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<mods:title id="892470464C2CF4DBDB4C8870DE6E3FC1">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="89F1A597952D1424885C1ABA9425BE41">Hookabe, Natsumi</mods:namePart>
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<mods:namePart id="C3E0A44AEC737AFC119FF7FC16C1F27E">Schwartz, Megan L.</mods:namePart>
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<mods:namePart id="075C665658535AFF10CA0CE94A5ABEBE">Kajihara, Hiroshi</mods:namePart>
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<mods:namePart id="C8D1221FAE5A75DDDA109D78001FE0E8">Norenburg, Jon L.</mods:namePart>
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<paragraph id="8BA536B2FFF7CC24FF73C4D9FCB9FA81" blockId="6.[151,779,1254,1316]" box="[151,779,1254,1281]" pageId="6" pageNumber="339">
<heading id="D0ED81DEFFF7CC24FF73C4D9FCB9FA81" bold="true" box="[151,779,1254,1281]" fontSize="11" level="1" pageId="6" pageNumber="339" reason="1">
<emphasis id="B96EEAA0FFF7CC24FF73C4D9FCB9FA81" bold="true" box="[151,779,1254,1281]" pageId="6" pageNumber="339">
<taxonomicName id="4C1A4D31FFF7CC24FF73C4D9FD14FA81" authority="Schwartz &amp; Norenburg" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[151,678,1254,1281]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">
<emphasis id="B96EEAA0FFF7CC24FF73C4D9FECBFA80" bold="true" box="[151,377,1254,1281]" italics="true" pageId="6" pageNumber="339">Dushia wijnhoffae</emphasis>
Schwartz &amp; Norenburg
</taxonomicName>
<taxonomicNameLabel id="A25D57DBFFF7CC24FD49C4D9FCB9FA81" box="[685,779,1254,1280]" pageId="6" pageNumber="339" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C3006539FFF7CC24FF73C535FE17FAA5" box="[151,421,1290,1316]" pageId="6" pageNumber="339" type="description">
<paragraph id="8BA536B2FFF7CC24FF73C535FE17FAA5" blockId="6.[151,779,1254,1316]" box="[151,421,1290,1316]" pageId="6" pageNumber="339">
(
<figureCitation id="13212A37FFF7CC24FF7BC535FF4EFAA5" box="[159,252,1290,1316]" captionStart="FIGURE 1" captionStartId="7.[151,250,1282,1307]" captionTargetBox="[201,1387,306,1253]" captionTargetId="figure@7.[200,1387,306,1254]" captionTargetPageId="7" captionText="FIGURE 1. Images of Dushia spp., taken in life. (A) Dushia wijnhoffae Schwartz &amp; Norenburg sp. nov. from Puerto Rico. (BF) Dushia nigra (Stimpson, 1855) species complex comb. nov.: (B) black form, ICHUM 3457, from Onna, Okinawa, Japan; (C) orange form, from Nakijin, Okinawa, caudal cirrus not pictured; (D) brown form, from Kakeroma, Japan; (E, F) black form from Okinawa, showing body fragments regenerating caudal cirrus, 3 days (E) and 10 days (F) after amputation. Scale bars: AD = 1 cm, E, F = 500 µm. Abbreviations: ao, apical organs; cc, caudal cirrus; cf, horizontal cephalic furrow." figureDoi="http://doi.org/10.5281/zenodo.3527359" httpUri="https://zenodo.org/record/3527359/files/figure.png" pageId="6" pageNumber="339">Figs 1A</figureCitation>
,
<figureCitation id="13212A37FFF7CC24FEECC535FEA7FAA5" box="[264,277,1290,1316]" 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="6" pageNumber="339">2</figureCitation>
AE, 3AD)
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF7CC24FF73C571FC57FA60" pageId="6" pageNumber="339" type="reference_group">
<paragraph id="8BA536B2FFF7CC24FF73C571FDBBFAE7" blockId="6.[151,997,1357,1505]" box="[151,521,1357,1382]" pageId="6" pageNumber="339">
<taxonomicName id="4C1A4D31FFF7CC24FF73C571FE6FFAE7" authority=": Correa 1963, p. 44" authorityName="Correa" authorityPageNumber="44" authorityYear="1963" box="[151,477,1357,1382]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="atra">
<emphasis id="B96EEAA0FFF7CC24FF73C571FEA3FAE7" box="[151,273,1358,1382]" italics="true" pageId="6" pageNumber="339">Dushia atra</emphasis>
:
<bibRefCitation id="EF8B4B43FFF7CC24FEFAC571FE2DFAE7" author="Correa, D. D." box="[286,415,1357,1382]" pageId="6" pageNumber="339" 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>
, p. 44
</taxonomicName>
48.
</paragraph>
<paragraph id="8BA536B2FFF7CC24FF73C552FD63FA04" blockId="6.[151,997,1357,1505]" box="[151,721,1388,1413]" pageId="6" pageNumber="339">
?
<taxonomicName id="4C1A4D31FFF7CC24FF46C552FD25FA04" authority=": Stiasny-Wijnhoff 1925, p. 108" authorityName="Stiasny-Wijnhoff" authorityPageNumber="108" authorityYear="1925" box="[162,663,1388,1413]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="ater">
<emphasis id="B96EEAA0FFF7CC24FF46C552FEE5FA04" box="[162,343,1389,1413]" italics="true" pageId="6" pageNumber="339">Cerebratulus ater</emphasis>
:
<bibRefCitation id="EF8B4B43FFF7CC24FE80C553FDFFFA04" author="Stiasny-Wijnhoff, G." box="[356,589,1388,1413]" pageId="6" pageNumber="339" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoff 1925</bibRefCitation>
, p. 108
</taxonomicName>
109.
</paragraph>
<paragraph id="8BA536B2FFF7CC24FF73C5B3FDC9FA25" blockId="6.[151,997,1357,1505]" box="[151,635,1419,1444]" pageId="6" pageNumber="339">
?
<taxonomicName id="4C1A4D31FFF7CC24FF46C5B3FE6DFA25" authority=": Coe 1951 a, p. 181" authorityName=": Coe" authorityPageNumber="181" authorityYear="1951" box="[162,479,1419,1444]" class="Anopla" family="Lineidae" genus="Lineus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="ater">
<emphasis id="B96EEAA0FFF7CC24FF46C5B3FEA7FA22" box="[162,277,1420,1443]" italics="true" pageId="6" pageNumber="339">Lineus ater</emphasis>
:
<bibRefCitation id="EF8B4B43FFF7CC24FEC7C5B3FE20FA25" author="Coe, W. R." box="[291,402,1419,1444]" pageId="6" pageNumber="339" pagination="149 - 186" refId="ref14314" refString="Coe, W. R. (1951 a) The nemertean faunas of the Gulf of Mexico and of southern Florida. Bulletin of Marine Science of the Gulf and Caribbean, 1, 149 - 186." type="journal article" year="1951">Coe 1951a</bibRefCitation>
, p. 181
</taxonomicName>
; 1951b, p. 330.
</paragraph>
<paragraph id="8BA536B2FFF7CC24FF73C594FD0AFA43" blockId="6.[151,997,1357,1505]" box="[151,696,1449,1474]" pageId="6" pageNumber="339">
Not
<taxonomicName id="4C1A4D31FFF7CC24FF20C595FD9AFA43" authority="Girard, 1851, p. 291" authorityName="Girard" authorityPageNumber="291" authorityYear="1851" box="[196,552,1449,1474]" class="Pilidiophora" family="Lineidae" genus="Meckelia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="atra">
<emphasis id="B96EEAA0FFF7CC24FF20C595FEE3FA43" box="[196,337,1450,1474]" italics="true" pageId="6" pageNumber="339">Meckelia atra</emphasis>
<bibRefCitation id="EF8B4B43FFF7CC24FEB3C596FE69FA43" author="Girard, C." box="[343,475,1449,1474]" pageId="6" pageNumber="339" 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, 1851</bibRefCitation>
, p. 291
</taxonomicName>
; 1893, p. 163.
</paragraph>
<paragraph id="8BA536B2FFF7CC24FF73C5F6FC57FA60" blockId="6.[151,997,1357,1505]" box="[151,997,1480,1505]" pageId="6" pageNumber="339">
Not
<taxonomicName id="4C1A4D31FFF7CC24FF20C5F6FDFFFA60" authority=": Verrill 1895, p. 531" authorityName="Verrill" authorityPageNumber="531" authorityYear="1895" box="[196,589,1480,1505]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="ater">
<emphasis id="B96EEAA0FFF7CC24FF20C5F6FECAFA60" box="[196,376,1481,1505]" italics="true" pageId="6" pageNumber="339">Cerebratulus ater</emphasis>
:
<bibRefCitation id="EF8B4B43FFF7CC24FE60C5F7FDB2FA60" author="Verrill, A. E." box="[388,512,1480,1505]" pageId="6" pageNumber="339" pagination="523 - 534" refId="ref18018" refString="Verrill, A. E. (1895) Supplement to the marine nemerteans and planarians of New England. Transactions of the Connecticut Academy of Arts and Sciences, 9, 523 - 534." type="journal article" year="1895">Verrill 1895</bibRefCitation>
, p. 531
</taxonomicName>
;
<bibRefCitation id="EF8B4B43FFF7CC24FDB3C5F6FD6BFA60" author="Burger, O." box="[599,729,1481,1505]" pageId="6" pageNumber="339" pagination="1 - 151" refId="ref13853" refString="Burger, O. (1904) Nemertini. Das Tierreich, 20, 1 - 151. https: // doi. org / 10.1039 / pl 9042000151" type="journal article" year="1904">Bürger 1904</bibRefCitation>
, p. 124;
<bibRefCitation id="EF8B4B43FFF7CC24FCD4C5F6FC26FA60" author="Coe, W. R." box="[816,916,1481,1505]" pageId="6" pageNumber="339" pagination="129 - 328" refId="ref14244" refString="Coe, W. R. (1943) Biology of the nemerteans of the Atlantic coast of North America. Transactions of the Connecticut Academy of Arts and Sciences, 35, 129 - 328." type="journal article" year="1943">Coe 1943</bibRefCitation>
, p. 253.
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF7CC2BFF73C630FC31FDE2" lastPageId="9" lastPageNumber="342" pageId="6" pageNumber="339" type="etymology">
<paragraph id="8BA536B2FFF7CC24FF73C630FA30F9CC" blockId="6.[151,1437,1551,2045]" pageId="6" pageNumber="339">
<emphasis id="B96EEAA0FFF7CC24FF73C630FEA7F9A8" box="[151,277,1551,1577]" italics="true" pageId="6" pageNumber="339">Etymology.</emphasis>
The specific epithet
<taxonomicName id="4C1A4D31FFF7CC24FE1DC630FDDDF9A8" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[505,623,1551,1577]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="species" species="wijnhoffae">
<emphasis id="B96EEAA0FFF7CC24FE1DC630FDDDF9A8" box="[505,623,1551,1577]" italics="true" pageId="6" pageNumber="339">wijnhoffae</emphasis>
</taxonomicName>
is the latinized form of Gerarda Stiasny-Wijnhoffs maiden name, chosen by us to link her with the author of
<taxonomicName id="4C1A4D31FFF7CC24FDC7C60CFCBCF9CC" authority=", Diva Correa" authorityName=", Diva Correa" authorityYear="1963" box="[547,782,1587,1614]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="6" pageNumber="339" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF7CC24FDC7C60CFDC6F9CC" box="[547,628,1587,1613]" italics="true" pageId="6" pageNumber="339">Dushia</emphasis>
, Diva Corrêa
</taxonomicName>
, another of the great pioneers in nemertean systematics.
</paragraph>
<paragraph id="8BA536B2FFF7CC24FF23C668FC72F883" blockId="6.[151,1437,1551,2045]" pageId="6" pageNumber="339">
<emphasis id="B96EEAA0FFF7CC24FF23C668FE17F9F0" box="[199,421,1623,1649]" italics="true" pageId="6" pageNumber="339">Material examined.</emphasis>
<materialsCitation id="3B723CEFFFF7CC24FE4AC668FE0BF938" ID-GBIF-Occurrence="2446790585" collectingDate="2003-08" collectionCode="USNM" country="Saint Barthelemy" location="Anse de Grand Fond" pageId="6" pageNumber="339" specimenCode="USNM 1594812" specimenCount="1" typeStatus="holotype">
<typeStatus id="54A18810FFF7CC24FE4AC668FDA9F9F0" box="[430,539,1623,1649]" pageId="6" pageNumber="339" type="holotype">Holotype</typeStatus>
:
<specimenCode id="DBBC9EC9FFF7CC24FDC3C668FD54F9F0" box="[551,742,1622,1649]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="6" pageNumber="339" type="Museum">USNM 1594812</specimenCode>
, serial frontal sections of anterior to foregut (14 slides), collected from
<location id="8EC56069FFF7CC24FEF8C644FDB6F914" LSID="urn:lsid:plazi:treatment:03B387A4FFF7CC29FF73C4D9FAE8FDE5:8EC56069FFF7CC24FEF8C644FDB6F914" box="[284,516,1659,1685]" country="Saint Barthelemy" name="Anse de Grand Fond" pageId="6" pageNumber="339">Anse de Grand Fond</location>
,
<collectingCountry id="F30D7622FFF7CC24FDEBC644FD0AF914" box="[527,696,1659,1685]" name="Saint Barthelemy" pageId="6" pageNumber="339">St. Barthélemy</collectingCountry>
,
<collectingCountry id="F30D7622FFF7CC24FD26C643FCBCF917" box="[706,782,1660,1686]" name="France" pageId="6" pageNumber="339">France</collectingCountry>
from under stones on coarse shell hash and sand just at the water line in
<date id="FFA41072FFF7CC24FEC2C6A0FE0BF938" box="[294,441,1695,1721]" pageId="6" pageNumber="339" value="2003-08">
<collectingDate id="EFE0E99AFFF7CC24FEC2C6A0FE0BF938" box="[294,441,1695,1721]" pageId="6" pageNumber="339" value="2003-08">August 2003</collectingDate>
</date>
</materialsCitation>
.
<materialsCitation id="3B723CEFFFF7CC24FE26C69FFC0EF883" ID-GBIF-Occurrence="2446790581" collectionCode="USNM" country="Saint Barthelemy" pageId="6" pageNumber="339" specimenCode="USNM 1594813" specimenCount="1" typeStatus="paratype">
<typeStatus id="54A18810FFF7CC24FE26C69FFD81F93B" box="[450,563,1696,1722]" pageId="6" pageNumber="339" type="paratype">Paratypes</typeStatus>
:
<specimenCode id="DBBC9EC9FFF7CC24FDD9C6A0FD4BF938" box="[573,761,1694,1721]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="6" pageNumber="339" type="Museum">USNM 1594813</specimenCode>
, serial transverse sections of anterior, foregut, mid-body, and intestinal regions (99 slides)
<bibRefCitation id="EF8B4B43FFF7CC24FE3FC6FCFD39F95F" author="Correa, D. D." box="[475,651,1731,1758]" pageId="6" pageNumber="339" 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>
voucher slides and specimens are presumed lost (her entire collection known to have been in her laboratory has been missing for many years)
</materialsCitation>
.
