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<mods:title id="6133CA5F2D1A57193595541F0E18FB7D">The invertebrate host of salmonid fish parasites Ceratonova shasta and Parvicapsula minibicornis (Cnidaria: Myxozoa), is a novel fabriciid annelid, Manayunkia occidentalis sp. nov. (Sabellida: Fabriciidae)</mods:title>
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<taxonomicName id="4C6E4D7B4A22DB0C9EFEFD32C9FB8EDA" ID-CoL="8707aaf5-c47f-43f1-b328-80d73a8b8133" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[151,456,729,756]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="3" pageNumber="313" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A22DB0C9EFEFD32C9FB8EDA" bold="true" box="[151,456,729,756]" italics="true" pageId="3" pageNumber="313">Manayunkia occidentalis</emphasis>
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<figureCitation id="13552A7D4A22DB0C9EFEFD16C92F8F39" box="[151,284,765,791]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStart-2="FIGURE 4" captionStartId-0="4.[151,250,1752,1777]" captionStartId-1="5.[151,250,1663,1688]" captionStartId-2="6.[151,250,1804,1829]" captionTargetBox-0="[152,1435,956,1726]" captionTargetBox-1="[149,1438,309,1638]" captionTargetBox-2="[155,1433,561,1779]" captionTargetId-0="figure@4.[152,1435,956,1726]" captionTargetId-1="figure@5.[149,1438,309,1638]" captionTargetId-2="figure@6.[155,1433,561,1779]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionTargetPageId-2="6" captionText-0="FIGURE 2. Gross morphological features of Manayunkia occidentalis sp. nov. resolved by light microscopy: A, multiple individuals emerged and feeding from their tubes; B, Individual annelid with prominent vascularized appendages (va) and showing refractile spots on one feeding palp on each side (arrowheads); C, female in reflected light, showing white oocytes (o) in segments 46; D, male in reflected light showing bright spermatogonic tissue (sp) in multiple posterior segments; E, Male in transmitted light showing parallel vascularized appendages (va) and fanned-out feeding palps anteriorly with two dark eyespots in peristomium, and granular coelom posteriorly due to presence of developing sperm." captionText-1="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." captionText-2="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi-0="http://doi.org/10.5281/zenodo.3713132" figureDoi-1="http://doi.org/10.5281/zenodo.3713134" figureDoi-2="http://doi.org/10.5281/zenodo.3713136" httpUri-0="https://zenodo.org/record/3713132/files/figure.png" httpUri-1="https://zenodo.org/record/3713134/files/figure.png" httpUri-2="https://zenodo.org/record/3713136/files/figure.png" pageId="3" pageNumber="313">Figures 24</figureCitation>
</paragraph>
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<paragraph id="8BD136F84A22DB0C9EFEFCAACD178F56" blockId="3.[151,1437,832,888]" pageId="3" pageNumber="313">
<taxonomicName id="4C6E4D7B4A22DB0C9EFEFCAACA338F77" ID-CoL="3XSXD" authority=": Hazel, 1966" authorityName=": Hazel" authorityYear="1966" box="[151,512,832,857]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="3" pageNumber="313" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A22DB0C9EFEFCAAC9478F77" box="[151,372,833,857]" italics="true" pageId="3" pageNumber="313">Manayunkia speciosa</emphasis>
:
<bibRefCitation id="EFFF4B094A22DB0C9FEBFCABCA338F77" author="Hazel, C. R." box="[386,512,832,857]" pageId="3" pageNumber="313" pagination="533 - 535" refId="ref6323" refString="Hazel, C. R. (1966) A note on the freshwater polychaete Manayunkia speciosa Leidy, from California and Oregon. Ohio Journal of Science, 66 (5), 533 - 535." type="journal article" year="1966">Hazel, 1966</bibRefCitation>
</taxonomicName>
;
<bibRefCitation id="EFFF4B094A22DB0C9C62FCABCB3F8F77" author="Bartholomew, J. L. &amp; Whipple, M. J. &amp; Stevens, D. G. &amp; Fryer, J. L." box="[523,780,832,857]" pageId="3" pageNumber="313" pagination="859 - 868" refId="ref5690" refString="Bartholomew, J. L., Whipple, M. J., Stevens, D. G. &amp; Fryer, J. L. (1997) The life cycle of Ceratomyxa shasta, a myxosporean parasite of salmonids, requires a freshwater polychaete as an alternate host. Journal of Parasitology, 83, 859 - 868. https: // doi. org / 10.2307 / 3284281" type="journal article" year="1997">
Bartholomew
<emphasis id="B91AEAEA4A22DB0C9CF5FCA9CAFA8F77" box="[668,713,833,857]" italics="true" pageId="3" pageNumber="313">et al</emphasis>
. 1997
</bibRefCitation>
; 2006; 2007;
<bibRefCitation id="EFFF4B094A22DB0C9DFFFCABCC448F77" author="Alexander, J. D. &amp; Hallett, S. L. &amp; Stocking, R. &amp; Xue, L. &amp; Bartholomew, J. L." box="[918,1143,832,857]" pageId="3" pageNumber="313" pagination="219 - 233" refId="ref5475" refString="Alexander, J. D., Hallett, S. L., Stocking, R., Xue, L. &amp; Bartholomew, J. L. (2014) Host and parasite populations after a flood. Northwest Science, 88, 219 - 233. https: // doi. org / 10.3955 / 046.088.0305" type="journal article" year="2014">
