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<document id="0C050C939FF593FE01F5CE097E8E1413" ID-DOI="10.5281/zenodo.157389" ID-GBIF-Dataset="99344430-adda-4ae7-ad21-46c1118f3aa2" ID-ISSN="1175­5326" ID-Zenodo-Dep="157389" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461016794336" checkinUser="plazi" docAuthor="Brodie, Gilianne D." docDate="2004" docId="0390696A8C21FFF4615FFA2656ADBC2B" docLanguage="en" docName="zt00503.pdf" docOrigin="Zootaxa 503" docStyle="DocumentStyle:FA7E419B012A62B0FC3AC15A186C3DAF.3:Zootaxa.2001-2006.journal_article" docStyleId="FA7E419B012A62B0FC3AC15A186C3DAF" docStyleName="Zootaxa.2001-2006.journal_article" docStyleVersion="3" docTitle="Dendrodoris atromaculata Alder &amp; Hancock 1864" docType="treatment" docVersion="9" lastPageNumber="11" masterDocId="FFA911128C23FFFE6057FFEF5718BA75" masterDocTitle="An unusual dendrodorid: redescription of the tropical nudibranch Dendrodoris atromaculata (Alder &amp; Hancock, 1864) (Anthobranchia: Doridoidea: Dendrodorididae)" masterLastPageNumber="13" masterPageNumber="1" pageNumber="3" updateTime="1698568908377" updateUser="plazi">
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<mods:title id="9A2793600CF2B1A5C17CD1A089D7F1AD">An unusual dendrodorid: redescription of the tropical nudibranch Dendrodoris atromaculata (Alder &amp; Hancock, 1864) (Anthobranchia: Doridoidea: Dendrodorididae)</mods:title>
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<mods:namePart id="7804692E2EBA041D7CAEC9FCAD4BF0CB">Brodie, Gilianne D.</mods:namePart>
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<mods:date id="3095C2FACCFAF8BB0D98C3D781B9F713">2004</mods:date>
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<treatment id="0390696A8C21FFF4615FFA2656ADBC2B" ID-DOI="http://doi.org/10.5281/zenodo.6271983" ID-GBIF-Taxon="119601701" ID-Zenodo-Dep="6271983" LSID="urn:lsid:plazi:treatment:0390696A8C21FFF4615FFA2656ADBC2B" httpUri="http://treatment.plazi.org/id/0390696A8C21FFF4615FFA2656ADBC2B" lastPageId="10" lastPageNumber="11" pageId="2" pageNumber="3">
<subSubSection id="C3238BF78C21FFFC615FFA265538BC73" pageId="2" pageNumber="3" type="nomenclature">
<paragraph id="8B86D87C8C21FFFC615FFA265491BF96" blockId="2.[264,905,1481,1542]" box="[264,905,1481,1507]" pageId="2" pageNumber="3">
<heading id="D0CE6F108C21FFFC615FFA265491BF96" bold="true" box="[264,905,1481,1507]" fontSize="11" level="1" pageId="2" pageNumber="3" reason="1">
<taxonomicName id="4C39A3FF8C21FFFC615FFA265491BF96" ID-CoL="34Q5P" authority="Alder &amp; Hancock, 1864" authorityName="Alder &amp; Hancock" authorityYear="1864" box="[264,905,1481,1507]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="2" pageNumber="3" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C21FFFC615FFA265491BF96" bold="true" box="[264,905,1481,1507]" pageId="2" pageNumber="3">
<emphasis id="B94D046E8C21FFFC615FFA265550BF96" bold="true" box="[264,584,1481,1507]" italics="true" pageId="2" pageNumber="3">Dendrodoris atromaculata</emphasis>
(
<bibRefCitation id="EFA8A58D8C21FFFC6200FA265499BF96" author="Alder" box="[599,897,1481,1507]" pageId="2" pageNumber="12" refString="Alder, J. &amp; Hancock, A. (1864) Notice of a collection of nudibranchiate Mollusca made in India by Walter Elliot, Esq., with description of several new genera and species. Transactions of the Zoological Society of London, 5, 113 - 147." type="journal article" year="1864">Alder &amp; Hancock, 1864</bibRefCitation>
)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8B86D87C8C21FFFC615FFA035538BC73" blockId="2.[264,905,1481,1542]" box="[264,544,1516,1542]" pageId="2" pageNumber="3">
(
<figureCitation id="1302C4F98C21FFFC6147FA035662BC73" box="[272,378,1516,1542]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="2" pageNumber="3">Figures 1</figureCitation>
A­F &amp; 2A­D)
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C21FFFC615FF9D35667BC20" box="[264,383,1596,1621]" pageId="2" pageNumber="3" type="description">
<paragraph id="8B86D87C8C21FFFC615FF9D35667BC20" blockId="2.[264,1323,1596,1787]" box="[264,383,1596,1621]" pageId="2" pageNumber="3">
<emphasis id="B94D046E8C21FFFC615FF9D35667BC20" box="[264,383,1596,1621]" italics="true" pageId="2" pageNumber="3">Synonymy</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C21FFFC616FF98B5463BC8E" pageId="2" pageNumber="3" type="reference_group">
<paragraph id="8B86D87C8C21FFFC616FF98B56FFBCCE" blockId="2.[264,1323,1596,1787]" pageId="2" pageNumber="3">
<taxonomicName id="4C39A3FF8C21FFFC616FF98B5455BC0E" authority="Alder &amp; Hancock, 1864" authorityName="Alder &amp; Hancock" authorityYear="1864" box="[312,845,1636,1659]" class="Gastropoda" family="Dendrodorididae" genus="Doridopsis" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="2" pageNumber="3" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C21FFFC616FF98B555CBC0E" box="[312,580,1636,1659]" italics="true" pageId="2" pageNumber="3">Doridopsis atromaculata</emphasis>
<bibRefCitation id="EFA8A58D8C21FFFC621DF98B5455BC0E" author="Alder" box="[586,845,1636,1659]" pageId="2" pageNumber="12" refString="Alder, J. &amp; Hancock, A. (1864) Notice of a collection of nudibranchiate Mollusca made in India by Walter Elliot, Esq., with description of several new genera and species. Transactions of the Zoological Society of London, 5, 113 - 147." type="journal article" year="1864">Alder &amp; Hancock, 1864</bibRefCitation>
</taxonomicName>
, p. 129, Pl. XXXI. figs. 20 ­ 24.
<taxonomicName id="4C39A3FF8C21FFFC616FF96B5425BCEE" ID-CoL="34Q5P" authority="Alder &amp; Hancock" authorityName="Alder &amp; Hancock" box="[312,829,1668,1691]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="2" pageNumber="3" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C21FFFC616FF96B554FBCEE" box="[312,599,1668,1691]" italics="true" pageId="2" pageNumber="3">Dendrodoris atromaculata</emphasis>
(Alder &amp; Hancock)
</taxonomicName>
. —
<bibRefCitation id="EFA8A58D8C21FFFC6327F96B54EABCEE" author="Eliot" box="[880,1010,1668,1691]" pageId="2" pageNumber="12" refString="Eliot, C. (1906 a) On the nudibranchs of southern India and Ceylon, with special reference to the drawings by Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at Newcastle-on-Tyne. Proceedings of the Zoological Society of London, 636 - 691." type="proceedings paper" year="1906" yearSuffix="a">Eliot 1906a</bibRefCitation>
, p. 662;
<bibRefCitation id="EFA8A58D8C21FFFC640EF96B56B8BCCE" author="Willan" pageId="2" pageNumber="13" refString="Willan R. C. &amp; Coleman, N. (1984) Nudibranchs of Australasia. Australasian Marine Photographic Index, Caringbah, Sydney, 56 pp." type="book" year="1984">Willan &amp; Coleman 1984</bibRefCitation>
, p. 53.
</paragraph>
<paragraph id="8B86D87C8C21FFFC616FF92B5463BC8E" blockId="2.[264,1323,1596,1787]" pageId="2" pageNumber="3">
<taxonomicName id="4C39A3FF8C21FFFC616FF92B56A7BCAE" ID-CoL="7NVPG" box="[312,447,1732,1755]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="2" pageNumber="3" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C21FFFC616FF92B56A7BCAE" box="[312,447,1732,1755]" italics="true" pageId="2" pageNumber="3">Dendrodoris</emphasis>
</taxonomicName>
sp. 2
<bibRefCitation id="EFA8A58D8C21FFFC61A9F92B55DDBCAE" author="Lim" box="[510,709,1732,1755]" pageId="2" pageNumber="13" refString="Lim, C. F. &amp; Chou, L. M. (1970) The dendrodorid and doridid nudibranchs of Singapore. Malayan Nature Journal, 23, 92 - 117." type="journal article" year="1970">Lim &amp; Chou 1970</bibRefCitation>
, 96­97, figs. 4N &amp; 9C.
<taxonomicName id="4C39A3FF8C21FFFC616FF90B543EBC8E" ID-CoL="34Q7F" authority="Lim &amp; Chou 1970" authorityName="Lim &amp; Chou" authorityYear="1970" box="[312,806,1764,1787]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="2" pageNumber="3" phylum="Mollusca" rank="species" species="singaporensis">
<emphasis id="B94D046E8C21FFFC616FF90B5540BC8E" box="[312,600,1764,1787]" italics="true" pageId="2" pageNumber="3">Dendrodoris singaporensis</emphasis>
<bibRefCitation id="EFA8A58D8C21FFFC6208F90B543EBC8E" author="Lim" box="[607,806,1764,1787]" pageId="2" pageNumber="13" refString="Lim, C. F. &amp; Chou, L. M. (1970) The dendrodorid and doridid nudibranchs of Singapore. Malayan Nature Journal, 23, 92 - 117." type="journal article" year="1970">Lim &amp; Chou 1970</bibRefCitation>
</taxonomicName>
, p. 104.
