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<document id="104EC775B4A028A4B33C8198875E4C2D" ID-CLB-Dataset="24995" ID-DOI="10.11646/zootaxa.4691.4.2" ID-GBIF-Dataset="94e09348-a782-45ec-9c88-6584fc3da9c0" ID-ISSN="1175-5326" ID-Zenodo-Dep="3527357" ID-ZooBank="875FB4E7-148A-4AC2-904C-174416B11256" IM.bibliography_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1572857692114" checkinUser="plazi" docAuthor="Hookabe, Natsumi, Schwartz, Megan L., Kajihara, Hiroshi &amp; Norenburg, Jon L." docDate="2019" docId="03B387A4FFF3CC26FF73C6BEFC3FFEC2" docLanguage="en" docName="zootaxa.4691.4.2.pdf" docOrigin="Zootaxa 4691 (4)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Dushia , Diva Correa 1963" docType="treatment" docVersion="9" lastPageNumber="336" masterDocId="FF8AFFDCFFF1CC22FFE4C03FFFB2FF81" masterDocTitle="Molecular systematics of the heteronemertean genus Dushia (Nemertea Pilidiophora), with descriptions of D. wijnhoffae sp. nov. and D. nigra species complex comb. nov." masterLastPageNumber="358" masterPageNumber="333" pageNumber="335" updateTime="1698761824128" updateUser="ExternalLinkService">
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<mods:title id="FB96A744FCBC8621B1CBEFA6C5B9FA4E">Molecular systematics of the heteronemertean genus Dushia (Nemertea Pilidiophora), with descriptions of D. wijnhoffae sp. nov. and D. nigra species complex comb. nov.</mods:title>
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<mods:namePart id="0163AE15C14221724FD1C5EACD2F3AD4">Hookabe, Natsumi</mods:namePart>
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<mods:namePart id="A7737396F52448D347ADF72E999F414E">Schwartz, Megan L.</mods:namePart>
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<mods:namePart id="B2BEA3201DB89F7401B390DCC99F3A50">Kajihara, Hiroshi</mods:namePart>
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<mods:namePart id="6F95E761490B3A83C09EE3D6197121CE">Norenburg, Jon L.</mods:namePart>
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<paragraph id="8BA536B2FFF3CC20FF73C6BEFE43F91A" blockId="2.[151,497,1665,1692]" box="[151,497,1665,1692]" pageId="2" pageNumber="335">
<heading id="D0ED81DEFFF3CC20FF73C6BEFE43F91A" bold="true" box="[151,497,1665,1692]" fontSize="11" level="1" pageId="2" pageNumber="335" reason="1">
<emphasis id="B96EEAA0FFF3CC20FF73C6BEFE43F91A" bold="true" box="[151,497,1665,1692]" pageId="2" pageNumber="335">
Genus
<taxonomicName id="4C1A4D31FFF3CC20FF0AC6BEFE43F91A" authority="Correa, 1963" authorityName=", Diva Correa" authorityYear="1963" box="[238,497,1665,1692]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="2" pageNumber="335" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF3CC20FF0AC6BEFEF4F91D" bold="true" box="[238,326,1665,1692]" italics="true" pageId="2" pageNumber="335">Dushia</emphasis>
<bibRefCitation id="EF8B4B43FFF3CC20FEAAC6BEFE43F91A" author="Correa, D. D." box="[334,497,1665,1691]" pageId="2" pageNumber="335" pagination="41 - 56" refId="ref14499" refString="Correa, D. D. (1963) Nemerteans from Curacao. Studies on the Fauna of Curacao and other Caribbean Islands, 75, 41 - 56. Available from: http: // www. repository. naturalis. nl / document / 550005 (Accessed 31 Oct. 2019)" type="journal article" year="1963">Corrêa, 1963</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF3CC20FF73C6F5FAC0F83A" pageId="2" pageNumber="335" type="diagnosis">
<paragraph id="8BA536B2FFF3CC20FF73C6F5FAC0F83A" blockId="2.[151,1437,1737,1979]" pageId="2" pageNumber="335">
<emphasis id="B96EEAA0FFF3CC20FF73C6F5FEA2F962" box="[151,272,1738,1763]" italics="true" pageId="2" pageNumber="335">Diagnosis.</emphasis>
Heteronemerteans with uniformly colored body (typically black, but also brown and orange) and white cephalic tip on both dorsal and ventral surfaces; with horizontal, lateral cephalic furrows; with caudal cirrus; dermal circular muscles external to dermal longitudinal muscles, the two separated by some diagonal fibers present as a lattice; connective tissue between cutis and outer longitudinal musculature (olm) at a minimum; dorsoventral muscles in body-wall inner longitudinal muscle layer in intestinal region, but not in outer longitudinal muscle layer; proboscis with typical heterotype musculature (
<bibRefCitation id="EF8B4B43FFF3CC20FD80C742FC9CF816" author="Chernyshev, A. V." box="[612,814,1916,1943]" pageId="2" pageNumber="335" pagination="547 - 561" refId="ref13977" refString="Chernyshev, A. V. (2015) CLSM analysis of the phalloidin-stained muscle system of the nemertean proboscis and rhynchocoel. Zoological Science, 32, 547 - 561. https: // doi. org / 10.2108 / zs 140267" type="journal article" year="2015">Chernyshev 2015</bibRefCitation>
) with two unequal muscle crosses; two lateral and one dorsal precerebral blood lacunae; neurocord cells present in dorsal portion of dorsal cerebral ganglia; no ocelli.
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF2CC21FF23C0A6FB1AFF56" pageId="3" pageNumber="336" type="etymology">
<paragraph id="8BA536B2FFF2CC21FF23C0A6FB1AFF56" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
<emphasis id="B96EEAA0FFF2CC21FF23C0A6FE2EFF32" box="[199,412,153,179]" italics="true" pageId="3" pageNumber="336">
Etymology.
<taxonomicName id="4C1A4D31FFF2CC21FEAFC0A6FE2EFF32" authorityName=", Diva Correa" authorityYear="1963" box="[331,412,153,179]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">Dushia</taxonomicName>
</emphasis>
appears to be a feminine latinized form derived from “
<emphasis id="B96EEAA0FFF2CC21FBE6C0A6FB24FF32" box="[1026,1174,153,179]" italics="true" pageId="3" pageNumber="336">Dushi Korsu,</emphasis>
” which in Papiamentu, a local language in
<collectingCountry id="F30D7622FFF2CC21FE95C082FE63FF56" box="[369,465,189,215]" name="Curaçao" pageId="3" pageNumber="336">Curaçao</collectingCountry>
, means “sweet
<collectingCountry id="F30D7622FFF2CC21FD9AC082FD6CFF56" box="[638,734,189,215]" name="Curaçao" pageId="3" pageNumber="336">Curaçao</collectingCountry>
” (Cynthia Santos, pers. comm. to MLS).
