322 lines
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322 lines
68 KiB
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<mods:title id="26ADABB5A50DA9035FE9B6059A2E52EC">An inordinate fondness for Osedax (Siboglinidae: Annelida): Fourteen new species of bone worms from California</mods:title>
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<mods:namePart id="EE91120F2E10DE38E0F1C336A426A1EC">Johnson, Shannon B.</mods:namePart>
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<heading id="D0A2819FFFA5614D1CB1FF342FAEFF67" bold="true" box="[151,421,151,178]" fontSize="11" level="1" pageId="28" pageNumber="479" reason="1">
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<taxonomicName id="4C554D70FFA5614D1CB1FF342F50FF64" ID-CoL="74YW4" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[151,347,151,177]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="lehmani" status="sp. nov.">
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<emphasis id="B921EAE1FFA5614D1CB1FF342F50FF64" bold="true" box="[151,347,151,177]" italics="true" pageId="28" pageNumber="479">Osedax lehmani</emphasis>
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<taxonomicNameLabel id="A212579AFFA5614D1D44FF3B2FAEFF67" box="[354,421,152,178]" pageId="28" pageNumber="479" rank="species">n. sp.</taxonomicNameLabel>
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<subSubSection id="C34F6578FFA5614D1CB1FF192F32FF06" box="[151,313,186,211]" pageId="28" pageNumber="479" type="description">
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<paragraph id="8BEA36F3FFA5614D1CB1FF192F32FF06" blockId="28.[151,421,151,211]" box="[151,313,186,211]" pageId="28" pageNumber="479">
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<figureCitation id="136E2A76FFA5614D1CB1FF192F1BFF06" box="[151,272,186,211]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Figures 14</figureCitation>
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,
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<figureCitation id="136E2A76FFA5614D1D3AFF192F32FF06" box="[284,313,186,211]" captionStart="FIGURE 15" captionStartId="27.[151,250,1726,1748]" captionTargetBox="[151,1436,193,1705]" captionTargetId="figure@27.[151,1436,193,1705]" captionTargetPageId="27" captionText="FIGURE 15. Osedax lehmani n. sp., previously Osedax ‘yellow collar’, ‘sp. 3 SBJ-2006’. A. Transverse section (7 µm) of paratype 3 (SIO-BIC A7807), through crown showing pinnules of all palps oriented dorsally and oviduct lying between the dorsalmost pair of palps. B. Transverse section (7 µm) through trunk of paratype 3 (SIO-BIC A7807) showing extensive musculatureand glands.Prominent dorsal and ventral blood vesselsare alsovisible.C. Live males, allotypes (SIO-BICA7805) in tube of their female (her pinnules are visible). Males show small patches of pigment anteriorly and posteriorly. D. Differential interference micrograph of a male dissected from tube of paratype 2 (SIO-BIC A7806). Prototroch is visible anteriorly.Somehooksare visiblein expanded posterior region. Nomature sperm is visible, though thereare severalbundlesof spermatids." httpUri="https://zenodo.org/record/1165904/files/figure.png" pageId="28" pageNumber="479">15</figureCitation>
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</paragraph>
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<paragraph id="8BEA36F3FFA5614D1CB1FEA12D76FEE2" blockId="28.[151,1436,258,403]" pageId="28" pageNumber="479">
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‘yellow collar’, (
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<bibRefCitation id="EFC44B02FFA5614D1D6CFEA12C06FECD" author="Braby, C. E. & Rouse, G. W. & Johnson, S. B. & Jones, W. J & Vrijenhoek, R. C." box="[330,525,258,280]" journalOrPublisher="Deep-Sea Research Part I" pageId="28" pageNumber="479" pagination="1773 - 1791" part="54" publicationUrl="https://doi.org/10.1016/j.dsr.2007.05.014" refString="Braby, C. E., Rouse, G. W., Johnson, S. B., Jones, W. J. & Vrijenhoek, R. C. (2007) Bathymetric and temporal variation among Osedax boneworms and associated megafauna on whale-falls in Monterey Bay, California. Deep-Sea Research Part I, 54, 1773 - 1791. https: // doi. org / 10.1016 / j. dsr. 2007.05.014" title="Bathymetric and temporal variation among Osedax boneworms and associated megafauna on whale-falls in Monterey Bay, California" type="journal article" year="2007">
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Braby
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<emphasis id="B921EAE1FFA5614D1DB2FEA02FC6FECD" box="[404,461,258,280]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2007
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</bibRefCitation>
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;
