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<document ID-DOI="10.1111/zoj.12333" ID-ISSN="0024-4082" ID-Zenodo-Dep="4720622" checkinTime="1619444012732" checkinUser="carolina" docAuthor="Prantoni, Alessandro Lívio, Wit, Pierre De &amp; Erséus, Christer" docDate="2016" docId="7B5287F0DE55E471E3A83FEB05E2CDBA" 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 CF. LEVIS COATES &amp; ERSEUS 1985" docType="treatment" docVersion="1" 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="502" updateTime="1673465143273" updateUser="felipe">
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<mods:titleInfo>
<mods:title>First reports of Grania (Clitellata: Enchytraeidae) from Africa and South America: molecular phylogeny and descriptions of nine new species</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Prantoni, Alessandro Lívio</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wit, Pierre De</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Erséus, Christer</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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<mods:part>
<mods:date>2016</mods:date>
<mods:detail type="pubDate">
<mods:number>2016-02-10</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>176</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>3</mods:number>
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<mods:extent unit="page">
<mods:start>485</mods:start>
<mods:end>510</mods:end>
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<mods:url>http://dx.doi.org/10.1111/zoj.12333</mods:url>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1111/zoj.12333</mods:identifier>
<mods:identifier type="ISSN">0024-4082</mods:identifier>
<mods:identifier type="Zenodo-Dep">4720622</mods:identifier>
</mods:mods>
<treatment LSID="urn:lsid:plazi:treatment:7B5287F0DE55E471E3A83FEB05E2CDBA" httpUri="http://treatment.plazi.org/id/7B5287F0DE55E471E3A83FEB05E2CDBA" lastPageId="19" lastPageNumber="502" pageId="17" pageNumber="502">
<subSubSection box="[225,717,1716,1738]" pageId="17" pageNumber="502" type="discussion">
<paragraph blockId="17.[180,762,1716,1802]" box="[225,717,1716,1738]" pageId="17" pageNumber="502">
<taxonomicName authority="CF. LEVIS COATES &amp; ERSEUS, 1985" authorityName="CF. LEVIS COATES &amp; ERSEUS" authorityYear="1985" box="[225,717,1716,1738]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="genus">
<emphasis box="[225,312,1717,1738]" italics="true" pageId="17" pageNumber="502">GRANIA</emphasis>
CF.
<emphasis box="[361,424,1720,1736]" italics="true" pageId="17" pageNumber="502">LEVIS</emphasis>
<bibRefCitation author="Coates KA &amp; Erseus C" box="[433,717,1716,1738]" 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>
</paragraph>
</subSubSection>
<subSubSection pageId="17" pageNumber="502" type="nomenclature">
<paragraph blockId="17.[180,762,1716,1802]" pageId="17" pageNumber="502">
<heading centered="true" fontSize="8" level="2" pageId="17" pageNumber="502" reason="8">
PROBABLY
<taxonomicName ID-CoL="84NVD" ID-ENA="167816" authority="LEVIS COATES &amp; ERSEUS, 1985: 111 - 112" authorityName="CF. LEVIS COATES &amp; ERSEUS" authorityPageNumber="111 - 112" authorityYear="1985" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="genus">
<emphasis box="[304,462,1748,1769]" italics="true" pageId="17" pageNumber="502">GRANIA LEVIS</emphasis>
<bibRefCitation 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: 111112</bibRefCitation>