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF2CC21FF23C0DDFB69FEC2" pageId="3" pageNumber="336" type="type_taxon">
<paragraph id="8BA536B2FFF2CC21FF23C0DDFB69FEC2" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
<emphasis id="B96EEAA0FFF2CC21FF23C0DDFEE9FF7A" box="[199,347,226,251]" italics="true" pageId="3" pageNumber="336">
<typeStatus id="54A18810FFF2CC21FF23C0DDFF49FF7A" box="[199,251,226,251]" pageId="3" pageNumber="336">Type</typeStatus>
species.
</emphasis>
Herein fixed under Article 70.3.2 of the International Code of Zoological Nomenclature (
<bibRefCitation id="EF8B4B43FFF2CC21FAA6C0DEFD5CFE9E" author="International Commission on Zoological Nomenclature" pageId="3" pageNumber="336" refId="ref15327" refString="International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature, 4 th Edition. International Trust for Zoological Nomenclature, London, xxix + 306 pp." type="book" year="1999">International Commission on Zoological Nomenclature 1999</bibRefCitation>
) as
<taxonomicName id="4C1A4D31FFF2CC21FCFDC13AFB5FFE9E" authority="Schwartz &amp; Norenburg" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[793,1261,261,287]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">
<emphasis id="B96EEAA0FFF2CC21FCFDC13AFC57FE9E" box="[793,997,261,287]" italics="true" pageId="3" pageNumber="336">Dushia wijnhoffae</emphasis>
Schwartz &amp; Norenburg
</taxonomicName>
<emphasis id="B96EEAA0FFF2CC21FB17C13AFAF9FE9E" bold="true" box="[1267,1355,261,287]" pageId="3" pageNumber="336">
<taxonomicNameLabel id="A25D57DBFFF2CC21FB17C13AFAF9FE9E" box="[1267,1355,261,287]" pageId="3" pageNumber="336" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
, which was misidentified as
<taxonomicName id="4C1A4D31FFF2CC21FE67C116FD07FEC2" authority="Girard, 1851" authorityName="Girard" authorityYear="1851" box="[387,693,296,323]" class="Pilidiophora" family="Lineidae" genus="Meckelia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="atra">
<emphasis id="B96EEAA0FFF2CC21FE67C116FDADFEC2" box="[387,543,297,323]" italics="true" pageId="3" pageNumber="336">Meckelia atra</emphasis>
<bibRefCitation id="EF8B4B43FFF2CC21FDC2C116FD07FEC2" author="Girard, C." box="[550,693,296,323]" pageId="3" pageNumber="336" pagination="137" refId="ref14652" refString="Girard, C. (1851) Descriptions of a new Planaria and a new Nemertes from the coast of Florida. Proceedings of the Boston Society of Natural History, 4, 137. Available from: https: // www. biodiversitylibrary. org / item / 35742 # page / 145 / mode / 1 up (Accessed 31 Oct. 2019)" type="journal article" year="1851">
Girard,
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</taxonomicName>
in the original designation by
<bibRefCitation id="EF8B4B43FFF2CC21FBEFC116FB65FEC2" author="Correa, D. D." box="[1035,1239,297,323]" pageId="3" pageNumber="336" pagination="41 - 56" refId="ref14499" refString="Correa, D. D. (1963) Nemerteans from Curacao. Studies on the Fauna of Curacao and other Caribbean Islands, 75, 41 - 56. Available from: http: // www. repository. naturalis. nl / document / 550005 (Accessed 31 Oct. 2019)" type="journal article" year="1963">Corrêa (1963: 43)</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF2CC21FF23C172FE69FD9D" pageId="3" pageNumber="336" type="diagnosis">
<paragraph id="8BA536B2FFF2CC21FF23C172FE69FD9D" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
<emphasis id="B96EEAA0FFF2CC21FF23C172FEECFEE6" box="[199,350,333,359]" italics="true" pageId="3" pageNumber="336">Composition.</emphasis>
The genus currently includes
<taxonomicName id="4C1A4D31FFF2CC21FD24C172FB15FEE6" authority="Schwartz &amp; Norenburg" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[704,1191,333,359]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">
<emphasis id="B96EEAA0FFF2CC21FD24C172FC23FEE6" box="[704,913,333,359]" italics="true" pageId="3" pageNumber="336">Dushia wijnhoffae</emphasis>
Schwartz &amp; Norenburg
</taxonomicName>
<emphasis id="B96EEAA0FFF2CC21FB55C172FABCFEE6" bold="true" box="[1201,1294,333,359]" pageId="3" pageNumber="336">
<taxonomicNameLabel id="A25D57DBFFF2CC21FB55C172FABCFEE6" box="[1201,1294,333,359]" pageId="3" pageNumber="336" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
and
<taxonomicName id="4C1A4D31FFF2CC21FAAFC172FE2FFE0A" authority="(Stimpson, 1855)" baseAuthorityName="Stimpson" baseAuthorityYear="1855" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="nigra">
<emphasis id="B96EEAA0FFF2CC21FAAFC172FF66FE0A" italics="true" pageId="3" pageNumber="336">Dushia nigra</emphasis>
(
<bibRefCitation id="EF8B4B43FFF2CC21FF06C14EFE27FE0A" author="Stimpson, W." box="[226,405,368,395]" pageId="3" pageNumber="336" pagination="375 - 384" refId="ref17526" refString="Stimpson, W. (1855) Descriptions of some of the new marine Invertebrata from the Chinese and Japanese Seas. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 375 - 384. https: // doi. org / 10.5962 / bhl. title. 51444" type="journal article" year="1855">Stimpson, 1855</bibRefCitation>
)
</taxonomicName>
species complex
<emphasis id="B96EEAA0FFF2CC21FD87C14EFD56FE0A" bold="true" box="[611,740,369,395]" pageId="3" pageNumber="336">comb. nov.</emphasis>
The latter contains the following four nominal species:
<taxonomicName id="4C1A4D31FFF2CC21FAB2C14EFE75FE2E" authority="Stimpson, 1855" authorityName="Stimpson" authorityYear="1855" class="Pilidiophora" family="Lineidae" genus="Meckelia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="nigra">
<emphasis id="B96EEAA0FFF2CC21FAB2C14EFEBAFE2E" italics="true" pageId="3" pageNumber="336">Meckelia nigra</emphasis>