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<bibRefCitation id="EFC44B02FFA5614D1E3AFEA12D06FECD" author="Huusgaard, R. S. & Vismann, B. & Kuhl, M. & Macnaugton, M. & Colmander, V & Rouse, G. W." box="[540,781,258,280]" journalOrPublisher="The potent respiratory system of Osedax mucofloris (Siboglinidae, Annelida) - a prerequisite for the origin of bone-eating Osedax? PLoS ONE" pageId="28" pageNumber="479" pagination="35975" part="7" publicationUrl="https://doi.org/10.1371/journal.pone.0035975" refString="Huusgaard, R. S., Vismann, B., Kuhl, M., Macnaugton, M., Colmander, V., Rouse, G. W., Glover, A. G., Dahlgren, T. & Worsaae, K. (2012) The potent respiratory system of Osedax mucofloris (Siboglinidae, Annelida) - a prerequisite for the origin of bone-eating Osedax? PLoS ONE, 7, e 35975. https: // doi. org / 10.1371 / journal. pone. 0035975" title="Glover, A. G., Dahlgren, T. & Worsaae, K" type="journal article" year="2012">
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Huusgaard
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<emphasis id="B921EAE1FFA5614D1EB2FEA02CC6FECD" box="[660,717,258,280]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2012
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</bibRefCitation>
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;
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<bibRefCitation id="EFC44B02FFA5614D1F3AFEA02DFEFECD" author="Katz, S. & Rouse, G. W." box="[796,1013,258,280]" journalOrPublisher="Invertebrate Biology" pageId="28" pageNumber="479" pagination="368 - 385" part="132" publicationUrl="https://doi.org/10.1111/ivb.12037" refString="Katz, S. & Rouse, G. W. (2013) The reproductive system of Osedax (Annelida, Siboglinidae): sperm ultrastructure, ovary structure and fertilization. Invertebrate Biology, 132, 368 - 385. https: // doi. org / 10.1111 / ivb. 12037" title="The reproductive system of Osedax (Annelida, Siboglinidae): sperm ultrastructure, ovary structure and fertilization" type="journal article" year="2013">Katz & Rouse 2013</bibRefCitation>
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;
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<bibRefCitation id="EFC44B02FFA5614D1822FEA02ACCFECD" author="Rouse, G. W. & Wilson, N. G. & Goffredi, S. K. & Johnson, S. B. & Smart, T. & Widmer, C. & Young, C. M. & Vrijenhoek, R. C." box="[1028,1223,258,280]" journalOrPublisher="Marine Biology" pageId="28" pageNumber="479" 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. & 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">
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Rouse
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<emphasis id="B921EAE1FFA5614D1868FEA02A8DFECD" box="[1102,1158,258,280]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2009
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</bibRefCitation>
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;
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<bibRefCitation id="EFC44B02FFA5614D18F3FEA02B93FECD" author="Rouse, G. W. & Wilson, N. G. & Worsaae, K. & Vrijenhoek, R. C." box="[1237,1432,258,280]" journalOrPublisher="Current Biology" pageId="28" pageNumber="479" 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. & 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">
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Rouse
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<emphasis id="B921EAE1FFA5614D1906FEA02B53FECD" box="[1312,1368,258,280]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2015
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</bibRefCitation>
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;
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<bibRefCitation id="EFC44B02FFA5614D1CE1FE812FB8FEE2" author="Tresguerres, M. & Katz, S. & Rouse, G. W." box="[199,435,289,311]" journalOrPublisher="Proceedings of the Royal Society of London Series B-Biological Sciences" pageId="28" pageNumber="479" pagination="20130625" part="280" publicationUrl="https://doi.org/10.1098/rspb.2013.0625" refString="Tresguerres, M., Katz, S. & Rouse, G. W. (2013) How to get into bones: proton pump and carbonic anhydrase in Osedax boneworms. Proceedings of the Royal Society of London Series B-Biological Sciences, 280, 20130625. https: // doi. org / 10.1098 / rspb. 2013.0625" title="How to get into bones: proton pump and carbonic anhydrase in Osedax boneworms" type="journal article" year="2013">
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Tresguerres
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<emphasis id="B921EAE1FFA5614D1D62FE812F7CFEE2" box="[324,375,289,311]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2013
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;
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<bibRefCitation id="EFC44B02FFA5614D1D9AFE822CA9FEE2" author="Rouse, G. W. & Wilson, N. G. & Goffredi, S. K. & Johnson, S. B. & Smart, T. & Widmer, C. & Young, C. M. & Vrijenhoek, R. C." box="[444,674,289,311]" journalOrPublisher="Marine Biology" pageId="28" pageNumber="479" 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. & 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">
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Vrijenhoek