Alexander
<emphasis id="B91AEAEA4A22DB0C9A6FFCA9CC098F77" box="[1030,1082,833,857]" italics="true" pageId="3" pageNumber="313">et al.</emphasis>
2014
</bibRefCitation>
;
<bibRefCitation id="EFFF4B094A22DB0C9AE8FCABC8CE8F56" author="Stocking, R. W. &amp; Bartholomew, J. L." pageId="3" pageNumber="313" pagination="78 - 88" refId="ref7675" refString="Stocking, R. W. &amp; Bartholomew, J. L. (2007) Distribution and habitat characteristics of Manayunkia speciosa and infection prevalence with the parasite Ceratomyxa shasta in the Klamath River, Oregon-California. Journal of Parasitology, 93, 78 - 88. https: // doi. org / 10.1645 / GE- 939 R. 1" type="journal article" year="2007">Stocking and Bartholomew, 2007</bibRefCitation>
;
<bibRefCitation id="EFFF4B094A22DB0C9F6EFCB4CA1B8F56" author="Bjork, S. J. &amp; Bartholomew, J. L." box="[263,552,863,888]" pageId="3" pageNumber="313" pagination="29 - 37" refId="ref5903" refString="Bjork, S. J. &amp; Bartholomew, J. L. (2009) Effects of Ceratomyxa shasta dose on a susceptible strain of rainbow trout and comparatively resistant Chinook and coho salmon. Diseases of Aquatic Organisms, 86, 29 - 37. https: // doi. org / 10.3354 / dao 02092" type="journal article" year="2009">Bjork &amp; Bartholomew 2009</bibRefCitation>
;
<bibRefCitation id="EFFF4B094A22DB0C9C58FCB4CAC58F56" author="Willson, S. J. &amp; Wilzbach, M. A. &amp; Malakauskas, D. M. &amp; Cummins, K. W." box="[561,758,863,888]" pageId="3" pageNumber="313" pagination="183 - 191" refId="ref7800" refString="Willson, S. J., Wilzbach, M. A., Malakauskas, D. M. &amp; Cummins, K. W. (2010) Lab Rearing of a Freshwater Polychaete (Manayunkia speciosa, Sabellidae) Host for Salmon Pathogens. Northwest Science, 84 (1), 183 - 191. https: // doi. org / 10.3955 / 046.084.0207" type="journal article" year="2010">
Willson
<emphasis id="B91AEAEA4A22DB0C9CEEFC8ACA808F56" box="[647,691,864,888]" italics="true" pageId="3" pageNumber="313">et al</emphasis>
. 2010
</bibRefCitation>
;
<bibRefCitation id="EFFF4B094A22DB0C9C97FCB4CBE58F56" author="Atkinson, S. D. &amp; Jones, S. R. M. &amp; Adlard, R. &amp; Bartholomew, J. L." box="[766,982,863,888]" pageId="3" pageNumber="313" pagination="960 - 968" refId="ref5535" refString="Atkinson, S. D., Jones, S. R. M., Adlard, R. &amp; Bartholomew, J. L. (2011) Geographical and host distribution patterns of Parvicapsula minibicornis (Myxozoa) small subunit ribosomal RNA genetic types. Parasitology, 138 (8), 960 - 968. https: // doi. org / 10.1017 / S 0031182011000734" type="journal article" year="2011">
Atkinson
<emphasis id="B91AEAEA4A22DB0C9D0BFC8ACBBD8F56" box="[866,910,864,888]" italics="true" pageId="3" pageNumber="313">et al</emphasis>
., 2011
</bibRefCitation>
;
<bibRefCitation id="EFFF4B094A22DB0C9D89FCB4CCD38F56" author="Malakauskas, D. M. &amp; Willson, S. J. &amp; Wilzbach, M. A. &amp; Som, N. A." box="[992,1248,863,888]" pageId="3" pageNumber="313" pagination="862 - 873" refId="ref6698" refString="Malakauskas, D. M., Willson, S. J., Wilzbach, M. A. &amp; Som, N. A. (2013) Flow variation and substrate type affect dislodgement of the freshwater polychaete, Manayunkia speciosa. Freshwater Science, 32, 862 - 873. https: // doi. org / 10.1899 / 12 - 140.1" type="journal article" year="2013">
Malakauskas
<emphasis id="B91AEAEA4A22DB0C9A02FC8ACCA48F56" box="[1131,1175,864,888]" italics="true" pageId="3" pageNumber="313">et al</emphasis>
., 2013
</bibRefCitation>
;
<bibRefCitation id="EFFF4B094A22DB0C9A83FCB4CD138F56" author="Malakauskas, D. M. &amp; Altman, E. C. &amp; Malakauskas, S. J. &amp; Thiem, S. M. &amp; Schloesser, D. W." box="[1258,1312,863,888]" pageId="3" pageNumber="313" pagination="101 - 104" refId="ref6764" refString="Malakauskas, D. M., Altman, E. C., Malakauskas, S. J., Thiem, S. M. &amp; Schloesser, D. W. (2015) Ribosomal DNA identification of Nosema / Vairimorpha in freshwater polychaete, Manayunkia speciosa, from Oregon / California and the Laurentian Great Lakes. Journal of Invertebrate Pathology, 132, 101 - 104. https: // doi. org / 10.1016 / j. jip. 2015.09.004" type="journal article" year="2015">2015</bibRefCitation>
.
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<typeStatus id="54D5885A4A22DB0C9EFEFC4DC9378FEE" box="[151,260,934,960]" pageId="3" pageNumber="313" type="holotype">Holotype</typeStatus>
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(in 70% ethanol, initially fixed in 10% formalin).
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male from
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mainstem near confluence with
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,
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.