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C21FFFD615FF8C355A8B913" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3" type="discussion">
<paragraph id="8B86D87C8C21FFFC615FF8C3553EBD30" blockId="2.[264,1321,1836,1902]" box="[264,550,1836,1861]" pageId="2" pageNumber="3">
<emphasis id="B94D046E8C21FFFC615FF8C3553EBD30" box="[264,550,1836,1861]" italics="true" pageId="2" pageNumber="3">Comments on Synonymy</emphasis>
</paragraph>
<paragraph id="8B86D87C8C21FFFD616FF8BB5451B83B" blockId="2.[264,1321,1836,1902]" lastBlockId="3.[264,1325,284,1350]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
The genus
<taxonomicName id="4C39A3FF8C21FFFC61E8F8BB554CBD18" box="[447,596,1876,1901]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="2" pageNumber="3" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C21FFFC61E8F8BB554CBD18" box="[447,596,1876,1901]" italics="true" pageId="2" pageNumber="3">Dendrodoris</emphasis>
</taxonomicName>
was created by
<bibRefCitation id="EFA8A58D8C21FFFC6349F8BB54E8BD1B" author="Ehrenberg" box="[798,1008,1876,1902]" pageId="2" pageNumber="12" refString="Ehrenberg, C. G. (1831) Symbolae physicae seu icones et descriptiones animalium evertebratorum sepositis insectis quae exitinere per Africam Borealem et Asiam Occidentalem-novae aut illustratae redierunt. Decas 1 Mollusca, [note has no page numbers]." type="book chapter" year="1831">Ehrenberg (1831)</bibRefCitation>
for a dorid from the Red Sea, which was neither adequately described nor dissected, and thus the name was neglected for almost a century (
<bibRefCitation id="EFA8A58D8C20FFFD622DFEAB548BBB2B" author="Gohar" box="[634,915,324,350]" pageId="3" pageNumber="12" refString="Gohar, H. A. F. &amp; Soliman, G. N. (1967) The biology and development of Dendrodoris (= Doridopsis) fumata (Ruppell &amp; Leuckart) (Gastropoda, Nudibranchia). Publications of the Marine Biological Station, Al-Ghardaqa, Red Sea, 14, 31 - 54." type="journal article" year="1967">Gohar &amp; Soliman 1967</bibRefCitation>
). Later,
<bibRefCitation id="EFA8A58D8C20FFFD63A4FEAB5234BB2B" author="Alder" box="[1011,1324,324,350]" pageId="3" pageNumber="12" refString="Alder, J. &amp; Hancock, A. (1864) Notice of a collection of nudibranchiate Mollusca made in India by Walter Elliot, Esq., with description of several new genera and species. Transactions of the Zoological Society of London, 5, 113 - 147." type="journal article" year="1864">Alder and Hancock (1864)</bibRefCitation>
founded an equivalent genus
<taxonomicName id="4C39A3FF8C20FFFD6208FE8355FABBF0" box="[607,738,364,389]" class="Gastropoda" family="Dendrodorididae" genus="Doridopsis" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD6208FE8355FABBF0" box="[607,738,364,389]" italics="true" pageId="3" pageNumber="4">Doridopsis</emphasis>
</taxonomicName>
on Indian Ocean material. Although
<taxonomicName id="4C39A3FF8C20FFFD64C1FE835233BBF0" box="[1174,1323,364,389]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD64C1FE835233BBF0" box="[1174,1323,364,389]" italics="true" pageId="3" pageNumber="4">Dendrodoris</emphasis>
</taxonomicName>
was the first genus name used to collectively describe radula­less dorids, the name
<taxonomicName id="4C39A3FF8C20FFFD615FFE535692BBA0" box="[264,394,444,469]" class="Gastropoda" family="Dendrodorididae" genus="Doridopsis" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD615FFE535692BBA0" box="[264,394,444,469]" italics="true" pageId="3" pageNumber="4">Doridopsis</emphasis>
</taxonomicName>
was often used because its original description was superior.
<bibRefCitation id="EFA8A58D8C20FFFD6400FE535232BBA3" author="Pruvot-Fol" box="[1111,1322,444,470]" pageId="3" pageNumber="13" refString="Pruvot-Fol, A. (1930) Du genre Dendrodoris Ehrenberg et de ses rapports avec le genre Doriopsis Pease et avec quelques autres. Note sur la taxonomie des nudibranches. Bulletin du Museum des National d'Histoire Naturelle Paris, Series 2, 2, 291 - 297." type="journal article" year="1930">Pruvot­Fol (1930)</bibRefCitation>
provided clarification when she showed that
<taxonomicName id="4C39A3FF8C20FFFD6373FE0B54BFBB88" box="[804,935,484,509]" class="Gastropoda" family="Dendrodorididae" genus="Doridopsis" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD6373FE0B54BFBB88" box="[804,935,484,509]" italics="true" pageId="3" pageNumber="4">Doridopsis</emphasis>
</taxonomicName>
was unequivocally synonymous with
<emphasis id="B94D046E8C20FFFD611CFDE356F0B850" box="[331,488,524,549]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="4C39A3FF8C20FFFD611CFDE356FCB850" box="[331,484,524,549]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">Dendrodoris</taxonomicName>
.
</emphasis>
She then advocated the use of the name
<taxonomicName id="4C39A3FF8C20FFFD6453FDE35381B850" box="[1028,1177,524,549]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD6453FDE35381B850" box="[1028,1177,524,549]" italics="true" pageId="3" pageNumber="4">Dendrodoris</emphasis>
</taxonomicName>
(instead of
<taxonomicName id="4C39A3FF8C20FFFD615FFDDB5692B838" box="[264,394,564,589]" class="Gastropoda" family="Dendrodorididae" genus="Doridopsis" higherTaxonomySource="CoL" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD615FFDDB5692B838" box="[264,394,564,589]" italics="true" pageId="3" pageNumber="4">Doridopsis</emphasis>
</taxonomicName>
) because of its chronological priority.
</paragraph>
<paragraph id="8B86D87C8C20FFFD616FFDB355A8B913" blockId="3.[264,1325,284,1350]" pageId="3" pageNumber="4">
The original description of
<taxonomicName id="4C39A3FF8C20FFFD62D4FDB3545EB800" box="[643,838,604,629]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C20FFFD62D4FDB3545EB800" box="[643,838,604,629]" italics="true" pageId="3" pageNumber="4">D. atromaculata</emphasis>
</taxonomicName>
was short, based on preserved material, and contained information about external features only. However, the description was accompanied by excellent drawings that leave no doubt that the material examined in the current study is the same species. Features of particular note in the description, that clearly match it to the current material, are the high convex body, overall colour and markings and the irregular “star­like” surface of the tubercles. The combination of these three features being unique within the Doridoidea.
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C20FFFD616FFC9B56BABF33" pageId="3" pageNumber="4" type="reference_group">
<paragraph id="8B86D87C8C20FFFD616FFC9B523FB9AB" blockId="3.[264,1325,284,1350]" pageId="3" pageNumber="4">
<taxonomicName id="4C39A3FF8C20FFFD616FFC9B556AB9F8" ID-CoL="34Q5P" box="[312,626,884,909]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C20FFFD616FFC9B556AB9F8" box="[312,626,884,909]" italics="true" pageId="3" pageNumber="4">Dendrodoris atromaculata</emphasis>
</taxonomicName>
was redescribed by
<bibRefCitation id="EFA8A58D8C20FFFD6331FC9B531AB9FB" author="Eliot" box="[870,1026,884,910]" pageId="3" pageNumber="12" refString="Eliot, C. (1906 a) On the nudibranchs of southern India and Ceylon, with special reference to the drawings by Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at Newcastle-on-Tyne. Proceedings of the Zoological Society of London, 636 - 691." type="proceedings paper" year="1906" yearSuffix="a">Eliot (1906a)</bibRefCitation>
based on reinvestigation of the preserved
<typeStatus id="548266DE8C20FFFD619DFC7356E5B9C3" box="[458,509,924,950]" pageId="3" pageNumber="4">type</typeStatus>
specimen. In this description, mention was made of the “remarkable” gill, in which the branchial plumes were described as “not very ample”(
<bibRefCitation id="EFA8A58D8C20FFFD6418FC2B5202B9AB" author="Eliot" box="[1103,1306,964,990]" pageId="3" pageNumber="12" refString="Eliot, C. (1906 a) On the nudibranchs of southern India and Ceylon, with special reference to the drawings by Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at Newcastle-on-Tyne. Proceedings of the Zoological Society of London, 636 - 691." type="proceedings paper" year="1906" yearSuffix="a">Eliot 1906a: 662</bibRefCitation>
).
</paragraph>
<paragraph id="8B86D87C8C20FFFD616FFC0356BABF33" blockId="3.[264,1325,284,1350]" pageId="3" pageNumber="4">
<bibRefCitation id="EFA8A58D8C20FFFD616FFC035533BE73" author="Lim" box="[312,555,1004,1030]" pageId="3" pageNumber="13" refString="Lim, C. F. &amp; Chou, L. M. (1970) The dendrodorid and doridid nudibranchs of Singapore. Malayan Nature Journal, 23, 92 - 117." type="journal article" year="1970">Lim &amp; Chou (1970)</bibRefCitation>
depicted, but did not immediately identify, a dendrodorid from
<collectingCountry id="F32E98EC8C20FFFD615FFBFB569BBE5B" box="[264,387,1044,1070]" name="Singapore" pageId="3" pageNumber="4">Singapore</collectingCountry>
. Their figure 9C, is instantly recognisable as
<taxonomicName id="4C39A3FF8C20FFFD63C0FBFB5340BE58" ID-CoL="34Q5P" box="[919,1112,1044,1069]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C20FFFD63C0FBFB5340BE58" box="[919,1112,1044,1069]" italics="true" pageId="3" pageNumber="4">D. atromaculata</emphasis>
</taxonomicName>
and their description (as
<taxonomicName id="4C39A3FF8C20FFFD6131FBD356E3BE20" ID-CoL="7NVPG" box="[358,507,1084,1109]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD6131FBD356E3BE20" box="[358,507,1084,1109]" italics="true" pageId="3" pageNumber="4">Dendrodoris</emphasis>
</taxonomicName>
sp. 2) confirms this with the words “creamy­yellow with round black patches on the mantle” and “ Gills … consisting of 4 quadripinnate branches arranged at right angles to each other … ” (see
<bibRefCitation id="EFA8A58D8C20FFFD62E9FB6354BBBED3" author="Lim" box="[702,931,1164,1190]" pageId="3" pageNumber="13" refString="Lim, C. F. &amp; Chou, L. M. (1970) The dendrodorid and doridid nudibranchs of Singapore. Malayan Nature Journal, 23, 92 - 117." type="journal article" year="1970">Lim &amp; Chou 1970</bibRefCitation>
: 96­97). In an addendum to the manuscript,
<bibRefCitation id="EFA8A58D8C20FFFD61CBFB5B55C8BEBB" author="Lim" box="[412,720,1204,1230]" pageId="3" pageNumber="13" refString="Lim, C. F. &amp; Chou, L. M. (1970) The dendrodorid and doridid nudibranchs of Singapore. Malayan Nature Journal, 23, 92 - 117." type="journal article" year="1970">Lim &amp; Chou (1970: 104)</bibRefCitation>
state that their
<taxonomicName id="4C39A3FF8C20FFFD63D8FB5B533CBEB8" ID-CoL="7NVPG" box="[911,1060,1204,1229]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C20FFFD63D8FB5B533CBEB8" box="[911,1060,1204,1229]" italics="true" pageId="3" pageNumber="4">Dendrodoris</emphasis>
</taxonomicName>
sp. 2 equals
<taxonomicName id="4C39A3FF8C20FFFD649FFB5B56ECBE80" ID-CoL="34Q7F" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="3" pageNumber="4" phylum="Mollusca" rank="species" species="singaporensis" status="sp. nov.">
<emphasis id="B94D046E8C20FFFD649FFB5B56ECBE80" italics="true" pageId="3" pageNumber="4">Dendrodoris singaporensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27EB9158C20FFFD6256FB33557DBE83" box="[513,613,1244,1270]" pageId="3" pageNumber="4" rank="species">sp. nov.</taxonomicNameLabel>
, and that the
<typeStatus id="548266DE8C20FFFD634EFB335453BE83" box="[793,843,1244,1270]" pageId="3" pageNumber="4">type</typeStatus>
material was lodged in the
<collectingCountry id="F32E98EC8C20FFFD64E3FB335233BE83" box="[1204,1323,1244,1270]" name="Singapore" pageId="3" pageNumber="4">Singapore</collectingCountry>
National Museum. Unfortunately enquires at this institution did not result in location of this material.