</paragraph>
<paragraph id="8BA536B2FFF7CC24FF23C734FE8BF87C" blockId="6.[151,1437,1551,2045]" pageId="6" pageNumber="339">
<emphasis id="B96EEAA0FFF7CC24FF23C734FE1DF8A4" box="[199,431,1803,1829]" italics="true" pageId="6" pageNumber="339">Additional material.</emphasis>
Six whole specimens fixed in 10% seawater-buffered formalin and preserved in 70% EtOH, and four samples preserved in 95% EtOH were collected from Bahia de la Chiva and Blue Beach,
<collectingRegion id="49DEF850FFF7CC24FADFC710FA2AF8C8" box="[1339,1432,1839,1865]" country="Puerto Rico" name="Vieques" pageId="6" pageNumber="339">Vieques</collectingRegion>
,
<collectingCountry id="F30D7622FFF7CC24FF73C76BFEACF8EC" box="[151,286,1875,1902]" name="Puerto Rico" pageId="6" pageNumber="339">Puerto Rico</collectingCountry>
in
<date id="FFA41072FFF7CC24FEA1C76BFE0BF8EC" box="[325,441,1875,1902]" pageId="6" pageNumber="339" value="2007-06">June 2007</date>
and
<date id="FFA41072FFF7CC24FE15C76CFD37F8EC" box="[497,645,1875,1901]" pageId="6" pageNumber="339" value="2011-08">August 2011</date>
;
<materialsCitation id="3B723CEFFFF7CC24FD76C76BFB2BF810" ID-GBIF-Occurrence="2446790575" collectingDate="2003" collectingDateMax="2007" collectingDateMin="2003" collectionCode="USNM" country="Saint Barthelemy" location="Sixteen" pageId="6" pageNumber="339" specimenCode="JLN0657" specimenCount="1">
two of those DNA extracts are available at the
<collectionCode id="ED0BAE77FFF7CC24FB48C76CFB4FF8EC" box="[1196,1277,1875,1901]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="6" pageNumber="339" type="Museum">USNM</collectionCode>
as
<specimenCode id="DBBC9EC9FFF7CC24FAC2C76DFA3CF8EC" box="[1318,1422,1874,1901]" collectionCode="JLN" pageId="6" pageNumber="339">JLN0657</specimenCode>
_ 0611_001 and 002.
<location id="8EC56069FFF7CC24FE95C748FE77F810" LSID="urn:lsid:plazi:treatment:03B387A4FFF7CC29FF73C4D9FAE8FDE5:8EC56069FFF7CC24FE95C748FE77F810" box="[369,453,1911,1937]" country="Saint Barthelemy" name="Sixteen" pageId="6" pageNumber="339">Sixteen</location>
specimens collected in
<collectingCountry id="F30D7622FFF7CC24FD28C748FC3CF810" box="[716,910,1911,1937]" name="Saint Barthelemy" pageId="6" pageNumber="339">Saint Barthélemy</collectingCountry>
in 2002,
<date id="FFA41072FFF7CC24FC10C748FB2BF810" box="[1012,1177,1911,1937]" pageId="6" pageNumber="339" value="2003" valueMax="2007">
<collectingDate id="EFE0E99AFFF7CC24FC10C748FB2BF810" box="[1012,1177,1911,1937]" pageId="6" pageNumber="339" value="2003" valueMax="2007">2003 and 2007</collectingDate>
</date>
</materialsCitation>
<materialsCitation id="3B723CEFFFF7CC24FB7AC747FA9BF834" ID-GBIF-Occurrence="2446790579" collectingDate="02-03-05" collectionCode="ETOH, USNM" country="Saint Barthelemy" pageId="6" pageNumber="339" specimenCode="JLN0657" specimenCount="1">
were preserved in 95%
<collectionCode id="ED0BAE77FFF7CC24FF73C7A4FF53F834" box="[151,225,1947,1973]" pageId="6" pageNumber="339">ETOH</collectionCode>
and extracted for DNA. Those genomic extracts are available at the
<collectionCode id="ED0BAE77FFF7CC24FC00C7A4FB87F834" box="[996,1077,1947,1973]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="6" pageNumber="339" type="Museum">USNM</collectionCode>
as
<specimenCode id="DBBC9EC9FFF7CC24FBBFC7A5FB76F834" box="[1115,1220,1946,1973]" collectionCode="JLN" pageId="6" pageNumber="339">JLN0657</specimenCode>
_
<date id="FFA41072FFF7CC24FB37C7A5FA9BF834" box="[1235,1321,1946,1973]" pageId="6" pageNumber="339" value="02-03-05">
<collectingDate id="EFE0E99AFFF7CC24FB37C7A5FA9BF834" box="[1235,1321,1946,1973]" pageId="6" pageNumber="339" value="02-03-05">030502</collectingDate>
</date>
</materialsCitation>
_
<materialsCitation id="3B723CEFFFF7CC24FADCC7A5FD0EF858" ID-GBIF-Occurrence="2446790584" collectingDate="03-03-08" collectionCode="JLN" country="Saint Barthelemy" pageId="6" pageNumber="339" specimenCode="JLN0657" specimenCount="1">
001, 002, 008, 010, 011, 012, 013, 014;
<specimenCode id="DBBC9EC9FFF7CC24FE0BC781FDEAF858" box="[495,600,1982,2009]" collectionCode="JLN" pageId="6" pageNumber="339">JLN0657</specimenCode>
_
<date id="FFA41072FFF7CC24FD83C781FD0EF858" box="[615,700,1982,2009]" pageId="6" pageNumber="339" value="03-03-08">
<collectingDate id="EFE0E99AFFF7CC24FD83C781FD0EF858" box="[615,700,1982,2009]" pageId="6" pageNumber="339" value="03-03-08">030803</collectingDate>
</date>
</materialsCitation>
_
<materialsCitation id="3B723CEFFFF7CC24FD2FC781FA9AF858" ID-GBIF-Occurrence="2446790577" box="[715,1320,1982,2009]" collectingDate="07-03-12" collectionCode="JLN" country="Saint Barthelemy" pageId="6" pageNumber="339" specimenCode="JLN0657" specimenCount="1">
003, 004;
<specimenCode id="DBBC9EC9FFF7CC24FCDEC781FC16F858" box="[826,932,1982,2009]" collectionCode="JLN" pageId="6" pageNumber="339">JLN0657</specimenCode>
_0301103_004;
<specimenCode id="DBBC9EC9FFF7CC24FBBEC781FB71F858" box="[1114,1219,1982,2009]" collectionCode="JLN" pageId="6" pageNumber="339">JLN0657</specimenCode>
_
<date id="FFA41072FFF7CC24FB36C781FA9AF858" box="[1234,1320,1982,2009]" pageId="6" pageNumber="339" value="07-03-12">
<collectingDate id="EFE0E99AFFF7CC24FB36C781FA9AF858" box="[1234,1320,1982,2009]" pageId="6" pageNumber="339" value="07-03-12">031207</collectingDate>
</date>
</materialsCitation>
_001, 002, 003, 004, 005.
</paragraph>
<paragraph id="8BA536B2FFF6CC25FF23C0A6FC11FE9E" blockId="7.[151,1436,152,287]" pageId="7" pageNumber="340">
<bibRefCitation id="EF8B4B43FFF6CC25FF23C0A6FEF9FF32" author="Coe, W. R." box="[199,331,152,179]" pageId="7" pageNumber="340" pagination="149 - 186" refId="ref14314" refString="Coe, W. R. (1951 a) The nemertean faunas of the Gulf of Mexico and of southern Florida. Bulletin of Marine Science of the Gulf and Caribbean, 1, 149 - 186." type="journal article" year="1951">Coe (1951a</bibRefCitation>
, b) identified several specimens as
<taxonomicName id="4C1A4D31FFF6CC25FD33C0A5FCE4FF32" authorityName=": Coe" authorityYear="1951" box="[727,854,154,179]" class="Anopla" family="Lineidae" genus="Lineus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="7" pageNumber="340" phylum="Nemertea" rank="species" species="ater">
<emphasis id="B96EEAA0FFF6CC25FD33C0A5FCE4FF32" box="[727,854,154,179]" italics="true" pageId="7" pageNumber="340">Lineus ater</emphasis>
</taxonomicName>
that were deposited in the Smithsonian Institutions National Museum of Natural History and include the following: USNM 1263, collected in
<date id="FFA41072FFF6CC25FB6FC082FA9DFF56" box="[1163,1327,189,215]" pageId="7" pageNumber="340" value="1884-02">February 1884</date>
from
<collectingCountry id="F30D7622FFF6CC25FA96C082FF66FF7A" name="Curaçao" pageId="7" pageNumber="340">Curaçao</collectingCountry>
by the Steamer Albatross R/V; USNM 22085, collected by Henry Hemphill in 1885 from the Florida Keys at low tide; and USNM 24750, collected in Louisiana from under rocks.
