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<mods:title id="284C7510E0D80573C4D099F523D18EB6">Carlgrens hesitation allayed: redescription and systematics of Heteranthus verruculatus Klunzinger, 1877 (Cnidaria, Actiniaria), with a redefinition of Heteranthidae Carlgren, 1900</mods:title>
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<paragraph id="C046B41EB374FFF7EBDCB674FDC2FEA1" blockId="7.[98,610,207,1741]" box="[129,538,351,377]" pageId="7" pageNumber="162">
Family
<taxonomicName id="07F9CF9DB374FFF7EB8FB674FDC2FEA1" ID-CoL="84G88" authority="Carlgren, 1900" authorityName="Carlgren" authorityYear="1900" box="[210,538,351,377]" class="Anthozoa" family="Heteranthidae" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="family">
Heteranthidae
<bibRefCitation id="A468C9EFB374FFF7EA26B674FDC2FEA1" author="Carlgren, O." box="[379,538,351,377]" pageId="7" pageNumber="162" pagination="21 - 144" refId="ref13017" refString="Carlgren, O. (1900) Ostafrikanische Actinien. Gesammelt von Herrn Dr. F. Stuhlmann 1888 und 1889. Mitt. Naturh. Museum, 17, 21 - 144." type="journal article" year="1900">Carlgren, 1900</bibRefCitation>
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<paragraph id="C046B41EB374FFF7EBDCB6A8FE43FDAD" blockId="7.[98,610,207,1741]" pageId="7" pageNumber="162">
We revised
<bibRefCitation id="A468C9EFB374FFF7EA5AB6A8FE37FE45" author="Carlgren, O." box="[263,495,387,413]" pageId="7" pageNumber="162" pagination="21 - 144" refId="ref13017" refString="Carlgren, O. (1900) Ostafrikanische Actinien. Gesammelt von Herrn Dr. F. Stuhlmann 1888 und 1889. Mitt. Naturh. Museum, 17, 21 - 144." type="journal article" year="1900">Carlgrens (1900: 92)</bibRefCitation>
diagnosis written in German. Key changes interpreted from recent studies (e.g.,
<bibRefCitation id="A468C9EFB374FFF7EAFDB6E0FF05FDD1" author="Gonzalez-Munoz, R. &amp; Simoes, N. &amp; Mascaro, M. &amp; Tello-Musi, J. L. &amp; Brugler, M. R. &amp; Rodriguez, E." pageId="7" pageNumber="162" pagination="69 - 79" refId="ref13856" refString="Gonzalez-Munoz, R., Simoes, N., Mascaro, M., Tello-Musi, J. L., Brugler, M. R. &amp; Rodriguez, E. (2015) Morphological and molecular variability of the sea anemone Phymanthus crucifer (Cnidaria, Anthozoa, Actiniaria, Actinoidea). J. Mar. Biol. Assoc. UK., 95, 69 - 79." type="journal article" year="2015">González-Muñoz et al., 2015</bibRefCitation>
;
<bibRefCitation id="A468C9EFB374FFF7EBB4B6C4FE18FDD1" author="Brugler, M. R. &amp; Gonzalez-Munoz, R. E. &amp; Tessler, M. &amp; Rodriguez, E." box="[233,448,495,521]" pageId="7" pageNumber="162" pagination="756 - 776" refId="ref12968" refString="Brugler, M. R., Gonzalez-Munoz, R. E., Tessler, M. &amp; Rodriguez, E. (2018) An EPIC Journey to locate single-copy nuclear markers in sea anemones. Zool. Scr., 47, 756 - 776." type="journal article" year="2018">Brugler et al., 2018</bibRefCitation>
) and are here indicated in
<emphasis id="F28D680CB374FFF7EBACB538FEFDFDF5" bold="true" box="[241,293,531,557]" pageId="7" pageNumber="162">bold</emphasis>
; all minor additions (i.e., of word order, updated terminology, adjectives for clarity, etc.) are in
<emphasis id="F28D680CB374FFF7EA09B571FE4DFDAC" box="[340,405,602,628]" italics="true" pageId="7" pageNumber="162">italics</emphasis>
.
