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162 lines
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<mods:title id="10E78FA039B62224ACF424B97CB4A007">Phylogeny and revision of Leucaltis and Leucettusa (Porifera: Calcarea), with new classification proposals and description of a new type of aquiferous system</mods:title>
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<mods:namePart id="8018D5715F388E43B1E6FB443AB204CB">Lopes, Matheus Vieira</mods:namePart>
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<mods:namePart id="2F2362C921F23625B3005101439B6183">Klautau, Michelle</mods:namePart>
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<mods:title id="234BC4DD73F965526E93F287A355D4F6">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="EEE3A701EE62A098942FE0E7C03E7C8C">2023</mods:date>
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<mods:number id="BFAA5B08AABA876D58B22FAB1386290B">2023-07-31</mods:number>
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<mods:number id="B6FFA62EAEBE0BFDE568B97AF37458BC">198</mods:number>
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<mods:identifier id="8A8A89C34161128CA9EBEDD1D9E78B0B" type="DOI">10.1093/zoolinnean/zlad008</mods:identifier>
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<mods:identifier id="2A96B33CD4AA4D35005FBF8869218DEC" type="ISSN">0024-4082</mods:identifier>
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<subSubSection id="C36D657CFFB77F063211FD0EFA98FD0E" box="[870,1363,672,697]" pageId="6" pageNumber="697" type="nomenclature">
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<paragraph id="8BC836F7FFB77F063211FD0EFA98FD0E" blockId="6.[870,1363,672,697]" box="[870,1363,672,697]" pageId="6" pageNumber="697">
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<heading id="D080819BFFB77F063211FD0EFA98FD0E" box="[870,1363,672,697]" centered="true" fontSize="9" level="2" pageId="6" pageNumber="697" reason="8">
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<smallCapsWord id="8D2EA02BFFB77F063211FD0EFC70FD00" baselines="691,690" box="[870,955,672,697]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Family" pageId="6" pageNumber="697">FAMILY</smallCapsWord>
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<taxonomicName id="4C774D74FFB77F0632B5FD0EFA98FD0E" ID-CoL="84KDN" ID-ENA="196262" authority="DENDY & ROW, 1913" authorityName="sensu Dendy & Row" authorityYear="1913" box="[962,1363,672,697]" class="Calcarea" family="Leucaltidae" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="family">
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<smallCapsWord id="8D2EA02BFFB77F0632B5FD0EFB95FD00" baselines="691,690" box="[962,1118,672,697]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Leucaltidae" pageId="6" pageNumber="697">LEUCALTIDAE</smallCapsWord>
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<bibRefCitation id="EFE64B06FFB77F063512FD0EFA98FD0E" author="Dendy A & Row H" box="[1125,1363,672,697]" pageId="6" pageNumber="697" pagination="704 - 813" refId="ref31149" refString="Dendy A, Row H. 1913. The classification and phylogeny of the calcareous sponges, with a reference list of all the described species, systematically arranged. Proceedings of the Zoological Society of London 1913: 704 - 813." type="journal article" year="1913">
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<smallCapsWord id="8D2EA02BFFB77F063512FD0EFB7DFD00" baselines="691,690" box="[1125,1206,672,697]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Dendy" pageId="6" pageNumber="697">DENDY</smallCapsWord>
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&
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<smallCapsWord id="8D2EA02BFFB77F0635AFFD0EFAC5FD00" baselines="691,690" box="[1240,1294,672,697]" lowerCaseFontSize="8" mainFontSize="10" normCase="title" normString="Row" pageId="6" pageNumber="697">ROW</smallCapsWord>
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, 1913
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</bibRefCitation>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C36D657CFFB77F06325EFD66FAD7FD69" box="[809,1308,712,734]" pageId="6" pageNumber="697" type="discussion">
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<paragraph id="8BC836F7FFB77F06325EFD66FAD7FD69" blockId="6.[809,1308,712,734]" box="[809,1308,712,734]" pageId="6" pageNumber="697">
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The most recent diagnosis of
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<taxonomicName id="4C774D74FFB77F063502FD66FB30FD69" authorityName="sensu Dendy & Row" authorityYear="1913" box="[1141,1275,712,734]" class="Calcarea" family="Leucaltidae" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="family">Leucaltidae</taxonomicName>
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is:
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</paragraph>
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</subSubSection>
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<subSubSection id="C36D657CFFB77F073236FD59FB50FEE1" lastPageId="7" lastPageNumber="698" pageId="6" pageNumber="697" type="description">
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<paragraph id="8BC836F7FFB77F063236FD59FB0CFBB6" blockId="6.[833,1400,759,1026]" pageId="6" pageNumber="697">
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<taxonomicName id="4C774D74FFB77F063236FD59FC19FCBA" authorityName="HARTMAN" authorityYear="1958" box="[833,978,759,781]" class="Calcarea" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="order">Clathrinida</taxonomicName>
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with a tubular, ramified or even anastomosed cormus with many oscula, or individualized with a large central atrium and a single osculum. The sponge wall is comprised of a distinct cortex sustained by a well-developed skeleton, and a choanosome. The skeleton of the choanosome and the atrial wall may be absent or comprised of small and dispersed triactines and tetractines (
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<bibRefCitation id="EFE64B06FFB77F0632BBFC42FB73FBB6" author="Borojevic R & Boury-Esnault N & Manuel M & Vacelet J" box="[972,1208,1004,1026]" pageId="6" pageNumber="697" pagination="1141 - 1152" refId="ref30334" refString="Borojevic R, Boury-Esnault N, Manuel M, Vacelet J. 2002. Order Clathrinida Hartman, 1958. In: Hooper JNA, Van Soest RWM, eds. Systema Porifera: a guide to the classification of sponges. New York: Kluwer Academic / Plenum Publishers, 1141 - 1152." type="book chapter" year="2002">
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Borojević
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<emphasis id="B903EAE5FFB77F06354BFC43FBBFFBB6" box="[1084,1140,1004,1026]" italics="true" pageId="6" pageNumber="697">et al.</emphasis>
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, 2002
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</bibRefCitation>
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).