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<emphasis id="B921EAE1FFA5614D1E15FE812C6DFEE2" box="[563,614,289,311]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2009
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</bibRefCitation>
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;
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<bibRefCitation id="EFC44B02FFA5614D1E8AFE812D7DFEE2" author="Worsaae, K. & Rimskaya-Korsakova, N. N. & Rouse, G. W." box="[684,886,289,311]" journalOrPublisher="BMC Evolutionary Biology" pageId="28" pageNumber="479" pagination="83" part="16" publicationUrl="https://doi.org/10.1186/s12862-016-0639-7" refString="Worsaae, K., Rimskaya-Korsakova, N. N. & Rouse, G. W. (2016) Neural reconstruction of bone-eating Osedax spp. (Annelida) and evolution of the siboglinid nervous system. BMC Evolutionary Biology, 16, 83. https: // doi. org / 10.1186 / s 12862 - 016 - 0639 - 7" title="Neural reconstruction of bone-eating Osedax spp. (Annelida) and evolution of the siboglinid nervous system" type="journal article" year="2016">
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Worsaae
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<emphasis id="B921EAE1FFA5614D1F2FFE812D37FEE2" box="[777,828,289,311]" italics="true" pageId="28" pageNumber="479">et al.</emphasis>
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2016
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</bibRefCitation>
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)
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</paragraph>
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<paragraph id="8BEA36F3FFA5614D1CB1FEE32D31FE83" blockId="28.[151,1436,258,403]" box="[151,826,320,342]" pageId="28" pageNumber="479">
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‘sp.3 SBJ-2006’ (GenBank
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<emphasis id="B921EAE1FFA5614D1D96FEE22FD0FE83" box="[432,475,321,342]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
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sequences
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<accessionNumber id="9406AB10FFA5614D1E69FEE32CCAFE83" box="[591,705,320,342]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996629" pageId="28" pageNumber="479">DQ996629</accessionNumber>
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- DQ99638)
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‘sp.4 SBJ-2006’ (GenBank
|
||
<emphasis id="B921EAE1FFA5614D1D96FEFC2FD0FEA1" box="[432,475,351,372]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequences
|
||
<accessionNumber id="9406AB10FFA5614D1E69FEFD2CC9FEA1" box="[591,706,350,372]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996640" pageId="28" pageNumber="479">DQ996640</accessionNumber>
|
||
, DQ99643)
|
||
</paragraph>
|
||
<paragraph id="8BEA36F3FFA5614D1CB1FEDE2CA6FE46" blockId="28.[151,1436,258,403]" box="[151,685,381,403]" pageId="28" pageNumber="479">
|
||
‘orange collar’ (GenBank
|
||
<emphasis id="B921EAE1FFA5614D1D84FEDD2FC6FE46" box="[418,461,382,403]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequence
|
||
<accessionNumber id="9406AB10FFA5614D1E11FEDE2CADFE46" box="[567,678,381,403]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/EU267672" pageId="28" pageNumber="479">EU267672</accessionNumber>
|
||
)
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C34F6578FFA5614D1CB1FE1C2C8EFD2D" pageId="28" pageNumber="479" type="materials_examined">
|
||
<paragraph id="8BEA36F3FFA5614D1CB1FE1C2C8EFD2D" blockId="28.[151,1437,447,1696]" pageId="28" pageNumber="479">
|
||
<emphasis id="B921EAE1FFA5614D1CB1FE1C2C02FE0D" bold="true" box="[151,521,447,472]" pageId="28" pageNumber="479">Material examined. Holotype</emphasis>
|
||
: SIO-BIC A7804, Female (GenBank
|
||
<emphasis id="B921EAE1FFA5614D1FC2FE632A1FFE02" box="[996,1044,448,471]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequence
|
||
<accessionNumber id="9406AB10FFA5614D18B8FE1C2B15FE0D" box="[1182,1310,447,472]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996629" pageId="28" pageNumber="479">DQ996629</accessionNumber>
|
||
), fixed in glutaraldehyde, preserved in ethanol, collected from a whale carcass (
|
||
<taxonomicName id="4C554D70FFA5614D1FF4FE472AC2FE28" baseAuthorityName="Lilljeborg" baseAuthorityYear="1861" box="[978,1225,484,509]" class="Mammalia" family="Eschrichtiidae" genus="Eschrichtius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Cetacea" pageId="28" pageNumber="479" phylum="Chordata" rank="species" species="robustus">
|
||
<emphasis id="B921EAE1FFA5614D1FF4FE472AC2FE28" box="[978,1225,484,509]" italics="true" pageId="28" pageNumber="479">Eschrichtius robustus</emphasis>
|
||
</taxonomicName>
|
||
) deployed at 385 meters depth in Monterey Submarine Canyon, California (36°47.401’N; 122° 53.235’W), ROV
|
||
<emphasis id="B921EAE1FFA5614D192EFDA42B6BFDF5" box="[1288,1376,519,544]" italics="true" pageId="28" pageNumber="479">Tiburon</emphasis>
|
||
dive number 933, Jan. 6, 2006.