</taxonomicName>
,
<figureCitation box="[493,558,1780,1802]" captionStart="Figure 6" captionStartId="11.[164,243,791,810]" captionTargetBox="[167,1441,198,758]" captionTargetId="figure-549@11.[164,1444,195,761]" captionTargetPageId="11" captionText="Figure 6. Grania simonae sp. nov. A, chaetae. B, spermatheca. C, coelomocytes. D, penial bulb and sperm funnel. See text for abbreviations." figureDoi="http://doi.org/10.5281/zenodo.4720636" httpUri="https://zenodo.org/record/4720636/files/figure.png" pageId="17" pageNumber="502">FIG. 6</figureCitation>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="17" pageNumber="502" type="materials_examined">
<paragraph blockId="17.[164,779,1819,1902]" box="[164,378,1819,1840]" pageId="17" pageNumber="502">
<emphasis box="[164,378,1819,1840]" italics="true" pageId="17" pageNumber="502">Material examined</emphasis>
</paragraph>
<paragraph blockId="17.[164,779,1819,1902]" lastBlockId="17.[826,1441,196,311]" pageId="17" pageNumber="502">
<materialsCitation accessionNumber="KT428114" collectingDate="2011-05-20" collectionCode="USNM" collectorName="C. Erseus" country="United States of America" elevation="492" latitude="33.173058" location="North Carolina" longLatPrecision="20" longitude="-76.756386" pageId="17" pageNumber="502" specimenCode="USNM 1283176" specimenCount="1" stateProvince="North Carolina">
<specimenCode box="[164,340,1849,1871]" 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="17" pageNumber="502" type="Museum">USNM 1283176</specimenCode>
, CE11570, whole-mounted, sexually immature specimen, with some segments amputated, from off
<collectingRegion box="[861,1033,197,219]" country="United States of America" name="North Carolina" pageId="17" pageNumber="502">North Carolina</collectingRegion>
,
<collectingCountry box="[1044,1099,197,218]" name="United States of America" pageId="17" pageNumber="502">USA</collectingCountry>
,
<geoCoordinate box="[1111,1237,196,218]" degrees="33" direction="north" minutes="10" orientation="latitude" pageId="17" pageNumber="502" precision="15" seconds="23" value="33.173058">33°1023″N</geoCoordinate>
,
<geoCoordinate box="[1249,1379,196,219]" degrees="76" direction="west" minutes="45" orientation="longitude" pageId="17" pageNumber="502" precision="15" seconds="23" value="-76.756386">76°4523″W</geoCoordinate>
. Continental shelf slope,
<quantity box="[1068,1138,228,249]" metricMagnitude="2" metricUnit="m" metricValue="4.92" pageId="17" pageNumber="502" unit="m" value="492.0">
<elevation box="[1068,1138,228,249]" metricMagnitude="2" metricUnit="m" metricValue="4.92" pageId="17" pageNumber="502" unit="m" value="492.0">492 m</elevation>
</quantity>
in depth, sand. Collected by
<collectorName box="[860,969,258,280]" pageId="17" pageNumber="502">C. Erséus</collectorName>
,
<date box="[981,1125,258,280]" pageId="17" pageNumber="502" value="2011-05-20">
<collectingDate box="[981,1125,258,280]" pageId="17" pageNumber="502" value="2011-05-20">20 May 2011</collectingDate>
</date>
.
<emphasis box="[1136,1182,258,279]" italics="true" pageId="17" pageNumber="502">COI</emphasis>
barcode
<accessionNumber box="[1283,1399,258,279]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/KT428114" pageId="17" pageNumber="502">KT428114</accessionNumber>
; for other genes, see
<tableCitation box="[1023,1107,289,311]" 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" pageId="17" pageNumber="502">Table 1</tableCitation>
</materialsCitation>
.
</paragraph>
<paragraph blockId="17.[826,1442,352,711]" box="[826,927,352,373]" pageId="17" pageNumber="502">
<emphasis box="[826,927,352,373]" italics="true" pageId="17" pageNumber="502">Remarks</emphasis>
</paragraph>
<paragraph blockId="17.[826,1442,352,711]" pageId="17" pageNumber="502">
This barcoded, but immature specimen, and thus unsuitable for complete morphological description, was included in the phylogenetic analysis, to enlarge the taxonomic sampling from the north-western Atlantic region. Phylogenetically, this specimen came out as closely related to
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[1028,1195,536,557]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis box="[1028,1195,536,557]" italics="true" pageId="17" pageNumber="502">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1202,1286,536,557]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
(
<figureCitation box="[1302,1382,536,558]" captionStart="Figure 15" captionStartId="22.[144,223,1323,1342]" captionTargetBox="[146,1422,197,1291]" captionTargetId="figure-242@22.[144,1424,195,1293]" captionTargetPageId="22" captionText="Figure 15. Multi-locus phylogeny of all available genetic data in Grania to date, using three mitochondrial and three nuclear loci. New species from this study are marked in bold. Majority-rule consensus tree from Bayesian Markov chain Monte Carlo (MCMC) inference; clade posterior probabilities are given next to the nodes." figureDoi="http://doi.org/10.5281/zenodo.4720659" httpUri="https://zenodo.org/record/4720659/files/figure.png" pageId="17" pageNumber="502">Fig. 15</figureCitation>
), but it is morphologically distinct by its complete lack of chaetae. The latter trait suggests that this specimen could belong to
<taxonomicName authority="Coates &amp; Erseus, 1985" authorityName="Coates &amp; Erseus" authorityYear="1985" box="[1013,1436,627,650]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="levis">
<emphasis box="[1013,1156,627,649]" italics="true" pageId="17" pageNumber="502">Grania levis</emphasis>
<bibRefCitation author="Coates KA &amp; Erseus C" box="[1166,1436,627,650]" 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>
, originally described from somewhat further north, from Georges Bank, south-east of
<collectingRegion box="[1164,1335,689,711]" country="United States of America" name="Massachusetts" pageId="17" pageNumber="502">Massachusetts</collectingRegion>
,
<collectingCountry box="[1349,1406,689,710]" name="United States of America" pageId="17" pageNumber="502">USA</collectingCountry>
.
</paragraph>
<paragraph blockId="17.[1025,1243,766,787]" box="[1025,1243,766,787]" pageId="17" pageNumber="502">
<collectionCode box="[1025,1243,766,787]" pageId="17" pageNumber="502">GENETIC ANALYSES</collectionCode>
</paragraph>
</subSubSection>
<subSubSection lastPageId="19" lastPageNumber="504" pageId="17" pageNumber="502" type="description">
<paragraph blockId="17.[826,1442,804,1438]" box="[826,993,804,826]" pageId="17" pageNumber="502">
<emphasis box="[826,993,804,826]" italics="true" pageId="17" pageNumber="502">COI clustering</emphasis>
</paragraph>
<paragraph blockId="17.[826,1442,804,1438]" pageId="17" pageNumber="502">
The Bayesian inference of the
<emphasis box="[1189,1236,835,856]" italics="true" pageId="17" pageNumber="502">COI</emphasis>
sequences divide the 38 individuals into ten well-supported clades (
<figureCitation box="[834,912,896,918]" captionStart="Figure 11" captionStartId="18.[144,223,1626,1645]" captionTargetBox="[146,1422,195,1594]" captionTargetId="figure-92@18.[144,1424,193,1596]" captionTargetPageId="18" captionText="Figure 11. Majority-rule consensus gene tree of cytochrome c oxidase subunit I (COI) locus from all species described in this study. Posterior probabilities calculated from Bayesian Markov chain Monte Carlo (MCMC) inference are noted above the branches. Note the four clades within Grania chilensis sp. nov." figureDoi="http://doi.org/10.5281/zenodo.4720648" httpUri="https://zenodo.org/record/4720648/files/figure.png" pageId="17" pageNumber="502">Fig. 11</figureCitation>
), four of which are found in
<collectingCountry box="[1242,1388,896,918]" name="South Africa" pageId="17" pageNumber="502">South Africa</collectingCountry>
, two in
<collectingCountry box="[857,920,927,949]" name="Chile" pageId="17" pageNumber="502">Chile</collectingCountry>
, one in
<collectingCountry box="[1013,1085,927,949]" name="Brazil" pageId="17" pageNumber="502">Brazil</collectingCountry>
, and three in the North Atlantic. Within-clade variation is generally low, but in one clade, i.e. all specimens referred to the new taxon
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[826,957,1018,1040]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="chilensis" status="sp. nov.">
<emphasis box="[826,957,1018,1040]" italics="true" pageId="17" pageNumber="502">G. chilensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[964,1051,1019,1040]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
, there is a notable subclustering pattern, dividing this clade into four subclades. A haplotype network (
<figureCitation box="[1138,1226,1080,1102]" captionStart="Figure 12" captionStartId="19.[164,243,1208,1227]" captionTargetBox="[166,1442,197,1177]" captionTargetId="figure-299@19.[164,1444,195,1179]" captionTargetPageId="19" captionText="Figure 12. Haplotype network of individuals of Grania chilensis sp. nov. Note the long distance (35 steps) between the two groups found in Valdivia." figureDoi="http://doi.org/10.5281/zenodo.4720650" httpUri="https://zenodo.org/record/4720650/files/figure.png" pageId="17" pageNumber="502">Fig. 12</figureCitation>
) indicates that
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[826,957,1110,1131]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="chilensis" status="sp. nov.">
<emphasis box="[826,957,1110,1131]" italics="true" pageId="17" pageNumber="502">G. chilensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[964,1049,1111,1132]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
is structured geographically, with two subclades found in the southernmost site (Valdivia), one subclade in the northernmost site (Coquimbo), and an intermediate subclade in the intermediately located site (Concepcion). Pairwise genetic distances indicate that in general there is a strong barcoding gap present between lineages within this group. In the
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[826,951,1324,1346]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="chilensis" status="sp. nov.">
<emphasis box="[826,951,1324,1346]" italics="true" pageId="17" pageNumber="502">G. chilensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[958,1039,1325,1346]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
clade, however, there is higher than average within-species divergence, although not nearly as great as the lowest between-species differences (
<figureCitation box="[834,914,1416,1438]" captionStart="Figure 13" captionStartId="20.[144,223,985,1004]" captionTargetBox="[369,1197,196,952]" captionTargetId="figure-437@20.[367,1199,194,954]" captionTargetPageId="20" captionText="Figure 13. Distance histogram of pairwise uncorrected p distances between all individuals analysed genetically in this study. Within-species distances range from 0 to 8% (03% if not counting Grania chilensis sp. nov.); between-species distances range from 16 to 25%." figureDoi="http://doi.org/10.5281/zenodo.4720655" httpUri="https://zenodo.org/record/4720655/files/figure.png" pageId="17" pageNumber="502">Fig. 13</figureCitation>
).
</paragraph>
<paragraph blockId="17.[826,1442,1480,1716]" box="[826,990,1480,1501]" pageId="17" pageNumber="502">
<emphasis box="[826,990,1480,1501]" italics="true" pageId="17" pageNumber="502">ITS clustering</emphasis>
</paragraph>
<paragraph blockId="17.[826,1442,1480,1716]" pageId="17" pageNumber="502">
The Bayesian inference analysis of 23
<emphasis box="[1274,1317,1511,1532]" italics="true" pageId="17" pageNumber="502">ITS</emphasis>
sequences supports all ten clusters found in the mitochondrial data (
<figureCitation box="[897,977,1571,1594]" captionStart="Figure 14" captionStartId="21.[164,243,1355,1374]" captionTargetBox="[166,1442,197,1323]" captionTargetId="figure-234@21.[164,1444,195,1325]" captionTargetPageId="21" captionText="Figure 14. Majority-rule consensus gene tree of the ITS region (ITS1-5.8S rDNA-ITS2) from all species described in this study. Posterior probabilities calculated from Bayesian Markov chain Monte Carlo (MCMC) inference are noted above the branches. Note that the four Grania chilensis sp. nov. clades seen in the COI locus are absent here." figureDoi="http://doi.org/10.5281/zenodo.4720657" httpUri="https://zenodo.org/record/4720657/files/figure.png" pageId="17" pageNumber="502">Fig. 14</figureCitation>
); however, although there is also variation within the
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[1016,1144,1602,1623]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="17" pageNumber="502" phylum="Annelida" rank="species" species="chilensis" status="sp. nov.">
<emphasis box="[1016,1144,1602,1623]" italics="true" pageId="17" pageNumber="502">G. chilensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1150,1233,1602,1623]" pageId="17" pageNumber="502" rank="species">sp. nov.</taxonomicNameLabel>
cluster in the
<emphasis box="[1398,1441,1602,1623]" italics="true" pageId="17" pageNumber="502">ITS</emphasis>
region, the geographic substructuring is not seen here. Instead, the variation seems to be randomly distributed with respect to geography.
</paragraph>
<paragraph blockId="17.[826,1442,1757,1902]" box="[826,1259,1757,1779]" pageId="17" pageNumber="502">
<emphasis box="[826,1259,1757,1779]" italics="true" pageId="17" pageNumber="502">Phylogenetic placement of new species</emphasis>
</paragraph>
<paragraph blockId="17.[826,1442,1757,1902]" lastBlockId="19.[164,779,1329,1443]" lastPageId="19" lastPageNumber="504" pageId="17" pageNumber="502">
The updated phylogeny is completely congruent with that described in
<bibRefCitation author="De Wit P &amp; Rota E &amp; Erseus C" box="[1020,1248,1818,1840]" pageId="17" pageNumber="502" pagination="509 - 519" refId="ref17070" refString="De Wit P, Rota E, Erseus C. 2011 b. Phylogeny and character evolution in Grania (Annelida, Clitellata). Zoologica Scripta 40: 509 - 519." type="journal article" year="2011">
De Wit
<emphasis box="[1105,1151,1818,1840]" italics="true" pageId="17" pageNumber="502">et al</emphasis>
. (2011b)
</bibRefCitation>
, containing three main clades (A, B, C in
<figureCitation box="[1087,1163,1849,1871]" captionStart="Figure 15" captionStartId="22.[144,223,1323,1342]" captionTargetBox="[146,1422,197,1291]" captionTargetId="figure-242@22.[144,1424,195,1293]" captionTargetPageId="22" captionText="Figure 15. Multi-locus phylogeny of all available genetic data in Grania to date, using three mitochondrial and three nuclear loci. New species from this study are marked in bold. Majority-rule consensus tree from Bayesian Markov chain Monte Carlo (MCMC) inference; clade posterior probabilities are given next to the nodes." figureDoi="http://doi.org/10.5281/zenodo.4720659" httpUri="https://zenodo.org/record/4720659/files/figure.png" pageId="17" pageNumber="502">Fig. 15</figureCitation>
). All of the South African species form one strongly supported clade within clade A, together with all the North Atlantic species. By contrast, the Chilean and the Brazilian species are placed in clade B, together with
<taxonomicName authority="De Wit &amp; Erseus, 2007" authorityName="De Wit &amp; Erseus" authorityYear="2007" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="curta">
<emphasis box="[419,561,1819,1840]" italics="true" pageId="18" pageNumber="503">Grania curta</emphasis>
<bibRefCitation author="De Wit P &amp; Erseus C" pageId="18" pageNumber="503" pagination="27 - 50" refId="ref16899" refString="De Wit P, Erseus C. 2007. Seven new species of Grania (Annelida: Clitellata: Enchytraeidae) from New Caledonia, South Pacific Ocean. Zootaxa 50: 27 - 50." type="journal article" year="2007">De Wit &amp; Erséus, 2007</bibRefCitation>
</taxonomicName>
and
<taxonomicName authorityName="Kennedy" authorityYear="1966" box="[256,408,1850,1871]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="americana">
<emphasis box="[256,408,1850,1871]" italics="true" pageId="18" pageNumber="503">G. americana</emphasis>
</taxonomicName>
.
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[423,608,1849,1871]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="unitheca" status="sp. nov.">
<emphasis box="[423,608,1849,1871]" italics="true" pageId="18" pageNumber="503">Grania unitheca</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[615,699,1850,1871]" pageId="18" pageNumber="503" rank="species">sp. nov.</taxonomicNameLabel>
from shallow water in
<collectingRegion box="[353,531,1880,1902]" country="United States of America" name="North Carolina" pageId="18" pageNumber="503">North Carolina</collectingRegion>
(North Atlantic) is placed together with the other North American species
<taxonomicName authorityName="Erseus &amp; Lasserre" authorityYear="1976" box="[806,1039,1788,1809]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="monospermatheca">
<emphasis box="[806,1039,1788,1809]" italics="true" pageId="18" pageNumber="503">G. monospermatheca</emphasis>
</taxonomicName>
(its sister taxon) and
<taxonomicName authorityName="Locke &amp; Coates" authorityYear="1999" box="[1290,1415,1788,1809]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="laxartus">
<emphasis box="[1290,1415,1788,1809]" italics="true" pageId="18" pageNumber="503">G. laxartus</emphasis>
</taxonomicName>
; however,
<taxonomicName authority="Prantoni &amp; Wit &amp; Erséus, 2016" authorityName="Prantoni &amp; Wit &amp; Erséus" authorityYear="2016" box="[911,1074,1819,1840]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="carolinensis" status="sp. nov.">
<emphasis box="[911,1074,1819,1840]" italics="true" pageId="18" pageNumber="503">G. carolinensis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1080,1162,1819,1840]" pageId="18" pageNumber="503" rank="species">sp. nov.</taxonomicNameLabel>
and the closely related immature specimen of
<taxonomicName authorityName="Coates &amp; Erseus" authorityYear="1985" box="[1072,1249,1849,1871]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="18" pageNumber="503" phylum="Annelida" rank="species" species="levis">
<emphasis box="[1072,1154,1850,1871]" italics="true" pageId="18" pageNumber="503">Grania</emphasis>
cf.
<emphasis box="[1198,1249,1849,1870]" italics="true" pageId="18" pageNumber="503">levis</emphasis>
</taxonomicName>
, both found in deep water off the North Carolinian coast, are placed as the sister clade of
<taxonomicName baseAuthorityName="Knollner" baseAuthorityYear="1935" box="[394,608,1329,1350]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="504" phylum="Annelida" rank="species" species="postclitellochaeta">
<emphasis box="[394,608,1329,1350]" italics="true" pageId="19" pageNumber="504">G. postclitellochaeta</emphasis>
</taxonomicName>
and the cryptic
<taxonomicName authorityName="De Wit &amp; Erseus" authorityYear="2010" box="[164,277,1360,1381]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="504" phylum="Annelida" rank="species" species="occulta">
<emphasis box="[164,277,1360,1381]" italics="true" pageId="19" pageNumber="504">G. occulta</emphasis>
</taxonomicName>
, whereas
<taxonomicName authorityName="Erseus" authorityYear="1977" box="[396,522,1360,1381]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="504" phylum="Annelida" rank="species" species="ovitheca">
<emphasis box="[396,522,1360,1381]" italics="true" pageId="19" pageNumber="504">G. ovitheca</emphasis>
</taxonomicName>
, which is morphologically identical to
<taxonomicName authorityName="De Wit &amp; Erseus" authorityYear="2010" box="[368,480,1390,1412]" class="Clitellata" family="Enchytraeidae" genus="Grania" kingdom="Animalia" order="Enchytraeida" pageId="19" pageNumber="504" phylum="Annelida" rank="species" species="occulta">
<emphasis box="[368,480,1390,1412]" italics="true" pageId="19" pageNumber="504">G. occulta</emphasis>
</taxonomicName>
, is strongly supported as the sister to this four-taxon group (
<figureCitation box="[580,661,1421,1443]" captionStart="Figure 15" captionStartId="22.[144,223,1323,1342]" captionTargetBox="[146,1422,197,1291]" captionTargetId="figure-242@22.[144,1424,195,1293]" captionTargetPageId="22" captionText="Figure 15. Multi-locus phylogeny of all available genetic data in Grania to date, using three mitochondrial and three nuclear loci. New species from this study are marked in bold. Majority-rule consensus tree from Bayesian Markov chain Monte Carlo (MCMC) inference; clade posterior probabilities are given next to the nodes." figureDoi="http://doi.org/10.5281/zenodo.4720659" httpUri="https://zenodo.org/record/4720659/files/figure.png" pageId="19" pageNumber="504">Fig. 15</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>