<bibRefCitation id="EF8B4B43FFF2CC21FEF5C1AAFE75FE2E" author="Stimpson, W." box="[273,455,404,431]" pageId="3" pageNumber="336" pagination="375 - 384" refId="ref17526" refString="Stimpson, W. (1855) Descriptions of some of the new marine Invertebrata from the Chinese and Japanese Seas. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 375 - 384. https: // doi. org / 10.5962 / bhl. title. 51444" type="journal article" year="1855">Stimpson, 1855</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C1A4D31FFF2CC21FE31C1AAFCE7FE2E" authority="Stimpson, 1855" authorityName="Stimpson" authorityYear="1855" box="[469,853,404,431]" class="Pilidiophora" family="Lineidae" genus="Meckelia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="rubella">
<emphasis id="B96EEAA0FFF2CC21FE31C1AAFD24FE2E" box="[469,662,405,431]" italics="true" pageId="3" pageNumber="336">Meckelia rubella</emphasis>
<bibRefCitation id="EF8B4B43FFF2CC21FD44C1AAFCE7FE2E" author="Stimpson, W." box="[672,853,404,431]" pageId="3" pageNumber="336" pagination="375 - 384" refId="ref17526" refString="Stimpson, W. (1855) Descriptions of some of the new marine Invertebrata from the Chinese and Japanese Seas. Proceedings of the Academy of Natural Sciences of Philadelphia, 7, 375 - 384. https: // doi. org / 10.5962 / bhl. title. 51444" type="journal article" year="1855">Stimpson, 1855</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C1A4D31FFF2CC21FC87C1A9FB4DFE2E" authority="Yamaoka, 1939" authorityName="Yamaoka" authorityYear="1939" box="[867,1279,405,431]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="formosana">
<emphasis id="B96EEAA0FFF2CC21FC87C1A9FBF0FE2E" box="[867,1090,405,431]" italics="true" pageId="3" pageNumber="336">Micrura formosana</emphasis>
<bibRefCitation id="EF8B4B43FFF2CC21FBAFC1AAFB4DFE2E" author="Yamaoka T." box="[1099,1279,405,431]" pageId="3" pageNumber="336" pagination="283 - 289" refId="ref18054" refString="Yamaoka T. (1939) Two nemerteans from Formosa. Annotationes Zoologicae Japonenses, 18, 283 - 289." type="journal article" year="1939">Yamaoka, 1939</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C1A4D31FFF2CC21FADBC1A9FE3DFE52" authority="Iwata, 1952" authorityName="Iwata" authorityYear="1952" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="japonica">
<emphasis id="B96EEAA0FFF2CC21FADBC1A9FF4BFE52" italics="true" pageId="3" pageNumber="336">Micrura japonica</emphasis>
<bibRefCitation id="EF8B4B43FFF2CC21FEE0C186FE3DFE52" author="Iwata, F." box="[260,399,440,467]" pageId="3" pageNumber="336" pagination="126 - 148" refId="ref15358" refString="Iwata, F. (1952) Nemertini from the coasts of Kyusyu. Journal of the Faculty of Science, Hokkaido University, Series VI, Zoology, 11, 126 - 148." type="journal article" year="1952">Iwata, 1952</bibRefCitation>
</taxonomicName>
. Circumscription and name allocation within this species complex would require genetic analyses of specimens from wide geographic area in the Southwest Pacific, including the
<typeStatus id="54A18810FFF2CC21FB95C1E2FB13FE76" box="[1137,1185,477,503]" pageId="3" pageNumber="336">type</typeStatus>
localities of these four nominal species (see below).
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF2CC26FF23C21AFC3FFEC2" lastPageId="4" lastPageNumber="337" pageId="3" pageNumber="336" type="discussion">
<paragraph id="8BA536B2FFF2CC21FF23C21AFBA4FCBA" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
<emphasis id="B96EEAA0FFF2CC21FF23C21AFE83FDBE" box="[199,305,549,575]" italics="true" pageId="3" pageNumber="336">Remarks.</emphasis>
The genera
<taxonomicName id="4C1A4D31FFF2CC21FE5CC21AFDF9FDBE" authorityName="Reiner" authorityYear="1804" box="[440,587,549,575]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FE5CC21AFDF9FDBE" box="[440,587,549,575]" italics="true" pageId="3" pageNumber="336">Cerebratulus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1A4D31FFF2CC21FD61C219FD50FDBE" authorityName="Ehrenberg" authorityYear="1828" box="[645,738,550,575]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FD61C219FD50FDBE" box="[645,738,550,575]" italics="true" pageId="3" pageNumber="336">Micrura</emphasis>
</taxonomicName>
lack definitive diagnoses, which makes it problematic to accurately place heteronemertean taxa into either of these nominal genera. When
<bibRefCitation id="EF8B4B43FFF2CC21FBFDC276FB0AFDE5" author="Correa, D. D." box="[1049,1208,585,612]" pageId="3" pageNumber="336" pagination="41 - 56" refId="ref14499" refString="Correa, D. D. (1963) Nemerteans from Curacao. Studies on the Fauna of Curacao and other Caribbean Islands, 75, 41 - 56. Available from: http: // www. repository. naturalis. nl / document / 550005 (Accessed 31 Oct. 2019)" type="journal article" year="1963">Corrêa (1963)</bibRefCitation>
established
<taxonomicName id="4C1A4D31FFF2CC21FAA0C276FA27FDE2" authorityName=", Diva Correa" authorityYear="1963" box="[1348,1429,585,611]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FAA0C276FA27FDE2" box="[1348,1429,585,611]" italics="true" pageId="3" pageNumber="336">Dushia</emphasis>
</taxonomicName>
, she distinguished it from
<taxonomicName id="4C1A4D31FFF2CC21FE5AC252FDE3FD06" authorityName="Reiner" authorityYear="1804" box="[446,593,621,647]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FE5AC252FDE3FD06" box="[446,593,621,647]" italics="true" pageId="3" pageNumber="336">Cerebratulus</emphasis>
</taxonomicName>
on the basis of two features;
<taxonomicName id="4C1A4D31FFF2CC21FC4FC252FC4EFD06" authorityName=", Diva Correa" authorityYear="1963" box="[939,1020,621,647]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FC4FC252FC4EFD06" box="[939,1020,621,647]" italics="true" pageId="3" pageNumber="336">Dushia</emphasis>
</taxonomicName>
lacked a caudal cirrus and does not have sharp lateral margins as in many
<taxonomicName id="4C1A4D31FFF2CC21FDAEC2AEFD6FFD2A" authorityName="Reiner" authorityYear="1804" box="[586,733,657,683]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FDAEC2AEFD6FFD2A" box="[586,733,657,683]" italics="true" pageId="3" pageNumber="336">Cerebratulus</emphasis>
</taxonomicName>
. Our observation on additional specimens from the Caribbean strongly suggests that
<bibRefCitation id="EF8B4B43FFF2CC21FE71C28AFDF4FD51" author="Correa, D. D." box="[405,582,693,720]" pageId="3" pageNumber="336" pagination="41 - 56" refId="ref14499" refString="Correa, D. D. (1963) Nemerteans from Curacao. Studies on the Fauna of Curacao and other Caribbean Islands, 75, 41 - 56. Available from: http: // www. repository. naturalis. nl / document / 550005 (Accessed 31 Oct. 2019)" type="journal article" year="1963">Corrêas (1963)</bibRefCitation>
material was not intact, having lost the tail together with the caudal cirrus. While the presence or absence of a caudal cirrus often appears to be moderately clade-specific, current evidence suggests that it has either been gained or lost multiple times, and is not informative for generic placement as traditionally used (
<bibRefCitation id="EF8B4B43FFF2CC21FED2C31EFDCCFCBA" author="Schwartz, M. L. &amp; Norenburg, J. L." box="[310,638,801,827]" pageId="3" pageNumber="336" pagination="165 - 174" refId="ref17274" refString="Schwartz, M. L. &amp; Norenburg, J. L. (2001) Can we infer heteronemertean phylogeny from available morphological data? Hydrobiologia, 456, 165 - 174. https: // doi. org / 10.1023 / A: 1013093629108" type="journal article" year="2001">Schwartz &amp; Norenburg 2001</bibRefCitation>
;
<bibRefCitation id="EF8B4B43FFF2CC21FD6EC31EFC86FCBA" author="Schwartz, M. L." box="[650,820,801,827]" pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz 2009</bibRefCitation>
;
<bibRefCitation id="EF8B4B43FFF2CC21FCA4C31EFBB8FCBA" author="Puerta, P. &amp; Andrade, S. C. S. &amp; Junoy, J." box="[832,1034,801,827]" pageId="3" pageNumber="336" pagination="2363 - 2378" refId="ref16480" refString="Puerta, P., Andrade, S. C. S. &amp; Junoy, J. (2010) Redescription of Lineus acutifrons Southern, 1913 (Nemertea: Pilidiophora) and comments on its phylogenetic position. Journal of Natural History, 44, 2363 - 2378. https: // doi. org / 10.1080 / 00222933.2010.504895" type="journal article" year="2010">
Puerta
<emphasis id="B96EEAA0FFF2CC21FC6AC31DFC72FCBA" box="[910,960,801,827]" italics="true" pageId="3" pageNumber="336">et al</emphasis>
. 2010
</bibRefCitation>
).
</paragraph>
<paragraph id="8BA536B2FFF2CC21FF23C37AFACEFC26" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
Putative and nominal
<taxonomicName id="4C1A4D31FFF2CC21FE51C37AFDF3FCDE" authorityName="Reiner" authorityYear="1804" box="[437,577,837,863]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FE51C37AFDF3FCDE" box="[437,577,837,863]" italics="true" pageId="3" pageNumber="336">Cerebratulus</emphasis>
</taxonomicName>
spp. with sharp lateral margins were examined by
<bibRefCitation id="EF8B4B43FFF2CC21FB88C37AFA90FCDE" author="Schwartz, M. L." box="[1132,1314,837,863]" pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz (2009)</bibRefCitation>
. These had dorsoventral musculature in the outer longitudinal muscle layer (olm), i.e., external to the lateral nerve cords, or additional circular muscles below the cutis and even into the olm (
<bibRefCitation id="EF8B4B43FFF2CC21FD0CC3B2FC38FC26" author="Schwartz, M. L." box="[744,906,909,935]" pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz 2009</bibRefCitation>
). Neither of these features is found in
<taxonomicName id="4C1A4D31FFF2CC21FACCC3B2FAC4FC26" authorityName=", Diva Correa" authorityYear="1963" box="[1320,1398,909,935]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FACCC3B2FAC4FC26" box="[1320,1398,909,935]" italics="true" pageId="3" pageNumber="336">Dushia</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BA536B2FFF2CC21FF23C38EFAC6FB46" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
<taxonomicName id="4C1A4D31FFF2CC21FF23C38EFEAAFC4A" authorityName=", Diva Correa" authorityYear="1963" box="[199,280,945,971]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FF23C38EFEAAFC4A" box="[199,280,945,971]" italics="true" pageId="3" pageNumber="336">Dushia</emphasis>
</taxonomicName>
also can be separated from
<taxonomicName id="4C1A4D31FFF2CC21FD80C38DFD73FC4A" authorityName="Ehrenberg" authorityYear="1828" box="[612,705,946,971]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FD80C38DFD73FC4A" box="[612,705,946,971]" italics="true" pageId="3" pageNumber="336">Micrura</emphasis>
</taxonomicName>
, although there has not been a comprehensive redescription of
<taxonomicName id="4C1A4D31FFF2CC21FF73C3E9FD9DFC6E" authority="Ehrenberg, 1828" authorityName="Ehrenberg" authorityYear="1828" box="[151,559,981,1007]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="fasciolata">
<emphasis id="B96EEAA0FFF2CC21FF73C3E9FED8FC6E" box="[151,362,981,1007]" italics="true" pageId="3" pageNumber="336">Micrura fasciolata</emphasis>
Ehrenberg, 1828
</taxonomicName>
, the
<typeStatus id="54A18810FFF2CC21FD80C3EAFD26FC6E" box="[612,660,981,1007]" pageId="3" pageNumber="336">type</typeStatus>
for the genus. Serial transverse sections of
<taxonomicName id="4C1A4D31FFF2CC21FB9BC3E9FAA0FC6E" authorityName="Ehrenberg" authorityYear="1828" box="[1151,1298,981,1007]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="species" species="fasciolata">
<emphasis id="B96EEAA0FFF2CC21FB9BC3E9FAA0FC6E" box="[1151,1298,981,1007]" italics="true" pageId="3" pageNumber="336">M. fasciolata</emphasis>
</taxonomicName>
collected in