|
||
<emphasis id="B921EAE1FFA5614D1DF6FD8F2C43FD90" bold="true" box="[464,584,556,581]" pageId="28" pageNumber="479">Paratypes</emphasis>
|
||
: Females, fixed in formalin or glutaraldehyde, preserved in ethanol, same locality and date as holotype, SIO-BIC A1640 (GenBank
|
||
<emphasis id="B921EAE1FFA5614D1F10FDF32D6CFDB2" box="[822,871,592,615]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequence
|
||
<accessionNumber id="9406AB10FFA5614D1FC7FDEC2A69FDBD" box="[993,1122,591,616]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996630" pageId="28" pageNumber="479">DQ996630</accessionNumber>
|
||
), same locality and date as holotype, SIO-BIC A7806 (GenBank
|
||
<emphasis id="B921EAE1FFA5614D1E74FDD62C89FD59" box="[594,642,629,652]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequence
|
||
<accessionNumber id="9406AB10FFA5614D1F26FDD72D8AFD58" box="[768,897,628,653]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996631" pageId="28" pageNumber="479">DQ996631</accessionNumber>
|
||
); SIO-BIC A7807 and A7808 (GenBank
|
||
<emphasis id="B921EAE1FFA5614D194AFDD62B96FD59" box="[1388,1437,629,652]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequence
|
||
<accessionNumber id="9406AB10FFA5614D1D21FD3B2F8AFD65" box="[263,385,664,688]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/EU223332" pageId="28" pageNumber="479">EU223332</accessionNumber>
|
||
and
|
||
<accessionNumber id="9406AB10FFA5614D1D9BFD3B2C32FD65" box="[445,569,664,688]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/EU223337" pageId="28" pageNumber="479">EU223337</accessionNumber>
|
||
, respectively), same locality as holotype, ROV
|
||
<emphasis id="B921EAE1FFA5614D1844FD342AB1FD65" box="[1122,1210,663,688]" italics="true" pageId="28" pageNumber="479">Tiburon</emphasis>
|
||
dive number 1070, Jan. 11, 2007; dwarf males (Allotypes) SIO-BIC A7805 (no GenBank
|
||
<emphasis id="B921EAE1FFA5614D1F9DFD1E2DE7FD01" box="[955,1004,701,724]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequence), several in tubes of various females, same locality and date as holotype.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C34F6578FFA5614D1CE1FCA72DFBFB7D" pageId="28" pageNumber="479" type="diagnosis">
|
||
<paragraph id="8BEA36F3FFA5614D1CE1FCA72DFBFB7D" blockId="28.[151,1437,447,1696]" pageId="28" pageNumber="479">
|
||
<emphasis id="B921EAE1FFA5614D1CE1FCA72C1BFCC8" bold="true" box="[199,528,772,797]" pageId="28" pageNumber="479">Diagnosis and description.</emphasis>
|
||
Holotype female, in life trunk 8 mm long, 0.6 mm wide; crown of palps contracted, curled, 1.5 mm long (
|
||
<figureCitation id="136E2A76FFA5614D1E2AFC842C78FC95" box="[524,627,807,832]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14C</figureCitation>
|
||
). Palps up to 10 mm long when extended in other specimens (
|
||
<figureCitation id="136E2A76FFA5614D190AFC842EA4FCB1" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Figs 14A, D</figureCitation>
|
||
). Pinnules of all palps oriented dorsally and the oviduct lying between the dorsalmost pair of palps (
|
||
<figureCitation id="136E2A76FFA5614D192AFCEF2EA0FC52" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Figs 14B, D, E</figureCitation>
|
||
,
|
||
<figureCitation id="136E2A76FFA5614D1C91FCCC2EE0FC5D" box="[183,235,879,904]" captionStart="FIGURE 15" captionStartId="27.[151,250,1726,1748]" captionTargetBox="[151,1436,193,1705]" captionTargetId="figure@27.[151,1436,193,1705]" captionTargetPageId="27" captionText="FIGURE 15. Osedax lehmani n. sp., previously Osedax ‘yellow collar’, ‘sp. 3 SBJ-2006’. A. Transverse section (7 µm) of paratype 3 (SIO-BIC A7807), through crown showing pinnules of all palps oriented dorsally and oviduct lying between the dorsalmost pair of palps. B. Transverse section (7 µm) through trunk of paratype 3 (SIO-BIC A7807) showing extensive musculatureand glands.Prominent dorsal and ventral blood vesselsare alsovisible.C. Live males, allotypes (SIO-BICA7805) in tube of their female (her pinnules are visible). Males show small patches of pigment anteriorly and posteriorly. D. Differential interference micrograph of a male dissected from tube of paratype 2 (SIO-BIC A7806). Prototroch is visible anteriorly.Somehooksare visiblein expanded posterior region. Nomature sperm is visible, though thereare severalbundlesof spermatids." httpUri="https://zenodo.org/record/1165904/files/figure.png" pageId="28" pageNumber="479">15A</figureCitation>
|
||
). Oviduct extends from trunk into crown for 1.75 mm (
|
||
<figureCitation id="136E2A76FFA5614D1F79FCCC2DBEFC5D" box="[863,949,879,904]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Figs 14</figureCitation>
|
||
B-D). In life, palps reddish with two major blood vessels in each (
|
||
<figureCitation id="136E2A76FFA5614D1DB7FC372C2BFC79" box="[401,544,916,941]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Figs 14A, D</figureCitation>
|
||
). Tube a gelatinous loose mass around trunk (
|
||
<figureCitation id="136E2A76FFA5614D183BFC372A8EFC79" box="[1053,1157,916,941]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14D</figureCitation>
|
||
). Trunk with pale yellow ring around anterior margin; ring broken mid-dorsally by oviduct (
|
||
<figureCitation id="136E2A76FFA5614D1FB9FC142A3FFC05" box="[927,1076,951,976]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Figs 14B, D</figureCitation>
|
||
). and mid-ventrally by an oval unpigmented patch (
|
||
<figureCitation id="136E2A76FFA5614D1DA2FC7F2FE4FC21" box="[388,495,988,1013]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14C</figureCitation>
|
||
). Clear demarcation of upper and lower trunk with oviduct emerging onto outer surface of upper trunk (
|
||
<figureCitation id="136E2A76FFA5614D1D87FC5C2C03FBCD" box="[417,520,1023,1048]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14B</figureCitation>
|
||
). Internally trunk shows extensive musculature and glands with prominent dorsal and ventral blood vessels (
|
||
<figureCitation id="136E2A76FFA5614D1DE0FB872C26FBE8" box="[454,557,1060,1085]" captionStart="FIGURE 15" captionStartId="27.[151,250,1726,1748]" captionTargetBox="[151,1436,193,1705]" captionTargetId="figure@27.[151,1436,193,1705]" captionTargetPageId="27" captionText="FIGURE 15. Osedax lehmani n. sp., previously Osedax ‘yellow collar’, ‘sp. 3 SBJ-2006’. A. Transverse section (7 µm) of paratype 3 (SIO-BIC A7807), through crown showing pinnules of all palps oriented dorsally and oviduct lying between the dorsalmost pair of palps. B. Transverse section (7 µm) through trunk of paratype 3 (SIO-BIC A7807) showing extensive musculatureand glands.Prominent dorsal and ventral blood vesselsare alsovisible.C. Live males, allotypes (SIO-BICA7805) in tube of their female (her pinnules are visible). Males show small patches of pigment anteriorly and posteriorly. D. Differential interference micrograph of a male dissected from tube of paratype 2 (SIO-BIC A7806). Prototroch is visible anteriorly.Somehooksare visiblein expanded posterior region. Nomature sperm is visible, though thereare severalbundlesof spermatids." httpUri="https://zenodo.org/record/1165904/files/figure.png" pageId="28" pageNumber="479">Fig. 15B</figureCitation>
|
||
). Ovisac in holotype extending laterally as two masses; simple lobes in other specimens (
|
||
<figureCitation id="136E2A76FFA5614D1D3AFBE42F88FBB5" box="[284,387,1095,1120]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14C</figureCitation>
|
||
). Roots of holotype extend from ovisac lobes (
|
||
<figureCitation id="136E2A76FFA5614D1FB2FBE42DF7FBB5" box="[916,1020,1095,1120]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14C</figureCitation>
|
||
); in other specimens, roots as simple lobes or extending laterally in two lobe or lobes (
|
||
<figureCitation id="136E2A76FFA5614D1EF2FBCF2D36FB51" box="[724,829,1132,1157]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14B</figureCitation>
|
||
). Males dwarfs, with spermatids and sperm, chaetalbearing segments not inflated; found in tube lumen of females (
|
||
<figureCitation id="136E2A76FFA5614D1F78FB2C2DA6FB7D" box="[862,941,1167,1192]" captionStart="FIGURE 15" captionStartId="27.[151,250,1726,1748]" captionTargetBox="[151,1436,193,1705]" captionTargetId="figure@27.[151,1436,193,1705]" captionTargetPageId="27" captionText="FIGURE 15. Osedax lehmani n. sp., previously Osedax ‘yellow collar’, ‘sp. 3 SBJ-2006’. A. Transverse section (7 µm) of paratype 3 (SIO-BIC A7807), through crown showing pinnules of all palps oriented dorsally and oviduct lying between the dorsalmost pair of palps. B. Transverse section (7 µm) through trunk of paratype 3 (SIO-BIC A7807) showing extensive musculatureand glands.Prominent dorsal and ventral blood vesselsare alsovisible.C. Live males, allotypes (SIO-BICA7805) in tube of their female (her pinnules are visible). Males show small patches of pigment anteriorly and posteriorly. D. Differential interference micrograph of a male dissected from tube of paratype 2 (SIO-BIC A7806). Prototroch is visible anteriorly.Somehooksare visiblein expanded posterior region. Nomature sperm is visible, though thereare severalbundlesof spermatids." httpUri="https://zenodo.org/record/1165904/files/figure.png" pageId="28" pageNumber="479">Fig. 15</figureCitation>
|
||
C-D).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C34F6578FFA5614D1CE1FB172FA7FB25" pageId="28" pageNumber="479" type="distribution">
|
||
<paragraph id="8BEA36F3FFA5614D1CE1FB172FA7FB25" blockId="28.[151,1437,447,1696]" pageId="28" pageNumber="479">
|
||
<emphasis id="B921EAE1FFA5614D1CE1FB172F6BFB18" bold="true" box="[199,352,1204,1229]" pageId="28" pageNumber="479">Distribution.</emphasis>
|
||
Known from Monterey Bay, California only from 349 meters depth (
|
||
<tableCitation id="C6D70348FFA5614D184BFB172ACEFB19" box="[1133,1221,1204,1229]" 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="28" pageNumber="479">Table 2</tableCitation>
|
||
). It has been found in whale and cow bones.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C34F6578FFA5614D1CE1FB5F2B49FAED" pageId="28" pageNumber="479" type="etymology">
|
||
<paragraph id="8BEA36F3FFA5614D1CE1FB5F2B49FAED" blockId="28.[151,1437,447,1696]" pageId="28" pageNumber="479">
|
||
<emphasis id="B921EAE1FFA5614D1CE1FB5F2F45FAC0" bold="true" box="[199,334,1276,1301]" pageId="28" pageNumber="479">Etymology.</emphasis>
|
||
This species is named (noun in the genitive case) in memory of Alan George Lehman, father of Ellen Lehman, in recognition of her long and continued support of the Scripps Oceanographic Collections.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C34F6578FFA5614D1CE1FAE72F25F975" pageId="28" pageNumber="479" type="discussion">
|
||
<paragraph id="8BEA36F3FFA5614D1CE1FAE72F25F975" blockId="28.[151,1437,447,1696]" pageId="28" pageNumber="479">
|
||
<emphasis id="B921EAE1FFA5614D1CE1FAE72F30FA88" bold="true" box="[199,315,1348,1373]" pageId="28" pageNumber="479">Remarks.</emphasis>
|
||
<taxonomicName id="4C554D70FFA5614D1D64FAE72FFCFA88" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[322,503,1348,1373]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="lehmani" status="sp. nov.">
|
||
<emphasis id="B921EAE1FFA5614D1D64FAE72FFCFA88" box="[322,503,1348,1373]" italics="true" pageId="28" pageNumber="479">Osedax lehmani</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A212579AFFA5614D1DD8FAE72C35FA88" box="[510,574,1348,1373]" pageId="28" pageNumber="479" rank="species">
|
||
<emphasis id="B921EAE1FFA5614D1DD8FAE72C35FA88" bold="true" box="[510,574,1348,1373]" pageId="28" pageNumber="479">n. sp.</emphasis>
|
||
</taxonomicNameLabel>
|
||
is part of
|
||
<taxonomicName id="4C554D70FFA5614D1E96FAE72D0EFA88" authorityName="Rouse, Goffredi & Vrijenhoek" authorityYear="2004" box="[688,773,1348,1373]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="genus">
|
||
<emphasis id="B921EAE1FFA5614D1E96FAE72D0EFA88" box="[688,773,1348,1373]" italics="true" pageId="28" pageNumber="479">Osedax</emphasis>
|
||
</taxonomicName>
|
||
Clade IV and is closest relative to
|
||
<taxonomicName id="4C554D70FFA5614D18ABFAE72B75FA88" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[1165,1406,1348,1373]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="packardorum" status="sp. nov.">
|
||
<emphasis id="B921EAE1FFA5614D18ABFAE72B75FA88" box="[1165,1406,1348,1373]" italics="true" pageId="28" pageNumber="479">Osedax packardorum</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A212579AFFA5614D19A3FAE72EB3FA55" pageId="28" pageNumber="479" rank="species">
|
||
<emphasis id="B921EAE1FFA5614D19A3FAE72EB3FA55" bold="true" pageId="28" pageNumber="479">n. sp.</emphasis>
|
||
</taxonomicNameLabel>
|
||
, (
|
||
<figureCitation id="136E2A76FFA5614D1CF3FAC42F17FA55" box="[213,284,1383,1408]" captionStart="FIGURE 1" captionStartId="4.[151,250,1820,1842]" captionTargetBox="[204,1329,193,1779]" captionTargetId="figure@4.[204,1330,193,1789]" captionTargetPageId="4" captionText="FIGURE 1. Osedax phylogenetic analyses. Osedax multi-gene phylogeny with Monilifera as outgroup.Maximum likelihood tree topology based on a partitioned dataset of five gene segments (MAFFT-aligned) for data shown in Table 2. The Bayesian analyses gave the same topology. Bootstrap support % (BS) is at each node, followed by posterior probability (PP) from the MrBayes analyses. * indicates values were ± 95% (BS) and 0.95 (PP). Missing values indicate BS <50% and PP <0.7. Six major Osedax clades are distinguished, following Vrijenhoek et al. (2009), Rouse et al. (2015) and Taboada et al. (2015). The various forms of palps (or absence thereof in the case of O. jabba n. sp.) are indicated for the major clades but these are not intended to indicate apomorphic states." httpUri="https://zenodo.org/record/1165876/files/figure.png" pageId="28" pageNumber="479">Fig. 1</figureCitation>
|
||
), with a minimum uncorrected distance for
|
||
<emphasis id="B921EAE1FFA5614D1F13FACB2D6DFAAA" box="[821,870,1384,1407]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
