617 lines
119 KiB
XML
617 lines
119 KiB
XML
<document id="FE52A20D7400D72FD2672E725D44714A" ID-DOI="10.11646/zootaxa.3956.3.5" ID-GBIF-Dataset="fd562f8c-5541-478c-8827-6464cc13f053" ID-ISSN="1175-5326" ID-Zenodo-Dep="240004" ID-ZooBank="25F9FA82-28BA-4C94-8EF5-D0EE88B0B48E" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461056603918" checkinUser="plazi" docAuthor="Díaz, Maria C., Thacker, Robert W., Redmond, Niamh E., Perez, Thierry & Collins, Allen G." docDate="2015" docId="03AF87AD1138635484A1FC4EFA80529F" docLanguage="en" docName="zt03956p412.pdf" docOrigin="Zootaxa 3956 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Vansoestia Díaz, Thacker, Redmond, Perez & Collins, 2015, gen. nov." docType="treatment" docVersion="7" lastPageNumber="411" masterDocId="FF96FFD5113A635C8436FFE4FFF9513B" masterDocTitle="Vansoestia caribensis gen. nov., sp. nov.: first report of the family Ianthellidae (Verongida, Demospongiae) in the Caribbean" masterLastPageNumber="412" masterPageNumber="403" pageNumber="405" updateTime="1698581249652" updateUser="plazi">
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<mods:titleInfo id="73EDE3E85BC51263271B4425D0507678">
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<mods:title id="DFF6C0BD5E4D5A80EB78620F79376C85">Vansoestia caribensis gen. nov., sp. nov.: first report of the family Ianthellidae (Verongida, Demospongiae) in the Caribbean</mods:title>
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<mods:name id="1F92166AD8D3174B9A53212096BE226C" type="personal">
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<mods:roleTerm id="19F01C4D4C6510224B8B37891B71FF3D">Author</mods:roleTerm>
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<mods:namePart id="AB5BA3F41133AAE2265E1BC083B5C1E1">Díaz, Maria C.</mods:namePart>
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</mods:name>
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<mods:roleTerm id="657E5790C6FEC18EF2BD848CFAE31844">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="B7CFCE5F4E63E444E2D7CA09963DCAA4">Thacker, Robert W.</mods:namePart>
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</mods:name>
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<mods:name id="F43C97A88ACA206DA9BE355E927FF8EB" type="personal">
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<mods:role id="FD2C3BDEBDFAD17A285E71E3AA36D27F">
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<mods:roleTerm id="D217D9AF01383A6B33976CEB16458880">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="E0D77A27A6BA1E788C383A06346391FC">Redmond, Niamh E.</mods:namePart>
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</mods:name>
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<mods:name id="7DB792FE278CCD68D4375FE7EEB772E2" type="personal">
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<mods:roleTerm id="B3A233314709C3BDE4CCFCC970A49ACA">Author</mods:roleTerm>
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<mods:namePart id="68EC815FF4799E2222E05C23FB28335D">Perez, Thierry</mods:namePart>
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</mods:name>
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<mods:name id="1C75C39992C0DA9EAC21BAFD361FE051" type="personal">
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<mods:roleTerm id="37907C0618F3D67646AED1B2F057D1B4">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="35E8532346C3B3AC9543295B8A18768B">Collins, Allen G.</mods:namePart>
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<mods:title id="8A6D21BB1214FB3765F7517C0160DCB8">Zootaxa</mods:title>
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<mods:part id="16476D6576689893A060865EE2EF53A4">
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<mods:date id="1818E1B19C8A17AA8A094ADC5B168C43">2015</mods:date>
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<mods:number id="BC4FC9BF25BA7D8D1EBA326A94ED9E8A">3956</mods:number>
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<mods:classification id="22E141409A44562624B898B470617A0C">journal article</mods:classification>
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<mods:identifier id="AF5A7054D30EAB5713AF04519248D74A" type="DOI">10.11646/zootaxa.3956.3.5</mods:identifier>
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<mods:identifier id="9FDAC52D4C1CBB89A9746920FF5AB36C" type="Zenodo-Dep">240004</mods:identifier>
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<treatment id="03AF87AD1138635484A1FC4EFA80529F" ID-DOI="http://doi.org/10.5281/zenodo.6093271" ID-GBIF-Taxon="119613088" ID-Zenodo-Dep="6093271" LSID="urn:lsid:plazi:treatment:03AF87AD1138635484A1FC4EFA80529F" httpUri="http://treatment.plazi.org/id/03AF87AD1138635484A1FC4EFA80529F" lastPageId="8" lastPageNumber="411" pageId="2" pageNumber="405">
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<subSubSection id="C31C65301138635E84A1FC4EFE6D52FF" box="[151,404,938,964]" pageId="2" pageNumber="405" type="nomenclature">
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<paragraph id="8BB936BB1138635E84A1FC4EFE6D52FF" blockId="2.[151,404,938,964]" box="[151,404,938,964]" pageId="2" pageNumber="405">
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<heading id="D0F181D71138635E84A1FC4EFE6D52FF" bold="true" box="[151,404,938,964]" fontSize="11" level="1" pageId="2" pageNumber="405" reason="1">
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<emphasis id="B972EAA91138635E84A1FC4EFE6D52FF" bold="true" box="[151,404,938,964]" pageId="2" pageNumber="405">
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<taxonomicName id="4C064D381138635E84A1FC4EFEEC52FF" ID-CoL="84VM7" box="[151,277,938,964]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="2" pageNumber="405" phylum="Porifera" rank="genus" status="gen. nov.">
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<emphasis id="B972EAA91138635E84A1FC4EFEEC52FF" bold="true" box="[151,277,938,964]" italics="true" pageId="2" pageNumber="405">Vansoestia</emphasis>
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</taxonomicName>
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,
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<taxonomicNameLabel id="A24157D21138635E8510FC4EFE6D52FF" box="[294,404,938,964]" pageId="2" pageNumber="405" rank="genus">gen. nov.</taxonomicNameLabel>
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C31C65301138635E84A1FC15FD605585" pageId="2" pageNumber="405" type="diagnosis">
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<paragraph id="8BB936BB1138635E84A1FC15FD605585" blockId="2.[151,1437,1009,2006]" pageId="2" pageNumber="405">
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<emphasis id="B972EAA91138635E84A1FC15FEE85531" bold="true" box="[151,273,1009,1034]" pageId="2" pageNumber="405">Diagnosis.</emphasis>
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<taxonomicName id="4C064D381138635E852CFC15FE675531" box="[282,414,1009,1034]" class="Demospongiae" family="Ianthellidae" kingdom="Animalia" order="Verongida" pageId="2" pageNumber="405" phylum="Porifera" rank="family">Ianthellidae</taxonomicName>
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represented by individuals with a thin (
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) encrusting growth form that presents a cellularized cortex of variable thickness (10–200 µm) composed of a collagenous matrix and various
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<typeStatus id="54BD88191138635E813EFBF2FABA5515" box="[1288,1347,1046,1070]" pageId="2" pageNumber="405">types</typeStatus>
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of cells with inclusions (i.e. granular, spherulous, and/or vacuolar). Abundant subdermal canals and lacunae are found immediately underneath the cortex. The choanosome consists of numerous aquiferous canals, a mesohyl where two or three
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<typeStatus id="54BD88191138635E84C0FB66FEC855A1" box="[246,305,1154,1178]" pageId="2" pageNumber="405">types</typeStatus>
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of cells with inclusions can be distinguished, and oval eurypylous choanocyte chambers 20–50 µm, all supported with abundant fibrillar collagen.
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</paragraph>
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</subSubSection>
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<subSubSection id="C31C65301138635E84F1FB2DFADB5739" pageId="2" pageNumber="405" type="materials_examined">
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<paragraph id="8BB936BB1138635E84F1FB2DFCCB55D9" blockId="2.[151,1437,1009,2006]" box="[199,818,1225,1250]" pageId="2" pageNumber="405">
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<emphasis id="B972EAA91138635E84F1FB2DFCCB55D9" bold="true" box="[199,818,1225,1250]" pageId="2" pageNumber="405">
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<typeStatus id="54BD88191138635E84F1FB2DFEFB55D9" box="[199,258,1225,1250]" pageId="2" pageNumber="405">Type</typeStatus>
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species:
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<taxonomicName id="4C064D381138635E855DFB2DFD9855D9" box="[363,609,1225,1250]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="2" pageNumber="405" phylum="Porifera" rank="species" species="caribensis" status="gen. nov.">
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<emphasis id="B972EAA91138635E855DFB2DFD9855D9" bold="true" box="[363,609,1225,1250]" italics="true" pageId="2" pageNumber="405">Vansoestia caribensis</emphasis>
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</taxonomicName>
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<taxonomicNameLabel id="A24157D21138635E865FFB2DFD2B55D9" box="[617,722,1225,1250]" pageId="2" pageNumber="405" rank="genus">gen. nov.</taxonomicNameLabel>
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sp. nov.
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</emphasis>
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</paragraph>
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<paragraph id="8BB936BB1138635E84F1FB09FDA65449" blockId="2.[151,1437,1009,2006]" pageId="2" pageNumber="405">
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||
<emphasis id="B972EAA91138635E84F1FB09FE48543D" bold="true" box="[199,433,1261,1286]" pageId="2" pageNumber="405">Material examined.</emphasis>
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All specimens studied were found in coral reefs from Bocas del Toro,
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<collectingCountry id="F311762B1138635E80E1FB0AFACA543E" box="[1239,1331,1262,1285]" name="Panama" pageId="2" pageNumber="405">Panama</collectingCountry>
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, between
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||
<quantity id="4CFE9B5E1138635E84A1FAF6FF035412" box="[151,250,1298,1322]" metricMagnitude="1" metricUnit="m" metricValue="1.5" metricValueMax="2.0" metricValueMin="1.0" pageId="2" pageNumber="405" unit="m" value="15.0" valueMax="20.0" valueMin="10.0">10–20 m</quantity>
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||
in depth. This species grows over dead coral and other organisms, in open and cryptic reef habitats.