<collectingCountry id="F30D7622FFF2CC21FF73C3C6FF42FB92" box="[151,240,1017,1043]" name="Sweden" pageId="3" pageNumber="336">Sweden</collectingCountry>
reveal a proboscis with “two muscle layers” in a traditional sense (
<bibRefCitation id="EF8B4B43FFF2CC21FC04C3C6FA9AFB92" author="Schwartz, M. L. &amp; Norenburg, J. L." box="[992,1320,1017,1043]" pageId="3" pageNumber="336" pagination="165 - 174" refId="ref17274" refString="Schwartz, M. L. &amp; Norenburg, J. L. (2001) Can we infer heteronemertean phylogeny from available morphological data? Hydrobiologia, 456, 165 - 174. https: // doi. org / 10.1023 / A: 1013093629108" type="journal article" year="2001">Schwartz &amp; Norenburg 2001</bibRefCitation>
;
<bibRefCitation id="EF8B4B43FFF2CC21FAD0C3C6FF61FBB6" author="Schwartz, M. L." pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz 2009</bibRefCitation>
), which corresponds to the modified heterotype without external muscle crosses and outer longitudinal musculature (consisting of four muscle layers,
<bibRefCitation id="EF8B4B43FFF2CC21FDB9C47EFC94FBDA" author="Chernyshev, A. V." box="[605,806,1088,1115]" pageId="3" pageNumber="336" pagination="547 - 561" refId="ref13977" refString="Chernyshev, A. V. (2015) CLSM analysis of the phalloidin-stained muscle system of the nemertean proboscis and rhynchocoel. Zoological Science, 32, 547 - 561. https: // doi. org / 10.2108 / zs 140267" type="journal article" year="2015">Chernyshev 2015</bibRefCitation>
), whereas the
<taxonomicName id="4C1A4D31FFF2CC21FC22C47EFBA5FBDA" authorityName=", Diva Correa" authorityYear="1963" box="[966,1047,1089,1115]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FC22C47EFBA5FBDA" box="[966,1047,1089,1115]" italics="true" pageId="3" pageNumber="336">Dushia</emphasis>
</taxonomicName>
spp. examined here have a “threelayered” (in a traditional sense) proboscis with two muscle crosses, which actually consists of five layers (
<bibRefCitation id="EF8B4B43FFF2CC21FADBC45AFEBDFB22" author="Chernyshev, A. V." pageId="3" pageNumber="336" pagination="547 - 561" refId="ref13977" refString="Chernyshev, A. V. (2015) CLSM analysis of the phalloidin-stained muscle system of the nemertean proboscis and rhynchocoel. Zoological Science, 32, 547 - 561. https: // doi. org / 10.2108 / zs 140267" type="journal article" year="2015">Chernyshev 2015</bibRefCitation>
). The number of proboscis muscle layers and their positional relationship also appear to be homoplastic among
<taxonomicName id="4C1A4D31FFF2CC21FF0EC492FE2FFB46" authorityName="Burger" authorityYear="1892" box="[234,413,1197,1223]" class="Anopla" higherTaxonomySource="GBIF" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="order">Heteronemertea</taxonomicName>
but they can be phylogenetically informative for less inclusive clades (
<bibRefCitation id="EF8B4B43FFF2CC21FB5EC492FADAFB46" author="Schwartz, M. L." box="[1210,1384,1197,1223]" pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz 2009</bibRefCitation>
).
</paragraph>
<paragraph id="8BA536B2FFF2CC21FF23C4EEFEF4F9D2" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
The presence or absence of neurocord cells has been used to distinguish
<taxonomicName id="4C1A4D31FFF2CC21FC13C4EEFB38FB6A" authorityName="Reiner" authorityYear="1804" box="[1015,1162,1233,1259]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FC13C4EEFB38FB6A" box="[1015,1162,1233,1259]" italics="true" pageId="3" pageNumber="336">Cerebratulus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C1A4D31FFF2CC21FB24C4EDFAAFFB6A" authorityName="Ehrenberg" authorityYear="1828" box="[1216,1309,1234,1259]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FB24C4EDFAAFFB6A" box="[1216,1309,1234,1259]" italics="true" pageId="3" pageNumber="336">Micrura</emphasis>
</taxonomicName>
from other genera. Neurocord cells and other neural cell
<typeStatus id="54A18810FFF2CC21FD75C4CAFD7EFA8E" box="[657,716,1269,1295]" pageId="3" pageNumber="336">types</typeStatus>
were first described in detail by
<bibRefCitation id="EF8B4B43FFF2CC21FBD3C4CAFB7EFA8E" author="Burger, O." box="[1079,1228,1268,1295]" pageId="3" pageNumber="336" refId="ref13740" refString="Burger, O. (1895) Fauna und Flora des Golfes von Neapel und der angrenzenden Meeres-abschnitte herausgegeben von der Zoologischen Stazion zu Neapel, 22. Monographie: Nemertinen. R. Friedlander &amp; Sohn, Berlin, xvi + 743 pp., 31 pls. https: // doi. org / 10.5962 / bhl. title. 14030" type="book" year="1895">Bürger (1895</bibRefCitation>
,
<date id="FFA41072FFF2CC21FB04C4CAFAEFFA8E" box="[1248,1373,1269,1295]" pageId="3" pageNumber="336" value="1897" valueMax="1907">
<bibRefCitation id="EF8B4B43FFF2CC21FB04C4CAFAA5FA8E" author="Burger, O." box="[1248,1303,1269,1295]" pageId="3" pageNumber="336" pagination="1 - 542" refId="ref13803" refString="Burger, O. (1897 - 1907) Nemertini (Schnurwurmer). In: Bronn, H. G. (Ed.), Klassen und Ordnungen des Tier-Reichs. Supplement 4. C. F. Winter, Leipzig, pp. 1 - 542." type="book chapter" year="1897">1897</bibRefCitation>
1907
</date>
) and further refined by
<bibRefCitation id="EF8B4B43FFF2CC21FE87C526FD9DFAB2" author="Thompson, C. B." box="[355,559,1305,1331]" pageId="3" pageNumber="336" pagination="641 - 661" refId="ref17971" refString="Thompson, C. B. (1908) The commissures and the neurochord cells of the brain of Cerebratulus lacteus. Journal of Comparative Neurology and Psychology, 18, 641 - 661. https: // doi. org / 10.1002 / cne. 920180606" type="journal article" year="1908">Thompson (1908)</bibRefCitation>
; their definitions are used for identification here. At minimum, most heteronemertean species possess Bürger
<typeStatus id="54A18810FFF2CC21FE1DC502FD9BFAD6" box="[505,553,1341,1367]" pageId="3" pageNumber="336">type</typeStatus>