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, -
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, collected by
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(
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),
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(
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)
</materialsCitation>
</paragraph>
<caption id="DF1166704A22DB0C9EFEF91BC94D8B93" ID-DOI="http://doi.org/10.5281/zenodo.3713130" ID-Zenodo-Dep="3713130" httpUri="https://zenodo.org/record/3713130/files/figure.png" pageId="3" pageNumber="313" startId="3.[151,250,1776,1801]" targetBox="[155,1433,1052,1749]" targetPageId="3">
<paragraph id="8BD136F84A22DB0C9EFEF91BC94D8B93" blockId="3.[151,1437,1776,1981]" pageId="3" pageNumber="313">
<emphasis id="B91AEAEA4A22DB0C9EFEF91BC9258B27" bold="true" box="[151,278,1776,1801]" pageId="3" pageNumber="313">FIGURE 1.</emphasis>
<taxonomicName id="4C6E4D7B4A22DB0C9F77F91ACA138B27" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[286,544,1777,1801]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="3" pageNumber="313" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A22DB0C9F77F91ACA138B27" box="[286,544,1777,1801]" italics="true" pageId="3" pageNumber="313">Manayunkia occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A22DB0C9C40F91ACA4F8B27" bold="true" box="[553,636,1777,1801]" pageId="3" pageNumber="313">
<taxonomicNameLabel id="A22957914A22DB0C9C40F91ACA4F8B27" box="[553,636,1777,1801]" pageId="3" pageNumber="313" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
from the Klamath River, CA, infected with myxozoan parasites
<taxonomicName id="4C6E4D7B4A22DB0C9B4CF91AC8EB8B03" baseAuthorityName="Noble" baseAuthorityYear="1950" class="Myxosporea" family="Ceratomyxidae" genus="Ceratonova" higherTaxonomySource="GBIF" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="shasta">
<emphasis id="B91AEAEA4A22DB0C9B4CF91AC8EB8B03" italics="true" pageId="3" pageNumber="313">Ceratonova shasta</emphasis>
</taxonomicName>
(AC) and
<taxonomicName id="4C6E4D7B4A22DB0C9F38F8FECA6C8B03" baseAuthorityName="Hallett &amp; Bartholomew" baseAuthorityYear="2009" box="[337,607,1813,1837]" class="Myxosporea" family="Parvicapsulidae" genus="Parvicapsula" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="minibicornis">
<emphasis id="B91AEAEA4A22DB0C9F38F8FECA6C8B03" box="[337,607,1813,1837]" italics="true" pageId="3" pageNumber="313">Parvicapsula minibicornis</emphasis>
</taxonomicName>
(DF). A, bright field image of
<taxonomicName id="4C6E4D7B4A22DB0C9DCFF8FECC368B03" baseAuthorityName="Noble" baseAuthorityYear="1950" box="[934,1029,1813,1837]" class="Myxosporea" family="Ceratomyxidae" genus="Ceratonova" higherTaxonomySource="GBIF" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="shasta">
<emphasis id="B91AEAEA4A22DB0C9DCFF8FECC368B03" box="[934,1029,1813,1837]" italics="true" pageId="3" pageNumber="313">C. shasta</emphasis>
</taxonomicName>
actinospores (arrowheads) in tegument; B, Nomarski interference contrast image of
<taxonomicName id="4C6E4D7B4A22DB0C9C0BF8D2CAF18B7F" baseAuthorityName="Noble" baseAuthorityYear="1950" box="[610,706,1849,1873]" class="Myxosporea" family="Ceratomyxidae" genus="Ceratonova" higherTaxonomySource="GBIF" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="shasta">
<emphasis id="B91AEAEA4A22DB0C9C0BF8D2CAF18B7F" box="[610,706,1849,1873]" italics="true" pageId="3" pageNumber="313">C. shasta</emphasis>
</taxonomicName>
pansporocyst in tegument; C, Individual tetractinomyxon type actinospores of
<taxonomicName id="4C6E4D7B4A22DB0C9E90F8B6C9648B5B" baseAuthorityName="Noble" baseAuthorityYear="1950" box="[249,343,1885,1909]" class="Myxosporea" family="Ceratomyxidae" genus="Ceratonova" higherTaxonomySource="GBIF" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="shasta">
<emphasis id="B91AEAEA4A22DB0C9E90F8B6C9648B5B" box="[249,343,1885,1909]" italics="true" pageId="3" pageNumber="313">C. shasta</emphasis>
</taxonomicName>
; D, bright field image of
<taxonomicName id="4C6E4D7B4A22DB0C9C30F8B6CAC28B5B" baseAuthorityName="Hallett &amp; Bartholomew" baseAuthorityYear="2009" box="[601,753,1885,1909]" class="Myxosporea" family="Parvicapsulidae" genus="Parvicapsula" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="318" phylum="Myxozoa" rank="species" species="minibicornis">
P.
<emphasis id="B91AEAEA4A22DB0C9C18F8B6CAC28B5B" box="[625,753,1885,1909]" italics="true" pageId="3" pageNumber="313">minibicornis</emphasis>
</taxonomicName>
actinospores (arrowhead) in coelom of infected annelid; E, mass of developing
<taxonomicName id="4C6E4D7B4A22DB0C9F64F869C9958BB7" baseAuthorityName="Hallett &amp; Bartholomew" baseAuthorityYear="2009" box="[269,422,1921,1945]" class="Myxosporea" family="Parvicapsulidae" genus="Parvicapsula" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="minibicornis">
<emphasis id="B91AEAEA4A22DB0C9F64F869C9958BB7" box="[269,422,1921,1945]" italics="true" pageId="3" pageNumber="313">P. minibicornis</emphasis>
</taxonomicName>
spores in coelom; F, Individual saccimyxon type actinospores of
<taxonomicName id="4C6E4D7B4A22DB0C9A51F869CCE38BB7" baseAuthorityName="Hallett &amp; Bartholomew" baseAuthorityYear="2009" box="[1080,1232,1921,1945]" class="Myxosporea" family="Parvicapsulidae" genus="Parvicapsula" kingdom="Animalia" order="Bivalvulida" pageId="3" pageNumber="313" phylum="Myxozoa" rank="species" species="minibicornis">
<emphasis id="B91AEAEA4A22DB0C9A51F869CCE38BB7" box="[1080,1232,1921,1945]" italics="true" pageId="3" pageNumber="313">P. minibicornis</emphasis>
</taxonomicName>
; morphologies after
<bibRefCitation id="EFFF4B094A22DB0C9EFEF84FC9498B93" author="Atkinson, S. D. &amp; Hallett, S. L. &amp; Diaz Morales, D. &amp; Bartholomew, J. L. &amp; de Buron, I." box="[151,378,1956,1981]" pageId="3" pageNumber="313" pagination="252 - 262" refId="ref5605" refString="Atkinson, S. D., Hallett, S. L., Diaz Morales, D., Bartholomew, J. L. &amp; de Buron, I. (2019) First myxozoan infection (Cnidaria: Myxosporea) in a marine polychaete from North America, and erection of actinospore collective group Saccimyxon. Journal of Parasitology, 105 (2), 252 - 262. https: // doi. org / 10.1645 / 18 - 183." type="journal article" year="2019">
Atkinson
<emphasis id="B91AEAEA4A22DB0C9E93F84DC91D8B93" box="[250,302,1957,1981]" italics="true" pageId="3" pageNumber="313">et al.</emphasis>
(2019)
</bibRefCitation>
.
</paragraph>
</caption>
<paragraph id="8BD136F84A25DB0B9EAEFF72C9328CD5" blockId="4.[151,1437,152,935]" pageId="4" pageNumber="314">
<materialsCitation id="3B063CA54A25DB0B9EAEFF72C8CE8CD5" ID-GBIF-Occurrence="2577306241" collectionCode="SIO" pageId="4" pageNumber="314" specimenCount="3" typeStatus="paratype">
<emphasis id="B91AEAEA4A25DB0B9EAEFF72C90C8C9D" bold="true" box="[199,319,153,179]" pageId="4" pageNumber="314">
<typeStatus id="54D5885A4A25DB0B9EAEFF72C90C8C9D" box="[199,319,153,179]" pageId="4" pageNumber="314" type="paratype">Paratypes</typeStatus>
</emphasis>
(in 70% ethanol, fixed in 10% formalin); One specimen, adult female, same locality, collector and date as
<typeStatus id="54D5885A4A25DB0B9E9AFF57C9668CF8" box="[243,341,188,214]" pageId="4" pageNumber="314" type="holotype">holotype</typeStatus>
(
<collectionCode id="ED7FAE3D4A25DB0B9F01FF56C9A48CF9" box="[360,407,189,215]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:14844" lsid="urn:lsid:biocol.org:col:14844" name="Scripps Institution of Oceanography" pageId="4" pageNumber="314" type="University or college">SIO</collectionCode>
<accessionNumber id="943DAB1B4A25DB0B9FF6FF57C9CD8CF9" box="[415,510,188,215]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A12116" pageId="4" pageNumber="314" type="EnaNcbi">A12116</accessionNumber>
); One specimen, adult male, same locality, collector and date as
<typeStatus id="54D5885A4A25DB0B9A90FF57CD688CF8" box="[1273,1371,188,214]" pageId="4" pageNumber="314" type="holotype">holotype</typeStatus>
(
<collectionCode id="ED7FAE3D4A25DB0B9B07FF56CDAE8CF9" box="[1390,1437,189,215]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:14844" lsid="urn:lsid:biocol.org:col:14844" name="Scripps Institution of Oceanography" pageId="4" pageNumber="314" type="University or college">SIO</collectionCode>
<accessionNumber id="943DAB1B4A25DB0B9EFEFF0AC8C58CD5" box="[151,246,225,251]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A12117" pageId="4" pageNumber="314" type="EnaNcbi">A12117</accessionNumber>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C37465734A25DB0B9EAEFEEECB3F8D6C" pageId="4" pageNumber="314" type="etymology">
<paragraph id="8BD136F84A25DB0B9EAEFEEECB3F8D6C" blockId="4.[151,1437,152,935]" pageId="4" pageNumber="314">
<emphasis id="B91AEAEA4A25DB0B9EAEFEEEC97D8D31" bold="true" box="[199,334,261,287]" pageId="4" pageNumber="314">Etymology.</emphasis>
The specific epithet is derived from the Latin word for “west”, to contrast with
<taxonomicName id="4C6E4D7B4A25DB0B9AACFEEECD798D30" authorityName=": Hazel" authorityYear="1966" box="[1221,1354,261,286]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="4" pageNumber="314" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A25DB0B9AACFEEECD798D30" box="[1221,1354,261,286]" italics="true" pageId="4" pageNumber="314">M. speciosa</emphasis>
</taxonomicName>
, which occurs in eastern and central North American drainages.
</paragraph>
</subSubSection>
<subSubSection id="C37465734A25DB0A9EAEFEA7C9D48D31" lastPageId="5" lastPageNumber="315" pageId="4" pageNumber="314" type="description">
<paragraph id="8BD136F84A25DB0B9EAEFEA7CCA38F89" blockId="4.[151,1437,152,935]" pageId="4" pageNumber="314">
<emphasis id="B91AEAEA4A25DB0B9EAEFEA7C96A8D49" bold="true" box="[199,345,332,359]" pageId="4" pageNumber="314">Description.</emphasis>
<typeStatus id="54D5885A4A25DB0B9F36FEA7C9F48D48" box="[351,455,332,358]" pageId="4" pageNumber="314" type="holotype">Holotype</typeStatus>
(SIO-BIC
<accessionNumber id="943DAB1B4A25DB0B9C2BFEA7CAAE8D49" box="[578,669,332,359]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A12115" pageId="4" pageNumber="314" type="EnaNcbi">A12115</accessionNumber>
) a male with eight thoracic segments and three abdominal segments. Total length (unfixed) 3.0 mm, crown
<quantity id="4C969B1D4A25DB0B9C2EFE9ACA928DA5" box="[583,673,369,395]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="314" unit="mm" value="0.3">0.3 mm</quantity>
. Body cylindrical with tapering, dorso-ventrally flattened abdomen. Body wall translucent grey with minor brown pigmentation on peristomium and pygidium, and green pigmentation on chaetiger 6. Branchial crown ~10% of total body length. Two pairs of radioles, each with six pinnules. One ventral pinnule on radiole with 45 yellow-white spots in life (
<figureCitation id="13552A7D4A25DB0B9D48FE37CB518DD9" box="[801,866,476,503]" captionStart="FIGURE 2" captionStartId="4.[151,250,1752,1777]" captionTargetBox="[152,1435,956,1726]" captionTargetId="figure@4.[152,1435,956,1726]" captionTargetPageId="4" captionText="FIGURE 2. Gross morphological features of Manayunkia occidentalis sp. nov. resolved by light microscopy: A, multiple individuals emerged and feeding from their tubes; B, Individual annelid with prominent vascularized appendages (va) and showing refractile spots on one feeding palp on each side (arrowheads); C, female in reflected light, showing white oocytes (o) in segments 46; D, male in reflected light showing bright spermatogonic tissue (sp) in multiple posterior segments; E, Male in transmitted light showing parallel vascularized appendages (va) and fanned-out feeding palps anteriorly with two dark eyespots in peristomium, and granular coelom posteriorly due to presence of developing sperm." figureDoi="http://doi.org/10.5281/zenodo.3713132" httpUri="https://zenodo.org/record/3713132/files/figure.png" pageId="4" pageNumber="314">Fig. 2</figureCitation>
BC). One pair of vascularised, unbranched ventral filamentous appendages present, extending for about two-thirds length of radiolar crown, approximately same width as pinnules (
<figureCitation id="13552A7D4A25DB0B9F4CFDCFC95B8E11" box="[293,360,548,575]" captionStart="FIGURE 2" captionStartId="4.[151,250,1752,1777]" captionTargetBox="[152,1435,956,1726]" captionTargetId="figure@4.[152,1435,956,1726]" captionTargetPageId="4" captionText="FIGURE 2. Gross morphological features of Manayunkia occidentalis sp. nov. resolved by light microscopy: A, multiple individuals emerged and feeding from their tubes; B, Individual annelid with prominent vascularized appendages (va) and showing refractile spots on one feeding palp on each side (arrowheads); C, female in reflected light, showing white oocytes (o) in segments 46; D, male in reflected light showing bright spermatogonic tissue (sp) in multiple posterior segments; E, Male in transmitted light showing parallel vascularized appendages (va) and fanned-out feeding palps anteriorly with two dark eyespots in peristomium, and granular coelom posteriorly due to presence of developing sperm." figureDoi="http://doi.org/10.5281/zenodo.3713132" httpUri="https://zenodo.org/record/3713132/files/figure.png" pageId="4" pageNumber="314">Fig. 2</figureCitation>
BC). Anterior peristomial ring with membranous collar, followed by posterior peristomial ring (
<figureCitation id="13552A7D4A25DB0B9EC9FDA3C8D58E4D" box="[160,230,584,611]" captionStart="FIGURE 4" captionStartId="6.[151,250,1804,1829]" captionTargetBox="[155,1433,561,1779]" captionTargetId="figure@6.[155,1433,561,1779]" captionTargetPageId="6" captionText="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi="http://doi.org/10.5281/zenodo.3713136" httpUri="https://zenodo.org/record/3713136/files/figure.png" pageId="4" pageNumber="314">Fig. 4</figureCitation>
AB,D). Collar margin smooth, higher ventrally, separated by a narrow dorsal gap that corresponds with ciliated faecal groove (
<figureCitation id="13552A7D4A25DB0B9FF6FD87C9CF8EA9" box="[415,508,620,647]" captionStart="FIGURE 4" captionStartId="6.[151,250,1804,1829]" captionTargetBox="[155,1433,561,1779]" captionTargetId="figure@6.[155,1433,561,1779]" captionTargetPageId="6" captionText="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi="http://doi.org/10.5281/zenodo.3713136" httpUri="https://zenodo.org/record/3713136/files/figure.png" pageId="4" pageNumber="314">Fig. 4D</figureCitation>
). Peristomial eyes, black rounded spots (
<figureCitation id="13552A7D4A25DB0B9DBAFD87CC1F8EA9" box="[979,1068,620,647]" captionStart="FIGURE 2" captionStartId="4.[151,250,1752,1777]" captionTargetBox="[152,1435,956,1726]" captionTargetId="figure@4.[152,1435,956,1726]" captionTargetPageId="4" captionText="FIGURE 2. Gross morphological features of Manayunkia occidentalis sp. nov. resolved by light microscopy: A, multiple individuals emerged and feeding from their tubes; B, Individual annelid with prominent vascularized appendages (va) and showing refractile spots on one feeding palp on each side (arrowheads); C, female in reflected light, showing white oocytes (o) in segments 46; D, male in reflected light showing bright spermatogonic tissue (sp) in multiple posterior segments; E, Male in transmitted light showing parallel vascularized appendages (va) and fanned-out feeding palps anteriorly with two dark eyespots in peristomium, and granular coelom posteriorly due to presence of developing sperm." figureDoi="http://doi.org/10.5281/zenodo.3713132" httpUri="https://zenodo.org/record/3713132/files/figure.png" pageId="4" pageNumber="314">Fig. 2E</figureCitation>
). Pygidial eyes absent. Superior thoracic notochaetae elongate, narrowly hooded; 57 per fascicle (chaetigers 15), 45 per fascicle (chaetigers 68) (
<figureCitation id="13552A7D4A25DB0B9EF6FD5FC8CE8EE1" box="[159,253,692,719]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="4" pageNumber="314">Figs 3A</figureCitation>
;
<figureCitation id="13552A7D4A25DB0B9F60FD5EC9248EE1" box="[265,279,693,719]" captionStart="FIGURE 4" captionStartId="6.[151,250,1804,1829]" captionTargetBox="[155,1433,561,1779]" captionTargetId="figure@6.[155,1433,561,1779]" captionTargetPageId="6" captionText="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi="http://doi.org/10.5281/zenodo.3713136" httpUri="https://zenodo.org/record/3713136/files/figure.png" pageId="4" pageNumber="314">4</figureCitation>
AC). Inferior thoracic notochaetae on chaetigers 18 short; 34 per fascicle except chaetiger 2, which has 6 per fascicle. Thoracic neuropodia each with 47 uncini in chaetigers 28 (
<figureCitation id="13552A7D4A25DB0B9A64FD33CC578EDD" box="[1037,1124,728,755]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="4" pageNumber="314">Fig. 3B</figureCitation>
). Thoracic uncini with rows of evenly small teeth gradually decreasing in size away from main fang (
<figureCitation id="13552A7D4A25DB0B9DA3FD17CC148F39" box="[970,1063,764,791]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="4" pageNumber="314">Figs 3B</figureCitation>
;
<figureCitation id="13552A7D4A25DB0B9A5AFD16CC648F39" box="[1075,1111,765,791]" captionStart="FIGURE 4" captionStartId="6.[151,250,1804,1829]" captionTargetBox="[155,1433,561,1779]" captionTargetId="figure@6.[155,1433,561,1779]" captionTargetPageId="6" captionText="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi="http://doi.org/10.5281/zenodo.3713136" httpUri="https://zenodo.org/record/3713136/files/figure.png" pageId="4" pageNumber="314">4E</figureCitation>
). Position of chaetae and uncini reversed in abdominal segments; number of abdominal uncini per fascicle decreasing posteriorly from 22 to 9. Abdominal uncini each with multiple rows of teeth that are uniform in size (
<figureCitation id="13552A7D4A25DB0B9D86FCAFCC7F8F71" box="[1007,1100,836,863]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="4" pageNumber="314">Figs 3C</figureCitation>
;
<figureCitation id="13552A7D4A25DB0B9A31FCAECC4D8F71" box="[1112,1150,837,863]" captionStart="FIGURE 4" captionStartId="6.[151,250,1804,1829]" captionTargetBox="[155,1433,561,1779]" captionTargetId="figure@6.[155,1433,561,1779]" captionTargetPageId="6" captionText="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi="http://doi.org/10.5281/zenodo.3713136" httpUri="https://zenodo.org/record/3713136/files/figure.png" pageId="4" pageNumber="314">4H</figureCitation>
); manubrium at least five times longer than dentate region, with base about two thirds the width of the dentate region. Abdominal neuropodia are elongate, narrowly hooded, decreasing posteriorly from 46 to 2 per fascicle (
<figureCitation id="13552A7D4A25DB0B9A40FC67CCB08F89" box="[1065,1155,908,935]" captionStart="FIGURE 4" captionStartId="6.[151,250,1804,1829]" captionTargetBox="[155,1433,561,1779]" captionTargetId="figure@6.[155,1433,561,1779]" captionTargetPageId="6" captionText="FIGURE 4. Morphological features of Manayunkia occidentalis sp. nov. resolved by scanning electron microscopy: A, lateral view; B, dorsal view; C, ventral view; D, anterior showing membranous collar (c) of the peristomal ring (pr), margin with fecal groove (fg); mucus covers the pinnules and adheres diatoms; E, detail of typical thoracic uncini; F, abdominal segments and anus, showing ciliated fecal groove extending anteriorly and wrapping around to the ventral surface at boundary between abdominal and thoracic segments; G, lateral view of thoracic segments showing contracted uncini and prominent neuropodia; H, higher magnification view of retracted abdominal uncini." figureDoi="http://doi.org/10.5281/zenodo.3713136" httpUri="https://zenodo.org/record/3713136/files/figure.png" pageId="4" pageNumber="314">Fig. 4G</figureCitation>
).
</paragraph>
<caption id="DF1166704A25DB0B9EFEF933CC378B8A" ID-DOI="http://doi.org/10.5281/zenodo.3713132" ID-Zenodo-Dep="3713132" httpUri="https://zenodo.org/record/3713132/files/figure.png" pageId="4" pageNumber="314" startId="4.[151,250,1752,1777]" targetBox="[152,1435,956,1726]" targetPageId="4">
<paragraph id="8BD136F84A25DB0B9EFEF933CC378B8A" blockId="4.[151,1437,1752,1957]" pageId="4" pageNumber="314">
<emphasis id="B91AEAEA4A25DB0B9EFEF933C9258ADF" bold="true" box="[151,278,1752,1777]" pageId="4" pageNumber="314">FIGURE 2.</emphasis>
Gross morphological features of
<taxonomicName id="4C6E4D7B4A25DB0B9C12F932CB4E8ADF" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[635,893,1753,1777]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="4" pageNumber="314" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A25DB0B9C12F932CB4E8ADF" box="[635,893,1753,1777]" italics="true" pageId="4" pageNumber="314">Manayunkia occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A25DB0B9DEEF932CBE88ADF" bold="true" box="[903,987,1753,1777]" pageId="4" pageNumber="314">
<taxonomicNameLabel id="A22957914A25DB0B9DEEF932CBE88ADF" box="[903,987,1753,1777]" pageId="4" pageNumber="314" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
resolved by light microscopy: A, multiple individuals emerged and feeding from their tubes; B, Individual annelid with prominent vascularized appendages (va) and showing refractile spots on one feeding palp on each side (arrowheads); C, female in reflected light, showing white oocytes (o) in segments 46; D, male in reflected light showing bright spermatogonic tissue (sp) in multiple posterior segments; E, Male in transmitted light showing parallel vascularized appendages (va) and fanned-out feeding palps anteriorly with two dark eyespots in peristomium, and granular coelom posteriorly due to presence of developing sperm.
</paragraph>
</caption>
<paragraph id="8BD136F84A24DB0A9EAEFF73C9D48D31" blockId="5.[151,1436,152,287]" pageId="5" pageNumber="315">
<emphasis id="B91AEAEA4A24DB0A9EAEFF73C90C8C9D" bold="true" box="[199,319,152,179]" pageId="5" pageNumber="315">Variation.</emphasis>
<typeStatus id="54D5885A4A24DB0A9F21FF72C9868C9D" box="[328,437,153,179]" pageId="5" pageNumber="315" type="paratype">Paratypes</typeStatus>
are similar to the
<typeStatus id="54D5885A4A24DB0A9CE4FF72CAC08C9D" box="[653,755,153,179]" pageId="5" pageNumber="315" type="holotype">holotype</typeStatus>
. However, females have oocytes in thoracic chaetigers 45 (
<figureCitation id="13552A7D4A24DB0A9EC9FF56C8C48CF9" box="[160,247,189,215]" captionStart="FIGURE 2" captionStartId="4.[151,250,1752,1777]" captionTargetBox="[152,1435,956,1726]" captionTargetId="figure@4.[152,1435,956,1726]" captionTargetPageId="4" captionText="FIGURE 2. Gross morphological features of Manayunkia occidentalis sp. nov. resolved by light microscopy: A, multiple individuals emerged and feeding from their tubes; B, Individual annelid with prominent vascularized appendages (va) and showing refractile spots on one feeding palp on each side (arrowheads); C, female in reflected light, showing white oocytes (o) in segments 46; D, male in reflected light showing bright spermatogonic tissue (sp) in multiple posterior segments; E, Male in transmitted light showing parallel vascularized appendages (va) and fanned-out feeding palps anteriorly with two dark eyespots in peristomium, and granular coelom posteriorly due to presence of developing sperm." figureDoi="http://doi.org/10.5281/zenodo.3713132" httpUri="https://zenodo.org/record/3713132/files/figure.png" pageId="5" pageNumber="315">Fig. 2C</figureCitation>
) and a pair of pigmented spermathecae in the base of the radiolar crown. Females differ from males in having thoracic neuropodia with uncini in chaetigers 25 followed by 36 elongate hooded chaetae in neuropodia of chaetigers 68 (
<figureCitation id="13552A7D4A24DB0A9F0EFEEEC9EA8D31" box="[359,473,261,287]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="5" pageNumber="315">Fig. 3E,G</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="DF1166704A24DB0A9EFEF994CCF38B65" ID-DOI="http://doi.org/10.5281/zenodo.3713134" ID-Zenodo-Dep="3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="5" pageNumber="315" startId="5.[151,250,1663,1688]" targetBox="[149,1438,309,1638]" targetPageId="5">
<paragraph id="8BD136F84A24DB0A9EFEF994CCF38B65" blockId="5.[151,1437,1663,1868]" pageId="5" pageNumber="315">
<emphasis id="B91AEAEA4A24DB0A9EFEF994C9208AB6" bold="true" box="[151,275,1663,1688]" pageId="5" pageNumber="315">FIGURE 3.</emphasis>
Images showing chaetal characters of
<taxonomicName id="4C6E4D7B4A24DB0A9CF7F994CBAE8AB9" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[670,925,1663,1687]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="5" pageNumber="315" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A24DB0A9CF7F994CBAE8AB9" box="[670,925,1663,1687]" italics="true" pageId="5" pageNumber="315">Manayunkia occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A24DB0A9DCDF96BCBC68AB6" bold="true" box="[932,1013,1664,1688]" pageId="5" pageNumber="315">
<taxonomicNameLabel id="A22957914A24DB0A9DCDF96BCBC68AB6" box="[932,1013,1664,1688]" pageId="5" pageNumber="315" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes.
</paragraph>
</caption>
<subSubSection id="C37465734A24DB099EAEF891CC328E32" lastPageId="6" lastPageNumber="316" pageId="5" pageNumber="315" type="discussion">
<paragraph id="8BD136F84A24DB099EAEF891CC328E32" blockId="5.[151,1437,1914,2012]" lastBlockId="6.[151,1437,153,540]" lastPageId="6" lastPageNumber="316" pageId="5" pageNumber="315">
<emphasis id="B91AEAEA4A24DB0A9EAEF891C9088BBA" bold="true" box="[199,315,1914,1940]" pageId="5" pageNumber="315">Remarks.</emphasis>
<taxonomicName id="4C6E4D7B4A24DB0A9F2DF891CA528BBA" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[324,609,1914,1940]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="5" pageNumber="315" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A24DB0A9F2DF891CA528BBA" box="[324,609,1914,1940]" italics="true" pageId="5" pageNumber="315">Manayunkia occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A24DB0A9C03F891CAF58BBA" bold="true" box="[618,710,1914,1940]" pageId="5" pageNumber="315">
<taxonomicNameLabel id="A22957914A24DB0A9C03F891CAF58BBA" box="[618,710,1914,1940]" pageId="5" pageNumber="315" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is the sister group to
<taxonomicName id="4C6E4D7B4A24DB0A9DACF890CC7F8BBA" authorityName=": Hazel" authorityYear="1966" box="[965,1100,1915,1940]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="5" pageNumber="315" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A24DB0A9DACF890CC7F8BBA" box="[965,1100,1915,1940]" italics="true" pageId="5" pageNumber="315">M. speciosa</emphasis>
</taxonomicName>
based on COI sequence data with a minimum pairwise distance (simple distance) to our
<taxonomicName id="4C6E4D7B4A24DB0A9D51F874CB8C8B96" authorityName=": Hazel" authorityYear="1966" box="[824,959,1951,1976]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="5" pageNumber="315" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A24DB0A9D51F874CB8C8B96" box="[824,959,1951,1976]" italics="true" pageId="5" pageNumber="315">M. speciosa</emphasis>
</taxonomicName>
samples of 11.0%, clearly supporting that they can be regarded as a separate taxon, although phylogenetic support for this relationship was weak (bootstrap 28%).
<taxonomicName id="4C6E4D7B4A27DB099E8AFF72C9CC8C9D" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[227,511,153,179]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A27DB099E8AFF72C9CC8C9D" box="[227,511,153,179]" italics="true" pageId="6" pageNumber="316">Manayunkia occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A27DB099C61FF72CA508C9D" bold="true" box="[520,611,153,179]" pageId="6" pageNumber="316">
<taxonomicNameLabel id="A22957914A27DB099C61FF72CA508C9D" box="[520,611,153,179]" pageId="6" pageNumber="316" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
and
<taxonomicName id="4C6E4D7B4A27DB099CF4FF71CB178C9D" authorityName=": Hazel" authorityYear="1966" box="[669,804,154,179]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A27DB099CF4FF71CB178C9D" box="[669,804,154,179]" italics="true" pageId="6" pageNumber="316">M. speciosa</emphasis>
</taxonomicName>
are the only species of
<emphasis id="B91AEAEA4A27DB099A5FFF72CCFA8C9D" box="[1078,1225,153,179]" italics="true" pageId="6" pageNumber="316">
<taxonomicName id="4C6E4D7B4A27DB099A5FFF72CCF68C9D" authorityName="Leidy" authorityYear="1859" box="[1078,1221,153,179]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="genus">Manayunkia</taxonomicName>
,
</emphasis>
and in fact of any
<taxonomicName id="4C6E4D7B4A27DB099EFEFF56C9268CF9" authorityName="Rioja" authorityYear="1923" box="[151,277,189,215]" class="Polychaeta" family="Fabriciidae" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="family">Fabriciidae</taxonomicName>
(see
<bibRefCitation id="EFFF4B094A27DB099F27FF56CA258CF9" author="Huang, D. &amp; Fitzhugh, K. &amp; Rouse, G. W." box="[334,534,189,215]" pageId="6" pageNumber="316" pagination="256 - 379" refId="ref6436" refString="Huang, D., Fitzhugh, K. &amp; Rouse, G. W. (2011) Inference of phylogenetic relationships within Fabriciidae Annelida) using molecular and morphological data. Cladistics, 27 (4), 256 - 379. https: // doi. org / 10.1111 / j. 1096 - 0031.2010.00343. x" type="journal article" year="2011">
Huang
<emphasis id="B91AEAEA4A27DB099FF7FF55C9FC8CF9" box="[414,463,189,215]" italics="true" pageId="6" pageNumber="316">et al</emphasis>
. 2011
</bibRefCitation>
), to show sexual dimorphism with regards to thoracic chaetal complement. Males of
<taxonomicName id="4C6E4D7B4A27DB099EDFFF09C9518CD5" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[182,354,225,251]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A27DB099EDFFF09C9518CD5" box="[182,354,225,251]" italics="true" pageId="6" pageNumber="316">M. occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A27DB099F00FF0AC9F08CD5" bold="true" box="[361,451,225,251]" pageId="6" pageNumber="316">
<taxonomicNameLabel id="A22957914A27DB099F00FF0AC9F08CD5" box="[361,451,225,251]" pageId="6" pageNumber="316" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, and
<taxonomicName id="4C6E4D7B4A27DB099C69FF09CAB58CD5" authorityName=": Hazel" authorityYear="1966" box="[512,646,226,251]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A27DB099C69FF09CAB58CD5" box="[512,646,226,251]" italics="true" pageId="6" pageNumber="316">M. speciosa</emphasis>
</taxonomicName>
(Rouse pers. obs.) on specimens from near the
<typeStatus id="54D5885A4A27DB099AF2FF0ACCF88CD5" box="[1179,1227,225,251]" pageId="6" pageNumber="316">type</typeStatus>