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C20FFFD615FFA9353FEBF93" pageId="3" pageNumber="4" type="materials_examined">
<paragraph id="8B86D87C8C20FFFD615FFA9356F3BFE0" blockId="3.[264,1325,1404,1870]" box="[264,491,1404,1429]" pageId="3" pageNumber="4">
<emphasis id="B94D046E8C20FFFD615FFA9356F3BFE0" box="[264,491,1404,1429]" italics="true" pageId="3" pageNumber="4">Material Examined</emphasis>
</paragraph>
<paragraph id="8B86D87C8C20FFFD616FFA4B53FEBF93" blockId="3.[264,1325,1404,1870]" pageId="3" pageNumber="4">
<emphasis id="B94D046E8C20FFFD616FFA4B569CBFCB" bold="true" box="[312,388,1444,1470]" pageId="3" pageNumber="4">
<collectingCountry id="F32E98EC8C20FFFD616FFA4B5667BFCB" box="[312,383,1444,1470]" name="India" pageId="3" pageNumber="4">India</collectingCountry>
:
</emphasis>
<typeStatus id="548266DE8C20FFFD61DBFA4B553EBFCB" box="[396,550,1444,1470]" pageId="3" pageNumber="4" type="holotype">HOLOTYPE</typeStatus>
Madras, Waltair;
<specimenCount id="9D3F13F58C20FFFD62ACFA4B5498BFCB" box="[763,896,1444,1470]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD63C7FA4B5309BFCB" box="[912,1041,1444,1470]" metricMagnitude="-2" metricUnit="m" metricValue="3.75" metricValueMax="4.0" metricValueMin="3.5" pageId="3" pageNumber="4" unit="mm" value="37.5" valueMax="40.0" valueMin="35.0">35–40 mm</quantity>
living length, unknown date in 1853­4, coll. Sir Walter Elliot, (
<collectionCode id="ED2840B98C20FFFD6286FA235451BF93" box="[721,841,1484,1510]" httpUri="http://grbio.org/cool/4g9v-dppp" name="Hancock Museum" pageId="3" pageNumber="4">NEWHM</collectionCode>
: 1998. H4379), probably intertidal.
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C20FFFD616FFA1B5586BD3B" pageId="3" pageNumber="4" type="description">
<paragraph id="8B86D87C8C20FFFD616FFA1B5403BC8B" blockId="3.[264,1325,1404,1870]" pageId="3" pageNumber="4">
<emphasis id="B94D046E8C20FFFD616FFA1B5544BC7B" bold="true" box="[312,604,1524,1550]" pageId="3" pageNumber="4">
<collectingCountry id="F32E98EC8C20FFFD616FFA1B56B7BC7B" box="[312,431,1524,1550]" name="Australia" pageId="3" pageNumber="4">Australia</collectingCountry>
, Queensland:
</emphasis>
Gulf of Carpentaria,
<quantity id="4CC175998C20FFFD6337FA1B54B4BC7B" box="[864,940,1524,1550]" metricMagnitude="4" metricUnit="m" metricValue="8.0" pageId="3" pageNumber="4" unit="km" value="80.0">80 km</quantity>
west of Weipa;
<specimenCount id="9D3F13F58C20FFFD6425FA1B53E1BC7B" box="[1138,1273,1524,1550]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD655CFA1B562EBC43" metricMagnitude="-2" metricUnit="m" metricValue="4.2" pageId="3" pageNumber="4" unit="mm" value="42.0">42 mm</quantity>
preserved length), unknown date, coll. M. Gormon &amp; C. Lu, (NMV Reg. No.
<accessionNumber id="946A459F8C20FFFD6490F9F3523ABC43" box="[1223,1314,1564,1590]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/F86791" pageId="3" pageNumber="4" type="EnaNcbi">F86791</accessionNumber>
) depth
<quantity id="4CC175998C20FFFD6107F9AB5697BC2B" box="[336,399,1604,1630]" metricMagnitude="1" metricUnit="m" metricValue="4.2" pageId="3" pageNumber="4" unit="m" value="42.0">42 m</quantity>
; Rowes Bay, Townsville;
<specimenCount id="9D3F13F58C20FFFD62E8F9AB545CBC2B" box="[703,836,1604,1630]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD6304F9AB54BFBC2B" box="[851,935,1604,1630]" metricMagnitude="-2" metricUnit="m" metricValue="6.7" pageId="3" pageNumber="4" unit="mm" value="67.0">67 mm</quantity>
long, live),
<date id="FF87FEBC8C20FFFD6462F9AB53BBBC2B" box="[1077,1187,1604,1630]" pageId="3" pageNumber="4">8 August</date>
, 1998, coll. G. Brodie (BC, J1);
<specimenCount id="9D3F13F58C20FFFD61A3F9835561BCF3" box="[500,633,1644,1670]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD62DEF98355C5BCF3" box="[649,733,1644,1670]" metricMagnitude="-2" metricUnit="m" metricValue="8.5" pageId="3" pageNumber="4" unit="mm" value="85.0">85 mm</quantity>
long, live),
<date id="FF87FEBC8C20FFFD633AF98354F1BCF3" box="[877,1001,1644,1670]" pageId="3" pageNumber="4" value="1927-08">27 August</date>
, 2000, coll. J. Lariscy, (BC Z1);
<specimenCount id="9D3F13F58C20FFFD6115F97B56DFBCDB" box="[322,455,1684,1710]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD6180F97B5533BCDB" box="[471,555,1684,1710]" metricMagnitude="-2" metricUnit="m" metricValue="5.4" pageId="3" pageNumber="4" unit="mm" value="54.0">54 mm</quantity>
preserved length),
<date id="FF87FEBC8C20FFFD6359F97B54AABCDB" box="[782,946,1684,1710]" pageId="3" pageNumber="4" value="1916-09">16 September</date>
, 2001, coll. F. Chouw (BC K2); Myora Spit, North Stradbroke Island;
<specimenCount id="9D3F13F58C20FFFD6295F953545EBCA3" box="[706,838,1724,1750]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD6302F95354B0BCA3" box="[853,936,1724,1750]" metricMagnitude="-2" metricUnit="m" metricValue="8.5" pageId="3" pageNumber="4" unit="mm" value="85.0">85 mm</quantity>
long, live),
<date id="FF87FEBC8C20FFFD6463F95353F3BCA3" box="[1076,1259,1724,1750]" pageId="3" pageNumber="4" value="1983-10-04">4 October 1983</date>
, coll. R. Willan &amp; O. Kelly (NTM Reg. No. P625).
</paragraph>
<paragraph id="8B86D87C8C20FFFD616FF8E35586BD3B" blockId="3.[264,1325,1404,1870]" pageId="3" pageNumber="4">
<emphasis id="B94D046E8C20FFFD616FF8E35675BD53" bold="true" box="[312,365,1804,1830]" pageId="3" pageNumber="4">
<collectingCountry id="F32E98EC8C20FFFD616FF8E35670BD53" box="[312,360,1804,1830]" name="Fiji" pageId="3" pageNumber="4">Fiji</collectingCountry>
:
</emphasis>
<collectingCountry id="F32E98EC8C20FFFD6123F8E356B8BD53" box="[372,416,1804,1830]" name="Fiji" pageId="3" pageNumber="4">Viti</collectingCountry>
Levu, Nasese, Suva,
<specimenCount id="9D3F13F58C20FFFD62CAF8E3543ABD53" box="[669,802,1804,1830]" pageId="3" pageNumber="4" type="generic">1 specimen</specimenCount>
(
<quantity id="4CC175998C20FFFD6366F8E3549DBD53" box="[817,901,1804,1830]" metricMagnitude="-2" metricUnit="m" metricValue="3.7" pageId="3" pageNumber="4" unit="mm" value="37.0">37 mm</quantity>
long live),
<date id="FF87FEBC8C20FFFD645CF8E353F5BD53" box="[1035,1261,1804,1830]" pageId="3" pageNumber="4" value="1987-12-26">26 December 1987</date>
, coll. G. &amp; J. Brodie, BC D1 (intertidal).
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C27FFF7615FFEF3553DB8B0" lastPageId="9" lastPageNumber="10" pageId="4" pageNumber="5" type="discussion">
<paragraph id="8B86D87C8C27FFFA615FFEF35527BB40" blockId="4.[264,1323,284,790]" box="[264,575,284,309]" pageId="4" pageNumber="5">
<emphasis id="B94D046E8C27FFFA615FFEF35527BB40" box="[264,575,284,309]" italics="true" pageId="4" pageNumber="5">
Remarks on
<typeStatus id="548266DE8C27FFFA61CDFEF356C9BB40" box="[410,465,284,309]" pageId="4" pageNumber="5">Type</typeStatus>
Material
</emphasis>
</paragraph>
<paragraph id="8B86D87C8C27FFFA616FFEAB53E6B963" blockId="4.[264,1323,284,790]" pageId="4" pageNumber="5">
As stated by
<bibRefCitation id="EFA8A58D8C27FFFA6180FEAB5560BB2B" author="Eliot" box="[471,632,324,350]" pageId="4" pageNumber="12" refString="Eliot, C. (1906 a) On the nudibranchs of southern India and Ceylon, with special reference to the drawings by Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at Newcastle-on-Tyne. Proceedings of the Zoological Society of London, 636 - 691." type="proceedings paper" year="1906" yearSuffix="a">Eliot (1906a)</bibRefCitation>
, the
<typeStatus id="548266DE8C27FFFA62E3FEAB5402BB2B" box="[692,794,324,350]" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
described by
<bibRefCitation id="EFA8A58D8C27FFFA6391FEAB53EBBB2B" author="Alder" box="[966,1267,324,350]" pageId="4" pageNumber="12" refString="Alder, J. &amp; Hancock, A. (1864) Notice of a collection of nudibranchiate Mollusca made in India by Walter Elliot, Esq., with description of several new genera and species. Transactions of the Zoological Society of London, 5, 113 - 147." type="journal article" year="1864">Alder &amp; Hancock (1864)</bibRefCitation>
was originally collected by Walter Elliot in 1853­4. This specimen is lodged in the Hancock Museum at Newcastle­on­Tyne and registered under the system described by
<bibRefCitation id="EFA8A58D8C27FFFA64A5FE7B5671BBA3" author="Eliot" pageId="4" pageNumber="12" refString="Eliot, C. (1906 b) On the nudibranchs of southern India and Ceylon, with special reference to the drawings by Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at Newcastle-on-Tyne. No. 2, Proceedings of the Zoological Society of London, 999 - 1008." type="book chapter" year="1906" yearSuffix="b">Eliot (1906b)</bibRefCitation>
. When the material was re­examined by
<bibRefCitation id="EFA8A58D8C27FFFA63D0FE535374BBA3" author="Eliot" box="[903,1132,444,470]" pageId="4" pageNumber="12" refString="Eliot, C. (1906 a) On the nudibranchs of southern India and Ceylon, with special reference to the drawings by Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at Newcastle-on-Tyne. Proceedings of the Zoological Society of London, 636 - 691." type="proceedings paper" year="1906" yearSuffix="a">Eliot (1906a: 662)</bibRefCitation>
he stated that, although many of the Alder &amp; Hancock specimens were in very poor condition, the
<typeStatus id="548266DE8C27FFFA64BAFE0B5622B853" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
of
<taxonomicName id="4C39A3FF8C27FFFA6134FDE3553DB850" box="[355,549,524,549]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="4" pageNumber="5" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C27FFFA6134FDE3553DB850" box="[355,549,524,549]" italics="true" pageId="4" pageNumber="5">D. atromaculata</emphasis>
</taxonomicName>
was “rather well preserved.” This
<typeStatus id="548266DE8C27FFFA639DFDE35329B853" box="[970,1073,524,550]" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
was examined in the current study. The specimen is indeed well preserved and the characteristic black notal markings can be clearly seen despite 150 years of preservation. The specimen has previously been dissected by an anterior dorsal incision. The internal organs are intact and appear in reasonable condition. Although the
<typeStatus id="548266DE8C27FFFA636DFD4354B9B8B3" box="[826,929,684,710]" pageId="4" pageNumber="5" type="holotype">holotype</typeStatus>
is small (only
<quantity id="4CC175998C27FFFA640DFD4353B7B8B3" box="[1114,1199,684,710]" metricMagnitude="-2" metricUnit="m" metricValue="1.5" pageId="4" pageNumber="5" unit="mm" value="15.0">15 mm</quantity>
preserved length) compared with the material examined in the current study, I have no doubt that they belong to the same species because of the distinct external features and markings.
