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<mods:title id="965F749446FCACE8A38F872536CCA976">Morphological description and DNA barcodes of shallow-water Tetractinellida (Porifera: Demospongiae) from Bocas del Toro, Panama, with description of a new species</mods:title>
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<mods:namePart id="BBB5FF5809620D6DBD2D0302AFD2175D">Cárdenas, Paco</mods:namePart>
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<mods:namePart id="7C2DE586626508347F1540C926EE3A73">Rapp, Hans Tore</mods:namePart>
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<mods:namePart id="1B0156F851CB76EE7B4FC939B2C9817D">Díaz, Maria Cristina</mods:namePart>
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<treatment id="03FD6D27AA55323EFF4B2F6CBC51EC4D" ID-DOI="http://doi.org/10.5281/zenodo.5689936" ID-GBIF-Taxon="119626238" ID-Zenodo-Dep="5689936" LSID="urn:lsid:plazi:treatment:03FD6D27AA55323EFF4B2F6CBC51EC4D" httpUri="http://treatment.plazi.org/id/03FD6D27AA55323EFF4B2F6CBC51EC4D" lastPageId="33" lastPageNumber="34" pageId="31" pageNumber="32">
<subSubSection id="C34E8FBAAA553200FF4B2F6CBF5CE8E8" pageId="31" pageNumber="32" type="nomenclature">
<paragraph id="8BEBDC31AA553200FF4B2F6CBC73E8CB" blockId="31.[151,564,882,943]" box="[151,564,882,908]" pageId="31" pageNumber="32">
<heading id="D0A36B5DAA553200FF4B2F6CBC73E8CB" bold="true" box="[151,564,882,908]" fontSize="11" level="1" pageId="31" pageNumber="32" reason="1">
<taxonomicName id="4C54A7B2AA553200FF4B2F6CBC73E8CB" authority="Hechtel, 1965" authorityName="Hechtel" authorityYear="1965" box="[151,564,882,908]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="31" pageNumber="32" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA553200FF4B2F6CBC73E8CB" bold="true" box="[151,564,882,908]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF4B2F6CBF37E8CB" bold="true" box="[151,368,882,908]" italics="true" pageId="31" pageNumber="32">Geodia papyracea</emphasis>
(
<bibRefCitation id="EFC5A1C0AA553200FEA32F6CBC6BE8CB" author="Hechtel" box="[383,556,882,908]" pageId="31" pageNumber="37" refString="Hechtel, G. J. (1965) A Systematic Study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History, 20, 1 - 103." type="journal article" year="1965">Hechtel, 1965</bibRefCitation>
)
</emphasis>
</taxonomicName>
</heading>
</paragraph>
<paragraph id="8BEBDC31AA553200FF4B2F8BBF5CE8E8" blockId="31.[151,564,882,943]" box="[151,283,917,943]" pageId="31" pageNumber="32">
(
<figureCitation id="136FC0B4AA553200FF432F8BBF55E8E8" box="[159,274,917,943]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="31" pageNumber="32">Figure 16</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C34E8FBAAA553200FF4B2FFCBF48E8BE" box="[151,271,994,1017]" pageId="31" pageNumber="32" type="description">
<paragraph id="8BEBDC31AA553200FF4B2FFCBF48E8BE" blockId="31.[151,1011,994,1081]" box="[151,271,994,1017]" pageId="31" pageNumber="32">Synonyms.</paragraph>
</subSubSection>
<subSubSection id="C34E8FBAAA553200FF4B281CBCA9EF7E" pageId="31" pageNumber="32" type="reference_group">
<paragraph id="8BEBDC31AA553200FF4B281CBCA9EF7E" blockId="31.[151,1011,994,1081]" pageId="31" pageNumber="32">
<treatmentCitationGroup id="AB44FB1FAA553200FF4B281CBCA9EF7E" pageId="31" pageNumber="32">
<taxonomicName id="4C54A7B2AA553200FF4B281CBCEDEF5E" authority="Hechtel, 1965: 71" authorityName="Hechtel" authorityPageNumber="71" authorityYear="1965" box="[151,682,1026,1049]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="31" pageNumber="32" phylum="Porifera" rank="species" species="papyracea" subGenus="Cydonium">
<emphasis id="B9200023AA553200FF4B281CBFA4EF5E" box="[151,483,1026,1049]" italics="true" pageId="31" pageNumber="32">Geodia (Cydonium) papyracea</emphasis>
<treatmentCitation id="0AF5FA20AA553200FE35281CBCEDEF5E" author="Hechtel" box="[489,682,1026,1049]" page="71" pageId="31" pageNumber="32" year="1965">
<bibRefCitation id="EFC5A1C0AA553200FE35281CBCC5EF5E" author="Hechtel" box="[489,642,1026,1049]" pageId="31" pageNumber="37" refString="Hechtel, G. J. (1965) A Systematic Study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History, 20, 1 - 103." type="journal article" year="1965">Hechtel, 1965</bibRefCitation>
: 71
</treatmentCitation>
</taxonomicName>
, text-fig. 13, pl. VIII, figs.12.