of 8-9% (
|
||
<tableCitation id="C6D70348FFA5614D1FCFFAC42A4DFA55" box="[1001,1094,1383,1408]" captionStart="TABLE4" captionStartId="7.[125,203,466,486]" captionText="TABLE4.COIpairwise uncorrecteđ đistance tablegenerateđ withPAUP*.Distances are expresseđ as %anđ are rounđeđ off.The sequences are as listeđin Table2, except for the followingađđitional sequences: O. knutei n.sp. B is GB sequence JF59952, O. lehmani n.sp. B is GB sequence DQ996629, Osedax lehmani n.sp. Cis GB sequence DQ996631.Sagami B isFM998090.Osedax Sagami 6is regarđeđ here as part of O. docricketts n. sp., Osedax Sagami 7 is regarđeđ here as part of O. randyi n.sp., Osedax Sagami8is regarđeđ here as part O. westernflyeri n.sp., Distanceshighlighteđ as bolđ are điscusseđin the text." pageId="28" pageNumber="479">Table 4</tableCitation>
|
||
). This is one of the smaller intraspecific distances among
|
||
<taxonomicName id="4C554D70FFA5614D1DD6FA2F2C4FFA70" authorityName="Rouse, Goffredi & Vrijenhoek" authorityYear="2004" box="[496,580,1420,1445]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="genus">
|
||
<emphasis id="B921EAE1FFA5614D1DD6FA2F2C4FFA70" box="[496,580,1420,1445]" italics="true" pageId="28" pageNumber="479">Osedax</emphasis>
|
||
</taxonomicName>
|
||
species, with others being between
|
||
<taxonomicName id="4C554D70FFA5614D1FC0FA2E2A83FA70" box="[998,1160,1420,1445]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="frankpressi">
|
||
<emphasis id="B921EAE1FFA5614D1FC0FA2E2A83FA70" box="[998,1160,1420,1445]" italics="true" pageId="28" pageNumber="479">O. frankpressi</emphasis>
|
||
</taxonomicName>
|
||
and the
|
||
<emphasis id="B921EAE1FFA5614D18C9FA2E2B00FA71" box="[1263,1291,1421,1444]" italics="true" pageId="28" pageNumber="479">O.</emphasis>
|
||
'1336_61_2' specimen from the Atlantic at ~10%; and
|
||
<taxonomicName id="4C554D70FFA5614D1E49FA132CD0FA1D" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[623,731,1455,1480]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="randyi" status="sp. nov.">
|
||
<emphasis id="B921EAE1FFA5614D1E49FA132CD0FA1D" box="[623,731,1455,1480]" italics="true" pageId="28" pageNumber="479">O. randyi</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A212579AFFA5614D1EC5FA0C2D29FA1D" box="[739,802,1455,1480]" pageId="28" pageNumber="479" rank="species">
|
||
<emphasis id="B921EAE1FFA5614D1EC5FA0C2D29FA1D" bold="true" box="[739,802,1455,1480]" pageId="28" pageNumber="479">n. sp.</emphasis>
|
||
</taxonomicNameLabel>
|
||
and O. ‘MB16 at ~ 6-7% (
|
||
<tableCitation id="C6D70348FFA5614D1872FA0C2AA7FA1D" box="[1108,1196,1455,1480]" captionStart="TABLE4" captionStartId="7.[125,203,466,486]" captionText="TABLE4.COIpairwise uncorrecteđ đistance tablegenerateđ withPAUP*.Distances are expresseđ as %anđ are rounđeđ off.The sequences are as listeđin Table2, except for the followingađđitional sequences: O. knutei n.sp. B is GB sequence JF59952, O. lehmani n.sp. B is GB sequence DQ996629, Osedax lehmani n.sp. Cis GB sequence DQ996631.Sagami B isFM998090.Osedax Sagami 6is regarđeđ here as part of O. docricketts n. sp., Osedax Sagami 7 is regarđeđ here as part of O. randyi n.sp., Osedax Sagami8is regarđeđ here as part O. westernflyeri n.sp., Distanceshighlighteđ as bolđ are điscusseđin the text." pageId="28" pageNumber="479">Table 4</tableCitation>
|
||
). The majority of the 31 available
|
||
<emphasis id="B921EAE1FFA5614D1D0BFA762F56FA39" box="[301,349,1493,1516]" italics="true" pageId="28" pageNumber="479">COI</emphasis>
|
||
sequences for
|
||
<taxonomicName id="4C554D70FFA5614D1E29FA772CCCFA38" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[527,711,1492,1517]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="lehmani" status="sp. nov.">
|
||
<emphasis id="B921EAE1FFA5614D1E29FA772CCCFA38" box="[527,711,1492,1517]" italics="true" pageId="28" pageNumber="479">Osedax lehmani</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A212579AFFA5614D1EF4FA772D1FFA38" box="[722,788,1492,1517]" pageId="28" pageNumber="479" rank="species">
|
||
<emphasis id="B921EAE1FFA5614D1EF4FA772D1FFA38" bold="true" box="[722,788,1492,1517]" pageId="28" pageNumber="479">n. sp.</emphasis>
|
||
</taxonomicNameLabel>
|
||
(
|
||
<tableCitation id="C6D70348FFA5614D1F00FA772D75FA39" box="[806,894,1492,1517]" captionStart="TABLE 3" captionStartId="5.[125,200,461,480]" captionText="TABLE 3. Details of Osedax species locations, substrates, depth, vouchers,types and GenBank sequences (COI). New species are bold." pageId="28" pageNumber="479">Table 3</tableCitation>
|
||
) show less than 1% divergence, though a few sequences e.g.