</paragraph>
<caption id="DF65663AFFF6CC25FF73C53DFB49FA2A" ID-DOI="http://doi.org/10.5281/zenodo.3527359" ID-Zenodo-Dep="3527359" httpUri="https://zenodo.org/record/3527359/files/figure.png" pageId="7" pageNumber="340" startId="7.[151,250,1282,1307]" targetBox="[201,1387,306,1253]" targetPageId="7">
<paragraph id="8BA536B2FFF6CC25FF73C53DFB49FA2A" blockId="7.[151,1437,1282,1451]" pageId="7" pageNumber="340">
<emphasis id="B96EEAA0FFF6CC25FF73C53DFEA7FA9A" bold="true" box="[151,277,1282,1307]" pageId="7" pageNumber="340">FIGURE 1.</emphasis>
Images of
<taxonomicName id="4C1A4D31FFF6CC25FE68C53DFE67FA9B" authorityName=", Diva Correa" authorityYear="1963" box="[396,469,1282,1306]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="7" pageNumber="340" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF6CC25FE68C53DFE67FA9B" box="[396,469,1282,1306]" italics="true" pageId="7" pageNumber="340">Dushia</emphasis>
</taxonomicName>
spp., taken in life. (A)
<taxonomicName id="4C1A4D31FFF6CC25FD2FC53DFBCDFA9A" authority="Schwartz &amp; Norenburg" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[715,1151,1282,1307]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="7" pageNumber="340" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">
<emphasis id="B96EEAA0FFF6CC25FD2FC53DFC3AFA9B" box="[715,904,1282,1306]" italics="true" pageId="7" pageNumber="340">Dushia wijnhoffae</emphasis>
Schwartz &amp; Norenburg
</taxonomicName>
<emphasis id="B96EEAA0FFF6CC25FB63C53DFB68FA9B" bold="true" box="[1159,1242,1282,1306]" pageId="7" pageNumber="340">
<taxonomicNameLabel id="A25D57DBFFF6CC25FB63C53DFB68FA9B" box="[1159,1242,1282,1306]" pageId="7" pageNumber="340" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
from Puerto Rico. (BF)
<taxonomicName id="4C1A4D31FFF6CC25FF3DC519FDA1FABE" authority="(Stimpson, 1855)" authorityName="Stimpson" authorityYear="1855" baseAuthorityName="Stimpson" baseAuthorityYear="1855" box="[217,531,1318,1343]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="7" pageNumber="340" phylum="Nemertea" rank="species" species="nigra">
<emphasis id="B96EEAA0FFF6CC25FF3DC519FEEDFABF" box="[217,351,1318,1342]" italics="true" pageId="7" pageNumber="340">Dushia nigra</emphasis>
(
<bibRefCitation id="EF8B4B43FFF6CC25FE8FC519FDBEFABE" author="Stimpson, W." box="[363,524,1318,1343]" pageId="7" pageNumber="340" 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="B96EEAA0FFF6CC25FD27C519FC84FABF" bold="true" box="[707,822,1318,1342]" pageId="7" pageNumber="340">comb. nov.</emphasis>
: (B) black form, ICHUM 3457, from Onna, Okinawa, Japan; (C) orange form, from Nakijin, Okinawa, caudal cirrus not pictured; (D) brown form, from Kakeroma, Japan; (E, F) black form from Okinawa, showing body fragments regenerating caudal cirrus, 3 days (E) and 10 days (F) after amputation. Scale bars: AD = 1 cm, E, F = 500 µm. Abbreviations: ao, apical organs; cc, caudal cirrus; cf, horizontal cephalic furrow.
</paragraph>
</caption>
<paragraph id="8BA536B2FFF6CC25FF23C5E6FE2FF9BA" blockId="7.[151,1437,1497,2027]" pageId="7" pageNumber="340">
<emphasis id="B96EEAA0FFF6CC25FF23C5E6FEF2FA73" box="[199,320,1497,1522]" italics="true" pageId="7" pageNumber="340">Diagnosis.</emphasis>
Heteronemertean with black body and dorso-ventraly white-tipped head; head shape retuse; ocelli absent; anterior foregut region moderately rounded becoming moderately dorsoventrally flattened but without sharp lateral margins present.
</paragraph>
<paragraph id="8BA536B2FFF6CC25FF23C67AFCD1F94A" blockId="7.[151,1437,1497,2027]" pageId="7" pageNumber="340">
<emphasis id="B96EEAA0FFF6CC25FF23C67AFEE1F9DF" box="[199,339,1605,1630]" italics="true" pageId="7" pageNumber="340">Description.</emphasis>
<emphasis id="B96EEAA0FFF6CC25FEB8C67AFD87F9DE" bold="true" box="[348,565,1604,1631]" pageId="7" pageNumber="340">External features.</emphasis>
The material examined agrees with
<bibRefCitation id="EF8B4B43FFF6CC25FC39C67AFB22F9DE" author="Correa, D. D." box="[989,1168,1604,1631]" pageId="7" pageNumber="340" 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>
description of
<taxonomicName id="4C1A4D31FFF6CC25FAA5C67AFA24F9DF" authorityName="Girard" authorityYear="1851" box="[1345,1430,1605,1630]" class="Pilidiophora" family="Lineidae" genus="Meckelia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="7" pageNumber="340" phylum="Nemertea" rank="species" species="atra">
<emphasis id="B96EEAA0FFF6CC25FAA5C67AFA24F9DF" box="[1345,1430,1605,1630]" italics="true" pageId="7" pageNumber="340">D. atra</emphasis>
</taxonomicName>
, except for her conclusion that a caudal cirrus is lacking. The cirrus is tiny and its presence is, in fact, extremely difficult to ascertain but can be observed in living animals as a discrete entity. Additional details are from examination of histological sections and observations of live specimens.
</paragraph>
<paragraph id="8BA536B2FFF6CC25FF23C6EAFDD3F822" blockId="7.[151,1437,1497,2027]" pageId="7" pageNumber="340">
The foregut region is slightly rounded in living specimens. The intestinal region is not distinctly flattened (
<figureCitation id="13212A37FFF6CC25FA8BC6EAFF0EF892" captionStart="FIGURE 1" captionStartId="7.[151,250,1282,1307]" captionTargetBox="[201,1387,306,1253]" captionTargetId="figure@7.[200,1387,306,1254]" captionTargetPageId="7" captionText="FIGURE 1. Images of Dushia spp., taken in life. (A) Dushia wijnhoffae Schwartz &amp; Norenburg sp. nov. from Puerto Rico. (BF) Dushia nigra (Stimpson, 1855) species complex comb. nov.: (B) black form, ICHUM 3457, from Onna, Okinawa, Japan; (C) orange form, from Nakijin, Okinawa, caudal cirrus not pictured; (D) brown form, from Kakeroma, Japan; (E, F) black form from Okinawa, showing body fragments regenerating caudal cirrus, 3 days (E) and 10 days (F) after amputation. Scale bars: AD = 1 cm, E, F = 500 µm. Abbreviations: ao, apical organs; cc, caudal cirrus; cf, horizontal cephalic furrow." figureDoi="http://doi.org/10.5281/zenodo.3527359" httpUri="https://zenodo.org/record/3527359/files/figure.png" pageId="7" pageNumber="340">Fig. 1A</figureCitation>
), and lacks sharp lateral margins or any internal dorsoventral musculature (dvm) external to the lateral nerve cords (lnc) that normally is associated with swimming cerebratulids.
<bibRefCitation id="EF8B4B43FFF6CC25FC7EC722FB8AF8B6" author="Girard, C." box="[922,1080,1820,1847]" pageId="7" pageNumber="340" 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 (1851)</bibRefCitation>
,
<bibRefCitation id="EF8B4B43FFF6CC25FBA7C722FB71F8B6" author="Coe, W. R." box="[1091,1219,1821,1847]" pageId="7" pageNumber="340" pagination="129 - 328" refId="ref14244" refString="Coe, W. R. (1943) Biology of the nemerteans of the Atlantic coast of North America. Transactions of the Connecticut Academy of Arts and Sciences, 35, 129 - 328." type="journal article" year="1943">Coe (1943)</bibRefCitation>
, and
<bibRefCitation id="EF8B4B43FFF6CC25FB1BC722FA2EF8B6" author="Correa, D. D." box="[1279,1436,1820,1847]" pageId="7" pageNumber="340" 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>
indicate that their specimens were between
<quantity id="4CE29B57FFF6CC25FD60C77FFD46F8DA" box="[644,756,1856,1883]" metricMagnitude="-1" metricUnit="m" metricValue="3.25" metricValueMax="5.0" metricValueMin="1.5" pageId="7" pageNumber="340" unit="cm" value="32.5" valueMax="50.0" valueMin="15.0">1550 cm</quantity>
long, whereas our samples are between 5 and
<quantity id="4CE29B57FFF6CC25FAE1C77FFAFDF8DA" box="[1285,1359,1856,1883]" metricMagnitude="-1" metricUnit="m" metricValue="1.5" pageId="7" pageNumber="340" unit="cm" value="15.0">15 cm</quantity>
. When living specimens are handled, they are sticky to touch. The worms have not been observed to swim even after prolonged nudging with a dissecting needle.
</paragraph>
<paragraph id="8BA536B2FFF6CC2AFF23C792FE05FE9E" blockId="7.[151,1437,1497,2027]" lastBlockId="8.[151,1436,153,287]" lastPageId="8" lastPageNumber="341" pageId="7" pageNumber="340">
<emphasis id="B96EEAA0FFF6CC25FF23C792FDA2F846" bold="true" box="[199,528,1965,1991]" pageId="7" pageNumber="340">Body wall and musculature.</emphasis>
The epidermis contains numerous red and yellow staining goblet-shaped cells and ciliated cells that rest on a distinct basal lamina (
<figureCitation id="13212A37FFF6CC25FD44C7EEFD4AF86A" box="[672,760,2001,2027]" 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="7" pageNumber="340">Fig. 2A</figureCitation>
). A thin layer of dermal musculature is present below the basal lamina of the epidermis. It includes a circular muscle layer only 23 fibers thick, a lattice-like arrangement of diagonal muscle fibers, and longitudinal muscle layer approximately 25 fibers thick (
<figureCitation id="13212A37FFF9CC2AFC17C082FBDFFF56" box="[1011,1133,189,215]" 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="8" pageNumber="341">Fig. 2A, B</figureCitation>
). Mucoid and serous gland cells are present in the cutis and define its border with the olm. Pigment granules are extracellular and are distributed in the epidermis and cutis.
</paragraph>
<caption id="DF65663AFFF9CC2AFF73C6FEFE31F87B" ID-DOI="http://doi.org/10.5281/zenodo.3527361" ID-Zenodo-Dep="3527361" httpUri="https://zenodo.org/record/3527361/files/figure.png" pageId="8" pageNumber="341" startId="8.[151,250,1729,1754]" targetBox="[183,1405,309,1703]" targetPageId="8">
<paragraph id="8BA536B2FFF9CC2AFF73C6FEFE31F87B" blockId="8.[151,1437,1729,2042]" pageId="8" pageNumber="341">
<emphasis id="B96EEAA0FFF9CC2AFF73C6FEFEA6F95B" bold="true" box="[151,276,1729,1754]" pageId="8" pageNumber="341">FIGURE 2.</emphasis>
<taxonomicName id="4C1A4D31FFF9CC2AFEFFC6FDFD7EF95B" authority="Schwartz &amp; Norenburg" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[283,716,1729,1754]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="8" pageNumber="341" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">
<emphasis id="B96EEAA0FFF9CC2AFEFFC6FDFE65F95B" box="[283,471,1730,1754]" italics="true" pageId="8" pageNumber="341">Dushia wijnhoffae</emphasis>
Schwartz &amp; Norenburg
</taxonomicName>
<emphasis id="B96EEAA0FFF9CC2AFD37C6FDFC97F95B" bold="true" box="[723,805,1730,1754]" pageId="8" pageNumber="341">
<taxonomicNameLabel id="A25D57DBFFF9CC2AFD37C6FDFC97F95B" box="[723,805,1730,1754]" pageId="8" pageNumber="341" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
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.
</paragraph>
</caption>
<paragraph id="8BA536B2FFF8CC2BFF23C0A6FBACFE9E" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">There are a few sparsely distributed mucoid gland cells present at the anterior most portions of the head near the apical organ pits. Acidophilic gland cells are located in a distinct arrangement as in a row across the ventral surface and dorsolaterally in two groups on either side of the rhynchodeal complex, just internal to the epidermis beginning in the anterior of the head and extending posterior to the cerebral ganglia region.</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C116FDE4FE52" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
The remaining body wall is typical, three-layer heteronemertean construction. Radial muscles run between the epithelium of the foregut and the epidermal basal lamina. The foregut and intestine are lined by a glandular epithelium with a few sparsely distributed muscle fibers.A plate of longitudinal muscles is present between the rhynchocoel musculature and the foregut epithelium in the region of the mouth (
<figureCitation id="13212A37FFF8CC2BFC7CC1AAFC43FE2E" box="[920,1009,405,431]" 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="9" pageNumber="342">Fig. 2E</figureCitation>
). The rhynchocoel musculature is not interwoven with the body wall muscles.
</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C1E3FC31FDE2" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C1E3FE72FE76" bold="true" box="[199,448,476,503]" pageId="9" pageNumber="342">Proboscis apparatus.</emphasis>
<bibRefCitation id="EF8B4B43FFF8CC2BFE22C1E2FDD0FE79" author="Correa, D. D." box="[454,610,477,504]" pageId="9" pageNumber="342" 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>
described the proboscis is as three layered with (terminology from everted form) a glandular epithelium, outer longitudinal muscle layer, a neural plexus, an inner circular and longitudinal layer, and two muscle crosses.
<bibRefCitation id="EF8B4B43FFF8CC2BFE0FC21AFD28FDC1" author="Correa, D. D." box="[491,666,549,576]" pageId="9" pageNumber="342" 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>
description does not indicate if pseudocnidae-like structures are present in the epithelium, and we did not observe them in our material.
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF8CC2BFF23C253FD52F9AE" pageId="9" pageNumber="342" type="description">
<paragraph id="8BA536B2FFF8CC2BFF23C253FE30FCBA" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C253FE12FD06" bold="true" box="[199,416,620,647]" pageId="9" pageNumber="342">Alimentary canal.</emphasis>
Intestinal diverticula can be identified by the presence of dorsoventral (dv) musculature that insert in the dorsal and ventral portions of the middle circular muscle (mcm) fibers passing through the inner longitudinal muscle (ilm) layer on either side of the intestine (
<figureCitation id="13212A37FFF8CC2BFCA0C28AFC2EFD4E" box="[836,924,693,719]" 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="9" pageNumber="342">Fig. 2C</figureCitation>
). Most of the dv musculature is external to the lateral blood vessels in the intestinal region with a few fibers passing internally to the vessel. The diverticula are also associated with a layer of longitudinally oriented muscles that are overlain by the dvm fibers and are held together by connective tissue.