<taxonomicName id="4C54A7B2AA553200FF4B283CBFBBEF7E" authority="Hechtel, 1965" authorityName="Hechtel" authorityYear="1965" box="[151,508,1058,1081]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="31" pageNumber="32" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA553200FF4B283CBF1AEF7E" box="[151,349,1058,1081]" italics="true" pageId="31" pageNumber="32">Geodia papyracea</emphasis>
<bibRefCitation id="EFC5A1C0AA553200FEBF283CBFBBEF7E" author="Hechtel" box="[355,508,1058,1081]" pageId="31" pageNumber="37" refString="Hechtel, G. J. (1965) A Systematic Study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History, 20, 1 - 103." type="journal article" year="1965">Hechtel, 1965</bibRefCitation>
</taxonomicName>
: Alcolado 1981, p. 38.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C34E8FBAAA553200FF4B2877BF37EE98" pageId="31" pageNumber="32" type="materials_examined">
<paragraph id="8BEBDC31AA553200FF4B2877BDF3EFC4" blockId="31.[151,1438,1129,1999]" box="[151,948,1129,1155]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF4B2877BF44EFC5" box="[151,259,1129,1154]" italics="true" pageId="31" pageNumber="32">
<typeStatus id="54EF6293AA553200FF4B2877BF44EFC5" box="[151,259,1129,1154]" pageId="31" pageNumber="32" type="holotype">Holotype</typeStatus>
</emphasis>
.
<collectionCode id="ED4544F4AA553200FECD2877BF16EFC4" LSID="urn:lsid:biocol.org:col:34880" box="[273,337,1129,1155]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34880" name="Peabody Museum of Natural History" pageId="31" pageNumber="32">YPM</collectionCode>
5045, mangrove boat channel, Port Royal,
<collectingCountry id="F3439CA1AA553200FC922877BDF7EFC4" box="[846,944,1129,1155]" name="Jamaica" pageId="31" pageNumber="32">Jamaica</collectingCountry>
.
</paragraph>
<paragraph id="8BEBDC31AA553200FF1A288EBA7EEFED" blockId="31.[151,1438,1129,1999]" box="[198,1081,1168,1194]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF1A288EBF72EFEE" box="[198,309,1168,1193]" italics="true" pageId="31" pageNumber="32">Material.</emphasis>
<collectionCode id="ED4544F4AA553200FEE0288EBFF7EFED" box="[316,432,1168,1194]" pageId="31" pageNumber="32">UMPCW</collectionCode>
921, Solarte Island, on a mangrove root,
<quantity id="4CAC71D4AA553200FC55288EBDAEEFED" box="[905,1001,1168,1194]" metricMagnitude="-1" metricUnit="m" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="31" pageNumber="32" unit="m" value="0.75" valueMax="1.0" valueMin="0.5">0.51 m</quantity>
depth.
</paragraph>
<paragraph id="8BEBDC31AA553200FF1A28A9BFBDEE59" blockId="31.[151,1438,1129,1999]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF1A28A9BC92EF97" box="[198,725,1207,1232]" italics="true" pageId="31" pageNumber="32">
Additional material examined.
<taxonomicName id="4C54A7B2AA553200FDE428A9BC92EF97" box="[568,725,1207,1232]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="31" pageNumber="32" phylum="Porifera" rank="species" species="papyracea">G. papyracea</taxonomicName>
</emphasis>
,
<collectionCode id="ED4544F4AA553200FD3A28A8BD61EF97" LSID="urn:lsid:biocol.org:col:34880" box="[742,806,1206,1232]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34880" name="Peabody Museum of Natural History" pageId="31" pageNumber="32">YPM</collectionCode>
5045,
<typeStatus id="54EF6293AA553200FCA528A8BDA5EF97" box="[889,994,1206,1232]" pageId="31" pageNumber="32" type="holotype">holotype</typeStatus>
, mangrove boat channel, Port Royal,
<collectingCountry id="F3439CA1AA553200FF4B28C3BF44EFB0" box="[151,259,1245,1271]" name="Jamaica" pageId="31" pageNumber="32">Jamaica</collectingCountry>
;
<collectionCode id="ED4544F4AA553200FECD28C3BF13EFB0" LSID="urn:lsid:biocol.org:col:34880" box="[273,340,1245,1271]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34880" name="Peabody Museum of Natural History" pageId="31" pageNumber="32">YPM</collectionCode>
5309, 5311,
<typeStatus id="54EF6293AA553200FE2328C3BC39EFB0" box="[511,638,1245,1271]" pageId="31" pageNumber="32" type="paratype">paratypes</typeStatus>
, mangrove boat channel, Port Royal,
<collectingCountry id="F3439CA1AA553200FBB428C3BA93EFB0" box="[1128,1236,1245,1271]" name="Jamaica" pageId="31" pageNumber="32">Jamaica</collectingCountry>
;
<collectionCode id="ED4544F4AA553200FB3E28C3BB7BEFB0" LSID="urn:lsid:biocol.org:col:34871" box="[1250,1340,1245,1271]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="31" pageNumber="32">USNM</collectionCode>
42662, mangrove, Twin Cays,
<collectingCountry id="F3439CA1AA553200FE7B291ABFB1EE59" box="[423,502,1284,1310]" name="Belize" pageId="31" pageNumber="32">Belize</collectingCountry>
.
</paragraph>
<paragraph id="8BEBDC31AA553200FF1A2935BF37EE98" blockId="31.[151,1438,1129,1999]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF1A2935BFE5EE03" box="[198,418,1323,1348]" italics="true" pageId="31" pageNumber="32">Outer morphology</emphasis>
(
<figureCitation id="136FC0B4AA553200FE682934BC4CEE03" box="[436,523,1322,1348]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="31" pageNumber="32">Fig. 16</figureCitation>
A). Whitish massive sponge with green tinges when alive. It is ca
<date id="FFEAFAF1AA553200FAF02934BB20EE03" box="[1324,1383,1322,1348]" pageId="31" pageNumber="32" value="1912-10-06">
6x
<quantity id="4CAC71D4AA553200FA962934BB20EE03" box="[1354,1383,1322,1348]" metricMagnitude="-1" metricUnit="m" metricValue="1.2" pageId="31" pageNumber="32" unit="cm" value="12.0">12</quantity>
</date>
cm. The sub-sample we have is whitish in alcohol; the cortex being lighter than the choanosome. Oscules are uniporal, each with a sphincter and grouped in a subcircular, slightly depressed plate. The oscule plate can be dark green. Pores are cribriporal and widely distributed over the entire surface. Each cribriporal plate has a diameter of
<quantity id="4CAC71D4AA553200FEF829DBBF2BEE98" box="[292,364,1477,1503]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="31" pageNumber="32" unit="mm" value="1.0">1 mm</quantity>
.
</paragraph>
</subSubSection>
<subSubSection id="C34E8FBAAA55323EFF1A29F2BA7FEADD" lastPageId="33" lastPageNumber="34" pageId="31" pageNumber="32" type="description">
<paragraph id="8BEBDC31AA553200FF1A29F2BDDEEDA9" blockId="31.[151,1438,1129,1999]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF1A29F2BF6DED42" box="[198,298,1516,1541]" italics="true" pageId="31" pageNumber="32">Skeleton</emphasis>
(
<figureCitation id="136FC0B4AA553200FEE629F2BFD7ED41" box="[314,400,1516,1542]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="31" pageNumber="32">Fig. 16</figureCitation>
B). The cortex is 480840 µm thick and composed of a thin ectocortex of strongylasters (072 µm) and a thicker endocortex made of sterrasters (480840 µm). The cortex is easily torn and detachable from the choanosome. Oxeas I and plagiotriaenes are positioned under the cortex, more or less radially. The plagiotrianes usually cross the cortex so that the cladomes end up in the ectocortex or at the surface of the cortex. The radial arrangement of megascleres is less obvious
<quantity id="4CAC71D4AA553200FBCD2A98BA12EDE7" box="[1041,1109,1670,1696]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="31" pageNumber="32" unit="mm" value="1.0">1 mm</quantity>
under the cortex. Oxyasters I, oxyasters II and strongylasters can be found in the whole choanosome, as well as developing sterrasters. Oxeas II, smaller than oxeas I, are only found in the choanosome.
</paragraph>
<paragraph id="8BEBDC31AA553200FF1A2AE5BB1EEC88" blockId="31.[151,1438,1129,1999]" pageId="31" pageNumber="32">
<emphasis id="B9200023AA553200FF1A2AE5BF6FEC53" box="[198,296,1787,1812]" italics="true" pageId="31" pageNumber="32">Spicules</emphasis>
(
<figureCitation id="136FC0B4AA553200FEE52AE4BFC8EC53" box="[313,399,1786,1812]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="31" pageNumber="32">Fig. 16</figureCitation>
CG).
<emphasis id="B9200023AA553200FE3D2AE5BC33EC53" box="[481,628,1787,1812]" italics="true" pageId="31" pageNumber="32">Megascleres</emphasis>
: (a) oxeas I, stout, straight or slightly bent, length: 651
<emphasis id="B9200023AA553200FAC42AE5BB14EC53" box="[1304,1363,1787,1812]" italics="true" pageId="31" pageNumber="32">
<date id="FFEAFAF1AA553200FAC42AE5BB14EC53" box="[1304,1363,1787,1812]" pageId="31" pageNumber="32" value="1039" valueMax="1248">1039</date>
</emphasis>
1248 µm; width: 7
<emphasis id="B9200023AA553200FE982B3EBF3FEC7E" box="[324,376,1824,1849]" italics="true" pageId="31" pageNumber="32">24.3</emphasis>
29 µm. (b) oxeas II, rare, sometimes with strongyle ends, usually straight, length: 95
<emphasis id="B9200023AA553200FF4B2B5BBE9DEC19" box="[151,218,1861,1886]" italics="true" pageId="31" pageNumber="32">124.3</emphasis>
244 µm; width: 1
<emphasis id="B9200023AA553200FE1B2B5BBF91EC19" box="[455,470,1861,1886]" italics="true" pageId="31" pageNumber="32">2</emphasis>
3 µm (N=14). (c) plagiotriaenes, rhabdome length: 641
<emphasis id="B9200023AA553200FB542B5BBA8CEC19" box="[1160,1227,1861,1886]" italics="true" pageId="31" pageNumber="32">962.7</emphasis>
1080; rhabdome width: 13
<emphasis id="B9200023AA553200FEC82B75BF00ECC3" box="[276,327,1899,1924]" italics="true" pageId="31" pageNumber="32">24.6</emphasis>
34 µm; clad length: 57
<emphasis id="B9200023AA553200FDB22B75BCE6ECC3" box="[622,673,1899,1924]" italics="true" pageId="31" pageNumber="32">97.1</emphasis>
132 µm. (d) anatriaenes, rare, rhabdome length: 354
<emphasis id="B9200023AA553200FAC32B75BB26ECC3" box="[1311,1377,1899,1924]" italics="true" pageId="31" pageNumber="32">633.2</emphasis>
885 µm (N=5); rhabdome width: 2.5
<emphasis id="B9200023AA553200FDF62B8EBC08ECEE" box="[554,591,1936,1961]" italics="true" pageId="31" pageNumber="32">3.2</emphasis>
4 µm (N=8); clad length: 5
<emphasis id="B9200023AA553200FC682B8EBDAFECEE" box="[948,1000,1936,1961]" italics="true" pageId="31" pageNumber="32">13.8</emphasis>
27 µm (N=8). (e) mesoprotriaenes, very rare, rhabdome length: 632 µm (N=1); rhabdome width: 3 µm (N=1); clad length: 916 µm (N=2).
</paragraph>
<caption id="DF2B8CB9AA6A323FFF4B2B1FBC69ECB4" httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="32" pageNumber="33" targetBox="[204,1371,183,1770]" targetPageId="32">
<paragraph id="8BEBDC31AA6A323FFF4B2B1FBC69ECB4" blockId="32.[151,1437,1793,2035]" pageId="32" pageNumber="33">
<emphasis id="B9200023AA6A323FFF4B2B1FBF6CEC5E" bold="true" box="[151,299,1793,1817]" pageId="32" pageNumber="33">FIGURE 16.</emphasis>
<taxonomicName id="4C54A7B2AA6A323FFEEF2B1CBCF2EC5F" authority="(Hechtel, 1965)" authorityName="Hechtel" authorityYear="1965" baseAuthorityName="Hechtel" baseAuthorityYear="1965" box="[307,693,1793,1817]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="32" pageNumber="33" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6A323FFEEF2B1CBFB8EC5E" box="[307,511,1794,1817]" italics="true" pageId="32" pageNumber="33">Geodia papyracea</emphasis>
(Hechtel, 1965)
</taxonomicName>
[UMPCW921]: A. Specimen
<emphasis id="B9200023AA6A323FFBD92B1CBA0DEC5E" box="[1029,1098,1794,1817]" italics="true" pageId="32" pageNumber="33">in situ</emphasis>
, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Crosssection showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Starshaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm;
<taxonomicName id="4C54A7B2AA6A323FFD082BA8BD25EC8A" box="[724,866,1974,1997]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="32" pageNumber="33" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6A323FFD082BA8BD25EC8A" box="[724,866,1974,1997]" italics="true" pageId="32" pageNumber="33">G. papyracea</emphasis>
</taxonomicName>
[YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm.
</paragraph>
</caption>
<paragraph id="8BEBDC31AA6B323EFF4B2C86BFFFEA34" blockId="33.[151,1437,152,1802]" pageId="33" pageNumber="34">
<emphasis id="B9200023AA6B323EFF4B2C86BF73EBF6" box="[151,308,152,177]" italics="true" pageId="33" pageNumber="34">Microscleres</emphasis>
: (f) sterrasters (
<figureCitation id="136FC0B4AA6B323EFE262C86BC15EBF5" box="[506,594,152,178]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
CD), subglobular, with warty 47 branched rosettes at their surface (diameter: ca 4 µm), length: 65
<emphasis id="B9200023AA6B323EFDEA2CA1BC2AEB9F" box="[566,621,191,216]" italics="true" pageId="33" pageNumber="34">71.6</emphasis>
77 µm; width: 63
<emphasis id="B9200023AA6B323EFCB72CA1BDE5EB9F" box="[875,930,191,216]" italics="true" pageId="33" pageNumber="34">70.3</emphasis>
75 µm; thickness: 50
<emphasis id="B9200023AA6B323EFB112CA1BB43EB9F" box="[1229,1284,191,216]" italics="true" pageId="33" pageNumber="34">55.9</emphasis>
61 µm. (g) acanthoxyasters I (
<figureCitation id="136FC0B4AA6B323EFEA82CFBBF8EEBB8" box="[372,457,229,255]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
E), 912 thin actines, center more or less developed, diameter: 22
<emphasis id="B9200023AA6B323EFB002CFBBB48EBB9" box="[1244,1295,229,254]" italics="true" pageId="33" pageNumber="34">27.5</emphasis>
36 µm. (h) acanthoxyasters II (
<figureCitation id="136FC0B4AA6B323EFEA02D12BF97EA61" box="[380,464,268,294]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
F), with large centrum and thicker actines than acanthoxyasters I, 917 actines, less common than acanthoxyasters I. diameter: 14
<emphasis id="B9200023AA6B323EFD662D2CBC90EA0C" box="[698,727,306,331]" italics="true" pageId="33" pageNumber="34">23</emphasis>
29 µm. (i) acanthostrongylasters (
<figureCitation id="136FC0B4AA6B323EFBAD2D2CBA81EA0B" box="[1137,1222,306,332]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
G), 1420 actines, diameter: 3
<emphasis id="B9200023AA6B323EFEF62D47BF08EA35" box="[298,335,345,370]" italics="true" pageId="33" pageNumber="34">4.9</emphasis>
8.8 µm.
</paragraph>
<paragraph id="8BEBDC31AA6B323EFF1A2D9EBA7FEADD" blockId="33.[151,1437,152,1802]" box="[198,1080,384,410]" pageId="33" pageNumber="34">
<emphasis id="B9200023AA6B323EFF1A2D9EBC3DEADE" box="[198,634,384,409]" italics="true" pageId="33" pageNumber="34">Habitat in the Bocas del Toro region.</emphasis>
On mangrove roots,
<quantity id="4CAC71D4AA6B323EFCAE2D9EBDAFEADD" box="[882,1000,384,410]" metricMagnitude="0" metricUnit="m" metricValue="1.0" metricValueMax="1.5" metricValueMin="0.5" pageId="33" pageNumber="34" unit="m" value="1.0" valueMax="1.5" valueMin="0.5">0.51.5 m</quantity>
depth.
</paragraph>
</subSubSection>
<subSubSection id="C34E8FBAAA6B323EFF1A2DB9BD54E949" pageId="33" pageNumber="34" type="distribution">
<paragraph id="8BEBDC31AA6B323EFF1A2DB9BD54E949" blockId="33.[151,1437,152,1802]" pageId="33" pageNumber="34">
<emphasis id="B9200023AA6B323EFF1A2DB9BF12EA87" box="[198,341,423,448]" italics="true" pageId="33" pageNumber="34">Distribution</emphasis>
.
<collectingCountry id="F3439CA1AA6B323EFEB42DB8BFE1EA87" box="[360,422,422,448]" name="Cuba" pageId="33" pageNumber="34">Cuba</collectingCountry>
(
<bibRefCitation id="EFC5A1C0AA6B323EFE652DB8BC36EA87" author="Alcolado" box="[441,625,422,448]" pageId="33" pageNumber="35" refString="Alcolado, P. M. (2002) Catalogo de las esponjas de Cuba. Avicennia, 15, 53 - 72." type="journal article" year="2002">Alcolado 2002</bibRefCitation>
);
<collectingCountry id="F3439CA1AA6B323EFD552DB8BC93EA87" box="[649,724,422,448]" name="Belize" pageId="33" pageNumber="34">Belize</collectingCountry>
(
<bibRefCitation id="EFC5A1C0AA6B323EFD3B2DB8BDCFEA87" author="Rutzler" box="[743,904,422,448]" pageId="33" pageNumber="38" refString="Rutzler, K. (1988) Mangrove sponge disease induced by cyanobacterial symbionts: failure of a primitive immune system? Diseases of Aquatic Organisms, 5, 143 - 149." type="journal article" year="1988">Rützler 1988</bibRefCitation>
;
<bibRefCitation id="EFC5A1C0AA6B323EFC442DB8BAC3EA87" author="Rutzler" box="[920,1156,422,448]" pageId="33" pageNumber="38" refString="Rutzler, K., Diaz, M. C., van Soest, R. W. M., Zea, S., Smith, K. P., Alvarez, B. &amp; Wulff, J. (2000) Diversity of sponge fauna in mangrove ponds, Pelican Cays, Belize. Atoll Research Bulletin, 476," type="book chapter" year="2000">
Rützler
<emphasis id="B9200023AA6B323EFC262DB9BA7FEA87" box="[1018,1080,423,448]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2000
</bibRefCitation>
);
<collectingCountry id="F3439CA1AA6B323EFB472DB8BABFEA87" box="[1179,1272,422,448]" name="Panama" pageId="33" pageNumber="34">Panama</collectingCountry>
(Díaz 2005);
<collectingCountry id="F3439CA1AA6B323EFF4B2DD3BEBFEAA0" box="[151,248,461,487]" name="Jamaica" pageId="33" pageNumber="34">Jamaica</collectingCountry>
(
<bibRefCitation id="EFC5A1C0AA6B323EFED42DD3BFE8EAA0" author="Hechtel" box="[264,431,461,487]" pageId="33" pageNumber="37" refString="Hechtel, G. J. (1965) A Systematic Study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History, 20, 1 - 103." type="journal article" year="1965">Hechtel 1965</bibRefCitation>
);
<collectingCountry id="F3439CA1AA6B323EFE1A2DD3BC7BEAA0" box="[454,572,461,487]" name="Colombia" pageId="33" pageNumber="34">Colombia</collectingCountry>
(
<bibRefCitation id="EFC5A1C0AA6B323EFD902DD3BDB3EAA0" author="Wintermann-Kilian" box="[588,1012,461,487]" pageId="33" pageNumber="39" refString="Wintermann-Kilian, G. &amp; Kilian, E. F. (1984) Marine sponges of the region of Santa Marta (Colombia). Part II. Homoslerophorida, Choristida, Spirophorida, Hadromerida, Axinellida, Halichondrida, Poecilosclerida. Studies on Neotropical Fauna and Environment, 19, 121 - 135." type="journal article" year="1984">Wintermann-Kilian &amp; Kilian 1984</bibRefCitation>
);
<collectingCountry id="F3439CA1AA6B323EFBD52DD3BA2AEAA0" box="[1033,1133,461,487]" name="Curaçao" pageId="33" pageNumber="34">Curaçao</collectingCountry>
(van Soest 1981);
<collectingCountry id="F3439CA1AA6B323EFA8F2DD3BBDBEAA0" box="[1363,1436,461,487]" name="Brazil" pageId="33" pageNumber="34">Brazil</collectingCountry>
(
<bibRefCitation id="EFC5A1C0AA6B323EFF432DEABF70E949" author="Burton" box="[159,311,500,526]" pageId="33" pageNumber="36" refString="Burton, M. (1940) Las esponjas marinas del Museo Argentino de Ciencias Naturales. Anales del Museo Argentino de Ciencias Naturales &quot; Bernardino Rivadavia &quot;, 40, 95 - 121." type="journal article" year="1940">Burton 1940</bibRefCitation>
; da Silva
<emphasis id="B9200023AA6B323EFE762DEABFA3E94A" box="[426,484,500,525]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2004;
<bibRefCitation id="EFC5A1C0AA6B323EFDEA2DEABD40E949" author="Cedro" box="[566,775,500,526]" pageId="33" pageNumber="36" refString="Cedro, V. R., Hajdu, E., Sovierzosky, H. H. &amp; Correia, M. D. (2007) Demospongiae (Porifera) of the shallow coral reefs of Maceio, Alagoas State, Brazil. In: Custodio, M. R., Lobo-Hajdu, G., Hajdu, E. &amp; Muricy, G. (Eds.). Porifera research: biodiversity, innovation and sustainability. Serie Livros 28, Museu Nacional, 233 - 237." type="journal article" year="2007">
Cedro
<emphasis id="B9200023AA6B323EFD592DEABCF8E94A" box="[645,703,500,525]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2007
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C34E8FBAAA6B323EFF1A2E04BC51EC4D" pageId="33" pageNumber="34" type="discussion">
<paragraph id="8BEBDC31AA6B323EFF1A2E04BC36E899" blockId="33.[151,1437,152,1802]" pageId="33" pageNumber="34">
<emphasis id="B9200023AA6B323EFF1A2E04BFAFE974" box="[198,488,538,563]" italics="true" pageId="33" pageNumber="34">Remarks and discussion.</emphasis>
The fragile and easily peeled cortex of
<taxonomicName id="4C54A7B2AA6B323EFC682E04BA17E974" box="[948,1104,538,563]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFC682E04BA17E974" box="[948,1104,538,563]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
might be due to the peculiar disposition of the triaenes, with the cladomes not supporting the cortex as in most
<taxonomicName id="4C54A7B2AA6B323EFB432E5FBABEE91D" box="[1183,1273,577,602]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="genus">
<emphasis id="B9200023AA6B323EFB432E5FBABEE91D" box="[1183,1273,577,602]" italics="true" pageId="33" pageNumber="34">Geodia</emphasis>
</taxonomicName>
species. The ectocortex of the Bocas del Toro specimen was much thinner (072 µm) than in the
<collectingCountry id="F3439CA1AA6B323EFBA12E76BA8FE9C5" box="[1149,1224,616,642]" name="Belize" pageId="33" pageNumber="34">Belize</collectingCountry>
sample (120325 µm) and the
<typeStatus id="54EF6293AA6B323EFEED2E90BFDFE9EF" box="[305,408,654,680]" pageId="33" pageNumber="34" type="holotype">holotype</typeStatus>
/
<typeStatus id="54EF6293AA6B323EFE7D2E90BC55E9EF" box="[417,530,654,680]" pageId="33" pageNumber="34" type="paratype">paratypes</typeStatus>
(93232 µm thick). Conversely, the sterrastreal layer in the Bocas del Toro specimens was thicker (480840 µm) than in the
<collectingCountry id="F3439CA1AA6B323EFD2A2EABBD04E988" box="[758,835,693,719]" name="Belize" pageId="33" pageNumber="34">Belize</collectingCountry>
sample (93279 µm) and the
<typeStatus id="54EF6293AA6B323EFB692EABBB58E988" box="[1205,1311,693,719]" pageId="33" pageNumber="34" type="holotype">holotype</typeStatus>
/
<typeStatus id="54EF6293AA6B323EFAF42EABBBDBE988" box="[1320,1436,693,719]" pageId="33" pageNumber="34" type="paratype">paratypes</typeStatus>
(186418 µm). This is partly a consequence of the larger sterrasters in comparison with the
<typeStatus id="54EF6293AA6B323EFB0C2EC2BB44E9B1" box="[1232,1283,732,758]" pageId="33" pageNumber="34">type</typeStatus>
material and previous
<taxonomicName id="4C54A7B2AA6B323EFEDB2F1CBFE0E85C" box="[263,423,770,795]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFEDB2F1CBFE0E85C" box="[263,423,770,795]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
records (da Silva
<emphasis id="B9200023AA6B323EFD592F1CBC84E85C" box="[645,707,770,795]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2004). We also noted that the surface of the Bocas del Toro specimen was less folded, the parchmentlike appearance characteristic of this species was therefore not as obvious as in the samples from
<collectingCountry id="F3439CA1AA6B323EFDDA2F4EBC23E82D" box="[518,612,848,874]" name="Jamaica" pageId="33" pageNumber="34">Jamaica</collectingCountry>
and
<collectingCountry id="F3439CA1AA6B323EFD402F4EBCACE82D" box="[668,747,848,874]" name="Belize" pageId="33" pageNumber="34">Belize</collectingCountry>
. The rest of our measurements were in accordance with the species description. This species was very common in Solarte lagoon in
<date id="FFEAFAF1AA6B323EFC342F68BAD1E8D7" box="[1000,1174,886,912]" pageId="33" pageNumber="34" value="2005" valueMax="2006">2005 and 2006</date>
(M. C. Díaz, personal observation). We were not able to find it anymore in
<date id="FFEAFAF1AA6B323EFCD32F83BD87E8F0" box="[783,960,925,951]" pageId="33" pageNumber="34" value="2007" valueMax="2008">2007 and 2008</date>
as it appeared to have been replaced by the sympatric species
<taxonomicName id="4C54A7B2AA6B323EFE452FDABC2DE89A" box="[409,618,964,989]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFE452FDABC2DE89A" box="[409,618,964,989]" italics="true" pageId="33" pageNumber="34">Geodia gibberosa</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BEBDC31AA6B323EFF1A2FF4BFE4EDD1" blockId="33.[151,1437,152,1802]" pageId="33" pageNumber="34">
In the field,
<taxonomicName id="4C54A7B2AA6B323EFE8A2FF4BFB4EF44" box="[342,499,1002,1027]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFE8A2FF4BFB4EF44" box="[342,499,1002,1027]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
can be mistaken with the previously described
<taxonomicName id="4C54A7B2AA6B323EFBFE2FF4BAF1EF44" box="[1058,1206,1002,1027]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFBFE2FF4BAF1EF44" box="[1058,1206,1002,1027]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
since they are both found living on mangrove roots and can even be found growing on one another (
<bibRefCitation id="EFC5A1C0AA6B323EFB92280FBAB4EF6C" author="Hechtel" box="[1102,1267,1041,1067]" pageId="33" pageNumber="37" refString="Hechtel, G. J. (1965) A Systematic Study of the Demospongiae of Port Royal, Jamaica. Bulletin of the Peabody Museum of Natural History, 20, 1 - 103." type="journal article" year="1965">Hechtel 1965</bibRefCitation>
). Again, gross morphology is here important:
<taxonomicName id="4C54A7B2AA6B323EFDD42826BCE2EF16" box="[520,677,1080,1105]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFDD42826BCE2EF16" box="[520,677,1080,1105]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
has very poor relief and a fragile cortex while
<taxonomicName id="4C54A7B2AA6B323EFB0B2826BB2AEF16" box="[1239,1389,1080,1105]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFB0B2826BB2AEF16" box="[1239,1389,1080,1105]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
has characteristic lobes and a tough and thicker cortex.
<taxonomicName id="4C54A7B2AA6B323EFCFA2841BD81EF3F" box="[806,966,1119,1144]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFCFA2841BD81EF3F" box="[806,966,1119,1144]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
seems to be more overgrown than
<taxonomicName id="4C54A7B2AA6B323EFA5E2841BF5EEFD9" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFA5E2841BF5EEFD9" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
. In thick sections, the skeleton was more radially organized in
<taxonomicName id="4C54A7B2AA6B323EFBF3289BBA8BEFD9" box="[1071,1228,1157,1182]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFBF3289BBA8BEFD9" box="[1071,1228,1157,1182]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
(
<figureCitation id="136FC0B4AA6B323EFB02289BBB70EFD8" box="[1246,1335,1157,1183]" captionStart="FIGURE 14" captionStartId="28.[151,260,1822,1846]" captionTargetBox="[151,1420,197,1793]" captionTargetId="figure@28.[151,1436,197,1796]" captionTargetPageId="28" captionText="FIGURE 14. Geodia gibberosa Lamarck, 1815: A. Large specimen hanging at the tip of a mangrove root. It is overgrown by the purple sponge Chalinula molibta, algae and oysters. Scale: 6 cm; B. Specimen with characteristic tumulose shape; C. Close up of oscular plates of an overgrown specimen in situ. Each oscule is uniporal. Scale: 4 cm; G. gibberosa: [ZMBN 81780]: D. Close up of the cribriporal pores. Scale: 1 mm; E. Cross section showing the skeletal architecture: cortex and choanosome. Note the radial arrangement of the triaenes right under the cortex. Scale: 1 mm; F. Cross section of a uniporal oscule. mx: cortical microxeas. s: sphincter. plgt: plagiotriaene. Scale: 0.5 mm." httpUri="https://zenodo.org/record/191102/files/figure.png" pageId="33" pageNumber="34">Fig. 14</figureCitation>
E), with plagiotriaenes supporting the cortex and not crossing it like in
<taxonomicName id="4C54A7B2AA6B323EFC7028B2BA14EF82" box="[940,1107,1196,1221]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFC7028B2BA14EF82" box="[940,1107,1196,1221]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
(
<figureCitation id="136FC0B4AA6B323EFBB828B2BA86EF81" box="[1124,1217,1196,1222]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
B). This probably accounts for the firm cortex in the former. Then,
<taxonomicName id="4C54A7B2AA6B323EFCD828CDBDE4EFAB" box="[772,931,1235,1260]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFCD828CDBDE4EFAB" box="[772,931,1235,1260]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
has larger oxeas and plagiotriaenes; the sterrasters are also a bit larger and more oval shaped. SEM pictures of asters revealed additional microstructural differences. When comparing the surface of sterrasters, the
<taxonomicName id="4C54A7B2AA6B323EFBC0293EBAF4EE7E" box="[1052,1203,1312,1337]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFBC0293EBAF4EE7E" box="[1052,1203,1312,1337]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
rosettes (
<figureCitation id="136FC0B4AA6B323EFAFB293EBB3BEE7D" box="[1319,1404,1312,1338]" captionStart="FIGURE 15" captionStartId="29.[151,257,1864,1888]" captionTargetBox="[168,1414,197,1835]" captionTargetId="figure@29.[168,1429,197,1836]" captionTargetPageId="29" captionText="FIGURE 15. Geodia gibberosa Lamarck, 1815 [ZMBN 81780]: A. Plagiotriaene and sterrasters. Scale: 200 µm; B. Anatriaene cladome. Scale: 20 µm; C. Sterraster. Scale: 20 µm; D. Star shaped smooth rosettes and hilum from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I. Scale: 4 µm; F. Acanthoxyaster II. Scale: 2 µm; G. Acanthoxyasters III. Scale: 2 µm; G. gibberosa [MNHN DT 608, holotype]: H. Smooth rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191103/files/figure.png" pageId="33" pageNumber="34">Fig. 15</figureCitation>
D) had less rays than those of
<taxonomicName id="4C54A7B2AA6B323EFE0F2959BC29EE27" box="[467,622,1351,1376]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFE0F2959BC29EE27" box="[467,622,1351,1376]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
(
<figureCitation id="136FC0B4AA6B323EFDA22958BC93EE27" box="[638,724,1350,1376]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
D). In
<taxonomicName id="4C54A7B2AA6B323EFCFC2959BDF3EE27" box="[800,948,1351,1376]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFCFC2959BDF3EE27" box="[800,948,1351,1376]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
, rosettes also had a smaller center and no warts on the rays. Seemingly, the hilum of the sterrasters were different (the hilum is a circular depression, a remnant of where the nucleus of the sclerocyte used to be): smooth in
<taxonomicName id="4C54A7B2AA6B323EFC38298ABA3CEEEA" box="[996,1147,1428,1453]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFC38298ABA3CEEEA" box="[996,1147,1428,1453]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
, warty in
<taxonomicName id="4C54A7B2AA6B323EFB2A298ABBD3EEEA" box="[1270,1428,1428,1453]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFB2A298ABBD3EEEA" box="[1270,1428,1428,1453]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
. These differences were confirmed by SEM observation of the holotypes sterrasters (
<figureCitation id="136FC0B4AA6B323EFBAF29A5BA95EE92" box="[1139,1234,1467,1493]" captionStart="FIGURE 15" captionStartId="29.[151,257,1864,1888]" captionTargetBox="[168,1414,197,1835]" captionTargetId="figure@29.[168,1429,197,1836]" captionTargetPageId="29" captionText="FIGURE 15. Geodia gibberosa Lamarck, 1815 [ZMBN 81780]: A. Plagiotriaene and sterrasters. Scale: 200 µm; B. Anatriaene cladome. Scale: 20 µm; C. Sterraster. Scale: 20 µm; D. Star shaped smooth rosettes and hilum from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I. Scale: 4 µm; F. Acanthoxyaster II. Scale: 2 µm; G. Acanthoxyasters III. Scale: 2 µm; G. gibberosa [MNHN DT 608, holotype]: H. Smooth rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191103/files/figure.png" pageId="33" pageNumber="34">Figs. 15</figureCitation>
H, 16H). We also noticed that the actines of the acanthoxyasters III in
<taxonomicName id="4C54A7B2AA6B323EFCD829FFBDDDEEBD" box="[772,922,1505,1530]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="gibberosa">
<emphasis id="B9200023AA6B323EFCD829FFBDDDEEBD" box="[772,922,1505,1530]" italics="true" pageId="33" pageNumber="34">G. gibberosa</emphasis>
</taxonomicName>
(
<figureCitation id="136FC0B4AA6B323EFC7629FFBDB9EEBC" box="[938,1022,1505,1531]" captionStart="FIGURE 15" captionStartId="29.[151,257,1864,1888]" captionTargetBox="[168,1414,197,1835]" captionTargetId="figure@29.[168,1429,197,1836]" captionTargetPageId="29" captionText="FIGURE 15. Geodia gibberosa Lamarck, 1815 [ZMBN 81780]: A. Plagiotriaene and sterrasters. Scale: 200 µm; B. Anatriaene cladome. Scale: 20 µm; C. Sterraster. Scale: 20 µm; D. Star shaped smooth rosettes and hilum from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I. Scale: 4 µm; F. Acanthoxyaster II. Scale: 2 µm; G. Acanthoxyasters III. Scale: 2 µm; G. gibberosa [MNHN DT 608, holotype]: H. Smooth rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191103/files/figure.png" pageId="33" pageNumber="34">Fig. 15</figureCitation>
G) have a more prominent spine at their tip when compared with the acanthostrongylasters of
<taxonomicName id="4C54A7B2AA6B323EFC992A16BDA6ED66" box="[837,993,1544,1569]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="species" species="papyracea">
<emphasis id="B9200023AA6B323EFC992A16BDA6ED66" box="[837,993,1544,1569]" italics="true" pageId="33" pageNumber="34">G. papyracea</emphasis>
</taxonomicName>
(
<figureCitation id="136FC0B4AA6B323EFC2C2A16BA02ED65" box="[1008,1093,1544,1570]" captionStart="FIGURE 16" captionStartId="32.[151,257,1793,1817]" captionTargetBox="[204,1371,183,1770]" captionTargetId="figure@32.[204,1386,183,1774]" captionTargetPageId="32" captionText="FIGURE 16. Geodia papyracea (Hechtel, 1965) [UMPCW 921]: A. Specimen in situ, growing on a mangrove root, arrow points to the oscule plate with uniporal oscules (Picture: Scott Nichols). Scale: 3 cm; B. Cross section showing the skeletal architecture: cortex and choanosome. Note the confused arrangement of the megascleres in the choanosome. Scale: 1 mm; C. Sterraster, with an acanthostrongylaster by the warty hilum. Scale: 20 µm; D. Star shaped warty rosettes from a sterraster surface. Scale: 4 µm; E. Acanthoxyaster I and acanthostrongylasters. Scale: 4 µm; F. Acanthoxyaster II. Scale: 4 µm; G. Acanthostrongylasters. Scale: 2 µm; G. papyracea [YPM 5045, holotype]: H. Warty rosettes and hilum from a sterraster surface. Scale: 4 µm." httpUri="https://zenodo.org/record/191104/files/figure.png" pageId="33" pageNumber="34">Fig. 16</figureCitation>
G). Finally, these two species had significantly different Folmer COI sequences (60 bp. difference) and 28S partial sequence (63 bp. difference). A molecular phylogenetic study shows that they belong to two clearly seperated
<taxonomicName id="4C54A7B2AA6B323EFB2E2A4BBB0DED29" box="[1266,1354,1621,1646]" class="Demospongiae" family="Geodiidae" genus="Geodia" kingdom="Animalia" order="Astrophorida" pageId="33" pageNumber="34" phylum="Porifera" rank="genus">
<emphasis id="B9200023AA6B323EFB2E2A4BBB0DED29" box="[1266,1354,1621,1646]" italics="true" pageId="33" pageNumber="34">Geodia</emphasis>
</taxonomicName>
clades (
<bibRefCitation id="EFC5A1C0AA6B323EFF432A62BFD0EDD1" author="Cardenas" box="[159,407,1660,1686]" pageId="33" pageNumber="36" refString="Cardenas, P., Rapp, H. T., Schander, C. &amp; Tendal, O. S. (2009) Molecular taxonomy and phylogeny of the Geodiidae (Porifera, Demospongiae, Astrophorida) - combining phylogenetic and Linnaean classification. Zoologica Scripta, doi: 10.1111 / j. 1463 - 6409.2009.00402. x." type="book" year="2009">
Cárdenas
<emphasis id="B9200023AA6B323EFEC92A62BF08EDD2" box="[277,335,1660,1685]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
2009
</bibRefCitation>
).
</paragraph>
<paragraph id="8BEBDC31AA6B323EFF1A2ABCBC51EC4D" blockId="33.[151,1437,152,1802]" pageId="33" pageNumber="34">
Although the
<typeStatus id="54EF6293AA6B323EFEB42ABCBF89EDFB" box="[360,462,1698,1724]" pageId="33" pageNumber="34" type="holotype">holotype</typeStatus>
(YPM 5045) has been fixed and preserved in alcohol, the DNA we extracted from it was poorly preserved. Consequently, we were unable to amplify the Folmer COI fragment or a 28S sequence from the
<typeStatus id="54EF6293AA6B323EFEA92AEEBFEFEC4D" box="[373,424,1776,1802]" pageId="33" pageNumber="34">type</typeStatus>
material.
</paragraph>
</subSubSection>
</treatment>
</document>