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</paragraph>
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<paragraph id="8BC836F7FFB77F063236FBB5FBAAF9C0" blockId="6.[809,1426,1051,1901]" pageId="6" pageNumber="697">
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According to our molecular and morphological results, only
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<taxonomicName id="4C774D74FFB77F063216FB94FC01FBF8" authorityName="Haeckel" authorityYear="1872" box="[865,970,1082,1103]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="genus">
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<emphasis id="B903EAE5FFB77F063216FB94FC01FBF8" box="[865,970,1082,1103]" italics="true" pageId="6" pageNumber="697">Leucaltis</emphasis>
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</taxonomicName>
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should be included in
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<taxonomicName id="4C774D74FFB77F0635A7FB94FA91FBE7" authorityName="sensu Dendy & Row" authorityYear="1913" box="[1232,1370,1082,1104]" class="Calcarea" family="Leucaltidae" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="family">Leucaltidae</taxonomicName>
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. The aquiferous system of
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<taxonomicName id="4C774D74FFB77F063569FBF6FB4CFBDA" authorityName="Haeckel" authorityYear="1872" box="[1054,1159,1112,1133]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="genus">
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<emphasis id="B903EAE5FFB77F063569FBF6FB4CFBDA" box="[1054,1159,1112,1133]" italics="true" pageId="6" pageNumber="697">Leucaltis</emphasis>
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</taxonomicName>
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does not fit into any of the
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<specimenCount id="9D71FD7EFFB77F06322FFBD9FC00FB3B" box="[856,971,1143,1165]" pageId="6" pageNumber="697" type="generic" typeStatus="types">five types</specimenCount>
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accepted up to date. Past works have considered it analogous to the leuconoid (e.g.
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<bibRefCitation id="EFE64B06FFB77F063432FB38FC78FB7E" author="Lanna E & Cavalcanti FF & Cardoso L & Muricy G & Klautau M" pageId="6" pageNumber="697" pagination="1 - 27" refId="ref32290" refString="Lanna E, Cavalcanti FF, Cardoso L, Muricy G, Klautau M. 2009. Taxonomy of calcareous sponges (Porifera, Calcarea) from Potiguar Basin, NE Brazil. Zootaxa 1973: 1 - 27." type="journal article" year="2009">
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Lanna
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<emphasis id="B903EAE5FFB77F06325EFB1BFCACFB7E" box="[809,871,1204,1226]" italics="true" pageId="6" pageNumber="697">et al.</emphasis>
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, 2009
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</bibRefCitation>
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) or syconoid (e.g.
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<bibRefCitation id="EFE64B06FFB77F0635EFFB1AFCA9FB5F" author="Worheide G & Hooper JNA" pageId="6" pageNumber="697" pagination="859 - 891" refId="ref33996" refString="Worheide G, Hooper JNA. 1999. Calcarea from the Great Barrier Reef. I: cryptic Calcinea from Heron Island and Wistari Reef (Capricorn-Bunker group). Memoirs of the Queensland Museum 43: 859 - 891." type="journal article" year="1999">Wörheide & Hooper, 1999</bibRefCitation>
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) aquiferous systems, although the choanocyte chambers are not spherical nor do they have a regular radial orientation around the atrium. Pinacocytes are restricted to the external and atrial surfaces.
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<bibRefCitation id="EFE64B06FFB77F063432FA81FCB9FAD4" author="Dendy A" pageId="6" pageNumber="697" pagination="159 - 257" refId="ref30997" refString="Dendy A. 1893. Studies on the comparative anatomy of sponges. V. Observations on the structure and classification of the Calcarea Heterocoela. Quarterly Journal of Microscopical Science 35: 159 - 257." type="journal article" year="1893">Dendy (1893)</bibRefCitation>
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described the ‘
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<typeStatus id="54CC8855FFB77F063553FAE0FB93FAD4" box="[1060,1112,1358,1379]" pageId="6" pageNumber="697">type</typeStatus>
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C’ for this organization of the choanocyte chambers, which are mostly elongated and ramified, commonly fusing with each other (
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<figureCitation id="134C2A72FFB77F063432FA25FA48FA17" box="[1349,1411,1419,1441]" captionStart="Figure 3" captionStartId="7.[164,242,1800,1822]" captionTargetBox="[191,1414,756,1766]" captionTargetId="figure-243@7.[187,1419,752,1771]" captionTargetPageId="7" captionText="Figure 3. Organization of the aquiferous system and anatomy of Leucaltis spp. A, body wall of Leucaltis nodusgordii after Poléjaeff (1883). B, body wall of Leucaltis clathria after Dendy (1893). C, body wall of Leucaltis nodusgordii from La Réunion (UFRJPOR 8930), with the ramified choanocyte chambers outlined in yellow. D, body wall of Leucaltis latitubulata comb. nov. (SME 503), with pinacocytes in pink and choanocytes in blue." pageId="6" pageNumber="697">Fig. 3</figureCitation>
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). It must not be mistaken with the solenoid aquiferous system because, in
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<taxonomicName id="4C774D74FFB77F06328DFA66FBA9FA6A" authorityName="Haeckel" authorityYear="1872" box="[1018,1122,1480,1501]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="genus">
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<emphasis id="B903EAE5FFB77F06328DFA66FBA9FA6A" box="[1018,1122,1480,1501]" italics="true" pageId="6" pageNumber="697">Leucaltis</emphasis>
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</taxonomicName>
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, the choanoderm is present inside the ramified chambers, instead of revesting the internal part of the anastomosed choanocyte tubes. Finally, it differs from the sylleibid aquiferous system because choanocyte chambers do not group around a common exhalant chamber.
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</paragraph>
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<paragraph id="8BC836F7FFB77F073236F92EFD40FD9B" blockId="6.[809,1426,1051,1901]" lastBlockId="7.[163,779,197,679]" lastPageId="7" lastPageNumber="698" pageId="6" pageNumber="697">
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This kind of aquiferous system is present in five out of the six accepted species (following this work).
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<taxonomicName id="4C774D74FFB77F06345EF931FC57F965" authorityName="Lopes & Klautau" authorityYear="2023" baseAuthorityName="Preiwisch" baseAuthorityYear="1904" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="species" species="sambucus">
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<emphasis id="B903EAE5FFB77F06345EF931FC57F965" italics="true" pageId="6" pageNumber="697">Leucaltis sambucus</emphasis>
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</taxonomicName>
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is said to be leuconoid, but samples of this species were not found, hence we cannot corroborate if it has indeed a leuconoid aquiferous system. Considering that the aquiferous system of
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<taxonomicName id="4C774D74FFB77F063452F8B7FA5BF899" authorityName="Haeckel" authorityYear="1872" box="[1317,1424,1817,1838]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="genus">
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<emphasis id="B903EAE5FFB77F063452F8B7FA5BF899" box="[1317,1424,1817,1838]" italics="true" pageId="6" pageNumber="697">Leucaltis</emphasis>
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</taxonomicName>
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has been referred to as leuconoid in past descriptions, it is possible that
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<taxonomicName id="4C774D74FFB77F063575F8F9FB5EF8DB" authorityName="Lopes & Klautau" authorityYear="2023" baseAuthorityName="Preiwisch" baseAuthorityYear="1904" box="[1026,1173,1879,1900]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="6" pageNumber="697" phylum="Porifera" rank="species" species="sambucus">
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<emphasis id="B903EAE5FFB77F063575F8F9FB5EF8DB" box="[1026,1173,1879,1900]" italics="true" pageId="6" pageNumber="697">L. sambucus</emphasis>
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</taxonomicName>
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is not leuconoid, but has the distinct aquiferous system of the other
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<taxonomicName id="4C774D74FFB67F0731D4FF4AFEBCFF4D" box="[163,375,228,250]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="species" species="undetermined">
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<emphasis id="B903EAE5FFB67F0731D4FF4AFED9FF4E" box="[163,274,228,249]" italics="true" pageId="7" pageNumber="698">Leucaltis</emphasis>
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species.
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</taxonomicName>
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<taxonomicName id="4C774D74FFB67F0730F6FF4AFDA8FF4E" authorityName="Lopes & Klautau" authorityYear="2023" baseAuthorityName="Haeckel" baseAuthorityYear="1872" box="[385,611,228,249]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="species" species="corticata">
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<emphasis id="B903EAE5FFB67F0730F6FF4AFDA8FF4E" box="[385,611,228,249]" italics="true" pageId="7" pageNumber="698">Leucaltis corticata</emphasis>
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</taxonomicName>
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is mentioned as having a similar aquiferous system to
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<taxonomicName id="4C774D74FFB67F0733EAFEACFE28FE81" authority="(Polejaeff, 1883: 10)" baseAuthorityName="Polejaeff" baseAuthorityPageNumber="10" baseAuthorityYear="1883" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="species" species="clathria">
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<emphasis id="B903EAE5FFB67F0733EAFEACFF36FE81" italics="true" pageId="7" pageNumber="698">Leucaltis clathria</emphasis>
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(
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<bibRefCitation id="EFE64B06FFB67F07307BFE8FFE11FE81" author="Polejaeff N" box="[268,474,289,311]" pageId="7" pageNumber="698" pagination="1 - 76" refId="ref32971" refString="Polejaeff N. 1883. Report on the Calcarea dredged by the HMS Challenger during the years 1873 - 1876. Report on the Scientific Results of the Voyage Challenger (Zoology) 8: 1 - 76." type="journal article" year="1883">Poléjaeff, 1883: 10</bibRefCitation>
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)
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</taxonomicName>
|
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|
: ‘with the extension of the flagellated chambers into larger cavities of irregular outline, which come into contact with each other and anastomose, thus forming still larger sinus-like spaces’. For that reason, we draw attention to the presence of the new aquiferous system of
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<taxonomicName id="4C774D74FFB67F0733F4FE14FD27FE78" authorityName="Haeckel" authorityYear="1872" box="[643,748,442,463]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="genus">
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<emphasis id="B903EAE5FFB67F0733F4FE14FD27FE78" box="[643,748,442,463]" italics="true" pageId="7" pageNumber="698">Leucaltis</emphasis>
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</taxonomicName>
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in
|
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<taxonomicName id="4C774D74FFB67F0731D4FE77FEBFFE58" authorityName="Lopes & Klautau" authorityYear="2023" baseAuthorityName="Haeckel" baseAuthorityYear="1872" box="[163,372,473,495]" class="Calcarea" family="Leucaltidae" genus="Leucaltis" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="species" species="corticata">
|
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<emphasis id="B903EAE5FFB67F0731D4FE77FEBFFE58" box="[163,372,473,495]" italics="true" pageId="7" pageNumber="698">Leucaltis corticata</emphasis>
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</taxonomicName>
|
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|
. We suggest the name kladonoid for this new aquiferous system, derived from the Greek
|
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<emphasis id="B903EAE5FFB67F0731D4FDB8FEC9FD9C" box="[163,258,534,555]" italics="true" pageId="7" pageNumber="698">kladotós</emphasis>
|
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(=
|
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|
<emphasis id="B903EAE5FFB67F073050FDB8FE53FD9B" box="[295,408,534,556]" italics="true" pageId="7" pageNumber="698">ΚΛαδωτός</emphasis>
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), branched, ramified.
|
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|
</paragraph>
|
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<paragraph id="8BC836F7FFB67F0731CCFD98FB50FEE1" blockId="7.[163,779,197,679]" lastBlockId="7.[827,1443,197,342]" pageId="7" pageNumber="698">
|
|||
|
For
|
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|
<taxonomicName id="4C774D74FFB67F07319EFD9BFEBBFDFC" authorityName="sensu Dendy & Row" authorityYear="1913" box="[233,368,565,587]" class="Calcarea" family="Leucaltidae" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="family">Leucaltidae</taxonomicName>
|
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|
, we propose to modify the diagnosis to:
|
|||
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<taxonomicName id="4C774D74FFB67F0731B3FDFAFE82FDDD" authorityName="HARTMAN" authorityYear="1958" box="[196,329,596,618]" class="Calcarea" kingdom="Animalia" order="Clathrinida" pageId="7" pageNumber="698" phylum="Porifera" rank="order">Clathrinida</taxonomicName>
|
|||
|
with a body comprised of large, ramified and anastomosed tubes; never clathroid. Each tube has a distinct cortex, a choanoderm and a central atrium. The skeleton of the choanosome and atrium is comprised of regular and/or sagittal triactines and tetractines much smaller than the cortical ones. Aquiferous system kladonoid, with elongated and ramified choanocyte chambers.
|
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</paragraph>
|
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</subSubSection>
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</treatment>
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</document>
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