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<typeStatus id="54BD88191138635E84A1FAD1FF065475" box="[151,255,1333,1358]" pageId="2" pageNumber="405" type="holotype">Holotype</typeStatus>
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||
:
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<collectionCode id="ED17AE7E1138635E852FFAD2FE935475" LSID="urn:lsid:biocol.org:col:34871" box="[281,362,1334,1358]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="405">USNM</collectionCode>
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1133773, Buoy 19 (N9°18.11, W 82°17.66), Bocas del Toro,
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<collectingCountry id="F311762B1138635E8004FAD2FB725476" box="[1074,1163,1334,1357]" name="Panama" pageId="2" pageNumber="405">Panama</collectingCountry>
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(P67),
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<quantity id="4CFE9B5E1138635E80D4FAD2FAE45476" box="[1250,1309,1334,1358]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="2" pageNumber="405" unit="m" value="10.0">10 m</quantity>
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deep, coll: M.C. Díaz and R.W. Thacker;
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<date id="FFB8107B1138635E85DBFABEFDA65449" box="[493,607,1370,1394]" pageId="2" pageNumber="405" value="2009-07-18">7/18/2009</date>
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</paragraph>
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||
<paragraph id="8BB936BB1138635E84F1FA9AFC6A54E5" blockId="2.[151,1437,1009,2006]" pageId="2" pageNumber="405">
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<typeStatus id="54BD88191138635E84F1FA9AFEC054AD" box="[199,313,1406,1430]" pageId="2" pageNumber="405" type="paratype">Paratypes</typeStatus>
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:
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<collectionCode id="ED17AE7E1138635E8573FA9AFE6F54AD" LSID="urn:lsid:biocol.org:col:34871" box="[325,406,1406,1430]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="405">USNM</collectionCode>
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1133782, Buoy 19 (N9°18.11, W 82°17.66), Bocas del Toro,
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<collectingCountry id="F311762B1138635E805FFA9AFB3B54AE" box="[1129,1218,1406,1429]" name="Panama" pageId="2" pageNumber="405">Panama</collectingCountry>
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(P76),
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<quantity id="4CFE9B5E1138635E812DFA99FAAE54AE" box="[1307,1367,1405,1430]" metricMagnitude="1" metricUnit="m" metricValue="1.5" pageId="2" pageNumber="405" unit="m" value="15.0">15 m</quantity>
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deep, coll: M.C. Díaz and R.W. Thacker;
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<date id="FFB8107B1138635E866EFA46FD375481" box="[600,718,1442,1466]" pageId="2" pageNumber="405" value="2009-07-18">7/18/2009</date>
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||
;
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<collectionCode id="ED17AE7E1138635E86D7FA46FCCB5481" LSID="urn:lsid:biocol.org:col:34871" box="[737,818,1442,1466]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="2" pageNumber="405">USNM</collectionCode>
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1204851 (P
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<date id="FFB8107B1138635E87FAFA46FC0F5481" box="[972,1014,1442,1466]" pageId="2" pageNumber="405" value="2012-10">12x</date>
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342), Punta Caracol (N9°22.66, W 82°18.19),
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<quantity id="4CFE9B5E1138635E8522FA21FEB454E6" box="[276,333,1477,1502]" metricMagnitude="1" metricUnit="m" metricValue="1.5" pageId="2" pageNumber="405" unit="m" value="15.0">15 m</quantity>
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deep, coll: M.C. Díaz and S.A. Pomponi,
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<date id="FFB8107B1138635E871EFA22FC7654E5" box="[808,911,1478,1502]" pageId="2" pageNumber="405" value="2012-08-08">8/8/2012</date>
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||
.
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||
</paragraph>
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||
<paragraph id="8BB936BB1138635E84F1FA0DFADB5739" blockId="2.[151,1437,1009,2006]" box="[199,1314,1513,1538]" pageId="2" pageNumber="405">
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||
<emphasis id="B972EAA91138635E84F1FA0DFE405739" bold="true" box="[199,441,1513,1538]" pageId="2" pageNumber="405">Additional material.</emphasis>
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P
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||
<date id="FFB8107B1138635E85E7FA0EFE005739" box="[465,505,1514,1538]" pageId="2" pageNumber="405" value="2012-10">12x</date>
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||
402, and P
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||
<date id="FFB8107B1138635E8647FA0EFD635739" box="[625,666,1514,1538]" pageId="2" pageNumber="405" value="2012-10">12x</date>
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403 were consumed in histological, or TEM preparations.
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</paragraph>
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</subSubSection>
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<subSubSection id="C31C65301138635E84F1F9E9FC355771" pageId="2" pageNumber="405" type="etymology">
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<paragraph id="8BB936BB1138635E84F1F9E9FC355771" blockId="2.[151,1437,1009,2006]" pageId="2" pageNumber="405">
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<emphasis id="B972EAA91138635E84F1F9E9FEB7571D" bold="true" box="[199,334,1549,1574]" pageId="2" pageNumber="405">Etymology.</emphasis>
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The species is named in honor of Dr. Rob van Soest, an essential contributor to the understanding of Demospongiae, including
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<taxonomicName id="4C064D381138635E85EAF9D5FDAB5771" box="[476,594,1585,1610]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="2" pageNumber="405" phylum="Porifera" rank="order">Verongida</taxonomicName>
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, in the Caribbean and worldwide.
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</paragraph>
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</subSubSection>
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||
<subSubSection id="C31C65301138635F84F1F9B1FDE3578B" lastPageId="3" lastPageNumber="406" pageId="2" pageNumber="405" type="description">
|
||
<paragraph id="8BB936BB1138635E84F1F9B1FDF75619" blockId="2.[151,1437,1009,2006]" pageId="2" pageNumber="405">
|
||
<emphasis id="B972EAA91138635E84F1F9B1FDA65755" bold="true" box="[199,607,1621,1646]" pageId="2" pageNumber="405">Description. External morphology</emphasis>
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: The sponge is soft, thinly encrusting (
|
||
<quantity id="4CFE9B5E1138635E8024F9B2FB965756" box="[1042,1135,1622,1646]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" metricValueMax="4.0" metricValueMin="1.0" pageId="2" pageNumber="405" unit="mm" value="2.5" valueMax="4.0" valueMin="1.0">1–4 mm</quantity>
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thick), orange to yellow in color internally and externally. Color turns slowly brown to purple in ethanol. The sponge grows over dead coral or over other organisms, particularly other sponges (
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<figureCitation id="133D2A3E1138635E86F0F979FCDC578D" box="[710,805,1693,1718]" captionStart="FIGURE 1" captionStartId="3.[151,250,1030,1052]" captionTargetBox="[151,1436,481,1004]" captionTargetId="figure@3.[151,1436,481,1009]" captionTargetPageId="3" captionText="FIGURE 1. Vansoestia caribensis gen. nov. sp. nov. growing on the edge of a dead plate coral. A. General habit showing its overall growth form, with algae and polychaete calcareous tubes mingled within the body. Scale bar = 2 cm. B. Close-up view of a more relaxed specimen showing the subdermal canals (Sc), 3 – 8 mm wide, and the profusely abundant pores. Scale bar = 1.2 cm" httpUri="https://zenodo.org/record/240005/files/figure.png" pageId="2" pageNumber="405">Figure 1</figureCitation>
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). Its surface is dominated by thin reticulated subdermal canals (
|
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<quantity id="4CFE9B5E1138635E84DAF926FEB357E2" box="[236,330,1730,1754]" metricMagnitude="-3" metricUnit="m" metricValue="5.5" metricValueMax="8.0" metricValueMin="3.0" pageId="2" pageNumber="405" unit="mm" value="5.5" valueMax="8.0" valueMin="3.0">3–8 mm</quantity>
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wide) that branch and decrease in width away from the oscules (
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||
<quantity id="4CFE9B5E1138635E8016F925FB7057E2" box="[1056,1161,1729,1754]" metricMagnitude="-3" metricUnit="m" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="2" pageNumber="405" unit="cm" value="0.75" valueMax="1.0" valueMin="0.5">0.5–1 cm</quantity>
|
||
wide). Small ostia (
|
||
<quantity id="4CFE9B5E1138635E815CF926FF1457C6" metricMagnitude="-4" metricUnit="m" metricValue="6" metricValueMax="8.0" metricValueMin="4.0" pageId="2" pageNumber="405" unit="mm" value="0.6" valueMax="0.8" valueMin="0.4">0.4– 0.8 mm</quantity>
|
||
wide) are dispersed between subdermal canals. The soft and fragile consistency of this sponge reflects the lack of a reticulate fiber skeleton.