</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C37BFB3CFBB6" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C37BFE00FCDE" bold="true" box="[199,434,836,863]" pageId="9" pageNumber="342">Circulatory system.</emphasis>
<bibRefCitation id="EF8B4B43FFF8CC2BFE5EC37AFDEBFCE1" author="Correa, D. D." box="[442,601,837,864]" pageId="9" pageNumber="342" 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>
identified four cephalic vessels in the circulatory system; a larger pair arranged laterally to the rhynchodaeum and a smaller pair arranged dorsal and laterally to the rhynchodaeum. The two smaller vessels have perforations in the septum separating them. In our material, the two small vessels are arranged as a single dorsal vessel for most of its length but does contain several partitions (
<figureCitation id="13212A37FFF8CC2BFBD4C38EFB38FC4A" box="[1072,1162,945,971]" 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="9" pageNumber="342">Fig. 2D</figureCitation>
).
<bibRefCitation id="EF8B4B43FFF8CC2BFB7AC38EFA8EFC4D" author="Correa, D. D." box="[1182,1340,945,972]" pageId="9" pageNumber="342" 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>
also did not observe a rhynchocoel vessel or villus. In our material, a vessel is present along the ventral portion of the rhynchocoel from the mouth and foregut regions to the intestinal region where it exits from the rhynchocoel ventrally and becomes the intestinal dorsal vessel positioned between the intestine and rhynchocoel.
</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C47EFBBEFB6A" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C47EFE25FBDA" bold="true" box="[199,407,1089,1115]" pageId="9" pageNumber="342">Excretory system</emphasis>
. The specimen sectioned was regenerating parts of foregut regions including the proboscis and rhynchocoel, near to the region where the nephridial system is usually found. The excretory system begins as a series of branching tubules in the region of the mouth. From our sample it is not clear if these branches are connected to the tubules in the foregut, although
<bibRefCitation id="EF8B4B43FFF8CC2BFD70C492FC80FB49" author="Correa, D. D." box="[660,818,1197,1224]" pageId="9" pageNumber="342" 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>
indicates that they are a single continuous system. The nerphridial ducts lead away from the tubules dorsolaterally to nephridial pores.
</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C4CAFAADFAD6" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C4CAFDEEFA8E" bold="true" box="[199,604,1269,1295]" pageId="9" pageNumber="342">Nervous system and sense organs.</emphasis>
Bürger
<typeStatus id="54A18810FFF8CC2BFD5CC4CAFD5AFA8E" box="[696,744,1269,1295]" pageId="9" pageNumber="342">type</typeStatus>
I, II, III and neurocord cells were observed in both the dorsal and ventral ganglia and the anterior most portions of the lateral nerve cord (lnc) (
<figureCitation id="13212A37FFF8CC2BFBF9C526FB27FAB2" box="[1053,1173,1305,1331]" 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="9" pageNumber="342">Fig. 3A, B</figureCitation>
). A few neurocords can be identified within the fibrous part of the lateral nerve cords, but only in the region near to the ganglia.
</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C55EFD52F9AE" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
A distinctive U-shaped ridge of cells with long cilia and interspersed gland cells is present in the cephalic furrows flanking the entrance to the cerebral organ (
<figureCitation id="13212A37FFF8CC2BFD1DC5BAFCC9FA1E" box="[761,891,1413,1439]" 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="9" pageNumber="342">Fig. 3C, D</figureCitation>
). This ridge can be observed in living, relaxed specimens by separating the cephalic furrows with needles. This feature is not usually noted in descriptions of heteronemerteans, yet is present in all heteronemerteans examined except those without cephalic furrows (MLS pers. obs.,
<figureCitation id="13212A37FFF8CC2BFF37C5CEFEE2F98A" box="[211,336,1521,1547]" 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="9" pageNumber="342">Figs 3C, D</figureCitation>
,
<figureCitation id="13212A37FFF8CC2BFEB8C5CEFE30F98D" box="[348,386,1521,1548]" 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="9" pageNumber="342">6D</figureCitation>
).
<bibRefCitation id="EF8B4B43FFF8CC2BFE72C5CEFD90F98A" author="Riser, N. W." box="[406,546,1521,1547]" pageId="9" pageNumber="342" pagination="597 - 606" refId="ref16825" refString="Riser, N. W. (1990) New Zealand nemertines from kelp holdfasts: Heteronemertinea I. Adenorhagas aurantiafrons gen. n., sp. n. New Zealand Journal of Zoology, 17, 597 - 606. https: // doi. org / 10.1080 / 03014223.1990.10422957" type="journal article" year="1990">Riser (1990)</bibRefCitation>
was the first to notice the structure, and named the genus
<taxonomicName id="4C1A4D31FFF8CC2BFB4BC5CEFAF6F98A" authorityName="Riser" authorityYear="1990" box="[1199,1348,1521,1547]" class="Anopla" family="Lineidae" genus="Adenorhagas" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="9" pageNumber="342" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFF8CC2BFB4BC5CEFAF6F98A" box="[1199,1348,1521,1547]" italics="true" pageId="9" pageNumber="342">Adenorhagas</emphasis>
</taxonomicName>
after it, however, it is not a structure restricted to this genus.
</paragraph>
</subSubSection>
<subSubSection id="C3006539FFF8CC29FF23C607FAE8FDE5" lastPageId="11" lastPageNumber="344" pageId="9" pageNumber="342" type="materials_examined">
<paragraph id="8BA536B2FFF8CC2BFF23C607FEDEF91A" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C607FE73F9D2" bold="true" box="[199,449,1592,1619]" pageId="9" pageNumber="342">Reproductive system</emphasis>
. The material examined was spawned out and the gonads appeared to be in the process of regressing. The gonads are arranged serially between the intestinal diverticula walls. Gonoducts and gonopores were not observed.
</paragraph>
<paragraph id="8BA536B2FFF8CC2BFF23C69AFDE9F83A" blockId="9.[151,1437,153,1979]" pageId="9" pageNumber="342">
<emphasis id="B96EEAA0FFF8CC2BFF23C69AFD1DF93E" box="[199,687,1701,1727]" italics="true" pageId="9" pageNumber="342">
Habitat and distribution.
<taxonomicName id="4C1A4D31FFF8CC2BFE06C69AFD1DF93E" authority="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg, 2019" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[482,687,1701,1727]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="9" pageNumber="342" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">Dushia wijnhoffae</taxonomicName>
</emphasis>
<emphasis id="B96EEAA0FFF8CC2BFD51C69AFCBFF93E" bold="true" box="[693,781,1701,1727]" pageId="9" pageNumber="342">
<taxonomicNameLabel id="A25D57DBFFF8CC2BFD51C69AFCBFF93E" box="[693,781,1701,1727]" pageId="9" pageNumber="342" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
is littoral, found under stones partially embedded in course shell hash and sand at the water line. Their particular habitat is very clean without much additional organic material and has few other inhabitants. They are easy to identify in the field since their black pigmentation makes the worms conspicuous against the white and brown of the shell and sand when turning rocks. Specimens have been collected or observed from the following locations in the Caribbean:
<collectingCountry id="F30D7622FFF8CC2BFCDBC70AFC58F8CE" box="[831,1002,1845,1871]" name="Saint Barthelemy" pageId="9" pageNumber="342">St. Barthélemy</collectingCountry>
,
<collectingCountry id="F30D7622FFF8CC2BFC13C70AFBF1F8CE" box="[1015,1091,1845,1871]" name="France" pageId="9" pageNumber="342">France</collectingCountry>
(
<collectionCode id="ED0BAE77FFF8CC2BFBB0C70AFB3BF8CE" box="[1108,1161,1845,1871]" pageId="9" pageNumber="342">JLN</collectionCode>
), Bahia de la Chiva and Blue Beach,
<collectingRegion id="49DEF850FFF8CC2BFEC3C766FE36F8F2" box="[295,388,1881,1907]" country="Puerto Rico" name="Vieques" pageId="9" pageNumber="342">Vieques</collectingRegion>
,
<collectingCountry id="F30D7622FFF8CC2BFE6BC766FDA5F8F2" box="[399,535,1881,1907]" name="Puerto Rico" pageId="9" pageNumber="342">Puerto Rico</collectingCountry>
,
<collectingCountry id="F30D7622FFF8CC2BFDC6C766FDEAF8F2" box="[546,600,1881,1907]" name="United States of America" pageId="9" pageNumber="342">USA</collectingCountry>
(
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),
<collectingCountry id="F30D7622FFF8CC2BFD5EC766FC9AF8F2" box="[698,808,1881,1907]" name="Dominica" pageId="9" pageNumber="342">Dominica</collectingCountry>
(Leslie Harris, pers. comm.), and
<collectingCountry id="F30D7622FFF8CC2BFB4CC766FAB7F8F2" box="[1192,1285,1881,1907]" name="Curaçao" pageId="9" pageNumber="342">Curaçao</collectingCountry>
(
<bibRefCitation id="EF8B4B43FFF8CC2BFAF0C766FEACF816" author="Stiasny-Wijnhoff, G." pageId="9" pageNumber="342" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoff 1925</bibRefCitation>
;
<bibRefCitation id="EF8B4B43FFF8CC2BFECEC742FE08F819" author="Correa, D. D." box="[298,442,1917,1944]" pageId="9" pageNumber="342" 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>
). Material deposited at
<collectionCode id="ED0BAE77FFF8CC2BFD20C742FCA8F816" box="[708,794,1917,1943]" country="USA" httpUri="http://grbio.org/cool/142r-0w94" name="Smithsonian Institution, National Museum of Natural History" pageId="9" pageNumber="342" type="Museum">NMNH</collectionCode>
implies that they are also found in littoral habitats in the Gulf of
<collectingCountry id="F30D7622FFF8CC2BFF14C79EFEF4F83A" box="[240,326,1953,1979]" name="Mexico" pageId="9" pageNumber="342">Mexico</collectingCountry>
and in the Florida Keys.