</paragraph>
<paragraph id="8B86D87C8C27FFFA615FFCA356B2B910" blockId="4.[264,1325,844,1870]" box="[264,426,844,869]" pageId="4" pageNumber="5">
<emphasis id="B94D046E8C27FFFA615FFCA356B2B910" box="[264,426,844,869]" italics="true" pageId="4" pageNumber="5">Redescription</emphasis>
</paragraph>
<paragraph id="8B86D87C8C27FFFA616FFC9B54F1BE73" blockId="4.[264,1325,844,1870]" pageId="4" pageNumber="5">
The following description is based on the examination of four live specimens [BC ­ D1, J1, Z1 &amp; K2], preserved material (including three specimens dissected and selectively sectioned [BC – J1, Z1 &amp; K2] and two additional specimens [NTM P625, NMV
<accessionNumber id="946A459F8C27FFFA649EFC2B523EB9AB" box="[1225,1318,964,990]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/F86791" pageId="4" pageNumber="5" type="EnaNcbi">F86791</accessionNumber>
] used to supplement the histological investigation of the notum.
</paragraph>
<paragraph id="8B86D87C8C27FFFA616FFBFB5399BF1B" blockId="4.[264,1325,844,1870]" pageId="4" pageNumber="5">
<taxonomicName id="4C39A3FF8C27FFFA616FFBFB556ABE58" box="[312,626,1044,1069]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="4" pageNumber="5" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C27FFFA616FFBFB556ABE58" box="[312,626,1044,1069]" italics="true" pageId="4" pageNumber="5">Dendrodoris atromaculata</emphasis>
</taxonomicName>
is tuberculate, possesses a relatively high body profile and is large and broad in comparison to several other members of the genus that are relatively smooth and elongate e.g.
<taxonomicName id="4C39A3FF8C27FFFA6262FB8B5468BE0B" authority="Stimpson, 1855" authorityName="Stimpson" authorityYear="1855" box="[565,880,1124,1150]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="4" pageNumber="5" phylum="Mollusca" rank="species" species="nigra">
<emphasis id="B94D046E8C27FFFA6262FB8B5581BE08" box="[565,665,1124,1149]" italics="true" pageId="4" pageNumber="5">D. nigra</emphasis>
(Stimpson, 1855)
</taxonomicName>
. The extended crawling length of living adults is up to
<quantity id="4CC175998C27FFFA61B1FB635525BED3" box="[486,573,1164,1190]" metricMagnitude="-2" metricUnit="m" metricValue="8.5" pageId="4" pageNumber="5" unit="mm" value="85.0">85 mm</quantity>
. The body is soft but firm. The dorsum is covered in irregular “star­like” compound tubercules, each consisting of a cluster of finger­like papillae of different heights that taper to rounded apices (
<figureCitation id="1302C4F98C27FFFA62A8FB335463BE83" box="[767,891,1244,1270]" captionStart="FIGURE 2 A – D. 0 0" captionStartId="7.[264,368,1577,1601]" captionTargetBox="[323,1280,313,1555]" captionTargetId="figure@7.[298,1299,300,1578]" captionTargetPageId="7" captionText="FIGURE 2 A – D. 0 0 Composite drawings of Dendrodoris atromaculata (preserved material) showing: A. compound dorsal tubercule. B. anterior part of the digestive system. C. posterior part of the digestive system. D. arrangement of the post­gonadial reproductive system. amp = ampulla, an = anus, bc = bursa copulatrix, bw = body wall, es = exit to stomach, hd = hermaphroditic duct, int = intestine, mb = muscular bulb, mo = mouth, ng = nidamental gland, np = notal papilla, oes = oesophagus, ot = oral tube, ov = oviduct, pg = prostate gland, ph = pharynx, pb = pyloric bulb (not present in all individuals), ptd = ptyaline duct, ptg = ptyaline gland, rm = retractor muscle, rs = receptaculum seminis, sg = salivary gland, va = vagina, vd = vas deferens, vg = vestibular gland. Scale bars = 2 mm." httpUri="https://zenodo.org/record/157390/files/figure.png" pageId="4" pageNumber="5">Figure 2A</figureCitation>
). The apex of these subsidiary papillae is often pinkish in formalin­preserved material and the central papilla is generally larger than the rest. A microscopic inspection of the remainder of the notum, between the tubercules and along the mantle edge, reveals a covering of simple papillae.
</paragraph>
<paragraph id="8B86D87C8C27FFFA616FFA935453BCF3" blockId="4.[264,1325,844,1870]" pageId="4" pageNumber="5">
The notum is pale yellow, sometimes darkening to orange centrally. In the Fijian specimen, irregular ginger­brown patches were present, primarily around the notal margin (
<figureCitation id="1302C4F98C27FFFA6147FA235693BF93" box="[272,395,1484,1510]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="4" pageNumber="5">Figure 1B</figureCitation>
). These were not observed in the Australian material but replaced by a more even spread of large, irregularly­shaped black patches (
<figureCitation id="1302C4F98C27FFFA63ECFA1B5325BC7B" box="[955,1085,1524,1550]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="4" pageNumber="5">Figure 1A</figureCitation>
). When these dark patches occurred close to a compound tubercle the pale apices of the papilla became even more obvious (because of the dark background), thus giving a distorted impression of an uneven distribution of tubercules over the notum.
</paragraph>
<paragraph id="8B86D87C8C27FFFA616FF97B53B2BD53" blockId="4.[264,1325,844,1870]" pageId="4" pageNumber="5">In one specimen examined from Rowes Bay, small black spots were scattered across the ventral underside of the pale orange notum, but these markings were absent in the remaining specimens. Some dark markings from the dorsal surface were visible ventrally in some preserved material because of the translucent nature of the body tissue.</paragraph>
<paragraph id="8B86D87C8C27FFFB616FF8DB5649BBF3" blockId="4.[264,1325,844,1870]" lastBlockId="5.[264,1324,284,910]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">The fully retractable rhinophores are small and relatively inconspicuous for the size of the animal. The clavus is orange, diagonally lamellate, and quite long proportionally. The stalk is translucent and there is no change in width or angle where the claval lamellae begin.</paragraph>
<paragraph id="8B86D87C8C26FFFB616FFE7B545EB8B3" blockId="5.[264,1324,284,910]" pageId="5" pageNumber="6">
In all of the material examined, the cream and grey gill consists of four, large branchiae. When the living animal is at rest, these branchiae form a large equilateral cross over the rear of the body (see
<figureCitation id="1302C4F98C26FFFB6270FE0B55B8BB8B" box="[551,672,484,510]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="5" pageNumber="6">Figure 1A</figureCitation>
). The edge of the gill pocket is simple, without a raised rim. The anal papilla is not large or prominent and there is no sign of a gap between the posterior gill branchiae as seen in some other
<taxonomicName id="4C39A3FF8C26FFFB636FFDDB54D5B838" box="[824,973,564,589]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="5" pageNumber="6" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C26FFFB636FFDDB54D5B838" box="[824,973,564,589]" italics="true" pageId="5" pageNumber="6">Dendrodoris</emphasis>
</taxonomicName>
species. Internally, large gill retractor muscles are found at the gill base and, as documented for
<taxonomicName id="4C39A3FF8C26FFFB6472FDB35391B800" box="[1061,1161,604,629]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="5" pageNumber="6" phylum="Mollusca" rank="species" species="nigra">
<emphasis id="B94D046E8C26FFFB6472FDB35391B800" box="[1061,1161,604,629]" italics="true" pageId="5" pageNumber="6">D. nigra</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFA8A58D8C26FFFB6486FDB35691B8EB" author="Wagele" pageId="5" pageNumber="13" refString="Wagele, H., Brodie, G. &amp; Klussmann-Kolb, A. (1999) Histological investigations of Dendrodoris nigra (Gastropoda, Nudibranchia, Dendrodorididae). Molluscan Research, 20, 79 - 94." type="journal article" year="1999">
Wägele
<emphasis id="B94D046E8C26FFFB615FFD6B5624B8E8" box="[264,316,644,669]" italics="true" pageId="5" pageNumber="6">et al</emphasis>
. 1999
</bibRefCitation>
), these extend down between the posterior lobes of digestive gland before running along the ventral surface of the body cavity.
</paragraph>
<paragraph id="8B86D87C8C26FFFB616FFD3B562EB9FB" blockId="5.[264,1324,284,910]" pageId="5" pageNumber="6">
The notum overlaps the vivid orange foot by a considerable margin. However, the foot itself is large and quite wide (
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wide in
<quantity id="4CC175998C26FFFB6370FD135463B963" box="[807,891,764,790]" metricMagnitude="-2" metricUnit="m" metricValue="3.7" pageId="5" pageNumber="6" unit="mm" value="37.0">37 mm</quantity>
long animal), sometimes with small brown spots on the dorsal surface. The mouth is ventral, within a notch in the antero­ventral surface of the foot. No oral tentacles were observed in either live or preserved material.
</paragraph>
<paragraph id="8B86D87C8C26FFFB615FFC2B56F9B9A8" blockId="5.[264,1323,964,1590]" box="[264,481,964,989]" pageId="5" pageNumber="6">
<emphasis id="B94D046E8C26FFFB615FFC2B56F9B9A8" box="[264,481,964,989]" italics="true" pageId="5" pageNumber="6">General Histology</emphasis>
</paragraph>
<paragraph id="8B86D87C8C26FFFB616FFC035507BF93" blockId="5.[264,1323,964,1590]" pageId="5" pageNumber="6">
The large size of specimens precluded serial sectioning of the whole animal. However, sections of the notal tissue were examined from three specimens (two collected fresh at Rowes Bay one year apart, and another from North Stradbroke Island). Bundles of large (~0.3 x 0.04 mm), elongate spicules (
<figureCitation id="1302C4F98C26FFFB6293FB8B5425BE0B" box="[708,829,1124,1150]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="5" pageNumber="6">Figure 1C</figureCitation>
) were located in the notal connective tissue. The notal tissue of all three specimens also contained unusual, microscopic, darkstaining structures forming thin, irregular lines (
<figureCitation id="1302C4F98C26FFFB6317FB5B54A6BEBB" box="[832,958,1204,1230]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="5" pageNumber="6">Figure 1D</figureCitation>
). At first these structures were thought to be fungi or bacteria. However, further analysis (using wax histology) gave a negative result for fungi (PAS stain, see methods section) and an inconclusive result for bacteria (Gram stain). In light of these results, I believe these structures to be black pigment within the connective tissue. However, this arrangement of pigment is quite different (very wide­spread throughout the tissue rather than concentrated along the notal epithelial boundary) to that seen in other members of
<taxonomicName id="4C39A3FF8C26FFFB6352FA4B5482BFC8" box="[773,922,1444,1469]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="5" pageNumber="6" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C26FFFB6352FA4B5482BFC8" box="[773,922,1444,1469]" italics="true" pageId="5" pageNumber="6">Dendrodoris</emphasis>
</taxonomicName>
(e.g. see
<taxonomicName id="4C39A3FF8C26FFFB645FFA4B5373BFC8" box="[1032,1131,1444,1469]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="5" pageNumber="6" phylum="Mollusca" rank="species" species="nigra">
<emphasis id="B94D046E8C26FFFB645FFA4B5373BFC8" box="[1032,1131,1444,1469]" italics="true" pageId="5" pageNumber="6">D. nigra</emphasis>
</taxonomicName>
in
<bibRefCitation id="EFA8A58D8C26FFFB64C7FA4B565FBF93" author="Wagele" pageId="5" pageNumber="13" refString="Wagele, H., Brodie, G. &amp; Klussmann-Kolb, A. (1999) Histological investigations of Dendrodoris nigra (Gastropoda, Nudibranchia, Dendrodorididae). Molluscan Research, 20, 79 - 94." type="journal article" year="1999">
Wägele
<emphasis id="B94D046E8C26FFFB64A7FA4B523CBFC8" box="[1264,1316,1444,1469]" italics="true" pageId="5" pageNumber="6">et al</emphasis>
. 1999
</bibRefCitation>
:
<figureCitation id="1302C4F98C26FFFB6105FA235509BF93" box="[338,529,1484,1510]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="5" pageNumber="6">Figures 1A &amp; B</figureCitation>
).
</paragraph>
<paragraph id="8B86D87C8C26FFFB616FFA1B53CDBC43" blockId="5.[264,1323,964,1590]" pageId="5" pageNumber="6">
Ciliated cells were not detected along the notal epithelium, nor were copiously vacuolated epithelial cells, as defined by (
<bibRefCitation id="EFA8A58D8C26FFFB62EDF9F35443BC43" author="Wagele" box="[698,859,1564,1590]" pageId="5" pageNumber="13" refString="Wagele, H. (1998) Histological investigation of some organs and specialised cellular structures in Opisthobranchia (Gastropoda) with the potential to yield phylogenetically significant characters. Zoologischer Anzeiger, 236, 119 - 131" type="journal article" year="1998">Wägele 1998</bibRefCitation>
), visible in the notal epithelium.
</paragraph>
<caption id="DF4688F48C26FFFB615FF9845232BD48" pageId="5" pageNumber="6">
<paragraph id="8B86D87C8C26FFFB615FF9845232BD48" blockId="5.[264,1323,1643,1893]" pageId="5" pageNumber="6">
<emphasis id="B94D046E8C26FFFB615FF98456A1BCF6" bold="true" box="[264,441,1643,1667]" pageId="5" pageNumber="6">FIGURE 1A–F.</emphasis>
Photographs of
<taxonomicName id="4C39A3FF8C26FFFB623CF98354E2BCF6" authority="A. Living" authorityName="A. Living" box="[619,1018,1643,1667]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="5" pageNumber="6" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C26FFFB623CF9835492BCF6" box="[619,906,1644,1667]" italics="true" pageId="5" pageNumber="6">Dendrodoris atromaculata</emphasis>
<emphasis id="B94D046E8C26FFFB63C6F98454B2BCF6" bold="true" box="[913,938,1643,1667]" pageId="5" pageNumber="6">A.</emphasis>
Living
</taxonomicName>
animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie.
<emphasis id="B94D046E8C26FFFB642BF97E538CBCDC" bold="true" box="[1148,1172,1681,1705]" pageId="5" pageNumber="6">B.</emphasis>
living animal (37 mm live length) found intertidally at Nasese, Fiji.
<emphasis id="B94D046E8C26FFFB633AF959549FBCBB" bold="true" box="[877,903,1718,1742]" pageId="5" pageNumber="6">C.</emphasis>
histological section showing spicules located in the notal tissue.
<emphasis id="B94D046E8C26FFFB6278F9345550BC86" bold="true" box="[559,584,1755,1779]" pageId="5" pageNumber="6">D.</emphasis>
histological section showing the unusual distribution of pigment in the notal tissue.
<emphasis id="B94D046E8C26FFFB6190F8EE56C7BD6C" bold="true" box="[455,479,1793,1817]" pageId="5" pageNumber="6">E.</emphasis>
confocal microscope scan of a semi­everted penis.
<emphasis id="B94D046E8C26FFFB6462F8EE535FBD6C" bold="true" box="[1077,1095,1793,1817]" pageId="5" pageNumber="6">F.</emphasis>
histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =
</paragraph>
</caption>
<paragraph id="8B86D87C8C26FFFB615FF8A154AABD10" blockId="5.[264,1323,1643,1893]" box="[264,946,1870,1893]" pageId="5" pageNumber="6">notal epithelium, pg = pigment, ps = penial spines, sp = spicule.</paragraph>
<caption id="DF4688F48C24FFF9615FF9C656CEBD20" httpUri="https://zenodo.org/record/157390/files/figure.png" pageId="7" pageNumber="8" targetBox="[323,1280,313,1555]" targetPageId="7">
<paragraph id="8B86D87C8C24FFF9615FF9C656CEBD20" blockId="7.[264,1323,1577,1877]" pageId="7" pageNumber="8">
<emphasis id="B94D046E8C24FFF9615FF9C656DFBC34" bold="true" box="[264,455,1577,1601]" pageId="7" pageNumber="8">FIGURE 2 A–D.</emphasis>
0 0Composite drawings of
<taxonomicName id="4C39A3FF8C24FFF962B3F9C6531ABC35" box="[740,1026,1577,1600]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="7" pageNumber="8" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C24FFF962B3F9C6531ABC35" box="[740,1026,1577,1600]" italics="true" pageId="7" pageNumber="8">Dendrodoris atromaculata</emphasis>
</taxonomicName>
(preserved material) showing:
<emphasis id="B94D046E8C24FFF9616FF9A4564ABC16" bold="true" box="[312,338,1611,1635]" pageId="7" pageNumber="8">A.</emphasis>
compound dorsal tubercule.
<emphasis id="B94D046E8C24FFF962DEF9A455B9BC16" bold="true" box="[649,673,1611,1635]" pageId="7" pageNumber="8">B.</emphasis>
anterior part of the digestive system.
<emphasis id="B94D046E8C24FFF96461F9A45348BC16" bold="true" box="[1078,1104,1611,1635]" pageId="7" pageNumber="8">C.</emphasis>
posterior part of the digestive system.
<emphasis id="B94D046E8C24FFF9619CF98156FCBCF3" bold="true" box="[459,484,1646,1670]" pageId="7" pageNumber="8">D.</emphasis>
arrangement of the post­gonadial reproductive system. amp = ampulla, an = anus, bc = bursa copulatrix, bw = body wall, es = exit to stomach, hd = hermaphroditic duct, int = intestine, mb = muscular bulb, mo = mouth, ng = nidamental gland, np = notal papilla, oes = oesophagus, ot = oral tube, ov = oviduct, pg = prostate gland, ph = pharynx, pb = pyloric bulb (not present in all individuals), ptd = ptyaline duct, ptg = ptyaline gland, rm = retractor muscle, rs = receptaculum seminis, sg = salivary gland, va = vagina, vd = vas deferens, vg = vestibular gland. Scale bars = 2 mm.
</paragraph>
</caption>
<paragraph id="8B86D87C8C2BFFF6615FFEF356D7BB40" blockId="8.[264,1324,284,1030]" box="[264,463,284,309]" pageId="8" pageNumber="9">
<emphasis id="B94D046E8C2BFFF6615FFEF356D7BB40" box="[264,463,284,309]" italics="true" pageId="8" pageNumber="9">Digestive System</emphasis>
</paragraph>
<paragraph id="8B86D87C8C2BFFF6616FFEAB53C6B89B" blockId="8.[264,1324,284,1030]" pageId="8" pageNumber="9">
A diagram of the anterior part of the digestive system is given in
<figureCitation id="1302C4F98C2BFFF66478FEAB53B0BB2B" box="[1071,1192,324,350]" captionStart="FIGURE 2 A – D. 0 0" captionStartId="7.[264,368,1577,1601]" captionTargetBox="[323,1280,313,1555]" captionTargetId="figure@7.[298,1299,300,1578]" captionTargetPageId="7" captionText="FIGURE 2 A – D. 0 0 Composite drawings of Dendrodoris atromaculata (preserved material) showing: A. compound dorsal tubercule. B. anterior part of the digestive system. C. posterior part of the digestive system. D. arrangement of the post­gonadial reproductive system. amp = ampulla, an = anus, bc = bursa copulatrix, bw = body wall, es = exit to stomach, hd = hermaphroditic duct, int = intestine, mb = muscular bulb, mo = mouth, ng = nidamental gland, np = notal papilla, oes = oesophagus, ot = oral tube, ov = oviduct, pg = prostate gland, ph = pharynx, pb = pyloric bulb (not present in all individuals), ptd = ptyaline duct, ptg = ptyaline gland, rm = retractor muscle, rs = receptaculum seminis, sg = salivary gland, va = vagina, vd = vas deferens, vg = vestibular gland. Scale bars = 2 mm." httpUri="https://zenodo.org/record/157390/files/figure.png" pageId="8" pageNumber="9">Figure 2B</figureCitation>
. The ptyaline gland is not distinctly bilobed and the ptyaline duct is relatively long and thin, becoming very narrow distally. Distally, the ptyaline duct and the oral tube are both surrounded by a muscular bulb. The smooth pharynx is relatively long and a pair of rounded, compact salivary glands is present, at the junction of the pharynx and oesophagus, one on either side. The salivary glands are difficult to find in preserved material because their coloration is very similar to the pharynx on which they sit. The oesophagus is broad and bumpy in external appearance, particularly distally. It curves around on itself several times before entering the stomach within the pale brown digestive gland. As is typical of the genus, the stomach is indistinct and totally embedded within the digestive gland. The digestive gland is by far the largest organ in the body cavity, and very distinctly bilobed posteriorly.
</paragraph>
<paragraph id="8B86D87C8C2BFFF6616FFD13558DBE73" blockId="8.[264,1324,284,1030]" pageId="8" pageNumber="9">
The posterior part of the digestive system is simple with the intestine arising from the embedded stomach to emerge from the posterior half of the digestive gland. In only one of the three animals dissected, a pyloric bulb with several folds could be seen proximally (
<figureCitation id="1302C4F98C2BFFF66147FC9B5694B9FB" box="[272,396,884,910]" captionStart="FIGURE 2 A – D. 0 0" captionStartId="7.[264,368,1577,1601]" captionTargetBox="[323,1280,313,1555]" captionTargetId="figure@7.[298,1299,300,1578]" captionTargetPageId="7" captionText="FIGURE 2 A – D. 0 0 Composite drawings of Dendrodoris atromaculata (preserved material) showing: A. compound dorsal tubercule. B. anterior part of the digestive system. C. posterior part of the digestive system. D. arrangement of the post­gonadial reproductive system. amp = ampulla, an = anus, bc = bursa copulatrix, bw = body wall, es = exit to stomach, hd = hermaphroditic duct, int = intestine, mb = muscular bulb, mo = mouth, ng = nidamental gland, np = notal papilla, oes = oesophagus, ot = oral tube, ov = oviduct, pg = prostate gland, ph = pharynx, pb = pyloric bulb (not present in all individuals), ptd = ptyaline duct, ptg = ptyaline gland, rm = retractor muscle, rs = receptaculum seminis, sg = salivary gland, va = vagina, vd = vas deferens, vg = vestibular gland. Scale bars = 2 mm." httpUri="https://zenodo.org/record/157390/files/figure.png" pageId="8" pageNumber="9">Figure 2C</figureCitation>
). No bulb or folds were seen in the two other specimens. The intestine is narrow, dorso­ventrally flattened and thin walled, it extends over the digestive gland before terminating at the anus. The anal papilla is not swollen and lying among the large gill branchiae remains inconspicuous.
</paragraph>
<paragraph id="8B86D87C8C2BFFF6615FFBD356E3BE20" blockId="8.[264,1324,1084,1430]" box="[264,507,1084,1109]" pageId="8" pageNumber="9">
<emphasis id="B94D046E8C2BFFF6615FFBD356E3BE20" box="[264,507,1084,1109]" italics="true" pageId="8" pageNumber="9">Reproductive System</emphasis>
</paragraph>
<paragraph id="8B86D87C8C2BFFF6616FFB8B536EBFE3" blockId="8.[264,1324,1084,1430]" pageId="8" pageNumber="9">
The vivid orange gonad was well developed in all three specimens dissected and covers most of the digestive gland. The post­gonadial reproductive system is shown in
<figureCitation id="1302C4F98C2BFFF66489FB635628BEBB" captionStart="FIGURE 2 A – D. 0 0" captionStartId="7.[264,368,1577,1601]" captionTargetBox="[323,1280,313,1555]" captionTargetId="figure@7.[298,1299,300,1578]" captionTargetPageId="7" captionText="FIGURE 2 A – D. 0 0 Composite drawings of Dendrodoris atromaculata (preserved material) showing: A. compound dorsal tubercule. B. anterior part of the digestive system. C. posterior part of the digestive system. D. arrangement of the post­gonadial reproductive system. amp = ampulla, an = anus, bc = bursa copulatrix, bw = body wall, es = exit to stomach, hd = hermaphroditic duct, int = intestine, mb = muscular bulb, mo = mouth, ng = nidamental gland, np = notal papilla, oes = oesophagus, ot = oral tube, ov = oviduct, pg = prostate gland, ph = pharynx, pb = pyloric bulb (not present in all individuals), ptd = ptyaline duct, ptg = ptyaline gland, rm = retractor muscle, rs = receptaculum seminis, sg = salivary gland, va = vagina, vd = vas deferens, vg = vestibular gland. Scale bars = 2 mm." httpUri="https://zenodo.org/record/157390/files/figure.png" pageId="8" pageNumber="9">Figure 2D</figureCitation>
. The system is triaulic with the rounded bursa copulatrix (= gametolytic gland) and elongate receptaculum seminis leading from the vaginal duct. The ampulla is simple and pyriform. The prostate gland is tubular and uniform in appearance. A rounded vestibular gland (~
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in diameter) is present within the body cavity and attached close to the base of the distal oviduct. The vas deferens is smooth and uniform in width; while the penis is armed with numerous rows of closely packed, elongate spines (
<figureCitation id="1302C4F98C2BFFF663B8FA935370BFE3" box="[1007,1128,1404,1430]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="8" pageNumber="9">Figure 1E</figureCitation>
).
</paragraph>
<paragraph id="8B86D87C8C2BFFF6615FFA23557FBF90" blockId="8.[264,1323,1484,1710]" box="[264,615,1484,1509]" pageId="8" pageNumber="9">
<emphasis id="B94D046E8C2BFFF6615FFA23557FBF90" box="[264,615,1484,1509]" italics="true" pageId="8" pageNumber="9">Histology of Vestibular Gland</emphasis>
</paragraph>
<paragraph id="8B86D87C8C2BFFF6616FFA1B569ABCDB" blockId="8.[264,1323,1484,1710]" pageId="8" pageNumber="9">
The vestibular gland contains large, rounded acini (
<figureCitation id="1302C4F98C2BFFF663C8FA1B530EBC7B" box="[927,1046,1524,1550]" captionStart="FIGURE 1 A – F" captionStartId="5.[264,367,1643,1667]" captionText="FIGURE 1 A – F. Photographs of Dendrodoris atromaculata A. Living animal (67 mm live length) found intertidally at Rowes Bay, Townsville, Australia. Photograph by J. Brodie. B. living animal (37 mm live length) found intertidally at Nasese, Fiji. C. histological section showing spicules located in the notal tissue. D. histological section showing the unusual distribution of pigment in the notal tissue. E. confocal microscope scan of a semi­everted penis. F. histological section through the vestibular gland of the reproductive system showing rounded acini. ac = acinus, ne =" pageId="8" pageNumber="9">Figure 1F</figureCitation>
) and the lumen has no epithelial fringe of deep staining microvili known to house bacteria in some other
<taxonomicName id="4C39A3FF8C2BFFF6649FF9F3565CBC28" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="8" pageNumber="9" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C2BFFF6649FF9F3565CBC28" italics="true" pageId="8" pageNumber="9">Dendrodoris</emphasis>
</taxonomicName>
species (see Klussmann­Kolb &amp; Brodie 1999). Additional wax sections of the vestibular gland were examined for bacteria, but none were detected by the Gram stain technique.
</paragraph>
<paragraph id="8B86D87C8C2BFFF6615FF90B56DABC88" blockId="8.[264,1323,1764,1870]" box="[264,450,1764,1789]" pageId="8" pageNumber="9">
<emphasis id="B94D046E8C2BFFF6615FF90B56DABC88" box="[264,450,1764,1789]" italics="true" pageId="8" pageNumber="9">Nervous System</emphasis>
</paragraph>
<paragraph id="8B86D87C8C2BFFF7616FF8E354CEBBF3" blockId="8.[264,1323,1764,1870]" lastBlockId="9.[264,1322,284,390]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">The nervous system is well developed with a ring of fused ganglia surrounding the distal pharynx. The smooth surfaced pharynx can move freely backwards and forwards through this ring. A separate pair of small, cream ganglia is also located well behind this principal ring, very close to the salivary glands. Without close inspection, it is possible that these ganglia could be confused with the salivary glands.</paragraph>
<paragraph id="8B86D87C8C2AFFF7615FFE535593BBA0" blockId="9.[264,1322,444,630]" box="[264,651,444,469]" pageId="9" pageNumber="10">
<emphasis id="B94D046E8C2AFFF7615FFE535593BBA0" box="[264,651,444,469]" italics="true" pageId="9" pageNumber="10">Circulatory &amp; Excretory Systems</emphasis>
</paragraph>
<paragraph id="8B86D87C8C2AFFF7616FFE0B55F5B803" blockId="9.[264,1322,444,630]" pageId="9" pageNumber="10">
The blood gland is irregular in shape and flattened dorso­ventrally.
<emphasis id="B94D046E8C2AFFF7640FFE0B53BABB88" box="[1112,1186,484,509]" italics="true" pageId="9" pageNumber="10">In situ</emphasis>
, it lies dorsally over the anterior section of the oesophagus, just posterior to the ring of ganglia described above. A rounded renal syrinx is present and located on the right side, in the posterior­most section of the body cavity.
</paragraph>
<paragraph id="8B86D87C8C2AFFF7615FFD43553DB8B0" blockId="9.[264,1323,684,990]" box="[264,549,684,709]" pageId="9" pageNumber="10">
<emphasis id="B94D046E8C2AFFF7615FFD43553DB8B0" box="[264,549,684,709]" italics="true" pageId="9" pageNumber="10">Occurrence and Habitat</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C2AFFF7616FFD3B56DABE58" pageId="9" pageNumber="10" type="reference_group">
<paragraph id="8B86D87C8C2AFFF7616FFD3B53E5B9AB" blockId="9.[264,1323,684,990]" pageId="9" pageNumber="10">
<taxonomicName id="4C39A3FF8C2AFFF7616FFD3B5568B898" ID-CoL="34Q5P" box="[312,624,724,749]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C2AFFF7616FFD3B5568B898" box="[312,624,724,749]" italics="true" pageId="9" pageNumber="10">Dendrodoris atromaculata</emphasis>
</taxonomicName>
appears to be relatively rare, a fact also noted by
<bibRefCitation id="EFA8A58D8C2AFFF764E2FD3B56D0B963" author="Alder" pageId="9" pageNumber="12" refString="Alder, J. &amp; Hancock, A. (1864) Notice of a collection of nudibranchiate Mollusca made in India by Walter Elliot, Esq., with description of several new genera and species. Transactions of the Zoological Society of London, 5, 113 - 147." type="journal article" year="1864">Alder and Hancock (1864)</bibRefCitation>
. The
<typeStatus id="548266DE8C2AFFF76259FD135558B963" box="[526,576,764,790]" pageId="9" pageNumber="10">type</typeStatus>
locality is Waltair,
<collectingCountry id="F32E98EC8C2AFFF76375FD13547AB963" box="[802,866,764,790]" name="India" pageId="9" pageNumber="10">India</collectingCountry>
. The field guide of
<bibRefCitation id="EFA8A58D8C2AFFF7641CFD13564EB94B" author="Willan" pageId="9" pageNumber="13" refString="Willan R. C. &amp; Coleman, N. (1984) Nudibranchs of Australasia. Australasian Marine Photographic Index, Caringbah, Sydney, 56 pp." type="book" year="1984">Willan &amp; Coleman (1984)</bibRefCitation>
records
<taxonomicName id="4C39A3FF8C2AFFF761E9FCCB5599B948" ID-CoL="34Q5P" box="[446,641,804,829]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C2AFFF761E9FCCB5599B948" box="[446,641,804,829]" italics="true" pageId="9" pageNumber="10">D. atromaculata</emphasis>
</taxonomicName>
as occurring in Queensland and Western
<collectingCountry id="F32E98EC8C2AFFF76425FCCB53F8B94B" box="[1138,1248,804,830]" name="Australia" pageId="9" pageNumber="10">Australia</collectingCountry>
, however, the current study documents for the first time the presence of
<taxonomicName id="4C39A3FF8C2AFFF76413FCA35211B910" ID-CoL="34Q5P" box="[1092,1289,844,869]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C2AFFF76413FCA35211B910" box="[1092,1289,844,869]" italics="true" pageId="9" pageNumber="10">D. atromaculata</emphasis>
</taxonomicName>
in tropical
<collectingCountry id="F32E98EC8C2AFFF7613CFC9B56C1B9FB" box="[363,473,884,910]" name="Australia" pageId="9" pageNumber="10">Australia</collectingCountry>
,
<collectingCountry id="F32E98EC8C2AFFF761B0FC9B5508B9FB" box="[487,528,884,910]" name="Fiji" pageId="9" pageNumber="10">Fiji</collectingCountry>
and
<collectingCountry id="F32E98EC8C2AFFF7621AFC9B55D0B9FB" box="[589,712,884,910]" name="Singapore" pageId="9" pageNumber="10">Singapore</collectingCountry>
. The four living specimens observed in the current study were found in very shallow water at the lower section of the intertidal zone, either out in the open within sponge­gardens or under rocks on sheltered muddy reefal areas.
</paragraph>
<paragraph id="8B86D87C8C2AFFF7615FFBFB56DABE58" blockId="9.[264,1323,1044,1430]" box="[264,450,1044,1069]" pageId="9" pageNumber="10">
<emphasis id="B94D046E8C2AFFF7615FFBFB56DABE58" box="[264,450,1044,1069]" italics="true" pageId="9" pageNumber="10">Natural History</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C2AFFF7616FFBD35531BD3B" pageId="9" pageNumber="10" type="description">
<paragraph id="8B86D87C8C2AFFF7616FFBD3530ABFE3" blockId="9.[264,1323,1044,1430]" pageId="9" pageNumber="10">
No details of egg masses or developmental biology are currently known for this species. The Townsville specimens were found in consecutive years during the month of August. This may well suggest a seasonal occurrence, particularly since the large size and conspicuous appearance of this species should make it relatively easy to find. However, these collections may also be the result of increased accessibility to intertidal sponge gardens during the very low tides at this time of year. One individual found in Rowes Bay was observed to excrete orange faeces, strongly suggesting it had been feeding on an orange sponge. Compared with several other dendrodorids (e.g.,
<taxonomicName id="4C39A3FF8C2AFFF76472FABB56A3BFE3" authority="Ruppell &amp; Leuckart, 1828" authorityName="Ruppell &amp; Leuckart" authorityYear="1828" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="fumata">
<emphasis id="B94D046E8C2AFFF76472FABB5387BF18" box="[1061,1183,1364,1389]" italics="true" pageId="9" pageNumber="10">D. fumata</emphasis>
Rüppell &amp; Leuckart, 1828
</taxonomicName>
)
<taxonomicName id="4C39A3FF8C2AFFF7619CFA935593BFE0" box="[459,651,1404,1429]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C2AFFF7619CFA935593BFE0" box="[459,651,1404,1429]" italics="true" pageId="9" pageNumber="10">D. atromaculata</emphasis>
</taxonomicName>
is relatively inactive in captivity.
</paragraph>
<paragraph id="8B86D87C8C2AFFF7615FFA2356EFBF90" blockId="9.[264,1323,1484,1870]" box="[264,503,1484,1509]" pageId="9" pageNumber="10">
<emphasis id="B94D046E8C2AFFF7615FFA2356EFBF90" box="[264,503,1484,1509]" italics="true" pageId="9" pageNumber="10">Diagnostic Features</emphasis>
</paragraph>
<paragraph id="8B86D87C8C2AFFF7616FFA1B5531BD3B" blockId="9.[264,1323,1484,1870]" pageId="9" pageNumber="10">
<taxonomicName id="4C39A3FF8C2AFFF7616FFA1B5569BC78" box="[312,625,1524,1549]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C2AFFF7616FFA1B5569BC78" box="[312,625,1524,1549]" italics="true" pageId="9" pageNumber="10">Dendrodoris atromaculata</emphasis>
</taxonomicName>
is unique among described members of
<taxonomicName id="4C39A3FF8C2AFFF7641AFA1B53FABC78" box="[1101,1250,1524,1549]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C2AFFF7641AFA1B53FABC78" box="[1101,1250,1524,1549]" italics="true" pageId="9" pageNumber="10">Dendrodoris</emphasis>
</taxonomicName>
being the only member of the genus known to possess finger­like notal papillae, large dense notal spicules and a gill plume that is cross­like. The distinguishing external features of
<taxonomicName id="4C39A3FF8C2AFFF76559F9AB56BDBCF0" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="9" pageNumber="10" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C2AFFF76559F9AB56BDBCF0" italics="true" pageId="9" pageNumber="10">D. atromaculata</emphasis>
</taxonomicName>
are its: (a) large size, (b) high body profile, (c) yellow ground colour (generally with black patches), (d) lack of oral tentacles, (e) finger­like papillae distributed over the notum, (f) “star­like” dorsal tubercles and (g) large cross­like gill plume. Internally, the distinctive features are: (h) an unusual distribution of black pigment within the connective tissue, (i) large, dense notal spicules, and (j) distinctive rounded acini within the reproductive vestibular gland.
</paragraph>
</subSubSection>
<subSubSection id="C3238BF78C29FFF4615FFEF356ADBC2B" pageId="10" pageNumber="11" type="discussion">
<paragraph id="8B86D87C8C29FFF4615FFEF35232BB40" blockId="10.[264,1323,284,1630]" box="[264,1322,284,309]" pageId="10" pageNumber="11">
<emphasis id="B94D046E8C29FFF4615FFEF35232BB40" box="[264,1322,284,309]" italics="true" pageId="10" pageNumber="11">Discussion of some important and previously used generic features and phylogenetic characters</emphasis>
</paragraph>
<paragraph id="8B86D87C8C29FFF4616FFEAB55DCBB28" blockId="10.[264,1323,284,1630]" box="[312,708,324,349]" pageId="10" pageNumber="11">
<emphasis id="B94D046E8C29FFF4616FFEAB55DCBB28" box="[312,708,324,349]" italics="true" pageId="10" pageNumber="11">Ptyaline Glands &amp; Notal Spicules</emphasis>
</paragraph>
<paragraph id="8B86D87C8C29FFF4616FFE83559BB803" blockId="10.[264,1323,284,1630]" pageId="10" pageNumber="11">
Possession of ptyaline glands, unique digestive structures found in all members of
<taxonomicName id="4C39A3FF8C29FFF4615FFE7B5684BBD8" box="[264,412,404,429]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C29FFF4615FFE7B5684BBD8" box="[264,412,404,429]" italics="true" pageId="10" pageNumber="11">Dendrodoris</emphasis>
</taxonomicName>
, confirmed the generic placement of
<taxonomicName id="4C39A3FF8C29FFF46300FE7B5300BBD8" box="[855,1048,404,429]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C29FFF46300FE7B5300BBD8" box="[855,1048,404,429]" italics="true" pageId="10" pageNumber="11">D. atromaculata</emphasis>
</taxonomicName>
. However, the discovery of prominent notal spicules was unexpected since
<bibRefCitation id="EFA8A58D8C29FFF463E6FE5353E8BBA3" author="Valdes" box="[945,1264,444,470]" pageId="10" pageNumber="13" refString="Valdes, A. &amp; Gosliner, T. M. (1999) Phylogeny of the radula-less dorids (Mollusca, Nudibranchia), with the description of a new genus and a new family. Zoologica Scripta, 28, 315 - 360." type="journal article" year="1999">Valdés &amp; Gosliner (1999)</bibRefCitation>
and
<bibRefCitation id="EFA8A58D8C29FFF4615FFE0B56B5BB8B" author="Valdes" box="[264,429,484,510]" pageId="10" pageNumber="13" refString="Valdes, A. (2002) A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society, 136, 535 - 636." type="journal article" year="2002">Valdés (2002)</bibRefCitation>
found “integumentary” spicules were absent in all species of
<taxonomicName id="4C39A3FF8C29FFF464C1FE0B5233BB88" box="[1174,1323,484,509]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C29FFF464C1FE0B5233BB88" box="[1174,1323,484,509]" italics="true" pageId="10" pageNumber="11">Dendrodoris</emphasis>
</taxonomicName>
included in their phylogenetic analyses of radula­less dorids and cryptobranch dorids respectively. An absence of such spicules can therefore no longer be considered as a generic feature for
<taxonomicName id="4C39A3FF8C29FFF461BFFDB35565B800" box="[488,637,604,629]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C29FFF461BFFDB35565B800" box="[488,637,604,629]" italics="true" pageId="10" pageNumber="11">Dendrodoris</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B86D87C8C29FFF4616FFD6B5507B8E8" blockId="10.[264,1323,284,1630]" box="[312,543,644,669]" pageId="10" pageNumber="11">
<emphasis id="B94D046E8C29FFF4616FFD6B5507B8E8" box="[312,543,644,669]" italics="true" pageId="10" pageNumber="11">Pyloric Bulb or Sac</emphasis>
</paragraph>
<paragraph id="8B86D87C8C29FFF4616FFD435388B913" blockId="10.[264,1323,284,1630]" pageId="10" pageNumber="11">
In only one of the three animals dissected was a pyloric bulb (as defined by
<bibRefCitation id="EFA8A58D8C29FFF464E4FD435675B89B" author="Wagele" pageId="10" pageNumber="13" refString="Wagele, H., Brodie, G. &amp; Klussmann-Kolb, A. (1999) Histological investigations of Dendrodoris nigra (Gastropoda, Nudibranchia, Dendrodorididae). Molluscan Research, 20, 79 - 94." type="journal article" year="1999">
Wägele
<emphasis id="B94D046E8C29FFF46542FD435606B898" italics="true" pageId="10" pageNumber="11">et al</emphasis>
. 1999
</bibRefCitation>
: 92) present (see
<figureCitation id="1302C4F98C29FFF4626EFD3B55ABB89B" box="[569,691,724,750]" captionStart="FIGURE 2 A – D. 0 0" captionStartId="7.[264,368,1577,1601]" captionTargetBox="[323,1280,313,1555]" captionTargetId="figure@7.[298,1299,300,1578]" captionTargetPageId="7" captionText="FIGURE 2 A – D. 0 0 Composite drawings of Dendrodoris atromaculata (preserved material) showing: A. compound dorsal tubercule. B. anterior part of the digestive system. C. posterior part of the digestive system. D. arrangement of the post­gonadial reproductive system. amp = ampulla, an = anus, bc = bursa copulatrix, bw = body wall, es = exit to stomach, hd = hermaphroditic duct, int = intestine, mb = muscular bulb, mo = mouth, ng = nidamental gland, np = notal papilla, oes = oesophagus, ot = oral tube, ov = oviduct, pg = prostate gland, ph = pharynx, pb = pyloric bulb (not present in all individuals), ptd = ptyaline duct, ptg = ptyaline gland, rm = retractor muscle, rs = receptaculum seminis, sg = salivary gland, va = vagina, vd = vas deferens, vg = vestibular gland. Scale bars = 2 mm." httpUri="https://zenodo.org/record/157390/files/figure.png" pageId="10" pageNumber="11">Figure 2C</figureCitation>
). These findings confirm the intraspecific variation of this intestinal structure, as previously found by
<bibRefCitation id="EFA8A58D8C29FFF46315FD135337B963" box="[834,1071,764,790]" pageId="10" pageNumber="12" refString="Brodie, G. D., Collins, J. D. &amp; Willan, R. C. (1997) Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata in the Indo-west Pacific region. Journal of Molluscan Studies, 63, 407 - 423." type="journal article">
Brodie
<emphasis id="B94D046E8C29FFF463CCFD1354C8B960" box="[923,976,764,789]" italics="true" pageId="10" pageNumber="11">et al</emphasis>
. (1997)
</bibRefCitation>
[described under the incorrect term pyloric gland], and confirms that the presence and form of a pyloric bulb (or sac) is unsuitable for separating species and for use in phylogenetic analyses.