I and II neural cells, whereas the presence of Bürger
<typeStatus id="54A18810FFF2CC21FB89C502FB2FFAD6" box="[1133,1181,1341,1367]" pageId="3" pageNumber="336">type</typeStatus>
III and neurocord cells is considered unique to certain taxa including
<taxonomicName id="4C1A4D31FFF2CC21FD50C55EFCF5FAFA" authorityName="Reiner" authorityYear="1804" box="[692,839,1377,1403]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FD50C55EFCF5FAFA" box="[692,839,1377,1403]" italics="true" pageId="3" pageNumber="336">Cerebratulus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C1A4D31FFF2CC21FCB3C55DFC6AFAFA" authorityName="Ris" authorityYear="1993" box="[855,984,1378,1403]" class="Anopla" family="Lineidae" genus="Parvicirrus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FCB3C55DFC6AFAFA" box="[855,984,1378,1403]" italics="true" pageId="3" pageNumber="336">Parvicirrus</emphasis>
</taxonomicName>
, and
<emphasis id="B96EEAA0FFF2CC21FBFEC55EFAABFAFA" box="[1050,1305,1377,1403]" italics="true" pageId="3" pageNumber="336">Polybranchiorhynchus</emphasis>
(
<bibRefCitation id="EF8B4B43FFF2CC21FAD6C55EFF66FA1E" author="Schwartz, M. L." pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz 2009</bibRefCitation>
). Neurocord cells in heteronemerteans are identified by their large size, over
<quantity id="4CE29B57FFF2CC21FBDCC5BAFB36FA1E" box="[1080,1156,1413,1439]" metricMagnitude="-2" metricUnit="m" metricValue="2.0" pageId="3" pageNumber="336" unit="mm" value="20.0">20 µm</quantity>
, a pyriform shape with a substantial spherical to sub-spherical nucleus situated proximally in the cell (
<bibRefCitation id="EF8B4B43FFF2CC21FBE4C596FB0CFA42" author="Thompson, C. B." box="[1024,1214,1449,1475]" pageId="3" pageNumber="336" pagination="641 - 661" refId="ref17971" refString="Thompson, C. B. (1908) The commissures and the neurochord cells of the brain of Cerebratulus lacteus. Journal of Comparative Neurology and Psychology, 18, 641 - 661. https: // doi. org / 10.1002 / cne. 920180606" type="journal article" year="1908">Thompson 1908</bibRefCitation>
).
<bibRefCitation id="EF8B4B43FFF2CC21FB36C596FA2EFA42" author="Thompson, C. B." box="[1234,1436,1449,1475]" pageId="3" pageNumber="336" pagination="641 - 661" refId="ref17971" refString="Thompson, C. B. (1908) The commissures and the neurochord cells of the brain of Cerebratulus lacteus. Journal of Comparative Neurology and Psychology, 18, 641 - 661. https: // doi. org / 10.1002 / cne. 920180606" type="journal article" year="1908">Thompson (1908)</bibRefCitation>
observed that
<emphasis id="B96EEAA0FFF2CC21FED1C5F1FE8FFA66" box="[309,317,1486,1511]" italics="true" pageId="3" pageNumber="336">i</emphasis>
) the cytoplasm tends to be clear when stained with hematoxylin and eosin;
<emphasis id="B96EEAA0FFF2CC21FB77C5F1FB11FA66" box="[1171,1187,1486,1511]" italics="true" pageId="3" pageNumber="336">ii</emphasis>
) they are found in the medial portions of the dorsal ganglia, and the dorsomedial portions of the ventral ganglia in one to seven pairs; and
<emphasis id="B96EEAA0FFF2CC21FF73C629FF1CF9AE" box="[151,174,1558,1583]" italics="true" pageId="3" pageNumber="336">iii</emphasis>
) occasionally, neurocord cells may also be found in proximal portions of the lateral nerve cords as they leave the ventral ganglia.
</paragraph>
<paragraph id="8BA536B2FFF2CC21FF23C662FECCF816" blockId="3.[151,1437,153,2015]" pageId="3" pageNumber="336">
Large cells found in the dorsal ganglia of the
<taxonomicName id="4C1A4D31FFF2CC21FD2FC662FCAEF9F6" authorityName=", Diva Correa" authorityYear="1963" box="[715,796,1629,1655]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="3" pageNumber="336" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF2CC21FD2FC662FCAEF9F6" box="[715,796,1629,1655]" italics="true" pageId="3" pageNumber="336">Dushia</emphasis>
</taxonomicName>
spp. (
<figureCitation id="13212A37FFF2CC21FC85C662FC0CF9F6" box="[865,958,1629,1655]" captionStart="FIGURE 3" captionStartId="10.[151,246,1384,1409]" captionTargetBox="[164,1422,182,1358]" captionTargetId="figure@10.[164,1423,181,1358]" captionTargetPageId="10" captionText="FIGURE 3. Dushia wijnhoffae Schwartz &amp; Norenburg sp. nov. (A, B, D) Photomicrographs of transverse sections (paratype, USNM 1594813); (C) photomicrograph of anesthetized animal with cephalic furrows reflected back to expose ciliated glandular ridge; the specimen was destroyed after examination and not extant. (A) Organization of the cerebral ganglia region showing dorsal and ventral ganglia and positions of Bürger type III cells, neurocord cell and neurocord. (B) ×40 magnification of Bürger type III cell and neurocord cell in A. (C) Ciliated and gland cell ridge of cephalic furrow marked by arrow heads. (D) Transverse section through glandular ridge in C, arrow heads mark the cells with elongated cilia, asterisks indicate gland cells. Scale bars:A, B, D = 100 µm, C = 500 µm. Abbreviations: b3, Bürger type III neural cell; cf, cephalic furrow; lnc, lateral nerve cord; nc, neurocord cell; vg, ventral ganglion." figureDoi="http://doi.org/10.5281/zenodo.3527363" httpUri="https://zenodo.org/record/3527363/files/figure.png" pageId="3" pageNumber="336">Figs 3B</figureCitation>
,
<figureCitation id="13212A37FFF2CC21FC2EC662FC5EF9F9" box="[970,1004,1629,1656]" captionStart="FIGURE 6" captionStartId="15.[151,250,1704,1729]" captionTargetBox="[185,1400,181,1677]" captionTargetId="figure@15.[185,1401,181,1678]" captionTargetPageId="15" captionText="FIGURE 6. Dushia nigra (Stimpson, 1855) species complex comb. nov. Photomicrographs of transverse sections (AD) (ICHUM 3457) and frontal sections (E, F) (USNM 1107399). (A) Ventral rhynchocoel wall in foregut region. (B) Dorsal ganglion showing Bürger type III neural cells and neurocord cells. (C) Neurocord cells. (D) Transverse section of cephalic furrow posterior to cerebral organ opening showing ciliated cells with elongated cilia (arrow heads) and gland cells (asterisks). (E) Frontal section of intestinal regions showing arrangement of gonads and intestinal diverticula. (F) Spermatozoa. Scale bars: A, B, D, E = 100 µm, C = 50 µm, F = 3 µm. Abbreviations: b3, Bürger type III neural cell; cf, cephalic furrow; cu, cutis; dg, dorsal ganglion; dv, dorsal ventral muscle; fg, foregut lumen; go, gonad; ilm, inner longitudinal muscle; in, lumen of the intestine;; lnc, lateral nerve cord; mcm, middle circular muscle; nc, neurocord cell; olm, outer longitudinal muscle; pr, proboscis; rc, rhynchocoel; rcc, rhynchocoel circular muscle; rcl, rhynchocoel longitudinal muscle, rv, rhynchocoel villus." figureDoi="http://doi.org/10.5281/zenodo.3527373" httpUri="https://zenodo.org/record/3527373/files/figure.png" pageId="3" pageNumber="336">6C</figureCitation>
) fit
<bibRefCitation id="EF8B4B43FFF2CC21FBF8C662FB4BF9F6" author="Thompson, C. B." box="[1052,1273,1629,1655]" pageId="3" pageNumber="336" pagination="641 - 661" refId="ref17971" refString="Thompson, C. B. (1908) The commissures and the neurochord cells of the brain of Cerebratulus lacteus. Journal of Comparative Neurology and Psychology, 18, 641 - 661. https: // doi. org / 10.1002 / cne. 920180606" type="journal article" year="1908">Thompsons (1908)</bibRefCitation>
description of neurocord cells, except that the cytoplasm tends to be stained pink in this material. These cells, however, do not fit the definition of Bürger
<typeStatus id="54A18810FFF2CC21FE2FC69AFE49F93E" box="[459,507,1701,1727]" pageId="3" pageNumber="336">type</typeStatus>
III cells, which are smaller and can be seen adjacent to the neurocord cells. Many of the neurocord cells appear as though they suffer from shrinkage, perhaps accounting for their darker staining with hematoxylin. Caution should be taken when interpreting neural cell
<typeStatus id="54A18810FFF2CC21FC2AC6D2FBBBF886" box="[974,1033,1773,1799]" pageId="3" pageNumber="336">types</typeStatus>
because they are subject to fixation artifact and inherently stain poorly with general staining protocols. Neurocord cells also appear to be a homoplastic feature among heteronemerteans (
<bibRefCitation id="EF8B4B43FFF2CC21FDFEC70AFD7BF8CE" author="Schwartz, M. L." box="[538,713,1845,1871]" pageId="3" pageNumber="336" refId="ref17223" refString="Schwartz, M. L. (2009) Untying a Gordian Knot of Worms: Systematics and Taxonomy of the Pilidiophora (Phylum Nemertea) from Multiple Data Sets. Ph. D. Dissertation, George Washington University, Washington, D. C., 432 pp." type="book" year="2009">Schwartz 2009</bibRefCitation>
), but also among other nemertean groups; e.g., monostiliferans (
<bibRefCitation id="EF8B4B43FFF2CC21FF7BC766FE44F8F2" author="Maslakova, S. A. &amp; Norenburg, J. L." box="[159,502,1881,1907]" pageId="3" pageNumber="336" pagination="1689 - 1727" refId="ref16295" refString="Maslakova, S. A. &amp; Norenburg, J. L. (2008) Revision of the smiling worms, genera Prosadenoporus Burger, 1890 and Pantinonemertes Moore and Gibson, 1981 and description of a new species Prosadenoporus floridensis sp. nov. (Prosorhochmidae; Hoplonemertea; Nemertea) from Florida and Belize. Journal of Natural History, 42, 1689 - 1727. https: // doi. org / 10.1080 / 00222930802130286" type="journal article" year="2008">Maslakova &amp; Norenburg 2008</bibRefCitation>
) and some reptantic polystiliferans (Kirsteur 1973, and references therein) also have neurocord-like cells.
</paragraph>
<paragraph id="8BA536B2FFF2CC26FF23C79EFC3FFEC2" blockId="3.[151,1437,153,2015]" lastBlockId="4.[151,1436,153,323]" lastPageId="4" lastPageNumber="337" pageId="3" pageNumber="336">
Longitudinal and oblique sections of the cutis revealed between the dermal circular muscles and the dermal longitudinal muscles a layer of diagonal muscle fibers organized in a cross-hatch or lattice pattern (
<figureCitation id="13212A37FFF2CC21FAE5C7FAFAECF85E" box="[1281,1374,1989,2015]" captionStart="FIGURE 2" captionStartId="8.[151,250,1729,1754]" captionTargetBox="[183,1405,309,1703]" captionTargetId="figure@8.[183,1405,309,1704]" captionTargetPageId="8" captionText="FIGURE 2. Dushia wijnhoffae Schwartz &amp; Norenburg sp. nov. Photomicrographs of transverse sections (A, CE) (paratype, USNM 1594813), and tangential section (B) (holotype, USNM 1594812). (A) Epidermis and cutis. (B) Dermal muscle components; diagonal muscles are indicated by “&gt;”. (C) Section through gonad with dorsoventral muscles and horizontal muscle. (D) Precerebral blood vessels and rhynchodaeum. (E) Cross section through foregut region with prominent longitudinal muscle plate. Scale bars:A, C, D = 100 µm, B = 50 µm, E = 200 µm.Abbreviations: cu, cutis; cv, cephalic vessel; dcm, dermal circular muscle; dlm, dermal longitudinal muscle; dv, dorsoventral muscle fibers; ep, epidermis; fg, foregut; gc, gland cells; go, gonad; hm, horizontal muscle; ilm, body-wall inner longitudinal muscle layer; in, intestinal lumen; lmp, longitudinal muscle plate; lnc, lateral nerve cord; mcm, body-wall middle circular muscle layer; olm, body-wall outer longitudinal muscle layer; rc, rhynchocoel; rd, rhynchodaeum." figureDoi="http://doi.org/10.5281/zenodo.3527361" httpUri="https://zenodo.org/record/3527361/files/figure.png" pageId="3" pageNumber="336">Figs 2B</figureCitation>
,
<figureCitation id="13212A37FFF2CC21FA8FC7FAFA3DF85E" box="[1387,1423,1989,2015]" captionStart="FIGURE 4" captionStartId="13.[151,250,1678,1703]" captionTargetBox="[151,1436,182,1653]" captionTargetId="figure@13.[151,1436,181,1653]" captionTargetPageId="13" captionText="FIGURE 4. Dushia nigra (Stimpson, 1855) species complex comb. nov., transverse sections (A, CF) (ICHUM 3457), (B) (USNM 1107399). (A) Arrangement of the epidermis and cutis. (B) Oblique section through dermal muscle layers showing dermal gland cells, dermal longitudinal, circular, and associated lattice muscles, and epidermis. (C) Organization of body wall in foregut region, nephridial tubules, and one of the numerous efferent ducts. (D) Precerebral region, showing musculature, rhynchodaeum, and cephalic vessels. (E) Proboscis insertion. (F) Longitudinal muscular plate and nephridial tubules. Scale bars:A, C, F = 50 µm, B, D, E = 100 µm.Abbreviations: cu, cutis; cv, cephalic vessel; dc, dermal gland cells; dcm, dermal circular muscle; ddf, dermal diagonal muscle fibers; dlm, dermal longitudinal muscle; ed, efferent duct; ep, epidermis; ex, excretory collecting tubule; ilm, body-wall inner longitudinal muscle layer; lmp, longitudinal muscle plate; mcm, body-wall middle circular muscle layer; mo, mouth; olm, body-wall outer longitudinal muscle layer; nt, nephridial tubules; pr, proboscis; rd, rhynchodaeum." figureDoi="http://doi.org/10.5281/zenodo.3527365" httpUri="https://zenodo.org/record/3527365/files/figure.png" pageId="3" pageNumber="336">4B</figureCitation>
). It is possible that this layer has been overlooked in other heteronemerteans because internal examination usually only involves observations from transverse sections. Using confocal laser scanning microscopy,
<bibRefCitation id="EF8B4B43FFF5CC26FB22C082FA2EFF56" author="Chernyshev, A. V." box="[1222,1436,189,215]" pageId="4" pageNumber="337" pagination="2287 - 2302" refId="ref13935" refString="Chernyshev, A. V. (2010) Confocal laser scanning microscopy analysis of the phalloidin-labelled musculature in nemerteans. Journal of Natural History, 44, 2287 - 2302. https: // doi. org / 10.1080 / 00222933.2010.504890" type="journal article" year="2010">Chernyshev (2010)</bibRefCitation>
recorded such dermal diagonal muscle fibers in
<taxonomicName id="4C1A4D31FFF5CC26FD5EC0DEFC66FF7A" authorityName="Renier" authorityYear="1804" box="[698,980,225,251]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="4" pageNumber="337" phylum="Nemertea" rank="species" species="marginatus">
<emphasis id="B96EEAA0FFF5CC26FD5EC0DEFC66FF7A" box="[698,980,225,251]" italics="true" pageId="4" pageNumber="337">Cerebratulus marginatus</emphasis>
</taxonomicName>
auct. non Renier, 1804,
<taxonomicName id="4C1A4D31FFF5CC26FB08C0DEFE0AFE9E" authority="(Takakura, 1898)" baseAuthorityName="Takakura" baseAuthorityYear="1898" class="Pilidiophora" family="Lineidae" genus="Kulikovia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Heteronemertea" pageId="4" pageNumber="337" phylum="Nemertea" rank="species" species="alborostrata">
<emphasis id="B96EEAA0FFF5CC26FB08C0DEFF42FE9E" italics="true" pageId="4" pageNumber="337">Kulikovia alborostrata</emphasis>
(Takakura, 1898)
</taxonomicName>
,
<taxonomicName id="4C1A4D31FFF5CC26FE27C139FD7DFE9E" authority="(Coe, 1901)" baseAuthorityName="Coe" baseAuthorityYear="1901" box="[451,719,261,287]" class="Pilidiophora" family="Lineidae" genus="Kulikovia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Heteronemertea" pageId="4" pageNumber="337" phylum="Nemertea" rank="species" species="torquata">
<emphasis id="B96EEAA0FFF5CC26FE27C139FDF0FE9E" box="[451,578,262,287]" italics="true" pageId="4" pageNumber="337">K. torquata</emphasis>
(
<bibRefCitation id="EF8B4B43FFF5CC26FDB5C13AFD75FE9E" author="Coe, W. R." box="[593,711,261,287]" pageId="4" pageNumber="337" pagination="223 - 229" refId="ref14210" refString="Coe, W. R. (1901) The nemerteans of Porto Rico. Bulletin of the United States Commission of Fish and Fisheries for 1900, 2, 223 - 229." type="journal article" year="1901">Coe, 1901</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C1A4D31FFF5CC26FD3FC139FBCEFE9E" authority="Chernyshev, 1992" authorityName="Chernyshev" authorityYear="1992" box="[731,1148,261,287]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="4" pageNumber="337" phylum="Nemertea" rank="species" species="kulikovae">
<emphasis id="B96EEAA0FFF5CC26FD3FC139FC1BFE9E" box="[731,937,261,287]" italics="true" pageId="4" pageNumber="337">Micrura kulikovae</emphasis>
Chernyshev, 1992
</taxonomicName>
,
<taxonomicName id="4C1A4D31FFF5CC26FB62C139FB51FE9E" authorityName="Ehrenberg" authorityYear="1828" box="[1158,1251,262,287]" class="Anopla" family="Lineidae" genus="Micrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="4" pageNumber="337" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF5CC26FB62C139FB51FE9E" box="[1158,1251,262,287]" italics="true" pageId="4" pageNumber="337">Micrura</emphasis>
</taxonomicName>
sp.,
<taxonomicName id="4C1A4D31FFF5CC26FAF2C139FE49FEC2" authority="(Yamaoka, 1940)" baseAuthorityName="Yamaoka" baseAuthorityYear="1940" class="Anopla" family="Lineidae" genus="Nipponomicrura" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="4" pageNumber="337" phylum="Nemertea" rank="species" species="uchidai">
<emphasis id="B96EEAA0FFF5CC26FAF2C139FE82FEC2" italics="true" pageId="4" pageNumber="337">Nipponomicrura uchidai</emphasis>
(Yamaoka, 1940)
</taxonomicName>
, and
<taxonomicName id="4C1A4D31FFF5CC26FDD1C116FD7FFEC2" authorityName="Hubrecht" authorityYear="1879" box="[565,717,297,323]" class="Anopla" family="Valenciniidae" kingdom="Animalia" pageId="4" pageNumber="337" phylum="Nemertea" rank="family">Valenciniidae</taxonomicName>
gen. et sp. indet.
</paragraph>
</subSubSection>
</treatment>
</document>