</paragraph>
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<emphasis id="B96EEAA0FFFBCC28FF73C557FEBDFA00" bold="true" box="[151,271,1384,1409]" pageId="10" pageNumber="343">FIGURE 3.</emphasis>
<taxonomicName id="4C1A4D31FFFBCC28FEF0C557FD1EFA00" authority="Schwartz &amp; Norenburg" authorityName="Hookabe &amp; Schwartz &amp; Kajihara &amp; Norenburg" authorityYear="2019" box="[276,684,1384,1409]" class="Anopla" family="Lineidae" genus="Dushia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="10" pageNumber="343" phylum="Nemertea" rank="species" species="wijnhoffae" status="sp. nov.">
<emphasis id="B96EEAA0FFFBCC28FEF0C557FE77FA01" box="[276,453,1384,1408]" italics="true" pageId="10" pageNumber="343">Dushia wijnhoffae</emphasis>
Schwartz &amp; Norenburg
</taxonomicName>
<emphasis id="B96EEAA0FFFBCC28FD55C557FD4CFA01" bold="true" box="[689,766,1384,1408]" pageId="10" pageNumber="343">
<taxonomicNameLabel id="A25D57DBFFFBCC28FD55C557FD4CFA01" box="[689,766,1384,1408]" pageId="10" pageNumber="343" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(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.
</paragraph>
</caption>
<paragraph id="8BA536B2FFFBCC28FF23C6B8FE21F8E8" blockId="10.[151,1437,1670,2031]" pageId="10" pageNumber="343">
<emphasis id="B96EEAA0FFFBCC28FF23C6B8FE83F920" box="[199,305,1671,1697]" italics="true" pageId="10" pageNumber="343">Remarks.</emphasis>
<taxonomicName id="4C1A4D31FFFBCC28FED3C6B8FDD9F920" authority="Girard, 1851" authorityName="Girard" authorityYear="1851" box="[311,619,1670,1697]" class="Pilidiophora" family="Lineidae" genus="Meckelia" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="10" pageNumber="343" phylum="Nemertea" rank="species" species="atra">
<emphasis id="B96EEAA0FFFBCC28FED3C6B8FE66F920" box="[311,468,1671,1697]" italics="true" pageId="10" pageNumber="343">Meckelia atra</emphasis>
<bibRefCitation id="EF8B4B43FFFBCC28FE3FC6B8FDD9F920" author="Girard, C." box="[475,619,1670,1697]" pageId="10" pageNumber="343" 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, 1851</bibRefCitation>
</taxonomicName>
was originally described from a single specimen dredged in deep waters off Cape
<collectingRegion id="49DEF850FFFBCC28FEE7C697FEE6F943" box="[259,340,1704,1730]" country="United States of America" name="Florida" pageId="10" pageNumber="343">Florida</collectingRegion>
(
<bibRefCitation id="EF8B4B43FFFBCC28FE81C697FE41F943" author="Girard, C." box="[357,499,1703,1730]" pageId="10" pageNumber="343" 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 1851</bibRefCitation>
,
<bibRefCitation id="EF8B4B43FFFBCC28FDE5C697FD8FF943" author="Girard, C." box="[513,573,1704,1730]" pageId="10" pageNumber="343" pagination="145 - 310" refId="ref14718" refString="Girard, C. (1893) Recherches sur les Planaries et les Nemertiens de l'Amerique du Nord. Annals des Sciences Naturelles, Serie 7, Zoologie, 15, 145 - 310. Available from: https: // www. biodiversitylibrary. org / item / 111186 # page / 155 / mode / 1 up (Accessed 31 Oct. 2019)" type="journal article" year="1893">1893</bibRefCitation>
).
<bibRefCitation id="EF8B4B43FFFBCC28FDB7C697FCB0F943" author="Girard, C." box="[595,770,1703,1730]" pageId="10" pageNumber="343" 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">Girards (1851)</bibRefCitation>
description is short and indicates that the specimen had a flattened body, was black throughout except with a paler anterior, and was
<quantity id="4CE29B57FFFBCC28FC1CC6F6FBF2F965" box="[1016,1088,1737,1764]" metricMagnitude="-1" metricUnit="m" metricValue="1.5" pageId="10" pageNumber="343" unit="cm" value="15.0">15 cm</quantity>
long after preservation. Later,
<bibRefCitation id="EF8B4B43FFFBCC28FF73C6D4FE80F884" author="Verrill, A. E." box="[151,306,1770,1797]" pageId="10" pageNumber="343" pagination="523 - 534" refId="ref18018" refString="Verrill, A. E. (1895) Supplement to the marine nemerteans and planarians of New England. Transactions of the Connecticut Academy of Arts and Sciences, 9, 523 - 534." type="journal article" year="1895">Verrill (1895)</bibRefCitation>
made
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<emphasis id="B96EEAA0FFFBCC28FE64C6D4FE55F884" box="[384,487,1771,1797]" italics="true" pageId="10" pageNumber="343">Meckelia</emphasis>
</taxonomicName>
a junior synonym to
<taxonomicName id="4C1A4D31FFFBCC28FD39C6D4FCC2F884" authorityName="Reiner" authorityYear="1804" box="[733,880,1771,1797]" class="Anopla" family="Lineidae" genus="Cerebratulus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="10" pageNumber="343" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFFBCC28FD39C6D4FCC2F884" box="[733,880,1771,1797]" italics="true" pageId="10" pageNumber="343">Cerebratulus</emphasis>
</taxonomicName>
.
<bibRefCitation id="EF8B4B43FFFBCC28FC64C6D4FBBEF884" author="Coe, W. R." box="[896,1036,1770,1797]" pageId="10" pageNumber="343" pagination="149 - 186" refId="ref14314" refString="Coe, W. R. (1951 a) The nemertean faunas of the Gulf of Mexico and of southern Florida. Bulletin of Marine Science of the Gulf and Caribbean, 1, 149 - 186." type="journal article" year="1951">Coe (1951a)</bibRefCitation>
eventually transferred it to
<taxonomicName id="4C1A4D31FFFBCC28FAAFC6D3FA24F884" authorityName="J.Sowerby" authorityYear="1806" box="[1355,1430,1772,1797]" class="Anopla" family="Lineidae" genus="Lineus" higherTaxonomySource="CoL" kingdom="Animalia" order="Heteronemertea" pageId="10" pageNumber="343" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFFBCC28FAAFC6D3FA24F884" box="[1355,1430,1772,1797]" italics="true" pageId="10" pageNumber="343">Lineus</emphasis>
</taxonomicName>
, presumably because of the lack of a caudal cirrus. It is not clear if
<bibRefCitation id="EF8B4B43FFFBCC28FC93C733FBB0F8A7" author="Coe, W. R." box="[887,1026,1803,1830]" pageId="10" pageNumber="343" pagination="149 - 186" refId="ref14314" refString="Coe, W. R. (1951 a) The nemertean faunas of the Gulf of Mexico and of southern Florida. Bulletin of Marine Science of the Gulf and Caribbean, 1, 149 - 186." type="journal article" year="1951">Coe (1951a)</bibRefCitation>
examined any living samples and he may have been working with material from the Steamer Albatross collected in
<date id="FFA41072FFFBCC28FBF2C712FB0CF8C9" box="[1046,1214,1837,1864]" pageId="10" pageNumber="343" value="1884" valueMax="1885">1884 and 1885</date>
that is deposited in the
<collectionCode id="ED0BAE77FFFBCC28FF24C770FEA4F8E8" box="[192,278,1871,1897]" country="USA" httpUri="http://grbio.org/cool/142r-0w94" name="Smithsonian Institution, National Museum of Natural History" pageId="10" pageNumber="343" type="Museum">NMNH</collectionCode>
collection.
</paragraph>
<paragraph id="8BA536B2FFFBCC29FF23C74FFA26FEFA" blockId="10.[151,1437,1670,2031]" lastBlockId="11.[151,1437,152,613]" lastPageId="11" lastPageNumber="344" pageId="10" pageNumber="343">
<bibRefCitation id="EF8B4B43FFFBCC28FF23C74FFE69F80B" author="Stiasny-Wijnhoff, G." box="[199,475,1903,1930]" pageId="10" pageNumber="343" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoff (1925)</bibRefCitation>
collected two headless specimens littorally in “Spanish waters” off
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but was unsure that they were the same as
<bibRefCitation id="EF8B4B43FFFBCC28FDE9C7AEFD0BF82D" author="Girard, C." box="[525,697,1937,1964]" pageId="10" pageNumber="343" 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">Girards (1851)</bibRefCitation>
material and placed a “?” mark in her description of the fragments.
<bibRefCitation id="EF8B4B43FFFBCC28FF73C78CFE19F84C" author="Stiasny-Wijnhoff, G." box="[151,427,1970,1997]" pageId="10" pageNumber="343" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoff (1925)</bibRefCitation>
apparently dissected the animals or cut histological sections because she noted the presence of distinct connective tissue layer between the cutis and olm layers in her specimens.
<bibRefCitation id="EF8B4B43FFFBCC28FB7BC7EBFADBF86F" author="Correa, D. D." box="[1183,1385,2003,2030]" pageId="10" pageNumber="343" 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: 51)</bibRefCitation>
col- lected in
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and returned to
<bibRefCitation id="EF8B4B43FFFACC29FDF4C0A6FC84FF32" author="Stiasny-Wijnhoff, G." box="[528,822,152,179]" pageId="11" pageNumber="344" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoffs (1925)</bibRefCitation>
sites but was not able to find any specimens and stated that the habitat was not correct for this particular species. In her histological examination of the
<collectingCountry id="F30D7622FFFACC29FB30C085FA83FF55" box="[1236,1329,186,212]" name="Curaçao" pageId="11" pageNumber="344">Curaçao</collectingCountry>
material,
<bibRefCitation id="EF8B4B43FFFACC29FF73C0E3FE84FF77" author="Correa, D. D." box="[151,310,219,246]" pageId="11" pageNumber="344" 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>
found that they did not possess a connective tissue layer as
<bibRefCitation id="EF8B4B43FFFACC29FC06C0E4FB47FF77" author="Stiasny-Wijnhoff, G." box="[994,1269,219,246]" pageId="11" pageNumber="344" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoff (1925)</bibRefCitation>
observed. The material examined here does not possess a thickened connective tissue layer, it is of minimal thickness, and mostly associated with the proximal outer longitudinal muscle fibers rather than a discrete layer (
<figureCitation id="13212A37FFFACC29FB42C121FA97FEB8" box="[1190,1317,286,313]" 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="11" pageNumber="344">Fig. 2A, B</figureCitation>
). Whether
<bibRefCitation id="EF8B4B43FFFACC29FF73C100FE8FFEDB" author="Girard, C." box="[151,317,319,346]" pageId="11" pageNumber="344" 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">Girards (1851</bibRefCitation>
,
<bibRefCitation id="EF8B4B43FFFACC29FEACC17FFE33FEDB" author="Girard, C." box="[328,385,320,346]" pageId="11" pageNumber="344" pagination="145 - 310" refId="ref14718" refString="Girard, C. (1893) Recherches sur les Planaries et les Nemertiens de l'Amerique du Nord. Annals des Sciences Naturelles, Serie 7, Zoologie, 15, 145 - 310. Available from: https: // www. biodiversitylibrary. org / item / 111186 # page / 155 / mode / 1 up (Accessed 31 Oct. 2019)" type="journal article" year="1893">1893</bibRefCitation>
) and
<bibRefCitation id="EF8B4B43FFFACC29FE5AC100FD56FEDB" author="Stiasny-Wijnhoff, G." box="[446,740,319,346]" pageId="11" pageNumber="344" pagination="97 - 120" refId="ref17488" refString="Stiasny-Wijnhoff, G. (1925) On a collection of nemerteans from Curacao. Bijdragen tot de Dierkunde, 24, 97 - 120. https: // doi. org / 10.1163 / 26660644 - 02401007" type="journal article" year="1925">Stiasny-Wijnhoffs (1925)</bibRefCitation>
specimens are conspecific or not remains indecisive, but these are apparently different from
<bibRefCitation id="EF8B4B43FFFACC29FE06C15EFD20FEFD" author="Correa, D. D." box="[482,658,353,380]" pageId="11" pageNumber="344" 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>
and ours, and the latter two undoubtedly represent the same species.
</paragraph>
<paragraph id="8BA536B2FFFACC29FF01C1BCFAE8FDE5" blockId="11.[151,1437,152,613]" pageId="11" pageNumber="344">
In
<collectingCountry id="F30D7622FFFACC29FEE0C1BDFE02FE1D" box="[260,432,386,412]" name="Saint Barthelemy" pageId="11" pageNumber="344">St. Barthélemy</collectingCountry>
, despite extensive sampling of all sand habitats around the island, the worms were found exclusively at the west end of Grand Fond in the intertidal wash zone behind a fossil reef that is exposed at low tide. They are abundant, often in clusters of two or three but as many as 10 or more worms, under large rubble lying on coarse carbonate (coral) sand; the worms burrowed very rapidly into this sand when exposed. Associated fauna includes shelled micro-mollusks, small crustaceans and members of three species of
<taxonomicName id="4C1A4D31FFFACC29FBDBC237FABDFDA3" authorityName="Diesing" authorityYear="1863" box="[1087,1295,520,546]" class="Enopla" family="Ototyphlonemertidae" genus="Ototyphlonemertes" kingdom="Animalia" order="Monostilifera" pageId="11" pageNumber="344" phylum="Nemertea" rank="genus">
<emphasis id="B96EEAA0FFFACC29FBDBC237FABDFDA3" box="[1087,1295,520,546]" italics="true" pageId="11" pageNumber="344">Ototyphlonemertes</emphasis>
</taxonomicName>
. In captivity the worms enthusiastically react to the scent of and feed on several kinds of crustacean, mollusk and fish tissues but, despite a very large mouth, the tissues must be chopped into cubes of
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or less; these are swallowed whole.
</paragraph>
</subSubSection>
</treatment>
</document>