</paragraph>
<paragraph id="8B86D87C8C29FFF4616FFC9B5513B9F8" blockId="10.[264,1323,284,1630]" box="[312,523,884,909]" pageId="10" pageNumber="11">
<emphasis id="B94D046E8C29FFF4616FFC9B5513B9F8" box="[312,523,884,909]" italics="true" pageId="10" pageNumber="11">Vestibular Glands</emphasis>
</paragraph>
<paragraph id="8B86D87C8C29FFF4616FFC7356ADBC2B" blockId="10.[264,1323,284,1630]" pageId="10" pageNumber="11">
The vestibular gland microstructure found in
<taxonomicName id="4C39A3FF8C29FFF4631BFC735315B9C0" box="[844,1037,924,949]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="atromaculata">
<emphasis id="B94D046E8C29FFF4631BFC735315B9C0" box="[844,1037,924,949]" italics="true" pageId="10" pageNumber="11">D. atromaculata</emphasis>
</taxonomicName>
has, to date, been found only in an unnamed species of
<taxonomicName id="4C39A3FF8C29FFF46222FC2B5412B9A8" box="[629,778,964,989]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C29FFF46222FC2B5412B9A8" box="[629,778,964,989]" italics="true" pageId="10" pageNumber="11">Dendrodoris</emphasis>
</taxonomicName>
from
<collectingCountry id="F32E98EC8C29FFF46305FC2B548DB9AB" box="[850,917,964,990]" name="Japan" pageId="10" pageNumber="11">Japan</collectingCountry>
(see
<bibRefCitation id="EFA8A58D8C29FFF46383FC2B5373B9AB" author="Brodie" box="[980,1131,964,990]" pageId="10" pageNumber="12" refString="Brodie, G. D. (2002) The molluscan nudibranch family Dendrodorididae (Anthobranchia: Doridoidea) in Australia: Systematics and Phylogenetic Relationships. PhD Thesis, School of Marine Biology &amp; Aquaculture, James Cook University, 198 pp." type="book" year="2002">Brodie 2002</bibRefCitation>
). Relatively few
<taxonomicName id="4C39A3FF8C29FFF4615FFC035685BE70" box="[264,413,1004,1029]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="genus">
<emphasis id="B94D046E8C29FFF4615FFC035685BE70" box="[264,413,1004,1029]" italics="true" pageId="10" pageNumber="11">Dendrodoris</emphasis>
</taxonomicName>
are known to possess a vestibular gland (e.g.,
<taxonomicName id="4C39A3FF8C29FFF463B0FC035356BE70" box="[999,1102,1004,1029]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="nigra">
<emphasis id="B94D046E8C29FFF463B0FC035356BE70" box="[999,1102,1004,1029]" italics="true" pageId="10" pageNumber="11">D. nigra</emphasis>
</taxonomicName>
,
<taxonomicName id="4C39A3FF8C29FFF46435FC0356ACBE5B" authority="Rao &amp; Kumary, 1973" authorityName="Rao &amp; Kumary" authorityYear="1973" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="goani">
<emphasis id="B94D046E8C29FFF46435FC0353D5BE70" box="[1122,1229,1004,1029]" italics="true" pageId="10" pageNumber="11">D. goani</emphasis>
Rao &amp; Kumary, 1973
</taxonomicName>
;
<taxonomicName id="4C39A3FF8C29FFF46195FBFB55FABE5B" authority="Baba, 1936" authorityName="Baba" authorityYear="1936" box="[450,738,1044,1070]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="elongata">
<emphasis id="B94D046E8C29FFF46195FBFB5555BE58" box="[450,589,1044,1069]" italics="true" pageId="10" pageNumber="11">D. elongata</emphasis>
Baba, 1936
</taxonomicName>
). However, vaginal glands sometimes also called vestibular glands (see
<bibRefCitation id="EFA8A58D8C29FFF4625DFBD355AABE23" author="Gosliner" box="[522,690,1084,1110]" pageId="10" pageNumber="12" refString="Gosliner, T. (1994) Gastropoda: Opisthobranchia. In: Harrison, F. W. &amp; Kohn, A. J. Microscopic Anatomy of Invertebrates. Volume 5: Mollusca. Wiley-Liss, New York, pp. 253 - 355." type="journal article" year="1994">Gosliner 1994</bibRefCitation>
) are reported from many dorid nudibranchs. Description of these reproductive accessory organs has largely been undertaken at the gross morphological level (
<bibRefCitation id="EFA8A58D8C29FFF461BEFB635586BED3" author="Gosliner" box="[489,670,1164,1190]" pageId="10" pageNumber="12" refString="Gosliner, T. (1994) Gastropoda: Opisthobranchia. In: Harrison, F. W. &amp; Kohn, A. J. Microscopic Anatomy of Invertebrates. Volume 5: Mollusca. Wiley-Liss, New York, pp. 253 - 355." type="journal article" year="1994">Gosliner 1994</bibRefCitation>
) with little description of ultrastructural detail or microanatomy. This fact makes the collective usage of “presence or absence” of such glands, at the gross morphological level in phylogenetic studies, inadvisable because ultrastructural or microanatomical investigation is required to determine their similarity. This conclusion is reinforced by the vestibular gland investigations of
<taxonomicName id="4C39A3FF8C29FFF46420FAC353FFBF30" box="[1143,1255,1324,1349]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="nigra">
<emphasis id="B94D046E8C29FFF46420FAC353FFBF30" box="[1143,1255,1324,1349]" italics="true" pageId="10" pageNumber="11">D. nigra</emphasis>
</taxonomicName>
(see Klussmann­Kolb &amp; Brodie 1999), where an unusual microstructure was found to house symbiotic bacteria, and in similar investigations of
<taxonomicName id="4C39A3FF8C29FFF4633CFA9353EBBFE3" authority="Kay &amp; Young, 1969" authorityName="Kay &amp; Young" authorityYear="1969" box="[875,1267,1404,1430]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="coronata">
<emphasis id="B94D046E8C29FFF4633CFA9354E0BFE0" box="[875,1016,1404,1429]" italics="true" pageId="10" pageNumber="11">D. coronata</emphasis>
Kay &amp;
<bibRefCitation id="EFA8A58D8C29FFF4640CFA9353EBBFE3" author="Young" box="[1115,1267,1404,1430]" pageId="10" pageNumber="13" refString="Young, D. K. (1969) The functional morphology of the feeding apparatus of some Indo-west-Pacific dorid nudibranchs. Malacologia, 9, 421 - 445." type="journal article" year="1969">Young, 1969</bibRefCitation>
</taxonomicName>
(see
<bibRefCitation id="EFA8A58D8C29FFF4615FFA4B5594BFCB" author="Brodie" box="[264,652,1444,1470]" pageId="10" pageNumber="12" refString="Brodie, G. D. &amp; Klussmann-Kolb, A. (2000) Symbiotic bacteria in the reproductive system of tropical nudibranchs. In: Ponder, W. (Ed). Molluscs 2000: Understanding Molluscan Biodiversity in our region into the 21 st Century, Abstracts, University of Sydney, Australia, p. 15." type="journal article" year="2000">Brodie &amp; Klussmann­Kolb 2000</bibRefCitation>
) and
<taxonomicName id="4C39A3FF8C29FFF46287FA4B5382BFCB" authority="Ruppell &amp; Leuckart, 1828" authorityName="Ruppell &amp; Leuckart" authorityYear="1828" box="[720,1178,1444,1470]" class="Gastropoda" family="Dendrodorididae" genus="Dendrodoris" kingdom="Animalia" order="Nudibranchia" pageId="10" pageNumber="11" phylum="Mollusca" rank="species" species="fumata">
<emphasis id="B94D046E8C29FFF46287FA4B545EBFC8" box="[720,838,1444,1469]" italics="true" pageId="10" pageNumber="11">D. fumata</emphasis>
(Rüppell &amp; Leuckart, 1828)
</taxonomicName>
(see
<bibRefCitation id="EFA8A58D8C29FFF4648DFA4B565FBF93" author="Brodie" pageId="10" pageNumber="12" refString="Brodie, G. D. (2001) The systematics and phylogeny of the nudibranch family Dendrodorididae (Opisthobranchia, Doridoidea). pp. 40. In: Salvini-Plawen, L., Voltzow, J., Sattmann, H. &amp; Steiner, G. (Eds). Abstracts, World Congress of Malacology 2001, Vienna, Austria, 417 pp." type="book" year="2001">Brodie 2001</bibRefCitation>
). In the latter two species, the vestibular gland is embedded within the body wall making it non­observable by gross dissection. In future comparative studies it is essential that microstructural investigations of these organs, and the surrounding body wall tissues, be undertaken.
</paragraph>
</subSubSection>
</treatment>
</document>
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