locality (SIO-BIC
<accessionNumber id="943DAB1B4A27DB099EFEFEEEC8C58D0E" box="[151,246,261,288]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/A12106" pageId="6" pageNumber="316" type="EnaNcbi">A12106</accessionNumber>
), show the typical arrangement of thoracic uncini in each of chaetigers 28, while females have thoracic uncini in chaetigers 25 only (
<figureCitation id="13552A7D4A27DB099F9CFEC2CA628D6D" box="[501,593,297,323]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="6" pageNumber="316">Fig. 3H</figureCitation>
). The neuropodia of chaetigers 68 instead have elongate hooded chaetae (
<figureCitation id="13552A7D4A27DB099EC9FEA6C8C98D49" box="[160,250,333,359]" captionStart="FIGURE 3" captionStartId="5.[151,250,1663,1688]" captionTargetBox="[149,1438,309,1638]" captionTargetId="figure@5.[149,1438,309,1638]" captionTargetPageId="5" captionText="FIGURE 3. Images showing chaetal characters of Manayunkia occidentalis sp. nov.: A, inferior and superior notochaete typical of thoracic segments 28; B, thoracic uncini; C, abdominal uncini; D, dorsal view of posterior thoracic segments and abdomen of male showing typical uncini in chaetigers t7 and t8 (arrowheads) and in scanning electron micrograph shown in F; E, dorsal view of posterior thoracic segments and abdomen of female showing neuropodia of thoracic chaetigers 68 with elongate neuropodia instead of uncini, seen also in scanning electron micrograph in G (arrowhead). H, anterior thoracic segment of female showing notochaetae and uncini (arrowhead), characteristic of thoracic segments 25 in both sexes." figureDoi="http://doi.org/10.5281/zenodo.3713134" httpUri="https://zenodo.org/record/3713134/files/figure.png" pageId="6" pageNumber="316">Fig. 3G</figureCitation>
). These elongate hooded chaetae were seen in all females examined. Larval development would be worth studying to see if there is a loss of thoracic uncini or if they never develop in chaetigers 68. It is somewhat surprising that this major difference in chaetal complement was never noted previous in observations on
<emphasis id="B91AEAEA4A27DB099B66FE7DC9838DFD" italics="true" pageId="6" pageNumber="316">
<taxonomicName id="4C6E4D7B4A27DB099B66FE7DCDAA8D81" authorityName=": Hazel" authorityYear="1966" box="[1295,1433,406,431]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="speciosa">M. speciosa</taxonomicName>
.
<taxonomicName id="4C6E4D7B4A27DB099EFEFE52C9838DFD" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[151,432,441,467]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">Manayunkia occidentalis</taxonomicName>
</emphasis>
<emphasis id="B91AEAEA4A27DB099FDEFE52CA238DFD" bold="true" box="[439,528,441,467]" pageId="6" pageNumber="316">
<taxonomicNameLabel id="A22957914A27DB099FDEFE52CA238DFD" box="[439,528,441,467]" pageId="6" pageNumber="316" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
differs markedly from descriptions of
<taxonomicName id="4C6E4D7B4A27DB099DD4FE51CC718DFD" authorityName=": Hazel" authorityYear="1966" box="[957,1090,442,467]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="speciosa">
<emphasis id="B91AEAEA4A27DB099DD4FE51CC718DFD" box="[957,1090,442,467]" italics="true" pageId="6" pageNumber="316">M. speciosa</emphasis>
</taxonomicName>
by Leidy (1883) and
<bibRefCitation id="EFFF4B094A27DB099B5BFE52C8D18DD9" author="Pettibone, M. H." pageId="6" pageNumber="316" pagination="149 - 153" refId="ref6970" refString="Pettibone, M. H. (1953) Fresh-Water Polychaetous Annelid, Manayunkia speciosa Leidy, from Lake Erie. The Biological Bulletin, 105 (1), 149 - 153. https: // doi. org / 10.2307 / 1538563" type="journal article" year="1953">Pettibone (1953)</bibRefCitation>
in that it has far fewer pinnules on the radioles. In this regard
<taxonomicName id="4C6E4D7B4A27DB099DF1FE36CC828DD9" authority="Atkinson &amp; Bartholomew &amp; Rouse, 2020" authorityName="Atkinson &amp; Bartholomew &amp; Rouse" authorityYear="2020" box="[920,1201,477,503]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="species" species="occidentalis" status="sp. nov.">
<emphasis id="B91AEAEA4A27DB099DF1FE36CC828DD9" box="[920,1201,477,503]" italics="true" pageId="6" pageNumber="316">Manayunkia occidentalis</emphasis>
</taxonomicName>
<emphasis id="B91AEAEA4A27DB099AD0FE36CD218DD9" bold="true" box="[1209,1298,477,503]" pageId="6" pageNumber="316">
<taxonomicNameLabel id="A22957914A27DB099AD0FE36CD218DD9" box="[1209,1298,477,503]" pageId="6" pageNumber="316" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is similar to other
<taxonomicName id="4C6E4D7B4A27DB099EB1FDEAC9578E35" authorityName="Leidy" authorityYear="1859" box="[216,356,513,539]" class="Polychaeta" family="Fabriciidae" genus="Manayunkia" kingdom="Animalia" order="Sabellida" pageId="6" pageNumber="316" phylum="Annelida" rank="genus">
<emphasis id="B91AEAEA4A27DB099EB1FDEAC9578E35" box="[216,356,513,539]" italics="true" pageId="6" pageNumber="316">Manayunkia</emphasis>
</taxonomicName>
species that have relatively few pinnules (see
<bibRefCitation id="EFFF4B094A27DB099D02FDEACBC78E32" author="Rouse, G. W." box="[875,1012,513,540]" pageId="6" pageNumber="316" pagination="1761 - 1778" refId="ref7404" refString="Rouse, G. W. (1996) New Fabriciola and Manayunkia species (Fabriciinae: Sabellidae: Polychaeta) from Papua New Guinea. Journal of Natural History, 30, 1761 - 1778. https: // doi. org / 10.1080 / 00222939600771031" type="journal article" year="1996">Rouse 1996</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>