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<document id="5354D7FD1FB62167C4E4B454453B80A7" ID-DOI="10.1111/zoj.12333" ID-ISSN="0024-4082" ID-Zenodo-Dep="4720622" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1619444012732" checkinUser="carolina" docAuthor="Prantoni, Alessandro Lívio, Wit, Pierre De &amp; Erséus, Christer" docDate="2016" docId="7B5287F0DE54E473E2423A410547CE94" docLanguage="en" docName="ZoolJLinnSoc.176.485-510.pdf" docOrigin="Zoological Journal of the Linnean Society 176 (3)" docSource="http://dx.doi.org/10.1111/zoj.12333" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Grania carolinensis Prantoni &amp; Wit &amp; Erséus 2016, SP. NOV." docType="treatment" docVersion="3" lastPageNumber="502" masterDocId="876BFF88DE44E462E349395E0740C819" masterDocTitle="First reports of Grania (Clitellata: Enchytraeidae) from Africa and South America: molecular phylogeny and descriptions of nine new species" masterLastPageNumber="510" masterPageNumber="485" pageNumber="501" updateTime="1698910564440" updateUser="plazi">
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<mods:title id="7E503E3DEE1CADBDD9F9A260C2A21F12">First reports of Grania (Clitellata: Enchytraeidae) from Africa and South America: molecular phylogeny and descriptions of nine new species</mods:title>
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<mods:namePart id="D47324858F95E9E94B2E5FCE4EF7031C">Prantoni, Alessandro Lívio</mods:namePart>
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<mods:namePart id="C2C8ADC10406259FD2868349D1D20AEB">Wit, Pierre De</mods:namePart>
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<mods:namePart id="6556F85739B231F4E683E12D830E207C">Erséus, Christer</mods:namePart>
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<mods:title id="FF269F32C4A99C0969793F84F8F922F4">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="FA7E5ABE791AAB4CE7C5EA2792768C44">2016</mods:date>
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<mods:number id="205BBBFF7AD072F67002E81ADA450106">2016-02-10</mods:number>
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<mods:number id="5077B834C95E454F27C8403C9E83C86D">176</mods:number>
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<treatment id="7B5287F0DE54E473E2423A410547CE94" ID-DOI="http://doi.org/10.5281/zenodo.7526477" ID-Zenodo-Dep="7526477" LSID="urn:lsid:plazi:treatment:7B5287F0DE54E473E2423A410547CE94" httpUri="http://treatment.plazi.org/id/7B5287F0DE54E473E2423A410547CE94" lastPageId="17" lastPageNumber="502" pageId="16" pageNumber="501">
<subSubSection id="BBE1656DDE54E472E2423A41053BCB2C" box="[267,635,799,821]" pageId="16" pageNumber="501" type="nomenclature">
<paragraph id="F34436E6DE54E472E2423A41053BCB2C" blockId="16.[267,635,799,853]" box="[267,635,799,821]" pageId="16" pageNumber="501">
<heading id="A80C818ADE54E472E2423A41053BCB2C" box="[267,635,799,821]" centered="true" fontSize="9" level="2" pageId="16" pageNumber="501" reason="4">
<taxonomicName id="34FB4D65DE54E472E2423A410559CB2D" ID-CoL="3H7YN" ID-ENA="1702334" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[267,537,799,820]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="16" pageNumber="501" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">GRANIA CAROLINENSIS</taxonomicName>
<taxonomicNameLabel id="DABC578FDE54E472E1693A7D053BCB2C" box="[544,635,800,821]" pageId="16" pageNumber="501" rank="species">
<emphasis id="C18FEAF4DE54E472E1693A7D053BCB2C" bold="true" box="[544,635,800,821]" pageId="16" pageNumber="501">SP. NOV.</emphasis>
</taxonomicNameLabel>
</heading>
</paragraph>
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<subSubSection id="BBE1656DDE54E473E2CE3A1E0649C953" lastPageId="17" lastPageNumber="502" pageId="16" pageNumber="501" type="description">