|
||
<accessionNumber id="9406AB10FFA5614D1D66FA542FB7F9C5" box="[320,444,1527,1552]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996640" pageId="28" pageNumber="479">DQ996640</accessionNumber>
|
||
are around 2% divergent from the majority and
|
||
<accessionNumber id="9406AB10FFA5614D1FF9FA542A57F9C5" box="[991,1116,1527,1552]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/DQ996631" pageId="28" pageNumber="479">DQ996631</accessionNumber>
|
||
shows the greatest distance, at ~3% (
|
||
<tableCitation id="C6D70348FFA5614D1CDFF9BF2F59F9E1" box="[249,338,1564,1589]" captionStart="TABLE4" captionStartId="7.[125,203,466,486]" captionText="TABLE4.COIpairwise uncorrecteđ đistance tablegenerateđ withPAUP*.Distances are expresseđ as %anđ are rounđeđ off.The sequences are as listeđin Table2, except for the followingađđitional sequences: O. knutei n.sp. B is GB sequence JF59952, O. lehmani n.sp. B is GB sequence DQ996629, Osedax lehmani n.sp. Cis GB sequence DQ996631.Sagami B isFM998090.Osedax Sagami 6is regarđeđ here as part of O. docricketts n. sp., Osedax Sagami 7 is regarđeđ here as part of O. randyi n.sp., Osedax Sagami8is regarđeđ here as part O. westernflyeri n.sp., Distanceshighlighteđ as bolđ are điscusseđin the text." pageId="28" pageNumber="479">Table 4</tableCitation>
|
||
), so the possibility of there being a cryptic species complex for
|
||
<taxonomicName id="4C554D70FFA5614D1813F9BE2ABFF9E0" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[1077,1204,1564,1589]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="lehmani" status="sp. nov.">
|
||
<emphasis id="B921EAE1FFA5614D1813F9BE2ABFF9E0" box="[1077,1204,1564,1589]" italics="true" pageId="28" pageNumber="479">O. lehmani</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A212579AFFA5614D189BF9BF2AF5F9E0" box="[1213,1278,1564,1589]" pageId="28" pageNumber="479" rank="species">
|
||
<emphasis id="B921EAE1FFA5614D189BF9BF2AF5F9E0" bold="true" box="[1213,1278,1564,1589]" pageId="28" pageNumber="479">n. sp.</emphasis>
|
||
</taxonomicNameLabel>
|
||
needs further investigation. The most distinguishing feature of
|
||
<taxonomicName id="4C554D70FFA5614D1EE4F99C2D7CF98D" authority="Rouse & Goffredi & Johnson & Vrijenhoek, 2018" authorityName="Rouse & Goffredi & Johnson & Vrijenhoek" authorityYear="2018" box="[706,887,1599,1624]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="species" species="lehmani" status="sp. nov.">
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||
<emphasis id="B921EAE1FFA5614D1EE4F99C2D7CF98D" box="[706,887,1599,1624]" italics="true" pageId="28" pageNumber="479">Osedax lehmani</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A212579AFFA5614D1F59F99C2DB5F98D" box="[895,958,1599,1624]" pageId="28" pageNumber="479" rank="species">
|
||
<emphasis id="B921EAE1FFA5614D1F59F99C2DB5F98D" bold="true" box="[895,958,1599,1624]" pageId="28" pageNumber="479">n. sp.</emphasis>
|
||
</taxonomicNameLabel>
|
||
is the yellow ring around the anterior part of the trunk. (
|
||
<figureCitation id="136E2A76FFA5614D1D09F9C72F74F9A9" box="[303,383,1636,1661]" captionStart="FIGURE 14" captionStartId="26.[151,250,1590,1612]" captionTargetBox="[193,1361,193,1567]" captionTargetId="figure@26.[193,1361,193,1567]" captionTargetPageId="26" captionText="FIGURE 14. Osedax lehmani n. sp., previously Osedax ‘yellow collar’. A. Several adult females on rib bone of a gray whale calf (Eschrichtius robustus) that was deployed at385 meters inMonterey Canyon.Ribwas recovered tostudy the Osedax in an aquariumsetting.Note distinctive yellow ringaround most of the anteriorend of trunk.The massivecirculatory system intrunk and palps is evident from red color of worms. B. Micrograph of live paratype 4 (SIO-BIC A7808), a female, in dorsal view dissected from gray whale deployed at 385 meters. The ‘root’ mass is asingle lobe extending away to the right of ovisac. Palps are retracted and partly curled, with pinnules all oriented dorsally.Note mid-dorsal gap in yellow pigment ring, through which oviduct runs and extends into crown.Note demarcation of the lower and upper trunk (arrow). Oviduct emerges onto surface of dorsal trunk at this point. Oviduct is full of oocytes and highly contracted nature of the trunk is evident in zigzag shape of the oviduct on trunk. C. Live holotype (SIO-BIC A7804), a female, in ventral view. Holotype is partially dissected from the gray whale deployed at 385 meters.Root mass(also containing ovarian tissue) has been dissected out ofbone on right sideand more extends into the bone, away from ovisac. Palps are retracted and curled. Note midventral gap in yellow pigment ring. D. Live adultfemale inlateral view on same ribbone as in A.There is a gap in yellowpigment ring whereoviduct traverses thetrunk to become part of the crown. E. Scanning electron micrograph of a female dissected from whale bone deployed at 385 meters. Pinnules of all palps are oriented dorsally." httpUri="https://zenodo.org/record/1165902/files/figure.png" pageId="28" pageNumber="479">Fig. 14</figureCitation>
|
||
A-D). This is the only
|
||
<taxonomicName id="4C554D70FFA5614D1E5AF9C72CDBF9A8" authorityName="Rouse, Goffredi & Vrijenhoek" authorityYear="2004" box="[636,720,1636,1661]" class="Polychaeta" family="Siboglinidae" genus="Osedax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Sabellida" pageId="28" pageNumber="479" phylum="Annelida" rank="genus">
|
||
<emphasis id="B921EAE1FFA5614D1E5AF9C72CDBF9A8" box="[636,720,1636,1661]" italics="true" pageId="28" pageNumber="479">Osedax</emphasis>
|
||
</taxonomicName>
|
||
species from Monterey Bay that was not found deeper than 389 meters depth.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |