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<mods:title id="CF0D8E2A0D11960DE2C4F5D55A4C41D0">An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California</mods:title>
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<subSubSection id="C34F6578FFA861401CB1FAB42F85FAE7" box="[151,398,1303,1330]" pageId="17" pageNumber="468" type="nomenclature">
<paragraph id="8BEA36F3FFA861401CB1FAB42F85FAE7" blockId="17.[151,398,1303,1363]" box="[151,398,1303,1330]" pageId="17" pageNumber="468">
<heading id="D0A2819FFFA861401CB1FAB42F85FAE7" bold="true" box="[151,398,1303,1330]" fontSize="11" level="1" pageId="17" pageNumber="468" reason="1">
<emphasis id="B921EAE1FFA861401CB1FAB42F85FAE7" bold="true" box="[151,398,1303,1330]" pageId="17" pageNumber="468">
<taxonomicName id="4C554D70FFA861401CB1FAB42F4FFAE4" ID-CoL="74Z7S" authority="Rouse &amp; Goffredi &amp; Johnson &amp; Vrijenhoek, 2018" authorityName="Rouse &amp; Goffredi &amp; Johnson &amp; Vrijenhoek" authorityYear="2018" box="[151,324,1303,1329]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="17" pageNumber="468" phylum="Annelida" rank="species" species="lonnyi" status="sp. nov.">
<emphasis id="B921EAE1FFA861401CB1FAB42F4FFAE4" bold="true" box="[151,324,1303,1329]" italics="true" pageId="17" pageNumber="468">Osedax lonnyi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A212579AFFA861401D6AFABB2F85FAE7" box="[332,398,1304,1330]" pageId="17" pageNumber="468" rank="species">n. sp.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861401CB1FA992EFEFA87" box="[151,245,1338,1363]" pageId="17" pageNumber="468" type="description">
<paragraph id="8BEA36F3FFA861401CB1FA992EFEFA87" blockId="17.[151,398,1303,1363]" box="[151,245,1338,1363]" pageId="17" pageNumber="468">
<figureCitation id="136E2A76FFA861401CB1FA992EFEFA87" box="[151,245,1338,1363]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Figure 7</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861401CB1FA212CD0FA4D" box="[151,731,1410,1432]" pageId="17" pageNumber="468" type="reference_group">
<paragraph id="8BEA36F3FFA861401CB1FA212CD0FA4D" blockId="17.[151,731,1410,1432]" box="[151,731,1410,1432]" pageId="17" pageNumber="468">
nude-palp-F (
<bibRefCitation id="EFC44B02FFA861401D16FA212FECFA4D" author="Rouse, G. W. &amp; Wilson, N. G. &amp; Worsaae, K. &amp; Vrijenhoek, R. C." box="[304,487,1410,1432]" journalOrPublisher="Current Biology" pageId="17" pageNumber="468" pagination="236 - 241" part="25" publicationUrl="https://doi.org/10.1016/j.cub.2014.11.032" refString="Rouse, G. W., Wilson, N. G., Worsaae, K. &amp; Vrijenhoek, R. C. (2015) A dwarf male reversal in bone-eating worms. Current Biology, 25, 236 - 241. https: // doi. org / 10.1016 / j. cub. 2014.11.032" title="A dwarf male reversal in bone-eating worms" type="journal article" year="2015">
Rouse
<emphasis id="B921EAE1FFA861401D5EFA202FA0FA4D" box="[376,427,1410,1432]" italics="true" pageId="17" pageNumber="468">et al.</emphasis>
2015
</bibRefCitation>
;
<bibRefCitation id="EFC44B02FFA861401DD7FA212CDFFA4D" author="Rouse, G. W. &amp; Wilson, N. G. &amp; Goffredi, S. K. &amp; Johnson, S. B. &amp; Smart, T. &amp; Widmer, C. &amp; Young, C. M. &amp; Vrijenhoek, R. C." box="[497,724,1410,1432]" journalOrPublisher="Marine Biology" pageId="17" pageNumber="468" pagination="395 - 405" part="156" publicationUrl="https://doi.org/10.1007/s00227-008-1091-z" refString="Rouse, G. W., Wilson, N. G., Goffredi, S. K., Johnson, S. B., Smart, T., Widmer, C., Young, C. M. &amp; Vrijenhoek, R. C. (2009) Spawning and development in Osedax boneworms (Siboglinidae, Annelida). Marine Biology, 156, 395 - 405. https: // doi. org / 10.1007 / s 00227 - 008 - 1091 - z" title="Spawning and development in Osedax boneworms (Siboglinidae, Annelida)" type="journal article" year="2009">
Vrijenhoek
<emphasis id="B921EAE1FFA861401E41FA202C90FA4D" box="[615,667,1410,1432]" italics="true" pageId="17" pageNumber="468">et al.</emphasis>
2009
</bibRefCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861401CB1FA662FD5F99F" pageId="17" pageNumber="468" type="materials_examined">
<paragraph id="8BEA36F3FFA861401CB1FA662FD5F99F" blockId="17.[151,1436,1477,2006]" pageId="17" pageNumber="468">
<emphasis id="B921EAE1FFA861401CB1FA662C1FFA0B" bold="true" box="[151,532,1477,1502]" pageId="17" pageNumber="468">Material examined. Holotype:</emphasis>
Female, SIO-BIC A1647 (GenBank
<emphasis id="B921EAE1FFA861401FCBFA652A15FA08" box="[1005,1054,1478,1501]" italics="true" pageId="17" pageNumber="468">COI</emphasis>
sequence
<accessionNumber id="9406AB10FFA86140188FFA662B17FA0B" box="[1193,1308,1477,1502]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/FJ347643" pageId="17" pageNumber="468">FJ347643</accessionNumber>
), fixed in glutaraldehyde-preserved in ethanol, collected from natural whale fall (
<taxonomicName id="4C554D70FFA861401FC3FA4A2AD6F9D7" baseAuthorityName="Lilljeborg" baseAuthorityYear="1861" box="[997,1245,1513,1538]" class="Mammalia" family="Eschrichtiidae" genus="Eschrichtius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="17" pageNumber="468" phylum="Chordata" rank="species" species="robustus">
<emphasis id="B921EAE1FFA861401FC3FA4A2AD6F9D7" box="[997,1245,1513,1538]" italics="true" pageId="17" pageNumber="468">Eschrichtius robustus</emphasis>
</taxonomicName>
) found at 2898 meters depth in Monterey Submarine Canyon, California (36°36.606N; 122°26.122W) ROV
<emphasis id="B921EAE1FFA861401920F9AE2B55F9F3" box="[1286,1374,1549,1574]" italics="true" pageId="17" pageNumber="468">Tiburon</emphasis>
dive number 1162, Dec. 19, 2007.
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861401CE1F9F62C33F8F7" pageId="17" pageNumber="468" type="diagnosis">
<paragraph id="8BEA36F3FFA861401CE1F9F62C33F8F7" blockId="17.[151,1436,1477,2006]" pageId="17" pageNumber="468">
<emphasis id="B921EAE1FFA861401CE1F9F62C0FF9BB" bold="true" box="[199,516,1621,1646]" pageId="17" pageNumber="468">Diagnosis and description.</emphasis>
Holotype female; in life trunk 1 mm long, 0.5 mm wide (
<figureCitation id="136E2A76FFA8614018B4F9F62AE7F9BB" box="[1170,1260,1621,1646]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7A</figureCitation>
), crown of four palps, 2.5 cm long (
<figureCitation id="136E2A76FFA861401D5BF9DA2FD2F947" box="[381,473,1657,1682]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7A</figureCitation>
), likely contracted even at this length. Tube gelatinous, thin, fitting about trunk and base of crown, 4 cm long (
<figureCitation id="136E2A76FFA861401D99F93E2C13F963" box="[447,536,1693,1718]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7A</figureCitation>
). In life, palps with visible blood vessels and greenish tips (
<figureCitation id="136E2A76FFA861401896F93E2B02F963" box="[1200,1289,1693,1718]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7A</figureCitation>
, inset). Palps without obvious pinnules, oviduct extends slightly beyond palps (
<figureCitation id="136E2A76FFA861401F85F9622DF4F90F" box="[931,1023,1729,1754]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7A</figureCitation>
inset). Trunk with pigmented ring demarking upper and lower trunk (
<figureCitation id="136E2A76FFA861401E39F9462C73F92B" box="[543,632,1765,1790]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7B</figureCitation>
). Ovisac 2 mm by 1 mm with damaged lobate greenish roots extending outwards (
<figureCitation id="136E2A76FFA861401D2BF8AA2F6EF8F7" box="[269,357,1801,1826]" captionStart="FIGURE 7" captionStartId="18.[151,250,1521,1543]" captionTargetBox="[171,1407,375,1492]" captionTargetId="figure@18.[171,1407,375,1492]" captionTargetPageId="18" captionText="FIGURE 7. Osedax lonnyi n. sp., previously Osedax nude-palp-F. A. Image of live holotype (SIO-BIC A1647) still in tube and attached to whale bone lying at 2898 meters. Note very long palps occupying about 2/3 of tube. No part of trunk is apparent. B. Live holotype dissected from bone and tube. Note short trunk with pigmented ring separating upper and lower trunk.Little root tissue could be extracted, but there was an ovisac region." httpUri="https://zenodo.org/record/1165888/files/figure.png" pageId="17" pageNumber="468">Fig. 7B</figureCitation>
). Males not found.
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861401CE1F88E2D1BF8BF" pageId="17" pageNumber="468" type="distribution">
<paragraph id="8BEA36F3FFA861401CE1F88E2D1BF8BF" blockId="17.[151,1436,1477,2006]" pageId="17" pageNumber="468">
<emphasis id="B921EAE1FFA861401CE1F88E2F6BF893" bold="true" box="[199,352,1837,1862]" pageId="17" pageNumber="468">Distribution.</emphasis>
Known from Monterey Bay, California from 2898 meters depth (
<tableCitation id="C6D70348FFA86140181FF88E2A84F893" box="[1081,1167,1837,1862]" captionStart="TABLE 2" captionStartId="2.[151,239,152,175]" captionText="TABLE 2. Specimen voucher and GenBankaccession numbers for the DNA sequences used in this study. Bold indicates new sequences. COI = cytochrome c oxidase, H3 = Histone H3, other loci are for ribosomal RNA." pageId="17" pageNumber="468">Table 2</tableCitation>
). It has only been found in a whale bone fragment, adjacent to the main skeleton.
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861401CF3F8D62A6FF867" pageId="17" pageNumber="468" type="etymology">
<paragraph id="8BEA36F3FFA861401CF3F8D62A6FF867" blockId="17.[151,1436,1477,2006]" pageId="17" pageNumber="468">
<emphasis id="B921EAE1FFA861401CF3F8D62F57F85B" bold="true" box="[213,348,1909,1934]" pageId="17" pageNumber="468">Etymology.</emphasis>
This species is named (noun in the genitive case) for Lonny Lundsten, Senior Research Technician at MBARI, for his enthusiasm and assistance on many
<taxonomicName id="4C554D70FFA861401FA6F83A2DDFF867" authorityName="Rouse, Goffredi &amp; Vrijenhoek" authorityYear="2004" box="[896,980,1945,1970]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="17" pageNumber="468" phylum="Annelida" rank="genus">
<emphasis id="B921EAE1FFA861401FA6F83A2DDFF867" box="[896,980,1945,1970]" italics="true" pageId="17" pageNumber="468">Osedax</emphasis>
</taxonomicName>
expeditions.
</paragraph>
</subSubSection>
<subSubSection id="C34F6578FFA861431CE1F81E2FC5FE95" lastPageId="18" lastPageNumber="469" pageId="17" pageNumber="468" type="discussion">
<paragraph id="8BEA36F3FFA861431CE1F81E2FC5FE95" blockId="17.[151,1436,1477,2006]" lastBlockId="18.[151,1437,151,320]" lastPageId="18" lastPageNumber="469" pageId="17" pageNumber="468">
<emphasis id="B921EAE1FFA861401CE1F81E2F30F803" bold="true" box="[199,315,1981,2006]" pageId="17" pageNumber="468">Remarks.</emphasis>
<taxonomicName id="4C554D70FFA861401D62F81E2FE3F803" authority="Rouse &amp; Goffredi &amp; Johnson &amp; Vrijenhoek, 2018" authorityName="Rouse &amp; Goffredi &amp; Johnson &amp; Vrijenhoek" authorityYear="2018" box="[324,488,1981,2006]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="17" pageNumber="468" phylum="Annelida" rank="species" species="lonnyi" status="sp. nov.">
<emphasis id="B921EAE1FFA861401D62F81E2FE3F803" box="[324,488,1981,2006]" italics="true" pageId="17" pageNumber="468">Osedax lonnyi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A212579AFFA861401DD4F81E2C3FF803" box="[498,564,1981,2006]" pageId="17" pageNumber="468" rank="species">
<emphasis id="B921EAE1FFA861401DD4F81E2C3FF803" bold="true" box="[498,564,1981,2006]" pageId="17" pageNumber="468">n. sp.</emphasis>
</taxonomicNameLabel>
is part of
<taxonomicName id="4C554D70FFA861401E97F81E2D0EF803" authorityName="Rouse, Goffredi &amp; Vrijenhoek" authorityYear="2004" box="[689,773,1981,2006]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="17" pageNumber="468" phylum="Annelida" rank="genus">
<emphasis id="B921EAE1FFA861401E97F81E2D0EF803" box="[689,773,1981,2006]" italics="true" pageId="17" pageNumber="468">Osedax</emphasis>
</taxonomicName>
Clade II and is the well-supported sistergroup to
<emphasis id="B921EAE1FFA86143196EF81E2EE6FF65" italics="true" lastPageId="18" lastPageNumber="469" pageId="17" pageNumber="468">
<taxonomicName id="4C554D70FFA86143196EF81E2EE1FF65" authorityName="Amon, Wiklund, Dahlgren, Copley, Smith, Jamieson &amp;" authorityYear="2014" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" lastPageId="18" lastPageNumber="469" order="Sabellida" pageId="17" pageNumber="468" phylum="Annelida" rank="species" species="rogersi">Osedax rogersi</taxonomicName>
,
</emphasis>
an Antarctic species with an uncorrected divergence of 12%.
<taxonomicName id="4C554D70FFAB61431F91FF342A50FF65" authority="Rouse &amp; Goffredi &amp; Johnson &amp; Vrijenhoek, 2018" authorityName="Rouse &amp; Goffredi &amp; Johnson &amp; Vrijenhoek" authorityYear="2018" box="[951,1115,151,176]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="18" pageNumber="469" phylum="Annelida" rank="species" species="lonnyi" status="sp. nov.">
<emphasis id="B921EAE1FFAB61431F91FF342A50FF65" box="[951,1115,151,176]" italics="true" pageId="18" pageNumber="469">Osedax lonnyi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A212579AFFAB61431845FF342AAFFF65" box="[1123,1188,151,176]" pageId="18" pageNumber="469" rank="species">
<emphasis id="B921EAE1FFAB61431845FF342AAFFF65" bold="true" box="[1123,1188,151,176]" pageId="18" pageNumber="469">n. sp.</emphasis>
</taxonomicNameLabel>
is remarkable for the very long palps (2.5 cm) that it shows relative to the very short trunk (1.5 mm). The species is known from only a single specimen (holotype) collected in 2007 and no further specimens were found during any subsequent sampling at the same site. This is somewhat surprising as it is one of the larger species of Clade II and the whale fall was sampled a number of times.
</paragraph>
</subSubSection>
</treatment>
</document>