<paragraph id="F34436E6DE54E472E2CE3A1E06BFCB4C" blockId="16.[267,635,799,853]" box="[391,511,832,853]" pageId="16" pageNumber="501">
<figureCitation id="6BC02A63DE54E472E2CE3A1E06BFCB4C" box="[391,511,832,853]" captionStart="Figure 10" captionStartId="16.[144,223,1852,1871]" captionTargetBox="[147,1421,1269,1819]" captionTargetId="figure-506@16.[144,1424,1266,1822]" captionTargetPageId="16" captionText="Figure 10. Grania carolinensis sp. nov. A, chaetae. B, spermatheca. C, penial bulb and sperm funnel. See text for abbreviations." figureDoi="http://doi.org/10.5281/zenodo.4720644" httpUri="https://zenodo.org/record/4720644/files/figure.png" pageId="16" pageNumber="501">FIGURE 10</figureCitation>
</paragraph>
<paragraph id="F34436E6DE54E472E3D93A3807B3CB62" blockId="16.[144,759,870,1106]" box="[144,243,870,891]" pageId="16" pageNumber="501">
<emphasis id="C18FEAF4DE54E472E3D93A3807B3CB62" box="[144,243,870,891]" italics="true" pageId="16" pageNumber="501">
<typeStatus id="2C408844DE54E472E3D93A3807B3CB62" box="[144,243,870,891]" pageId="16" pageNumber="501" type="holotype">Holotype</typeStatus>
</emphasis>
</paragraph>
<paragraph id="F34436E6DE54E472E3D93ADB07AACC48" blockId="16.[144,759,870,1106]" pageId="16" pageNumber="501">
<materialsCitation id="43933CBBDE54E472E3D93ADB07AACC48" accessionNumber="KT428112" collectingDate="2011-05-20" collectionCode="USNM" collectorName="C. Erseus &amp; GenBank" country="United States of America" elevation="492" latitude="33.173058" location="North Carolina" longLatPrecision="20" longitude="-76.756386" pageId="16" pageNumber="501" specimenCode="USNM 1283174" specimenCount="1" stateProvince="North Carolina" typeStatus="holotype">
<specimenCode id="A35D9E9DDE54E472E3D93ADB0609CB83" box="[144,329,901,922]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="16" pageNumber="501" type="Museum">USNM 1283174</specimenCode>
, CE11569, whole-mounted, sexually mature specimen, with some segments amputated, from off
<collectingRegion id="313FF804DE54E472E3FB3A9C061ECBC1" box="[178,350,962,984]" country="United States of America" name="North Carolina" pageId="16" pageNumber="501">North Carolina</collectingRegion>
,
<collectingCountry id="8BEC7676DE54E472E2233A9C06E1CBCE" box="[362,417,962,983]" name="United States of America" pageId="16" pageNumber="501">USA</collectingCountry>
,
<geoCoordinate id="96CF5021DE54E472E2E53A9C056BCBCE" box="[428,555,961,983]" degrees="33" direction="north" minutes="10" orientation="latitude" pageId="16" pageNumber="501" precision="15" seconds="23" value="33.173058">33°10′23″N</geoCoordinate>
,
<geoCoordinate id="96CF5021DE54E472E17E3A9C05F8CBCE" box="[567,696,961,984]" degrees="76" direction="west" minutes="45" orientation="longitude" pageId="16" pageNumber="501" precision="15" seconds="23" value="-76.756386">76°45′23″W</geoCoordinate>
. Continental shelf slope,
<quantity id="34039B03DE54E472E2CB3ABF0687CBEF" box="[386,455,993,1014]" metricMagnitude="2" metricUnit="m" metricValue="4.92" pageId="16" pageNumber="501" unit="m" value="492.0">
<elevation id="78D6D1D5DE54E472E2CB3ABF0687CBEF" box="[386,455,993,1014]" metricMagnitude="2" metricUnit="m" metricValue="4.92" pageId="16" pageNumber="501" unit="m" value="492.0">492 m</elevation>
</quantity>
in depth, sand. Collected by
<collectorName id="5E0E5330DE54E472E3FE3AA1066BCC0C" box="[183,299,1023,1045]" pageId="16" pageNumber="501">C. Erséus</collectorName>
,
<date id="87451026DE54E472E2753AA10693CC0D" box="[316,467,1023,1044]" pageId="16" pageNumber="501" value="2011-05-20">
<collectingDate id="9701E9CEDE54E472E2753AA10693CC0D" box="[316,467,1023,1044]" pageId="16" pageNumber="501" value="2011-05-20">20 May 2011</collectingDate>
</date>
.
<emphasis id="C18FEAF4DE54E472E2AA3AA10553CC0D" box="[483,531,1023,1044]" italics="true" pageId="16" pageNumber="501">COI</emphasis>
barcode sequence,
<collectorName id="5E0E5330DE54E472E3D93D4007BCCC2D" box="[144,252,1054,1076]" pageId="16" pageNumber="501">GenBank</collectorName>
acc. no.
<accessionNumber id="ECA8AB05DE54E472E2283D400698CC2A" box="[353,472,1054,1075]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/KT428112" pageId="16" pageNumber="501">KT428112</accessionNumber>
; for other sequences, see
<tableCitation id="BE79035DDE54E472E3D93D6207A4CC48" box="[144,228,1084,1106]" captionStart="Table 1" captionStartId="3.[31,95,393,414]" captionTargetPageId="3" captionText="Table 1. Specimens of the new species used for genetic analysis, with places of origin, voucher numbers, and GenBank accession numbers" httpUri="http://table.plazi.org/id/A784666EDE47E461E35638D702B7C987" pageId="16" pageNumber="501" tableUuid="A784666EDE47E461E35638D702B7C987">Table 1</tableCitation>
.
</materialsCitation>
</paragraph>
<paragraph id="F34436E6DE54E472E3D93D270647CC97" blockId="16.[144,461,1145,1198]" box="[144,263,1145,1166]" pageId="16" pageNumber="501">
<emphasis id="C18FEAF4DE54E472E3D93D270647CC97" box="[144,263,1145,1166]" italics="true" pageId="16" pageNumber="501">Etymology</emphasis>
</paragraph>
<paragraph id="F34436E6DE54E472E3D93DC604E7C8C3" blockId="16.[144,461,1145,1198]" box="[144,935,197,1198]" lastBlockId="16.[806,1422,197,1198]" pageId="16" pageNumber="501">
Named for
<collectingRegion id="313FF804DE54E472E25C3DC60689CCB7" box="[277,457,1176,1198]" country="United States of America" name="North Carolina" pageId="16" pageNumber="501">North Carolina</collectingRegion>
.
<emphasis id="C18FEAF4DE54E472E06F399B04E7C8C3" box="[806,935,197,218]" italics="true" pageId="16" pageNumber="501">Description</emphasis>
</paragraph>
<paragraph id="F34436E6DE54E472E06F39BD0217CA70" blockId="16.[806,1422,197,1198]" pageId="16" pageNumber="501">
Body&gt;
<quantity id="34039B03DE54E472E03139BD0497C8E0" box="[888,983,227,249]" metricMagnitude="-3" metricUnit="m" metricValue="5.1" pageId="16" pageNumber="501" unit="mm" value="5.1">5.10 mm</quantity>
long,&gt; 30 segments (posterior end used for DNA extractions),
<quantity id="34039B03DE54E472E77F385C03DBC90E" box="[1078,1179,258,279]" metricMagnitude="-5" metricUnit="m" metricValue="8.0" pageId="16" pageNumber="501" unit="mm" value="0.08">0.08 mm</quantity>
wide at segment V,
<quantity id="34039B03DE54E472E06F387F04C9C92F" box="[806,905,289,310]" metricMagnitude="-5" metricUnit="m" metricValue="9.0" pageId="16" pageNumber="501" unit="mm" value="0.09">0.09 mm</quantity>
wide at segment XII. Prostomium rounded, 51 μm long, 66 μm wide, epidermis 10 μm thick, reduced to 4 μm at front tip. Ventral chaetae from segment XIII, lateral chaetae absent. Chaetae (
<figureCitation id="6BC02A63DE54E472E7E23822024CC98A" box="[1195,1292,380,403]" captionStart="Figure 10" captionStartId="16.[144,223,1852,1871]" captionTargetBox="[147,1421,1269,1819]" captionTargetId="figure-506@16.[144,1424,1266,1822]" captionTargetPageId="16" captionText="Figure 10. Grania carolinensis sp. nov. A, chaetae. B, spermatheca. C, penial bulb and sperm funnel. See text for abbreviations." figureDoi="http://doi.org/10.5281/zenodo.4720644" httpUri="https://zenodo.org/record/4720644/files/figure.png" pageId="16" pageNumber="501">Fig. 10A</figureCitation>
) 42–50 μm long, shaft straight, 3.7 μm thick at midpoint, L-shaped, proximally bent into a foot with low instep and slight heel. Chaetal index (
<bibRefCitation id="976A4B17DE54E472E75A38860242C9F7" author="Erseus C &amp; Rota E" box="[1043,1282,472,494]" pageId="16" pageNumber="501" pagination="892 - 900" refId="ref17321" refString="Erseus C, Rota E. 2003. New findings and an overview of the oligochaetous Clitellata (Annelida) of the North Atlantic deep sea. Proceedings of the Biological Society of Washington 116: 892 - 900." type="journal article" year="2003">Rota &amp; Erséus, 2003</bibRefCitation>
) 3.60 ± 0.33 (
<emphasis id="C18FEAF4DE54E472E06738A9047DCA15" box="[814,829,503,524]" italics="true" pageId="16" pageNumber="501">n</emphasis>
= 4). Epidermal gland cells inconspicuous. Clitellum not well developed. Spermathecal pores in lateral lines, immediately posterior to 4/5. Male pores ventrolateral in mid-segment XII. Female pores not observed.
</paragraph>
<paragraph id="F34436E6DE54E473E0763B2F044BC901" blockId="16.[806,1422,197,1198]" lastBlockId="17.[164,779,197,280]" lastPageId="17" lastPageNumber="502" pageId="16" pageNumber="501">
Brain posteriorly indented. Head organ (
<emphasis id="C18FEAF4DE54E472E65B3B2C0211CA9E" box="[1298,1361,626,647]" italics="true" pageId="16" pageNumber="501">sensu</emphasis>
<bibRefCitation id="976A4B17DE54E472E6113B2C04A1CABF" author="Rota E &amp; Erseus C" pageId="16" pageNumber="501" pagination="169 - 183" refId="ref18340" refString="Rota E, Erseus C. 1996. Six new species of Grania (Oligochaeta, Enchytraeidae) from the Ross Sea, Antarctica. Antarctic Science 2: 169 - 183." type="journal article" year="1996">Rota &amp; Erséus, 1996</bibRefCitation>
) absent. Pharyngeal glands in segments IV–VI, dorsal lobes in segment IV (one pair), in segment V (one pair), and in segment VI (one pair), ventral lobes present in segment IV (one pair), in segment V (two pairs), and in segment VI (two pairs); glands not connected dorsally. Nephridia not observed. Chloragogen cells not observed. Dorsal blood vessel commencing in segment XVII. Coelomocytes not observed. Sperm sac extending into half of segment XV, egg sac extending into segment XVII. Sperm funnels about six times longer than wide (
<figureCitation id="6BC02A63DE54E472E79C3A9C027CCBC1" box="[1237,1340,962,984]" captionStart="Figure 10" captionStartId="16.[144,223,1852,1871]" captionTargetBox="[147,1421,1269,1819]" captionTargetId="figure-506@16.[144,1424,1266,1822]" captionTargetPageId="16" captionText="Figure 10. Grania carolinensis sp. nov. A, chaetae. B, spermatheca. C, penial bulb and sperm funnel. See text for abbreviations." figureDoi="http://doi.org/10.5281/zenodo.4720644" httpUri="https://zenodo.org/record/4720644/files/figure.png" pageId="16" pageNumber="501">Fig. 10C</figureCitation>
). Vasa deferentia observed in segments XII–XIII, internally ciliated, coiled, 7 μm wide. Penial apparatus
<typeStatus id="2C408844DE54E472E60F3AA10238CC0D" box="[1350,1400,1023,1044]" pageId="16" pageNumber="501">type</typeStatus>
1 (
<emphasis id="C18FEAF4DE54E472E0673D40042BCC2A" box="[814,875,1054,1075]" italics="true" pageId="16" pageNumber="501">sensu</emphasis>
<bibRefCitation id="976A4B17DE54E472E03B3D400340CC2D" author="Coates KA" box="[882,1024,1054,1076]" pageId="16" pageNumber="501" pagination="45 - 50" refId="ref16484" refString="Coates KA. 1984. Specific criteria in Grania (Oligochaeta, Enchytraeidae). Hydrobiologia 115: 45 - 50." type="journal article" year="1984">Coates, 1984</bibRefCitation>
), small, glandular bulb, 40 μm long, 31 μm wide; stylet absent. Midventral copulatory gland (in segment XIV) not observed. Each spermatheca attached to oesophagus in posterior half of segment V. Ampulla large, oval, 100 μm long, 44 μm wide. Sperm rings maximally 11 μm wide, but few. Ectal duct slender, 53 μm long, 14 μm wide at midcourse, narrowing at distal end
<figureCitation id="6BC02A63DE55E473E253385C06D9C901" box="[282,409,258,280]" captionStart="Figure 10" captionStartId="16.[144,223,1852,1871]" captionTargetBox="[147,1421,1269,1819]" captionTargetId="figure-506@16.[144,1424,1266,1822]" captionTargetPageId="16" captionText="Figure 10. Grania carolinensis sp. nov. A, chaetae. B, spermatheca. C, penial bulb and sperm funnel. See text for abbreviations." figureDoi="http://doi.org/10.5281/zenodo.4720644" httpUri="https://zenodo.org/record/4720644/files/figure.png" pageId="17" pageNumber="502">Figure 10B</figureCitation>
). No gland at spermathecal pore.
</paragraph>
<caption id="A784666EDE54E472E3D93E620665CF74" ID-DOI="http://doi.org/10.5281/zenodo.4720644" ID-Zenodo-Dep="4720644" httpUri="https://zenodo.org/record/4720644/files/figure.png" pageId="16" pageNumber="501" startId="16.[144,223,1852,1871]" targetBox="[147,1421,1269,1819]" targetPageId="16">
<paragraph id="F34436E6DE54E472E3D93E620665CF74" blockId="16.[144,1421,1852,1901]" pageId="16" pageNumber="501">
<emphasis id="C18FEAF4DE54E472E3D93E62051ACF56" bold="true" box="[144,602,1852,1871]" pageId="16" pageNumber="501">
Figure 10.
<taxonomicName id="34FB4D65DE54E472E2593E6206BECF56" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[272,510,1852,1871]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="16" pageNumber="501" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE54E472E2593E6206BECF56" bold="true" box="[272,510,1852,1871]" italics="true" pageId="16" pageNumber="501">Grania carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE54E472E14D3E62051ACF56" box="[516,602,1852,1871]" pageId="16" pageNumber="501" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
A, chaetae. B, spermatheca. C, penial bulb and sperm funnel. See text for abbreviations.
</paragraph>
</caption>
<paragraph id="F34436E6DE55E473E3ED386B0649C953" blockId="17.[164,779,309,1677]" box="[164,265,309,330]" pageId="17" pageNumber="502">
<emphasis id="C18FEAF4DE55E473E3ED386B0649C953" box="[164,265,309,330]" italics="true" pageId="17" pageNumber="502">Remarks</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="BBE1656DDE55E473E3ED380A0547CE94" pageId="17" pageNumber="502" type="reference_group">
<paragraph id="F34436E6DE55E473E3ED380A0547CE94" blockId="17.[164,779,309,1677]" pageId="17" pageNumber="502">
<taxonomicName id="34FB4D65DE55E473E3ED380A06CBC970" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[164,395,340,361]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E3ED380A06CBC970" box="[164,395,340,361]" italics="true" pageId="17" pageNumber="502">Grania carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E2DB380A06A8C970" box="[402,488,340,361]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
is similar to the northeast Atlantic,
<taxonomicName id="34FB4D65DE55E473E274382D0447C991" ID-CoL="3H825" authority="(Knollner, 1935)" baseAuthorityName="Knollner" baseAuthorityYear="1935" box="[317,775,370,393]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="postclitellochaeta">
<emphasis id="C18FEAF4DE55E473E274382D050DC991" box="[317,589,371,392]" italics="true" pageId="17" pageNumber="502">Grania postclitellochaeta</emphasis>
(
<bibRefCitation id="976A4B17DE55E473E112382D05BFC991" author="Knollner FH" box="[603,767,370,393]" pageId="17" pageNumber="502" pagination="425 - 512" refId="ref17729" refString="Knollner FH. 1935. Okologische und systematische Untersuchungen uber litorale und marine Oligochaten der Kieler Bucht. Zoologische Jahrbucher, Abteilung fur Systematik, Okologie und Geographie der Tiere 66: 425 - 512." type="journal article" year="1935">Knöllner, 1935</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="34FB4D65DE55E473E3ED38CC06BCC9BF" ID-CoL="3H7ZX" authority="Erseus, 1977" authorityName="Erseus" authorityYear="1977" box="[164,508,401,423]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="ovitheca">
<emphasis id="C18FEAF4DE55E473E3ED38CC061CC9BF" box="[164,348,401,423]" italics="true" pageId="17" pageNumber="502">Grania ovitheca</emphasis>
Erséus, 1977
</taxonomicName>
, and the cryptic congener to
<taxonomicName id="34FB4D65DE55E473E3BC38EE0632C9DC" ID-CoL="3H7ZX" authorityName="Erseus" authorityYear="1977" box="[245,370,432,453]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="ovitheca">
<emphasis id="C18FEAF4DE55E473E3BC38EE0632C9DC" box="[245,370,432,453]" italics="true" pageId="17" pageNumber="502">G. ovitheca</emphasis>
</taxonomicName>
,
<taxonomicName id="34FB4D65DE55E473E2CD38EE079EC9FD" ID-CoL="3H7ZW" authority="De Wit &amp; Erseus, 2010" authorityName="De Wit &amp; Erseus" authorityYear="2010" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="occulta">
<emphasis id="C18FEAF4DE55E473E2CD38EE056FC9DC" box="[388,559,432,453]" italics="true" pageId="17" pageNumber="502">Grania occulta</emphasis>
<bibRefCitation id="976A4B17DE55E473E17338EE079EC9FD" author="De Wit P &amp; Erseus C" pageId="17" pageNumber="502" pagination="285 - 293" refId="ref16935" refString="De Wit P, Erseus C. 2010. Genetic variation and phylogeny of Scandinavian species of Grania (Annelida: Clitellata: Enchytraeidae), with the discovery of a cryptic species. Journal of Zoological Systematics and Evolutionary Research 48: 285 - 293." type="journal article" year="2010">De Wit &amp; Erséus, 2010</bibRefCitation>
</taxonomicName>
, a species constellation also supported by the genetic data (see ‘Phylogenetic analyses’ below). All of these four species have the same distribution of ventral chaetae (beginning in segment XIII), and they lack lateral chaetae and the midventral copulatory gland in segment XIV.
<taxonomicName id="34FB4D65DE55E473E21E3B36056CCA65" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[343,556,615,637]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E21E3B36056CCA65" box="[343,556,615,637]" italics="true" pageId="17" pageNumber="502">Grania carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E17B3B3605C4CA64" box="[562,644,616,637]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
,
<taxonomicName id="34FB4D65DE55E473E1C73B360444CA65" ID-CoL="3H7ZX" authorityName="Erseus" authorityYear="1977" box="[654,772,615,637]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="ovitheca">
<emphasis id="C18FEAF4DE55E473E1C73B360444CA65" box="[654,772,615,637]" italics="true" pageId="17" pageNumber="502">G. ovitheca</emphasis>
</taxonomicName>
, and
<taxonomicName id="34FB4D65DE55E473E39C3BD80601CA82" ID-CoL="3H7ZW" authorityName="De Wit &amp; Erseus" authorityYear="2010" box="[213,321,646,667]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="occulta">
<emphasis id="C18FEAF4DE55E473E39C3BD80601CA82" box="[213,321,646,667]" italics="true" pageId="17" pageNumber="502">G. occulta</emphasis>
</taxonomicName>
also share the shape of the spermatheca, which has a short and slender ectal duct and a large oval ampulla. In
<taxonomicName id="34FB4D65DE55E473E2223B9D06A9CAC1" ID-CoL="3H7ZX" authorityName="Erseus" authorityYear="1977" box="[363,489,707,728]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="ovitheca">
<emphasis id="C18FEAF4DE55E473E2223B9D06A9CAC1" box="[363,489,707,728]" italics="true" pageId="17" pageNumber="502">G. ovitheca</emphasis>
</taxonomicName>
and
<taxonomicName id="34FB4D65DE55E473E16F3B9D05D6CAC1" ID-CoL="3H7ZW" authorityName="De Wit &amp; Erseus" authorityYear="2010" box="[550,662,707,728]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="occulta">
<emphasis id="C18FEAF4DE55E473E16F3B9D05D6CAC1" box="[550,662,707,728]" italics="true" pageId="17" pageNumber="502">G. occulta</emphasis>
</taxonomicName>
, however, the sperm funnels are longer (between eight and 15 times longer than wide, as opposed to six times longer than wide in
<taxonomicName id="34FB4D65DE55E473E2733A41069ECB2D" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[314,478,799,820]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E2733A41069ECB2D" box="[314,478,799,820]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E2AC3A410578CB2D" box="[485,568,799,820]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
) and they lack the ventral lobes of the pharyngeal glands in segment IV that are present in
<taxonomicName id="34FB4D65DE55E473E2CD3A02056BCB68" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[388,555,860,881]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E2CD3A02056BCB68" box="[388,555,860,881]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E1783A0305C5CB6B" box="[561,645,861,882]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
and also in
<taxonomicName id="34FB4D65DE55E473E3ED3A2506C8CB89" ID-CoL="3H825" baseAuthorityName="Knollner" baseAuthorityYear="1935" box="[164,392,891,912]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="postclitellochaeta">
<emphasis id="C18FEAF4DE55E473E3ED3A2506C8CB89" box="[164,392,891,912]" italics="true" pageId="17" pageNumber="502">G. postclitellochaeta</emphasis>
</taxonomicName>
.
<taxonomicName id="34FB4D65DE55E473E2D03A2505F8CB89" ID-CoL="3H825" baseAuthorityName="Knollner" baseAuthorityYear="1935" box="[409,696,891,912]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="postclitellochaeta">
<emphasis id="C18FEAF4DE55E473E2D03A2505F8CB89" box="[409,696,891,912]" italics="true" pageId="17" pageNumber="502">Grania postclitellochaeta</emphasis>
</taxonomicName>
differs from
<taxonomicName id="34FB4D65DE55E473E3AC3AC406CECBB7" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[229,398,921,943]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E3AC3AC406CECBB7" box="[229,398,921,943]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E2DC3AC406ABCBB6" box="[405,491,922,943]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
by possessing a racketshaped diverticulum in the spermathecal ampulla, which is absent in
<taxonomicName id="34FB4D65DE55E473E2033A890542CBF2" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[330,514,982,1004]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E2033A890542CBF2" box="[330,514,982,1004]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E1423A890527CBF5" box="[523,615,983,1004]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
In addition,
<taxonomicName id="34FB4D65DE55E473E3ED3AAB0606CC13" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[164,326,1013,1034]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E3ED3AAB0606CC13" box="[164,326,1013,1034]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E2043AA806DFCC12" box="[333,415,1014,1035]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
differs from all the species mentioned above by its penial apparatus, with a simple invagination surrounded by a small and glandular bulb. In
<taxonomicName id="34FB4D65DE55E473E38D3D0F06E3CC7F" ID-CoL="3H825" baseAuthorityName="Knollner" baseAuthorityYear="1935" box="[196,419,1105,1126]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="postclitellochaeta">
<emphasis id="C18FEAF4DE55E473E38D3D0F06E3CC7F" box="[196,419,1105,1126]" italics="true" pageId="17" pageNumber="502">G. postclitellochaeta</emphasis>
</taxonomicName>
the penial bulbs are covered by a large supplementary glandular body, whereas in
<taxonomicName id="34FB4D65DE55E473E3ED3DD00662CCBA" ID-CoL="3H7ZX" authorityName="Erseus" authorityYear="1977" box="[164,290,1166,1187]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="ovitheca">
<emphasis id="C18FEAF4DE55E473E3ED3DD00662CCBA" box="[164,290,1166,1187]" italics="true" pageId="17" pageNumber="502">G. ovitheca</emphasis>
</taxonomicName>
the bulbs instead have lateral aglandular sacs directed posteriorly (
<bibRefCitation id="976A4B17DE55E473E2993DF3058ACCDA" author="Erseus C &amp; Rota E" box="[464,714,1197,1219]" pageId="17" pageNumber="502" pagination="892 - 900" refId="ref17321" refString="Erseus C, Rota E. 2003. New findings and an overview of the oligochaetous Clitellata (Annelida) of the North Atlantic deep sea. Proceedings of the Biological Society of Washington 116: 892 - 900." type="journal article" year="2003">Rota &amp; Erséus, 2003</bibRefCitation>
). The aglandular sacs were not mentioned in the original description of
<taxonomicName id="34FB4D65DE55E473E27A3DB406E3CCE6" ID-CoL="3H7ZW" authorityName="De Wit &amp; Erseus" authorityYear="2010" box="[307,419,1258,1279]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="occulta">
<emphasis id="C18FEAF4DE55E473E27A3DB406E3CCE6" box="[307,419,1258,1279]" italics="true" pageId="17" pageNumber="502">G. occulta</emphasis>
</taxonomicName>
(see
<bibRefCitation id="976A4B17DE55E473E2AF3DB405BFCD19" author="De Wit P &amp; Erseus C" box="[486,767,1258,1280]" pageId="17" pageNumber="502" pagination="285 - 293" refId="ref16935" refString="De Wit P, Erseus C. 2010. Genetic variation and phylogeny of Scandinavian species of Grania (Annelida: Clitellata: Enchytraeidae), with the discovery of a cryptic species. Journal of Zoological Systematics and Evolutionary Research 48: 285 - 293." type="journal article" year="2010">De Wit &amp; Erséus, 2010</bibRefCitation>
), but the authors noted that the single available specimen possessed oval structures next to the epidermal invaginations. No accessory glands or sacs could be observed near the male pores of
<taxonomicName id="34FB4D65DE55E473E14E3C3B05F0CD60" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[519,688,1380,1402]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E14E3C3B05F0CD60" box="[519,688,1380,1402]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E1FE3C3B044BCD63" box="[695,779,1381,1402]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
The deep-sea Atlantic
<taxonomicName id="34FB4D65DE55E473E2823CDD0676CDAE" ID-CoL="3H7YD" authority="Coates &amp; Erseus, 1985" authorityName="Coates &amp; Erseus" authorityYear="1985" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="atlantica">
<emphasis id="C18FEAF4DE55E473E2823CDD05D1CD81" box="[459,657,1411,1432]" italics="true" pageId="17" pageNumber="502">Grania atlantica</emphasis>
<bibRefCitation id="976A4B17DE55E473E1D43CDD0676CDAE" author="Coates KA &amp; Erseus C" pageId="17" pageNumber="502" pagination="103 - 116" refId="ref16570" refString="Coates KA, Erseus C. 1985. Marine Enchytraeids (Oligochaeta) of the Coastal Northwest Atlantic (Northern and Mid U. S. A.) Zoologica Scripta, 14: 103 - 116." type="journal article" year="1985">Coates &amp; Erséus, 1985</bibRefCitation>
</taxonomicName>
also resembles
<taxonomicName id="34FB4D65DE55E473E2A33CFC05CFCDAF" ID-CoL="3H7YN" authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[490,655,1441,1463]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis id="C18FEAF4DE55E473E2A33CFC05CFCDAF" box="[490,655,1441,1463]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="DABC578FDE55E473E1DF3CFC05A9CDAE" box="[662,745,1442,1463]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
by the proportions of the sperm funnel and the morphology of the penial apparatus, but the former differs from the latter by its spermathecal ampulla, which has a large sacciform diverticulum. Moreover,
<taxonomicName id="34FB4D65DE55E473E1103F420598CE28" ID-CoL="3H7YD" authorityName="Coates &amp; Erseus" authorityYear="1985" box="[601,728,1564,1585]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="atlantica">
<emphasis id="C18FEAF4DE55E473E1103F420598CE28" box="[601,728,1564,1585]" italics="true" pageId="17" pageNumber="502">G. atlantica</emphasis>
</taxonomicName>
possesses lateral chaetae, a head organ, as well as a midventral copulatory gland in segment XIV, all of which are absent in the new species.
</paragraph>
</subSubSection>
</treatment>
</document>
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