|
||
</paragraph>
|
||
<paragraph id="8BB936BB1138635F84F1F8C9FDF15097" blockId="2.[151,1437,1009,2006]" lastBlockId="3.[151,1436,151,429]" lastPageId="3" lastPageNumber="406" pageId="2" pageNumber="405">
|
||
Internal morphology (
|
||
<figureCitation id="133D2A3E1138635E85F5F8C9FDA8567D" box="[451,593,1837,1862]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStart-2="FIGURE 4" captionStartId-0="4.[151,250,1429,1451]" captionStartId-1="5.[151,250,1653,1675]" captionStartId-2="6.[151,250,1668,1690]" captionTargetBox-0="[153,1436,218,1385]" captionTargetBox-1="[167,1419,581,1583]" captionTargetBox-2="[151,1431,193,1640]" captionTargetId-0="figure@4.[151,1436,213,1391]" captionTargetId-1="figure@5.[151,1436,577,1592]" captionTargetId-2="figure@6.[151,1436,193,1646]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionTargetPageId-2="6" captionText-0="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." captionText-1="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." captionText-2="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri-0="https://zenodo.org/record/240006/files/figure.png" httpUri-1="https://zenodo.org/record/240007/files/figure.png" httpUri-2="https://zenodo.org/record/240008/files/figure.png" pageId="2" pageNumber="405">Figures 2–4</figureCitation>
|
||
). There is a clear separation between ectosome and choanosome (
|
||
<figureCitation id="133D2A3E1138635E8167F8C9FF5F5651" captionStart="FIGURE 2" captionStartId="4.[151,250,1429,1451]" captionTargetBox="[153,1436,218,1385]" captionTargetId="figure@4.[151,1436,213,1391]" captionTargetPageId="4" captionText="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="2" pageNumber="405">Figure 2</figureCitation>
|
||
A). The ectosome is a collagenous cortex of variable thickness (10–200 µm), with collagen in the form of an amorphous matrix containing loose cells with inclusions (
|
||
<figureCitation id="133D2A3E1138635E872DF891FC8256B5" box="[795,891,1909,1934]" captionStart="FIGURE 2" captionStartId="4.[151,250,1429,1451]" captionTargetBox="[153,1436,218,1385]" captionTargetId="figure@4.[151,1436,213,1391]" captionTargetPageId="4" captionText="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="2" pageNumber="405">Figure 2</figureCitation>
|
||
A, B), and an external cuticle (
|
||
<figureCitation id="133D2A3E1138635E80E6F891FAC956B5" box="[1232,1328,1909,1934]" captionStart="FIGURE 3" captionStartId="5.[151,250,1653,1675]" captionTargetBox="[167,1419,581,1583]" captionTargetId="figure@5.[151,1436,577,1592]" captionTargetPageId="5" captionText="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="2" pageNumber="405">Figure 3</figureCitation>
|
||
A). Large channels and lacunae (subdermal spaces) are observed immediately underneath the ectosome (
|
||
<figureCitation id="133D2A3E1138635E80E8F87DFABA5689" box="[1246,1347,1945,1970]" captionStart="FIGURE 2" captionStartId="4.[151,250,1429,1451]" captionTargetBox="[153,1436,218,1385]" captionTargetId="figure@4.[151,1436,213,1391]" captionTargetPageId="4" captionText="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="2" pageNumber="405">Figure 2</figureCitation>
|
||
A). The choanosome consists of densely arranged oval-shaped choanocyte chambers (20–40 µm in diameter) (
|
||
<figureCitation id="133D2A3E1138635E8129F859FA7856ED" box="[1311,1409,1981,2006]" captionStart="FIGURE 2" captionStartId="4.[151,250,1429,1451]" captionTargetBox="[153,1436,218,1385]" captionTargetId="figure@4.[151,1436,213,1391]" captionTargetPageId="4" captionText="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="2" pageNumber="405">Figure 2</figureCitation>
|
||
A, C), and a mesohyl reinforced by collagen fibrils arranged in strands (
|
||
<figureCitation id="133D2A3E1139635F8791FF73FBF1518B" box="[935,1032,151,176]" captionStart="FIGURE 3" captionStartId="5.[151,250,1653,1675]" captionTargetBox="[167,1419,581,1583]" captionTargetId="figure@5.[151,1436,577,1592]" captionTargetPageId="5" captionText="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figure 3</figureCitation>
|
||
B, C), packed with various
|
||
<typeStatus id="54BD88191139635F8177FF7CFA85518B" box="[1345,1404,152,176]" pageId="3" pageNumber="406">types</typeStatus>
|
||
of cells with inclusions, and with abundant aquiferous canals. The chambers are densely and homogeneously distributed in some areas (
|
||
<figureCitation id="133D2A3E1139635F85F2FF3BFDDF51C3" box="[452,550,223,248]" captionStart="FIGURE 2" captionStartId="4.[151,250,1429,1451]" captionTargetBox="[153,1436,218,1385]" captionTargetId="figure@4.[151,1436,213,1391]" captionTargetPageId="4" captionText="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="3" pageNumber="406">Figure 2</figureCitation>
|
||
A, C), and rare in others that are instead densely packed by conglomerates of cells (
|
||
<figureCitation id="133D2A3E1139635F84ECFEE0FEC25027" box="[218,315,260,285]" captionStart="FIGURE 3" captionStartId="5.[151,250,1653,1675]" captionTargetBox="[167,1419,581,1583]" captionTargetId="figure@5.[151,1436,577,1592]" captionTargetPageId="5" captionText="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figure 3</figureCitation>
|
||
D). Choanocytes (4–6 µm in diameter) appear oval to triangular in shape (
|
||
<figureCitation id="133D2A3E1139635F80BEFEE0FB125027" box="[1160,1259,260,285]" captionStart="FIGURE 2" captionStartId="4.[151,250,1429,1451]" captionTargetBox="[153,1436,218,1385]" captionTargetId="figure@4.[151,1436,213,1391]" captionTargetPageId="4" captionText="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="3" pageNumber="406">Figure 2</figureCitation>
|
||
D). Larger cells with inclusions of various shapes and sizes (4–12 µm in diameter) can be observed in both the mesohyl and the ectosome (
|
||
<figureCitation id="133D2A3E1139635F8527FEA8FE65505F" box="[273,412,332,357]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStartId-0="4.[151,250,1429,1451]" captionStartId-1="5.[151,250,1653,1675]" captionTargetBox-0="[153,1436,218,1385]" captionTargetBox-1="[167,1419,581,1583]" captionTargetId-0="figure@4.[151,1436,213,1391]" captionTargetId-1="figure@5.[151,1436,577,1592]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionText-0="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." captionText-1="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri-0="https://zenodo.org/record/240006/files/figure.png" httpUri-1="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figures 2–3</figureCitation>
|
||
). The largest cells with inclusions (Ci1) are widespread in the mesohyl, in the cortex, and lining canals (
|
||
<figureCitation id="133D2A3E1139635F8505FE8BFE6B50B3" box="[307,402,367,392]" captionStart="FIGURE 3" captionStartId="5.[151,250,1653,1675]" captionTargetBox="[167,1419,581,1583]" captionTargetId="figure@5.[151,1436,577,1592]" captionTargetPageId="5" captionText="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figure 3</figureCitation>
|
||
B and 3D). There are some canals lined by these large cells with inclusions (
|
||
<figureCitation id="133D2A3E1139635F80DEFE8BFABE50B3" box="[1256,1351,367,392]" captionStart="FIGURE 3" captionStartId="5.[151,250,1653,1675]" captionTargetBox="[167,1419,581,1583]" captionTargetId="figure@5.[151,1436,577,1592]" captionTargetPageId="5" captionText="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figure 3</figureCitation>
|
||
D), and others without them (
|
||
<figureCitation id="133D2A3E1139635F85B3FE70FE1C5097" box="[389,485,404,429]" captionStart="FIGURE 3" captionStartId="5.[151,250,1653,1675]" captionTargetBox="[167,1419,581,1583]" captionTargetId="figure@5.[151,1436,577,1592]" captionTargetPageId="5" captionText="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figure 3</figureCitation>
|
||
C).
|
||
</paragraph>
|
||
<caption id="DF7966331139635F84A1FBE2FF255543" httpUri="https://zenodo.org/record/240005/files/figure.png" pageId="3" pageNumber="406" targetBox="[151,1436,481,1004]" targetPageId="3">
|
||
<paragraph id="8BB936BB1139635F84A1FBE2FF255543" blockId="3.[151,1436,1030,1144]" pageId="3" pageNumber="406">
|
||
<emphasis id="B972EAA91139635F84A1FBE2FEED5527" bold="true" box="[151,276,1030,1052]" pageId="3" pageNumber="406">FIGURE 1.</emphasis>
|
||
<taxonomicName id="4C064D381139635F852DFBECFE0F5527" box="[283,502,1030,1053]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="3" pageNumber="406" phylum="Porifera" rank="species" species="caribensis" status="gen. nov.">
|
||
<emphasis id="B972EAA91139635F852DFBECFE0F5527" box="[283,502,1030,1053]" italics="true" pageId="3" pageNumber="406">Vansoestia caribensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B972EAA91139635F85CBFBE3FD4F5527" bold="true" box="[509,694,1031,1052]" pageId="3" pageNumber="406">
|
||
<taxonomicNameLabel id="A24157D21139635F85CBFBE3FDA45527" box="[509,605,1031,1052]" pageId="3" pageNumber="406" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
sp. nov.
|
||
</emphasis>
|
||
growing on the edge of a dead plate coral. A. General habit showing its overall growth form, with algae and polychaete calcareous tubes mingled within the body. Scale bar = 2 cm. B. Close-up view of a more relaxed specimen showing the subdermal canals (Sc), 3–8 mm wide, and the profusely abundant pores. Scale bar = 1.2 cm
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BB936BB1139635F84F1FB44FDE3578B" blockId="3.[151,1437,1183,2037]" pageId="3" pageNumber="406">
|
||
TEM sections allowed further distinction of the various
|
||
<typeStatus id="54BD88191139635F8773FB44FC795583" box="[837,896,1184,1208]" pageId="3" pageNumber="406">types</typeStatus>
|
||
of cells with inclusions, and the observation of archaeocytes, choanocytes, and collagen fibrils. Spherulous cells 1 (
|
||
<figureCitation id="133D2A3E1139635F8791FB20FBF355E7" box="[935,1034,1220,1245]" captionStart="FIGURE 4" captionStartId="6.[151,250,1668,1690]" captionTargetBox="[151,1431,193,1640]" captionTargetId="figure@6.[151,1436,193,1646]" captionTargetPageId="6" captionText="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="3" pageNumber="406">Figure 4</figureCitation>
|
||
A, B and D) are ovoid to roundish slightly deformed (6–10 µm in diameter) with spherules (0.7–2 µm wide), containing microgranular inclusions that are dense but grey or black in color (0.06–0.15 µm in diameter), with a clear round nucleus (1.3–2 µm in diameter). By the size and distribution of spherules, these cells probably correspond to the cells with inclusions 1 (Sc1) observed in the histological cross sections (
|
||
<figureCitation id="133D2A3E1139635F86BAFAB0FCE05457" box="[652,793,1364,1389]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStartId-0="4.[151,250,1429,1451]" captionStartId-1="5.[151,250,1653,1675]" captionTargetBox-0="[153,1436,218,1385]" captionTargetBox-1="[167,1419,581,1583]" captionTargetId-0="figure@4.[151,1436,213,1391]" captionTargetId-1="figure@5.[151,1436,577,1592]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionText-0="FIGURE 2. Photomicrographs of Vansoestia caribensis gen. nov. sp. nov. histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “ amorphous matrix ” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm." captionText-1="FIGURE 3. Detail of the internal morphology of Vansoestia caribensis gen. nov. sp. nov. at 400 – 600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci 1, Ci 2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “ cell conglomerates ” and other types of cells, Scale = 32 µm." httpUri-0="https://zenodo.org/record/240006/files/figure.png" httpUri-1="https://zenodo.org/record/240007/files/figure.png" pageId="3" pageNumber="406">Figures 2–3</figureCitation>
|
||
). Spherulous cells 2 (
|
||
<figureCitation id="133D2A3E1139635F8025FAB0FB8E5457" box="[1043,1143,1364,1389]" captionStart="FIGURE 4" captionStartId="6.[151,250,1668,1690]" captionTargetBox="[151,1431,193,1640]" captionTargetId="figure@6.[151,1436,193,1646]" captionTargetPageId="6" captionText="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="3" pageNumber="406">Figure 4</figureCitation>
|
||
B) are similar in size and shape of Sc1, but their spherules are larger and ellipsoid in shape (0.1–0.3 µm in diameter), and electron dense dark. These cells might represent a different stage of the spherulous cell 1. A third
|
||
<typeStatus id="54BD88191139635F8000FA79FB90548E" box="[1078,1129,1437,1461]" pageId="3" pageNumber="406">type</typeStatus>
|
||
, which seems like vacuolar cells (
|
||
<figureCitation id="133D2A3E1139635F84ECFA5BFEC454E3" box="[218,317,1471,1496]" captionStart="FIGURE 4" captionStartId="6.[151,250,1668,1690]" captionTargetBox="[151,1431,193,1640]" captionTargetId="figure@6.[151,1436,193,1646]" captionTargetPageId="6" captionText="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="3" pageNumber="406">Figure 4</figureCitation>
|
||
C), occurs with variable shapes, smaller size (5–8 µm in diameter), and containing a single round, dark dense granule (0.4–0.8 µm in diameter) within each vacuole (1–1.5 µm in diameter). By the shape and size of the granules, these cells likely correspond to the cell with inclusion 2 observed in the histological cross sections. Finally, a microgranular cell with membrane-bound inclusions with striated content, was observed once (
|
||
<figureCitation id="133D2A3E1139635F8167F9C8FF5E5753" captionStart="FIGURE 4" captionStartId="6.[151,250,1668,1690]" captionTargetBox="[151,1431,193,1640]" captionTargetId="figure@6.[151,1436,193,1646]" captionTargetPageId="6" captionText="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="3" pageNumber="406">Figure 4</figureCitation>
|
||
D). A thick layer of collagen fibrils (Co) with loose bacteria surrounds the choanocyte chambers and long cytoplasmic projections could be seen in some choanocytes (
|
||
<figureCitation id="133D2A3E1139635F8773F990FC5E57B7" box="[837,935,1652,1677]" captionStart="FIGURE 4" captionStartId="6.[151,250,1668,1690]" captionTargetBox="[151,1431,193,1640]" captionTargetId="figure@6.[151,1436,193,1646]" captionTargetPageId="6" captionText="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="3" pageNumber="406">Figure 4</figureCitation>
|
||
E). One choanocyte with a basal nucleolated nucleus was observed (
|
||
<figureCitation id="133D2A3E1139635F85ACF973FE02578B" box="[410,507,1687,1712]" captionStart="FIGURE 4" captionStartId="6.[151,250,1668,1690]" captionTargetBox="[151,1431,193,1640]" captionTargetId="figure@6.[151,1436,193,1646]" captionTargetPageId="6" captionText="FIGURE 4. TEM sections of Vansoestia caribensis gen. nov. sp. nov. A. Roundish to ovoid Spherulous cells type 1 (Sc 1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm." httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="3" pageNumber="406">Figure 4</figureCitation>
|
||
F).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C31C65301139635484F1F958FA80529F" lastPageId="8" lastPageNumber="411" pageId="3" pageNumber="406" type="discussion">
|
||
<paragraph id="8BB936BB1139635F84F1F958FCB156CF" blockId="3.[151,1437,1183,2037]" pageId="3" pageNumber="406">
|
||
<emphasis id="B972EAA91139635F84F1F958FEC257EE" bold="true" box="[199,315,1724,1749]" pageId="3" pageNumber="406">Remarks.</emphasis>
|
||
The delicate and thin specimens of
|
||
<taxonomicName id="4C064D381139635F86C6F959FC1F57EF" box="[752,998,1724,1748]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="3" pageNumber="406" phylum="Porifera" rank="species" species="caribensis">
|
||
<emphasis id="B972EAA91139635F86C6F959FC1F57EF" box="[752,998,1724,1748]" italics="true" pageId="3" pageNumber="406">Vansoestia caribensis</emphasis>
|
||
</taxonomicName>
|
||
have been found in Bocas del Toro,
|
||
<collectingCountry id="F311762B1139635F84A1F904FF0A57CC" box="[151,243,1760,1783]" name="Panama" pageId="3" pageNumber="406">Panama</collectingCountry>
|
||
, since 2006; however, only the slow oxidation observed when specimens were placed into alcohol suggested an affinity to verongids. Due to the lack of a fiber skeleton, we initially affiliated this sponge to the genus
|
||
<taxonomicName id="4C064D381139635F84A1F8CCFEF3567B" box="[151,266,1832,1856]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="3" pageNumber="406" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA91139635F84A1F8CCFEF3567B" box="[151,266,1832,1856]" italics="true" pageId="3" pageNumber="406">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
. Histological observations corroborated its ianthellid nature (large, sac-shaped choanocyte chambers), but its distinct identity and closest affinity to the known sequences of fiber-bearing ianthellid genera from the Western Tropical Pacific appeared after analyzing 18S and 28S ribosomal gene sequences (Redmond
|
||
<emphasis id="B972EAA91139635F8122F895FAB456B3" box="[1300,1357,1904,1928]" italics="true" pageId="3" pageNumber="406">et al.</emphasis>
|
||
, 2013; Thacker
|
||
<emphasis id="B972EAA91139635F84CAF871FECF5697" box="[252,310,1940,1964]" italics="true" pageId="3" pageNumber="406">et al.</emphasis>
|
||
, 2013). Its genetic distance from the only previously known fiber-less verongid genus,
|
||
<taxonomicName id="4C064D381139635F8114F870FA6C5697" box="[1314,1429,1940,1964]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="3" pageNumber="406" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA91139635F8114F870FA6C5697" box="[1314,1429,1940,1964]" italics="true" pageId="3" pageNumber="406">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
, prompted the need to erect a new genus to classify this species. This biological record constitutes the first member of family
|
||
<taxonomicName id="4C064D381139635F8533F838FE7056CE" box="[261,393,2012,2037]" class="Demospongiae" family="Ianthellidae" kingdom="Animalia" order="Verongida" pageId="3" pageNumber="406" phylum="Porifera" rank="family">Ianthellidae</taxonomicName>
|
||
ever encountered in the Caribbean Sea.
|
||
</paragraph>
|
||
<caption id="DF796633113E635884A1FA71FA785759" httpUri="https://zenodo.org/record/240006/files/figure.png" pageId="4" pageNumber="407" targetBox="[153,1436,218,1385]" targetPageId="4">
|
||
<paragraph id="8BB936BB113E635884A1FA71FA785759" blockId="4.[151,1436,1428,1634]" pageId="4" pageNumber="407">
|
||
<emphasis id="B972EAA9113E635884A1FA71FEED5491" bold="true" box="[151,276,1429,1451]" pageId="4" pageNumber="407">FIGURE 2.</emphasis>
|
||
Photomicrographs of
|
||
<taxonomicName id="4C064D38113E635885CCFA72FD2F5491" box="[506,726,1428,1451]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="4" pageNumber="407" phylum="Porifera" rank="species" species="caribensis" status="gen. nov.">
|
||
<emphasis id="B972EAA9113E635885CCFA72FD2F5491" box="[506,726,1428,1451]" italics="true" pageId="4" pageNumber="407">Vansoestia caribensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B972EAA9113E635886E8FA71FC615491" bold="true" box="[734,920,1429,1450]" pageId="4" pageNumber="407">
|
||
<taxonomicNameLabel id="A24157D2113E635886E8FA71FCC75491" box="[734,830,1429,1450]" pageId="4" pageNumber="407" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
sp. nov.
|
||
</emphasis>
|
||
histological cross sections. A. Upper body region showing a thin ectosome (Ec) followed by wide subdermal canals (Sc) and the choanoderm with densely arranged choanocyte chambers (Cc). Scale = 110 µm. B. Detail of an area of the ectosome showing a dense “amorphous matrix” (Am) of collagen with dispersed dark stained cells with inclusions (Ci), Scale = 30 µm; C. Basal region where the sponge is attached to the substrate. Note the corallites (Co) of the dead coral where the sponge was growing, the choanocyte chambers (Cc) and canals (Ca), Scale = 200 µm. D. Detail of choanocyte chamber with more than 20 choanocytes (oval to triangular in shape). Notice at least two types of cells with inclusions (Ci) and collagen strands (Cs) in between the chambers and the cells, Scale = 15 µm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BB936BB113E635984F1F96EFCAB51C3" blockId="4.[151,1436,1674,2023]" lastBlockId="5.[151,1436,151,501]" lastPageId="5" lastPageNumber="408" pageId="4" pageNumber="407">
|
||
There is a striking similarity between
|
||
<taxonomicName id="4C064D38113E63588647F96FFD065799" box="[625,767,1674,1698]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="4" pageNumber="407" phylum="Porifera" rank="species" species="caribensis">
|
||
<emphasis id="B972EAA9113E63588647F96FFD065799" box="[625,767,1674,1698]" italics="true" pageId="4" pageNumber="407">V. caribensis</emphasis>
|
||
</taxonomicName>
|
||
gen nov. sp. nov. and the external appearance of
|
||
<taxonomicName id="4C064D38113E6358811FF96EFD3657FC" authority="Topsent, 1896 (Reveillaud et al., 2012)" authorityName="Topsent, 1896 (Reveillaud et al." authorityYear="2012" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="4" pageNumber="407" phylum="Porifera" rank="species" species="pruvoti">
|
||
<emphasis id="B972EAA9113E6358811FF96EFF1157FD" italics="true" pageId="4" pageNumber="407">Hexadella pruvoti</emphasis>
|
||
<bibRefCitation id="EF974B4A113E635884C3F94BFE6357FC" author="Topsent" box="[245,410,1710,1735]" pageId="4" pageNumber="407" refString="Topsent, E. (1896 a) Materiaux pour servir a l'etude de la faune des spongiaires de France. Memoires de la Societe zoologique de France, 9, 113 - 133" type="journal volume" year="1896">Topsent, 1896</bibRefCitation>
|
||
(
|
||
<bibRefCitation id="EF974B4A113E63588599F94AFD3E57FC" author="Reveillaud" box="[431,711,1710,1735]" pageId="4" pageNumber="407" refString="Reveillaud, J., Allewaert, C., Perez, T., Vacelet, J., Banaigs, B. & Vanreusel, A. (2012) Relevance of an integrative approach for taxonomic revision in sponge taxa: case study of the shallow-water Atlanto-Mediterranean Hexadella species (Porifera: Ianthellidae: Verongida). Invertebrate Systematics, 26, 230 - 248. http: // dx. doi. org / 10.1071 / IS 11044" type="journal article" year="2012">
|
||
Reveillaud
|
||
<emphasis id="B972EAA9113E6358860EF94BFD8E57FD" box="[568,631,1710,1734]" italics="true" pageId="4" pageNumber="407">et al.</emphasis>
|
||
, 2012
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, particularly with respect to the abundant subdermal canals converging in a wide oscule, and its yellowish coloration. However, 18S and 28S gene sequence analyses confirmed the very distinct nature of these two species. A cellularized cortex of variable thickness (10–200 µm thick) composed of amorphous collagen and various
|
||
<typeStatus id="54BD8819113E635886D1F8FFFCDB5608" box="[743,802,1819,1843]" pageId="4" pageNumber="407">types</typeStatus>
|
||
of cells is present in
|
||
<taxonomicName id="4C064D38113E63588038F8FFFB635609" box="[1038,1178,1818,1842]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="4" pageNumber="407" phylum="Porifera" rank="subSpecies" species="cari" status="gen. nov." subSpecies="bensis">
|
||
<emphasis id="B972EAA9113E63588038F8FFFB635609" box="[1038,1178,1818,1842]" italics="true" pageId="4" pageNumber="407">V. cari bensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B972EAA9113E63588092F8FEFA945608" bold="true" box="[1188,1389,1818,1843]" pageId="4" pageNumber="407">
|
||
<taxonomicNameLabel id="A24157D2113E63588092F8FEFAF45608" box="[1188,1293,1818,1843]" pageId="4" pageNumber="407" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
sp. nov.
|
||
</emphasis>
|
||
and forms an important morphological difference between these two genera, since
|
||
<taxonomicName id="4C064D38113E63588037F8DAFB8D566D" box="[1025,1140,1854,1878]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="4" pageNumber="407" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113E63588037F8DAFB8D566D" box="[1025,1140,1854,1878]" italics="true" pageId="4" pageNumber="407">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
possesses a thinner cortex with a 1 µm thick cuticle. A well-developed cellularized cortex might represent a potential synapomorphy between
|
||
<taxonomicName id="4C064D38113E635884A1F863FEF556A5" box="[151,268,1927,1950]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="4" pageNumber="407" phylum="Porifera" rank="genus" status="gen. nov.">
|
||
<emphasis id="B972EAA9113E635884A1F863FEF556A5" box="[151,268,1927,1950]" italics="true" pageId="4" pageNumber="407">Vansoestia</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A24157D2113E6358852EF863FE7856A5" box="[280,385,1927,1951]" pageId="4" pageNumber="407" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
,
|
||
<taxonomicName id="4C064D38113E635885A7F862FE0F56A5" box="[401,502,1926,1950]" class="Demospongiae" family="Ianthellidae" genus="Ianthella" kingdom="Animalia" order="Verongida" pageId="4" pageNumber="407" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113E635885A7F862FE0F56A5" box="[401,502,1926,1950]" italics="true" pageId="4" pageNumber="407">Ianthella</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C064D38113E6358860FF862FD1556A5" box="[569,748,1926,1950]" class="Demospongiae" family="Ianthellidae" genus="Anomoianthella" kingdom="Animalia" order="Verongida" pageId="4" pageNumber="407" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113E6358860FF862FD1556A5" box="[569,748,1926,1950]" italics="true" pageId="4" pageNumber="407">Anomoianthella</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="EF974B4A113E635886CBF862FC4F56A4" author="Bergquist" box="[765,950,1926,1951]" pageId="4" pageNumber="407" refString="Bergquist, P. R. (1980) A revision of the supraspecific classification of the orders Dictyoceratida, Dendroceratida and Verongida (class Demospongiae). New Zealand Journal of Zoology, 7 (4), 443 - 503." type="journal article" year="1980">Bergquist, 1980</bibRefCitation>
|
||
, 1995;
|
||
<bibRefCitation id="EF974B4A113E6358803BF862FA6E56A4" author="Bergquist" box="[1037,1431,1926,1951]" pageId="4" pageNumber="407" refString="Bergquist, P. R. & Kelly-Borges, M. (1995) Systematics and biogeography of the genus Ianthella (Demospongiae: Verongida: Ianthellidae) in the South-West Pacific. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 12, 151 - 176." type="journal article" year="1995">Bergquist and Kelly-Borges, 1995</bibRefCitation>
|
||
; Bergquist and Cook, 2002; present study). Indeed, Bergquist, (1995) and
|
||
<bibRefCitation id="EF974B4A113E63588037F84EFA6556F8" author="Bergquist" box="[1025,1436,1962,1987]" pageId="4" pageNumber="407" refString="Bergquist, P. R. & Kelly-Borges, M. (1995) Systematics and biogeography of the genus Ianthella (Demospongiae: Verongida: Ianthellidae) in the South-West Pacific. The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 12, 151 - 176." type="journal article" year="1995">Bergquist and Kelly-Borges (1995)</bibRefCitation>
|
||
described a collagenous ectosome (70 to 300 µm thick) containing cells for all
|
||
<taxonomicName id="4C064D38113E63588027F82AFB8F56DD" box="[1041,1142,1998,2022]" class="Demospongiae" family="Ianthellidae" genus="Ianthella" kingdom="Animalia" order="Verongida" pageId="4" pageNumber="407" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113E63588027F82AFB8F56DD" box="[1041,1142,1998,2022]" italics="true" pageId="4" pageNumber="407">Ianthella</emphasis>
|
||
</taxonomicName>
|
||
species and a cellularized cuticle for
|
||
<taxonomicName id="4C064D38113F63598522FF7CFE3E518B" box="[276,455,152,176]" class="Demospongiae" family="Ianthellidae" genus="Anomoianthella" kingdom="Animalia" order="Verongida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113F63598522FF7CFE3E518B" box="[276,455,152,176]" italics="true" pageId="5" pageNumber="408">Anomoianthella</emphasis>
|
||
</taxonomicName>
|
||
species (
|
||
<bibRefCitation id="EF974B4A113F63598607FF73FD12518B" author="Bergquist" box="[561,747,151,176]" pageId="5" pageNumber="408" refString="Bergquist, P. R. (1980) A revision of the supraspecific classification of the orders Dictyoceratida, Dendroceratida and Verongida (class Demospongiae). New Zealand Journal of Zoology, 7 (4), 443 - 503." type="journal article" year="1980">Bergquist, 1980</bibRefCitation>
|
||
). Together with the presence of a collagen-reinforced cortex combined with a cuticle, the high degree of reinforcement of the choanosome with collagen is also a characteristic shared by
|
||
<taxonomicName id="4C064D38113F6359853CFF04FE7B51CC" box="[266,386,224,247]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus" status="gen. nov.">
|
||
<emphasis id="B972EAA9113F6359853CFF04FE7B51CC" box="[266,386,224,247]" italics="true" pageId="5" pageNumber="408">Vansoestia</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A24157D2113F635985BFFF04FE1651CC" box="[393,495,224,248]" pageId="5" pageNumber="408" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
,
|
||
<taxonomicName id="4C064D38113F635985CAFF04FD9851C3" box="[508,609,224,248]" class="Demospongiae" family="Ianthellidae" genus="Ianthella" kingdom="Animalia" order="Verongida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113F635985CAFF04FD9851C3" box="[508,609,224,248]" italics="true" pageId="5" pageNumber="408">Ianthella</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis id="B972EAA9113F635986AEFF04FCAB51C3" box="[664,850,224,248]" italics="true" pageId="5" pageNumber="408">
|
||
<taxonomicName id="4C064D38113F635986AEFF04FCB651C3" box="[664,847,224,248]" class="Demospongiae" family="Ianthellidae" genus="Anomoianthella" kingdom="Animalia" order="Verongida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus">Anomoianthella</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph id="8BB936BB113F635984F1FEE0FB0350CE" blockId="5.[151,1436,151,501]" pageId="5" pageNumber="408">
|
||
Some similarities among the spherulous cells of
|
||
<taxonomicName id="4C064D38113F635986D5FEE1FC885027" box="[739,881,260,284]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="5" pageNumber="408" phylum="Porifera" rank="species" species="caribensis" status="gen. nov.">
|
||
<emphasis id="B972EAA9113F635986D5FEE1FC885027" box="[739,881,260,284]" italics="true" pageId="5" pageNumber="408">V. caribensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B972EAA9113F6359874EFEE0FBB95026" bold="true" box="[888,1088,260,285]" pageId="5" pageNumber="408">
|
||
<taxonomicNameLabel id="A24157D2113F6359874EFEE0FC185026" box="[888,993,260,285]" pageId="5" pageNumber="408" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
sp. nov.
|
||
</emphasis>
|
||
, and other verongids were also noticed (
|
||
<bibRefCitation id="EF974B4A113F635984CBFEC3FE59507B" author="Vacelet" box="[253,416,295,320]" pageId="5" pageNumber="408" refString="Vacelet J. (1967) Les cellules a inclusions de l'eponge cornee Verongia cavernicola. Journal de Microscopie, 6, 237 - 240." type="journal article" year="1967">Vacelet, 1967</bibRefCitation>
|
||
). The cells with striated membrane-bound bodies are very similar to cells observed in
|
||
<taxonomicName id="4C064D38113F635984A1FEA8FE87505F" box="[151,382,332,356]" class="Demospongiae" family="Aplysinidae" genus="Aplysina" kingdom="Animalia" order="Verongida" pageId="5" pageNumber="408" phylum="Porifera" rank="species" species="aerophoba">
|
||
<emphasis id="B972EAA9113F635984A1FEA8FE87505F" box="[151,382,332,356]" italics="true" pageId="5" pageNumber="408">Aplysina aerophoba</emphasis>
|
||
</taxonomicName>
|
||
by
|
||
<bibRefCitation id="EF974B4A113F63598583FEA8FD69505F" author="Maldonado" box="[437,656,332,357]" pageId="5" pageNumber="408" refString="Maldonado, M. (2009) Embryonic development of verongid demosponges supports the independent acquisition of spongin skeletons as an alternative to the siliceous skeleton of sponges. Biological Journal of the Linnaean Society, 97, 427 - 447. http: // dx. doi. org / 10.1111 / j. 1095 - 8312.2009.01202. x" type="journal article" year="2009">Maldonado (2009)</bibRefCitation>
|
||
. The dark granules found in cells of
|
||
<taxonomicName id="4C064D38113F6359806AFEA9FB2D505F" box="[1116,1236,333,356]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus" status="gen. nov.">
|
||
<emphasis id="B972EAA9113F6359806AFEA9FB2D505F" box="[1116,1236,333,356]" italics="true" pageId="5" pageNumber="408">Vansoestia</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A24157D2113F635980D7FEA9FAB7505F" box="[1249,1358,333,357]" pageId="5" pageNumber="408" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
might represent glycogen granules (
|
||
<bibRefCitation id="EF974B4A113F635985D4FE8BFD5450B3" author="Maldonado" box="[482,685,367,392]" pageId="5" pageNumber="408" refString="Maldonado, M. (2009) Embryonic development of verongid demosponges supports the independent acquisition of spongin skeletons as an alternative to the siliceous skeleton of sponges. Biological Journal of the Linnaean Society, 97, 427 - 447. http: // dx. doi. org / 10.1111 / j. 1095 - 8312.2009.01202. x" type="journal article" year="2009">Maldonado, 2009</bibRefCitation>
|
||
), or they might contain brominated compounds (Thompson, 1986) like has been found for microgranular and spherulous cells of two different
|
||
<taxonomicName id="4C064D38113F635987D2FE70FBBE5097" box="[996,1095,404,428]" class="Demospongiae" family="Aplysinidae" genus="Aplysina" kingdom="Animalia" order="Verongida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113F635987D2FE70FBBE5097" box="[996,1095,404,428]" italics="true" pageId="5" pageNumber="408">Aplysina</emphasis>
|
||
</taxonomicName>
|
||
species. The large diversity of cells with inclusions, and their similarities with species of
|
||
<taxonomicName id="4C064D38113F63598719FE5CFC5B50EB" box="[815,930,440,464]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="5" pageNumber="408" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113F63598719FE5CFC5B50EB" box="[815,930,440,464]" italics="true" pageId="5" pageNumber="408">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
and other verongid genera invites continued research to understand the morphology and physiological functions of these cells and their products.
|
||
</paragraph>
|
||
<caption id="DF796633113F635984A1F991FE955678" httpUri="https://zenodo.org/record/240007/files/figure.png" pageId="5" pageNumber="408" targetBox="[167,1419,581,1583]" targetPageId="5">
|
||
<paragraph id="8BB936BB113F635984A1F991FE955678" blockId="5.[151,1436,1653,1859]" pageId="5" pageNumber="408">
|
||
<emphasis id="B972EAA9113F635984A1F991FEEE57B0" bold="true" box="[151,279,1653,1675]" pageId="5" pageNumber="408">FIGURE 3.</emphasis>
|
||
Detail of the internal morphology of
|
||
<taxonomicName id="4C064D38113F63598684F993FC6857B0" box="[690,913,1653,1676]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="5" pageNumber="408" phylum="Porifera" rank="species" species="caribensis" status="gen. nov.">
|
||
<emphasis id="B972EAA9113F63598684F993FC6857B0" box="[690,913,1653,1676]" italics="true" pageId="5" pageNumber="408">Vansoestia caribensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B972EAA9113F635987AAF992FBA657B0" bold="true" box="[924,1119,1654,1675]" pageId="5" pageNumber="408">
|
||
<taxonomicNameLabel id="A24157D2113F635987AAF992FC0657B0" box="[924,1023,1654,1675]" pageId="5" pageNumber="408" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
sp. nov.
|
||
</emphasis>
|
||
at 400–600 X. A. Ectosomal cellularized cortex, where a 1 µm cuticle (Cu) lining the outer surface of the sponge is evidenced, Scale = 13 µm. B. Various cells with inclusions (Ci1, Ci2) and smaller choanocytes (Ch) embedded in a mesohyl with abundant strands of collagen (Cs), Scale = 16 µm. C. Portion of the sponge body in contact with the substrate (Co), showing abundance of collagen strands (Cs), and cells. Notice an aquiferous canal (Ac) that is not lined by larger cells with inclusions, Scale = 25 µm. D. Detail of the mesohyl showing an aquiferous canal that is lined by cells with inclusions type 1, and “cell conglomerates” and other types of cells, Scale = 32 µm.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF796633113C635A84A1F960FB9456B4" httpUri="https://zenodo.org/record/240008/files/figure.png" pageId="6" pageNumber="409" targetBox="[151,1431,193,1640]" targetPageId="6">
|
||
<paragraph id="8BB936BB113C635A84A1F960FB9456B4" blockId="6.[151,1436,1668,1935]" pageId="6" pageNumber="409">
|
||
<emphasis id="B972EAA9113C635A84A1F960FEEB57A2" bold="true" box="[151,274,1668,1690]" pageId="6" pageNumber="409">FIGURE 4.</emphasis>
|
||
TEM sections of
|
||
<taxonomicName id="4C064D38113C635A85F0F961FD5857A1" box="[454,673,1668,1690]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="6" pageNumber="409" phylum="Porifera" rank="species" species="caribensis" status="gen. nov.">
|
||
<emphasis id="B972EAA9113C635A85F0F961FD5857A1" box="[454,673,1668,1690]" italics="true" pageId="6" pageNumber="409">Vansoestia caribensis</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B972EAA9113C635A8691F960FCA557A2" bold="true" box="[679,860,1668,1689]" pageId="6" pageNumber="409">
|
||
<taxonomicNameLabel id="A24157D2113C635A8691F960FCFC57A2" box="[679,773,1668,1689]" pageId="6" pageNumber="409" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
sp. nov.
|
||
</emphasis>
|
||
A. Roundish to ovoid Spherulous cells type 1 (Sc1) with nucleus (Nu) bearing round spherules containing light and dark inclusions, and abundant collagen fibrils (Co), Scale = 2.5 µm. B. Sperulous cell type 2 with electron dense dark granules, and bacteria within it, close to an aquiferous channel (Ac), Scale = 2 µm. Bacteria can be seen in the interior of the channel. C. Vacuolar cells (Sc 3) seen in the mesohyl close to an aquiferous channel, Scale = 2 µm. D. A microgranular cell (Mc) with striated membrane-bound bodies, among spherulous cells 1, and 2. Note the presence of abundant collagen fibrils (Co) between the cells, Scale = 1.3 µm. E. Portion of a choanocyte chamber where main body of the choanocytes can be seen, some with cytoplasmatic projections (Cy). A collagen fibril layer (Co) is found surrounding all choanocyte chambers, Scale = 1.6 µm. F. A triangular shaped choanocyte (Ch) with a basal nucleolated nucleus, contiguous to a layer of fibrillar collagen (Co) containing bacteria (Ba), Scale = 1.3 µm.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF796633113D635B84A1FB5AFBFA5758" httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="7" pageNumber="410" targetBox="[160,1428,204,1176]" targetPageId="7">
|
||
<paragraph id="8BB936BB113D635B84A1FB5AFBFA5758" blockId="7.[151,1436,1214,1635]" pageId="7" pageNumber="410">
|
||
<emphasis id="B972EAA9113D635B84A1FB5AFEE055EF" bold="true" box="[151,281,1214,1236]" pageId="7" pageNumber="410">FIGURE 5.</emphasis>
|
||
Schematic representation of the phylogenetic relationships of the genera of
|
||
<taxonomicName id="4C064D38113D635B8053FB5AFB2955EF" box="[1125,1232,1214,1236]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS2+28S]; Díaz
|
||
<emphasis id="B972EAA9113D635B8798FB3AFC1B55C8" box="[942,994,1245,1267]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013 [cox1]; Redmond
|
||
<emphasis id="B972EAA9113D635B80D4FB3AFAEF55C8" box="[1250,1302,1245,1267]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013 [18S]; Thacker
|
||
<emphasis id="B972EAA9113D635B84C5FB19FED35429" box="[243,298,1276,1298]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013 [28S];). Black circles indicate nodes with 100% maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100% maximum likelihood bootstrap support in 1 study, and none under 85%; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1.
|
||
<taxonomicName id="4C064D38113D635B857EFABCFE495455" box="[328,432,1368,1390]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family
|
||
<taxonomicName id="4C064D38113D635B84A1FA92FED254B7" box="[151,299,1398,1420]" class="Demospongiae" family="Chondrosiidae" kingdom="Animalia" order="Chondrosida" pageId="7" pageNumber="410" phylum="Porifera" rank="family">Chondrosiidae</taxonomicName>
|
||
(
|
||
<emphasis id="B972EAA9113D635B850CFA93FE8B54B7" box="[314,370,1399,1420]" italics="true" pageId="7" pageNumber="410">sensu</emphasis>
|
||
Redmond
|
||
<emphasis id="B972EAA9113D635B85D5FA93FDEF54B7" box="[483,534,1398,1420]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the
|
||
<taxonomicName id="4C064D38113D635B84A1FA71FEFB5490" box="[151,258,1429,1451]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by
|
||
<taxonomicName id="4C064D38113D635B80C1FA71FAA65490" box="[1271,1375,1429,1451]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113D635B80C1FA71FAA65490" box="[1271,1375,1429,1451]" italics="true" pageId="7" pageNumber="410">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62%) containing all the non
|
||
<emphasis id="B972EAA9113D635B81A2FA51FF0654D3" italics="true" pageId="7" pageNumber="410">
|
||
-
|
||
<taxonomicName id="4C064D38113D635B84A1FA36FF0654D3" box="[151,255,1490,1512]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">Hexadella</taxonomicName>
|
||
</emphasis>
|
||
classical genera of
|
||
<taxonomicName id="4C064D38113D635B85E5FA36FDC754D3" box="[467,574,1490,1512]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
; 6. Moderately supported clade (85%) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BB936BB113D635484F1F96EFAA15323" blockId="7.[151,1436,1674,2023]" lastBlockId="8.[151,1437,151,933]" lastPageId="8" lastPageNumber="411" pageId="7" pageNumber="410">
|
||
<emphasis id="B972EAA9113D635B84F1F96EFE345798" bold="true" box="[199,461,1674,1699]" pageId="7" pageNumber="410">Phylogenetic remarks</emphasis>
|
||
. 18S and 28S sequences demonstrate that this fiber-less member of
|
||
<taxonomicName id="4C064D38113D635B80D0F96EFA935798" box="[1254,1386,1674,1699]" class="Demospongiae" family="Ianthellidae" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="family">Ianthellidae</taxonomicName>
|
||
is a sister group to the clade that includes
|
||
<taxonomicName id="4C064D38113D635B8677F94AFD5F57FD" box="[577,678,1710,1734]" class="Demospongiae" family="Ianthellidae" genus="Ianthella" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113D635B8677F94AFD5F57FD" box="[577,678,1710,1734]" italics="true" pageId="7" pageNumber="410">Ianthella</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis id="B972EAA9113D635B86EBF94AFC6E57FD" box="[733,919,1710,1734]" italics="true" pageId="7" pageNumber="410">
|
||
<taxonomicName id="4C064D38113D635B86EBF94AFC6A57FD" box="[733,915,1710,1734]" class="Demospongiae" family="Ianthellidae" genus="Anomoianthella" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">Anomoianthella</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
(Redmond
|
||
<emphasis id="B972EAA9113D635B802AF94BFBAD57FD" box="[1052,1108,1710,1734]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013; Thacker
|
||
<emphasis id="B972EAA9113D635B813BF94BFABC57FD" box="[1293,1349,1710,1734]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013).
|
||
<figureCitation id="133D2A3E113D635B84A1F936FF0F57D0" box="[151,246,1746,1771]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="7" pageNumber="410">Figure 5</figureCitation>
|
||
summarizes the current phylogenetic interpretation of the relationships among verongid genera based on the latest molecular, and morphologic studies of the group (
|
||
<bibRefCitation id="EF974B4A113D635B8706F912FBA95634" author="Erwin" box="[816,1104,1782,1807]" pageId="7" pageNumber="410" refString="Erwin, P. M. & Thacker, R. W. (2007) Phylogenetic analyses of marine sponges within the order Verongida: a comparison of morphological and molecular data. Invertebrate Biology, 126, 220 - 234. http: // dx. doi. org / 10.1111 / j. 1744 - 7410.2007.00092. x" type="journal article" year="2007">Erwin and Thacker, 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EF974B4A113D635B806AF912FAA25634" author="Erpenbeck" box="[1116,1371,1782,1807]" pageId="7" pageNumber="410" refString="Erpenbeck, D., Sutcliffe, P., Cook, S., Dietzel, A., Maldonado, M., van Soest, R. W. M., Hooper, J. & Worheide, G. (2012) Horny sponges and their affairs on the phylogenetic relationships of keratose sponges. Molecular Phylogenetics and Evolution, 63, 809 - 816. http: // dx. doi. org / 10.1016 / j. ympev. 2012.02.024" type="journal article" year="2012">
|
||
Erpenbeck
|
||
<emphasis id="B972EAA9113D635B80ECF913FAEB5635" box="[1242,1298,1782,1806]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2012
|
||
</bibRefCitation>
|
||
; Díaz
|
||
<emphasis id="B972EAA9113D635B84A1F8FFFF215608" box="[151,216,1819,1843]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013; Redmond
|
||
<emphasis id="B972EAA9113D635B858DF8FFFE055608" box="[443,508,1819,1843]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013; Thacker
|
||
<emphasis id="B972EAA9113D635B86F8F8FFFCF65608" box="[718,783,1819,1843]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013). The two sister clades make up the subclass Myxospongiae, one containing the amended
|
||
<taxonomicName id="4C064D38113D635B8692F8DAFCEE566C" box="[676,791,1854,1879]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="7" pageNumber="410" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
(after Redmond
|
||
<emphasis id="B972EAA9113D635B87D5F8DBFBD9566D" box="[995,1056,1854,1878]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013) and the other containing
|
||
<taxonomicName id="4C064D38113D635B84A1F887FEEB5640" box="[151,274,1891,1915]" class="Demospongiae" family="Chondrillidae" genus="Chondrilla" kingdom="Animalia" order="Chondrosida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113D635B84A1F887FEEB5640" box="[151,274,1891,1915]" italics="true" pageId="7" pageNumber="410">Chondrilla</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C064D38113D635B8565F887FE3B5640" box="[339,450,1891,1915]" class="Demospongiae" family="Halisarcidae" genus="Halisarca" kingdom="Animalia" order="Chondrosida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113D635B8565F887FE3B5640" box="[339,450,1891,1915]" italics="true" pageId="7" pageNumber="410">Halisarca</emphasis>
|
||
</taxonomicName>
|
||
(Order Chondrillida
|
||
<emphasis id="B972EAA9113D635B86F6F887FD075641" box="[704,766,1891,1914]" italics="true" pageId="7" pageNumber="410">sensu</emphasis>
|
||
Redmond
|
||
<emphasis id="B972EAA9113D635B87B2F887FC3B5640" box="[900,962,1891,1915]" italics="true" pageId="7" pageNumber="410">et al.</emphasis>
|
||
, 2013). All verongids are grouped in a strongly supported monophyletic clade (
|
||
<figureCitation id="133D2A3E113D635B8657F862FD3D56A4" box="[609,708,1926,1951]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="7" pageNumber="410">Figure 5</figureCitation>
|
||
, clade 1), which shares the presence of collagen fibrils forming bundles either in the cortex, in the mesohyl, or within fibers.
|
||
<taxonomicName id="4C064D38113D635B8754F84FFC1C56F8" box="[866,997,1963,1987]" class="Demospongiae" family="Chondrosiidae" genus="Chondrosia" kingdom="Animalia" order="Chondrosida" pageId="7" pageNumber="410" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113D635B8754F84FFC1C56F8" box="[866,997,1963,1987]" italics="true" pageId="7" pageNumber="410">Chondrosia</emphasis>
|
||
</taxonomicName>
|
||
(family
|
||
<taxonomicName id="4C064D38113D635B807DF84EFB0956F8" box="[1099,1264,1962,1987]" class="Demospongiae" family="Chondrosiidae" kingdom="Animalia" order="Chondrosida" pageId="7" pageNumber="410" phylum="Porifera" rank="family">Chondrosiidae</taxonomicName>
|
||
) forms a wellsupported sister clade to all other verongids (
|
||
<figureCitation id="133D2A3E113D635B86B9F82AFD0956DC" box="[655,752,1998,2023]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="7" pageNumber="410">Figure 5</figureCitation>
|
||
, clade 2), and is represented by sponges with extremely high collagen content, a thick (>
|
||
<quantity id="4CFE9B5E1132635485F8FF7CFDF75194" box="[462,526,152,176]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="411" unit="mm" value="1.0">1 mm</quantity>
|
||
thick) “collagenous cortex”, and a fiber-less body. The rest of the verongids form a highly supported clade (
|
||
<figureCitation id="133D2A3E113263548580FF58FDEE51EE" box="[438,535,188,213]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
, clade 3), containing all taxa that produce brominated secondary metabolites and have abundant spherulous cells. Basal within this group, a highly supported clade (
|
||
<figureCitation id="133D2A3E113263548070FF3BFB5351C3" box="[1094,1194,223,248]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
, clade 4) contains all
|
||
<taxonomicName id="4C064D381132635484A1FEE0FEF35027" box="[151,266,260,284]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA91132635484A1FEE0FEF35027" box="[151,266,260,284]" italics="true" pageId="8" pageNumber="411">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
species which share eurypylous choanocyte chambers, a thin cuticle (1–2 µm) and a very thin ectosome. The next clade (
|
||
<figureCitation id="133D2A3E11326354857DFEC3FE54507B" box="[331,429,295,320]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
, clade 5) is a moderately supported clade containing all non-
|
||
<taxonomicName id="4C064D3811326354806EFECCFB32507B" box="[1112,1227,296,320]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA911326354806EFECCFB32507B" box="[1112,1227,296,320]" italics="true" pageId="8" pageNumber="411">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
verongids. Within clade 5, two clades are distinguished: a moderately (85%) supported clade (
|
||
<figureCitation id="133D2A3E11326354800FFEA8FB5B505E" box="[1081,1186,332,357]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
, clade 6) containing
|
||
<taxonomicName id="4C064D381132635484A1FE94FEF550BC" box="[151,268,368,391]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus" status="gen. nov.">
|
||
<emphasis id="B972EAA91132635484A1FE94FEF550BC" box="[151,268,368,391]" italics="true" pageId="8" pageNumber="411">Vansoestia</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A24157D2113263548520FE94FE8550BC" box="[278,380,368,392]" pageId="8" pageNumber="411" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
(skeleton less) and the fiber bearing Pacific taxa of genera
|
||
<taxonomicName id="4C064D38113263548020FE94FB8250B3" box="[1046,1147,368,392]" class="Demospongiae" family="Ianthellidae" genus="Ianthella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113263548020FE94FB8250B3" box="[1046,1147,368,392]" italics="true" pageId="8" pageNumber="411">Ianthella</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis id="B972EAA9113263548084FE94FA9C50B3" box="[1202,1381,368,392]" italics="true" pageId="8" pageNumber="411">Anamoianthella</emphasis>
|
||
, and a moderately to highly supported clade (
|
||
<figureCitation id="133D2A3E113263548650FE70FD335096" box="[614,714,404,429]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
, clade 7) including all the taxa possessing diplodal choanocyte chambers. Within clade 7 (
|
||
<figureCitation id="133D2A3E1132635485E4FE53FDCE50EB" box="[466,567,439,464]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
), a clearly monophyletic group of sponges (clade 8) contains all
|
||
<taxonomicName id="4C064D3811326354810CFE5CFA6550EB" box="[1338,1436,440,464]" class="Demospongiae" family="Aplysinidae" genus="Aplysina" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA911326354810CFE5CFA6550EB" box="[1338,1436,440,464]" italics="true" pageId="8" pageNumber="411">Aplysina</emphasis>
|
||
</taxonomicName>
|
||
species, whereas a poorly supported clade (clade 9) contains at least six genera (
|
||
<emphasis id="B972EAA911326354804AFE38FD485323" italics="true" pageId="8" pageNumber="411">
|
||
<taxonomicName id="4C064D3811326354804AFE38FAFD50CF" box="[1148,1284,476,500]" class="Demospongiae" family="Aplysinidae" genus="Aiolochroia" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Aiolochroia</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C064D38113263548121FE38FA6150CF" box="[1303,1432,476,500]" class="Demospongiae" family="Aplysinellidae" genus="Aplysinella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Aplysinella</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C064D381132635484A1FDE4FEF45323" box="[151,269,512,536]" class="Demospongiae" family="Aplysinellidae" genus="Porphyria" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Porphyria</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C064D3811326354852EFDE4FE325323" box="[280,459,512,536]" class="Demospongiae" family="Pseudoceratinidae" genus="Pseudoceratina" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Pseudoceratina</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C064D381132635485E0FDE4FDCA5323" box="[470,563,512,536]" class="Demospongiae" family="Aplysinellidae" genus="Suberea" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Suberea</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C064D38113263548609FDE4FD485323" box="[575,689,512,536]" class="Demospongiae" family="Aplysinidae" genus="Verongula" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Verongula</taxonomicName>
|
||
</emphasis>
|
||
) and an undescribed genus from Moorea (Díaz
|
||
<emphasis id="B972EAA91132635480FFFDE5FAF85323" box="[1225,1281,512,536]" italics="true" pageId="8" pageNumber="411">et al.</emphasis>
|
||
, 2013).
|
||
</paragraph>
|
||
<paragraph id="8BB936BB1132635484F1FDC0FB7F522F" blockId="8.[151,1437,151,933]" pageId="8" pageNumber="411">
|
||
The
|
||
<taxonomicName id="4C064D381132635484C9FDC0FE8B5306" box="[255,370,548,573]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
taxa with eurypylous, sac-shaped choanocyte chambers are placed in two distinct clades (
|
||
<figureCitation id="133D2A3E1132635484A8FDA3FEF9535B" box="[158,256,583,608]" captionStart="FIGURE 5" captionStartId="7.[151,250,1214,1236]" captionTargetBox="[160,1428,204,1176]" captionTargetId="figure@7.[151,1435,193,1193]" captionTargetPageId="7" captionText="FIGURE 5. Schematic representation of the phylogenetic relationships of the genera of Verongida, summarizing four previously published phylogenies (Erwin & Thacker 2007 [ITS 2 + 28 S]; Díaz et al., 2013 [cox 1]; Redmond et al., 2013 [18 S]; Thacker et al., 2013 [28 S];). Black circles indicate nodes with 100 % maximum likelihood bootstrap support in at least 2 studies; gray circles indicate nodes with 100 % maximum likelihood bootstrap support in 1 study, and none under 85 %; white circles indicate poorly resolved nodes, or phylogenies incongruent between studies. Proposed common characters are indicated with numbers: 1. Verongida with bundles of interstitial collagen fibrils either in the cortex, mesohyl, or within fibers; 2. Family Chondrosiidae (sensu Redmond et al., 2013) with a thick, acellular collagenous cortex; 3. Clade represented by the rest of the Verongida, all containing spherulous cells and brominated secondary metabolites; 4. Clade represented by Hexadella, with eurypylous choanocyte chambers, a thin cuticle, and lacking fibers; 5. Poorly supported clade (62 %) containing all the non - Hexadella classical genera of Verongida; 6. Moderately supported clade (85 %) containing all eurypylous verongids with a cellularized collagenous cortex. 7. Verongids with diplodal choanocyte chambers; 8. Verongids with an isotropic, welldeveloped fiber reticle, with oval to hexagonal meshes, and fibers with a distinct organic pith and a bark (concentrically layered) throughout the body. 9. Verongids with a variably developed reticulation of fibers, sometimes absent, with thick fibers with or without apparent bark; a morphological synapomorphy is yet to be discovered." httpUri="https://zenodo.org/record/240009/files/figure.png" pageId="8" pageNumber="411">Figure 5</figureCitation>
|
||
, clades 4 and 6), demonstrating that this feature is paraphyletic within
|
||
<taxonomicName id="4C064D38113263548020FDA3FB74535B" box="[1046,1165,583,608]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
. Current studies suggest the classification of
|
||
<taxonomicName id="4C064D3811326354854CFD88FE1453BE" box="[378,493,620,645]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
comprises four main clades: (1) a clear monophyletic group containing all
|
||
<taxonomicName id="4C064D3811326354810CFD88FA6553BF" box="[1338,1436,620,644]" class="Demospongiae" family="Aplysinidae" genus="Aplysina" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA911326354810CFD88FA6553BF" box="[1338,1436,620,644]" italics="true" pageId="8" pageNumber="411">Aplysina</emphasis>
|
||
</taxonomicName>
|
||
species studied, and representatives of all
|
||
<taxonomicName id="4C064D38113263548642FD6BFCE35393" box="[628,794,655,680]" class="Demospongiae" family="Aplysinellidae" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="family">Aplysinellidae</taxonomicName>
|
||
, and Pseudoceratiniidae genera; (2) a clade that contains
|
||
<emphasis id="B972EAA91132635484A1FD50FE3053F7" box="[151,457,692,716]" italics="true" pageId="8" pageNumber="411">
|
||
<taxonomicName id="4C064D381132635484A1FD50FF0653F7" box="[151,255,692,716]" class="Demospongiae" family="Ianthellidae" genus="Ianthella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Ianthella</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C064D38113263548539FD50FE3C53F7" box="[271,453,692,716]" class="Demospongiae" family="Ianthellidae" genus="Anomoianthella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">Anomoianthella</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName id="4C064D3811326354863FFD51FD8753F7" box="[521,638,693,716]" class="Demospongiae" family="Ianthellidae" genus="Vansoestia" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Verongiida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus" status="gen. nov.">
|
||
<emphasis id="B972EAA911326354863FFD51FD8753F7" box="[521,638,693,716]" italics="true" pageId="8" pageNumber="411">Vansoestia</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="A24157D21132635486BAFD51FD0353F7" box="[652,762,693,717]" pageId="8" pageNumber="411" rank="genus">gen. nov.</taxonomicNameLabel>
|
||
, but excludes
|
||
<taxonomicName id="4C064D38113263548791FD50FBE353F7" box="[935,1050,692,716]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA9113263548791FD50FBE353F7" box="[935,1050,692,716]" italics="true" pageId="8" pageNumber="411">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
, which should be defined by an amended family
|
||
<taxonomicName id="4C064D38113263548564FD33FE2353CB" box="[338,474,727,752]" class="Demospongiae" family="Ianthellidae" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="family">Ianthellidae</taxonomicName>
|
||
; (3) a clade that presently contains only
|
||
<taxonomicName id="4C064D381132635487ACFD3CFBF453CB" box="[922,1037,728,752]" class="Demospongiae" family="Ianthellidae" genus="Hexadella" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="genus">
|
||
<emphasis id="B972EAA91132635487ACFD3CFBF453CB" box="[922,1037,728,752]" italics="true" pageId="8" pageNumber="411">Hexadella</emphasis>
|
||
</taxonomicName>
|
||
and that should be erected as a new family; and (4) a clade containing the family
|
||
<taxonomicName id="4C064D381132635486A4FD18FB7B522F" authority="Redmond et al. (2013)" authorityName="Redmond et al." authorityYear="2013" box="[658,1154,764,789]" class="Demospongiae" family="Chondrosiidae" kingdom="Animalia" order="Chondrosida" pageId="8" pageNumber="411" phylum="Porifera" rank="family">
|
||
Chondrosiidae
|
||
<emphasis id="B972EAA911326354870AFD19FC83522F" box="[828,890,765,788]" italics="true" pageId="8" pageNumber="411">sensu</emphasis>
|
||
Redmond
|
||
<emphasis id="B972EAA91132635487C3FD19FBD4522F" box="[1013,1069,764,788]" italics="true" pageId="8" pageNumber="411">et al.</emphasis>
|
||
(2013)
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BB936BB1132635484F1FCC4FA80529F" blockId="8.[151,1437,151,933]" pageId="8" pageNumber="411">
|
||
We demonstrate here that the suprageneric classification of
|
||
<taxonomicName id="4C064D381132635487B3FCFBFC015203" box="[901,1016,799,824]" class="Demospongiae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Verongida" pageId="8" pageNumber="411" phylum="Porifera" rank="order">Verongida</taxonomicName>
|
||
must be further revised in order to reflect our current knowledge of phylogenetic relationships among the genera of Myxospongiae. Considering the diagnostic characters that we have in hand, a thorough comparative review of the histological and ultrastructural features of representatives of each genus is definitely needed to improve the taxonomic diagnoses of these taxa.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |