77" author="Benham WB" box="[1014,1207,595,618]" pageId="15" pageNumber="1118" pagination="251 - 294" refId="ref23163" refString="Benham WB. 1909. Report on Oligochaeta of the subantarctic islands of New Zealand. The Subantarctic Islands of New Zealand 1: 251 - 294." type="journal article" year="1909">Benham (1909)</bibRefCitation>
is similar to that described by
<bibRefCitation id="EFA50F90FFBFFFDB4EF0FDDFFB5B5D95" author="Stephenson J" box="[977,1190,626,648]" pageId="15" pageNumber="1118" pagination="233 - 264" refId="ref25867" refString="Stephenson J. 1932. Oligochaeta. Part. I. Microdrili. Discovery Reports 4: 233 - 264." type="journal article" year="1932">Stephenson (1932)</bibRefCitation>
; both authors showed a thicker epithelium in the ectal part, much like in our specimens. This change in height of the duct epithelium and its expansion to merge with the ampulla are also reminiscent of the spermathecae of the re-examined
<typeStatus id="548FCCC3FFBFFFDB4E1AFCA1FC855C3C" box="[827,888,780,801]" pageId="15" pageNumber="1118">types</typeStatus>
of
<taxonomicName id="4C3409E2FFBFFFDB4EBBFCA1FBDD5C3C" ID-CoL="72Q66" baseAuthorityName="Michaelsen" baseAuthorityYear="1888" box="[922,1056,780,801]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="maximus">
<emphasis id="B940AE73FFBFFFDB4EBBFCA1FBDD5C3C" box="[922,1056,780,801]" italics="true" pageId="15" pageNumber="1118">L. maximus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFA50F90FFBFFFDB490EFCA1FB555C3C" author="Rota E" box="[1071,1192,780,801]" pageId="15" pageNumber="1118" pagination="225 - 238" refId="ref25143" refString="Rota E. 2001. Oversized enchytraeids (Annelida, Clitellata): a comparative study, with a revised description of Lumbricillus maximus (Michaelsen). Organisms Diversity &amp; Evolution 1: 225 - 238." type="journal article" year="2001">Rota, 2001</bibRefCitation>
: fig. 1f). Furthermore, Stephenson and Benham noted pharyngeal glands as far back as VII, which is similar to what we observe and which distinguishes
<taxonomicName id="4C3409E2FFBFFFDB4905FCC5FB185C60" ID-CoL="72Q69" authorityName="Benham" authorityYear="1905" box="[1060,1253,872,893]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="macquariensis">
<emphasis id="B940AE73FFBFFFDB4905FCC5FB185C60" box="[1060,1253,872,893]" italics="true" pageId="15" pageNumber="1118">L. macquariensis</emphasis>
</taxonomicName>
from most other
<taxonomicName id="4C3409E2FFBFFFDB4E1AFC2BFC325C86" ID-CoL="84PT8" authorityName="Orsted" authorityYear="1844" box="[827,975,902,923]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="genus">
<emphasis id="B940AE73FFBFFFDB4E1AFC2BFC325C86" box="[827,975,902,923]" italics="true" pageId="15" pageNumber="1118">Lumbricillus</emphasis>
</taxonomicName>
species, except
<taxonomicName id="4C3409E2FFBFFFDB49A5FC2AFA625C81" ID-CoL="72Q66" authority="(Rota, 2001)" baseAuthorityName="Rota" baseAuthorityYear="2001" box="[1156,1439,902,924]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="maximus">
<emphasis id="B940AE73FFBFFFDB49A5FC2AFAF65C81" box="[1156,1291,903,924]" italics="true" pageId="15" pageNumber="1118">L. maximus</emphasis>
(
<bibRefCitation id="EFA50F90FFBFFFDB483AFC2AFA695C86" author="Rota E" box="[1307,1428,902,924]" pageId="15" pageNumber="1118" pagination="225 - 238" refId="ref25143" refString="Rota E. 2001. Oversized enchytraeids (Annelida, Clitellata): a comparative study, with a revised description of Lumbricillus maximus (Michaelsen). Organisms Diversity &amp; Evolution 1: 225 - 238." type="journal article" year="2001">Rota, 2001</bibRefCitation>
)
</taxonomicName>
. Such posteriad extension of the third pair of glands represents a distinct situation from the development of an extra pair of glands in VII, a character distributed erratically also in other genera otherwise characterized by pharyngeal glands in IVVI, for example,
<taxonomicName id="4C3409E2FFBFFFDB4812FBB2FBDE5B4E" ID-CoL="99CVF" authority="Michaelsen, 1889 a" authorityName="Michaelsen" authorityYear="1889" class="Magnoliopsida" family="Bignoniaceae" genus="Fridericia" kingdom="Plantae" order="Lamiales" pageId="15" pageNumber="1118" phylum="Tracheophyta" rank="genus">
<emphasis id="B940AE73FFBFFFDB4812FBB2FA5E5B29" box="[1331,1443,1055,1076]" italics="true" pageId="15" pageNumber="1118">Fridericia</emphasis>
<bibRefCitation id="EFA50F90FFBFFFDB4E1AFB93FBDE5B4E" author="Michaelsen W" box="[827,1059,1086,1108]" pageId="15" pageNumber="1118" pagination="1 - 17" refId="ref24432" refString="Michaelsen W. 1889 a. Oligochaeten des Naturhistorischen Museums in Hamburg. Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 6: 1 - 17." type="journal article" year="1889">Michaelsen, 1889a</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFA50F90FFBFFFDB4919FB92FB405B4E" author="Rota E" box="[1080,1213,1086,1108]" pageId="15" pageNumber="1118" pagination="225 - 238" refId="ref25143" refString="Rota E. 2001. Oversized enchytraeids (Annelida, Clitellata): a comparative study, with a revised description of Lumbricillus maximus (Michaelsen). Organisms Diversity &amp; Evolution 1: 225 - 238." type="journal article" year="2001">Rota, 2001</bibRefCitation>
, 2015;
<bibRefCitation id="EFA50F90FFBFFFDB483CFB93FC2F5B6F" author="Schmelz RM &amp; Collado R" pageId="15" pageNumber="1118" pagination="1 - 176" refId="ref25622" refString="Schmelz RM, Collado R. 2010. A guide to European terrestrial and freshwater species of Enchytraeidae (Oligochaeta). Soil Organisms 82: 1 - 176." type="journal article" year="2010">Schmelz &amp; Collado, 2010</bibRefCitation>
).
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4E72FBD6FAA35AD9" blockId="15.[827,1444,197,1905]" pageId="15" pageNumber="1118">
Another peculiar trait of our species is the structure of the testis sacs, which seemed at first sight to fall in the regularly lobed arrangement seen in most
<taxonomicName id="4C3409E2FFBFFFDB4E1AFB7AFC325BF1" authorityName="Orsted" authorityYear="1844" box="[827,975,1239,1260]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="species" species="undetermined">
<emphasis id="B940AE73FFBFFFDB4E1AFB7AFC325BF1" box="[827,975,1239,1260]" italics="true" pageId="15" pageNumber="1118">Lumbricillus</emphasis>
</taxonomicName>
species. However, the lobed sacs here are much thinner. This could perhaps be an indication that the sacs were not fully developed, but the developing testis sacs of, for example,
<emphasis id="B940AE73FFBFFFDB49A7FA99FB635A54" box="[1158,1182,1332,1353]" italics="true" pageId="15" pageNumber="1118">L.</emphasis>
sp. Marion Is., from Marion Island (reported above) look completely different. It is possible that both Stephenson and Benham saw this structure but did not consider it deviant and therefore made no remarks upon it.
</paragraph>
<paragraph id="8B8B7261FFBFFFDB4E72FA60FB975808" blockId="15.[827,1444,197,1905]" pageId="15" pageNumber="1118">
Our specimens might well belong to the same species as those studied by Stephenson from
<collectingCountry id="F32332F1FFBFFFDB49D9FA46FA5F591C" box="[1272,1442,1515,1537]" name="South Georgia and the South Sandwich Islands" pageId="15" pageNumber="1118">South Georgia</collectingCountry>
and, like Stephenson, we found a good correspondence with Benhams descriptions from
<collectingCountry id="F32332F1FFBFFFDB49EDF984FA635922" box="[1228,1438,1577,1599]" name="Australia" pageId="15" pageNumber="1118">Macquarie Island</collectingCountry>
, particularly in the deep invagination of the body wall where the penial bulb discharges. A circumpolar distribution of the species would not be implausible, because a corresponding pattern has been observed for
<taxonomicName id="4C3409E2FFBFFFDB4E1AF96FFC3259CA" ID-CoL="84PT8" authorityName="Orsted" authorityYear="1844" box="[827,975,1730,1751]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="genus">
<emphasis id="B940AE73FFBFFFDB4E1AF96FFC3259CA" box="[827,975,1730,1751]" italics="true" pageId="15" pageNumber="1118">Lumbricillus</emphasis>
</taxonomicName>
species in the Northern Hemisphere. The other possibility is that these are two or three separate but closely related species.
</paragraph>
<paragraph id="8B8B7261FFBFFFC44E72F8B3FEA95D56" blockId="15.[827,1444,197,1905]" lastBlockId="16.[145,761,197,587]" lastPageId="16" lastPageNumber="1119" pageId="15" pageNumber="1118">
Genetically, our
<taxonomicName id="4C3409E2FFBFFFDB4939F8B3FA7A5829" ID-CoL="72Q69" authorityName="Orsted" authorityYear="1844" box="[1048,1415,1822,1844]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" isUncertain="true" kingdom="Animalia" order="Enchytraeida" pageId="15" pageNumber="1118" phylum="Annelida" rank="genus">
<emphasis id="B940AE73FFBFFFDB4939F8B3FB4D582E" box="[1048,1200,1822,1843]" italics="true" pageId="15" pageNumber="1118">Lumbricillus</emphasis>
cf.
<emphasis id="B940AE73FFBFFFDB49FFF8B2FA7A5829" box="[1246,1415,1823,1844]" italics="true" pageId="15" pageNumber="1118">macquariensis</emphasis>
</taxonomicName>
is sister to
<taxonomicName id="4C3409E2FFBFFFDB4E86F890FC01584F" ID-CoL="4QS2" authorityName="Southern" authorityYear="1913" box="[935,1020,1853,1874]" class="Florideophyceae" family="Acrochaetiaceae" genus="Grania" kingdom="Plantae" order="Nemaliales" pageId="15" pageNumber="1118" phylum="Rhodophyta" rank="genus">
<emphasis id="B940AE73FFBFFFDB4E86F890FC01584F" box="[935,1020,1853,1874]" italics="true" pageId="15" pageNumber="1118">Grania</emphasis>
</taxonomicName>
in our phylogeny, but although its morphology is dissimilar in many ways to that of most
<taxonomicName id="4C3409E2FFA0FFC44DB0FF68FED05FC7" ID-CoL="84PT8" authorityName="Orsted" authorityYear="1844" box="[145,301,197,218]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="16" pageNumber="1119" phylum="Annelida" rank="genus">
<emphasis id="B940AE73FFA0FFC44DB0FF68FED05FC7" box="[145,301,197,218]" italics="true" pageId="16" pageNumber="1119">Lumbricillus</emphasis>
</taxonomicName>
(mentioned above), it is by no means closer to that of
<taxonomicName id="4C3409E2FFA0FFC44C7FFF49FE495FE4" ID-CoL="4QS2" authorityName="Southern" authorityYear="1913" box="[350,436,228,249]" class="Florideophyceae" family="Acrochaetiaceae" genus="Grania" kingdom="Plantae" order="Nemaliales" pageId="16" pageNumber="1119" phylum="Rhodophyta" rank="genus">
<emphasis id="B940AE73FFA0FFC44C7FFF49FE495FE4" box="[350,436,228,249]" italics="true" pageId="16" pageNumber="1119">Grania</emphasis>
</taxonomicName>
. Species of
<taxonomicName id="4C3409E2FFA0FFC44F69FF49FD635FE4" ID-CoL="4QS2" authorityName="Southern" authorityYear="1913" box="[584,670,228,249]" class="Florideophyceae" family="Acrochaetiaceae" genus="Grania" kingdom="Plantae" order="Nemaliales" pageId="16" pageNumber="1119" phylum="Rhodophyta" rank="genus">
<emphasis id="B940AE73FFA0FFC44F69FF49FD635FE4" box="[584,670,228,249]" italics="true" pageId="16" pageNumber="1119">Grania</emphasis>
</taxonomicName>
have a much more slender body, only one chaeta per bundle and most often lack chaetae in several segments, and have masses of developing sperm cells and oocytes enveloped in septal distensions (seminal vesicles and ovisacs, respectively) extending backwards. In order to retain the well-defined genera
<taxonomicName id="4C3409E2FFA0FFC44F4EFE31FD395EAC" ID-CoL="4QS2" authorityName="Southern" authorityYear="1913" box="[623,708,412,433]" class="Florideophyceae" family="Acrochaetiaceae" genus="Grania" kingdom="Plantae" order="Nemaliales" pageId="16" pageNumber="1119" phylum="Rhodophyta" rank="genus">
<emphasis id="B940AE73FFA0FFC44F4EFE31FD395EAC" box="[623,708,412,433]" italics="true" pageId="16" pageNumber="1119">Grania</emphasis>
</taxonomicName>
and
<emphasis id="B940AE73FFA0FFC44DB0FE17FEA05ECD" box="[145,349,442,464]" italics="true" pageId="16" pageNumber="1119">
<taxonomicName id="4C3409E2FFA0FFC44DB0FE17FECD5ED2" ID-CoL="84PT8" authorityName="Orsted" authorityYear="1844" box="[145,304,442,463]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" kingdom="Animalia" order="Enchytraeida" pageId="16" pageNumber="1119" phylum="Annelida" rank="genus" sensu="stricto">Lumbricillus</taxonomicName>
<taxonomicNameLabel id="A2731308FFA0FFC44C1BFE16FEA05ECD" box="[314,349,443,464]" pageId="16" pageNumber="1119" sensu="stricto">s.s.</taxonomicNameLabel>
</emphasis>
,
<taxonomicName id="4C3409E2FFA0FFC44C55FE17FD045ECD" ID-CoL="72Q69" authorityName="Orsted" authorityYear="1844" box="[372,761,442,464]" class="Clitellata" family="Enchytraeidae" genus="Lumbricillus" isUncertain="true" kingdom="Animalia" order="Enchytraeida" pageId="16" pageNumber="1119" phylum="Annelida" rank="genus">
<emphasis id="B940AE73FFA0FFC44C55FE17FDEE5ED2" box="[372,531,442,463]" italics="true" pageId="16" pageNumber="1119">Lumbricillus</emphasis>
cf.
<emphasis id="B940AE73FFA0FFC44F69FE16FD045ECD" box="[584,761,443,464]" italics="true" pageId="16" pageNumber="1119">macquariensis</emphasis>
</taxonomicName>
needs to be transferred to another, probably new, genus. However, owing to the unresolved taxonomy and limited number of mature specimens we leave this to future studies.
</paragraph>
</subSubSection>
</treatment>
</document>