</paragraph>
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<paragraph id="C046B41EB374FFF7EBDCB554FDBAFBFB" blockId="7.[98,610,207,1741]" pageId="7" pageNumber="162">
<emphasis id="F28D680CB374FFF7EBDCB554FF28FD40" box="[129,240,639,664]" italics="true" pageId="7" pageNumber="162">Diagnosis.</emphasis>
<taxonomicName id="07F9CF9DB374FFF7EA53B555FE4BFD40" authorityName="Rafinesque" authorityYear="1815" box="[270,403,638,664]" class="Anthozoa" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="superFamily" superFamily="Actinioidea">
<emphasis id="F28D680CB374FFF7EA53B555FE4BFD40" bold="true" box="[270,403,638,664]" pageId="7" pageNumber="162">Actinioidea</emphasis>
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with a well-developed
<emphasis id="F28D680CB374FFF7EBADB589FEF4FD64" box="[240,300,674,700]" italics="true" pageId="7" pageNumber="162">pedal</emphasis>
disc. Sphincter
<emphasis id="F28D680CB374FFF7E946B589FFAFFD38" italics="true" pageId="7" pageNumber="162">diffuse to</emphasis>
circumscript,
<emphasis id="F28D680CB374FFF7EA6FB5EDFE30FD38" box="[306,488,710,736]" italics="true" pageId="7" pageNumber="162">weak to strong</emphasis>
.
<emphasis id="F28D680CB374FFF7E95EB5EDFE1BFCDC" bold="true" pageId="7" pageNumber="162">Number of siphonoglyphs variable</emphasis>
.
<emphasis id="F28D680CB374FFF7EAB3B5C1FDD0FCF0" italics="true" pageId="7" pageNumber="162">Distalmost margin with marginal projections</emphasis>
, fosse
<emphasis id="F28D680CB374FFF7EB3FB419FEEBFC94" box="[98,307,818,844]" italics="true" pageId="7" pageNumber="162">generally shallow</emphasis>
.
<emphasis id="F28D680CB374FFF7EA05B41AFDB9FCB7" bold="true" pageId="7" pageNumber="162">Distal column with verrucae. Number of mesenteries variable,</emphasis>
<emphasis id="F28D680CB374FFF7EB3FB451FE5AFC4B" box="[98,386,889,915]" italics="true" pageId="7" pageNumber="162">retractor muscles strong</emphasis>
,
<emphasis id="F28D680CB374FFF7EAC3B452FF22FC6F" bold="true" pageId="7" pageNumber="162">typically diffuse circumscript.</emphasis>
Tentacles
<emphasis id="F28D680CB374FFF7EA2DB4B6FE40FC03" italics="true" pageId="7" pageNumber="162">without protuberances nor globose in appearance</emphasis>
.
<emphasis id="F28D680CB374FFF7EAF2B4EAFDBAFBFB" bold="true" pageId="7" pageNumber="162">
Zooxanthellate. Cnidom: spirocysts, basitrichs, microbasic amastigophores,microbasic
<emphasis id="F28D680CB374FFF7EAC3B322FE6DFBFB" bold="true" box="[414,437,1033,1059]" italics="true" pageId="7" pageNumber="162">p-</emphasis>
mastigophores.
</emphasis>
</paragraph>
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<emphasis id="F28D680CB374FFF7EBDCB305FED9FB9F" box="[129,257,1070,1095]" italics="true" pageId="7" pageNumber="162">
<typeStatus id="1F420ABCB374FFF7EBDCB305FF6DFB9F" box="[129,181,1070,1095]" pageId="7" pageNumber="162">Type</typeStatus>
genus.
</emphasis>
<taxonomicName id="07F9CF9DB374FFF7EA5BB306FD91FB9F" authority="Klunzinger, 1877" authorityName="Klunzinger" authorityYear="1877" box="[262,585,1069,1095]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="genus">
<emphasis id="F28D680CB374FFF7EA5BB306FE56FB9F" box="[262,398,1069,1095]" italics="true" pageId="7" pageNumber="162">Heteranthus</emphasis>
<bibRefCitation id="A468C9EFB374FFF7EAC9B306FD91FB9F" author="Klunzinger, C. B." box="[404,585,1069,1095]" pageId="7" pageNumber="162" refId="ref14097" refString="Klunzinger, C. B. (1877) Die Korallthiere des rothen Meeres. Erster Theil: Die Alcyonarien und Malacodermen. Verlag der Gutmann'schen Buchhandlung, Berlin." type="book" year="1877">Klunzinger, 1877</bibRefCitation>
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</paragraph>
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<paragraph id="C046B41EB374FFF7EBDCB37AFE41FBB3" blockId="7.[98,610,207,1741]" box="[129,409,1105,1131]" pageId="7" pageNumber="162">
<emphasis id="F28D680CB374FFF7EBDCB37AFE89FBB3" box="[129,337,1105,1131]" italics="true" pageId="7" pageNumber="162">Other valid genera.</emphasis>
None.
</paragraph>
</subSubSection>
<subSubSection id="88E3E795B374FFF7EBDCB35EFEA9F972" pageId="7" pageNumber="162" type="discussion">
<paragraph id="C046B41EB374FFF7EBDCB35EFEA9F972" blockId="7.[98,610,207,1741]" pageId="7" pageNumber="162">
<emphasis id="F28D680CB374FFF7EBDCB35EFF3FFB57" box="[129,231,1141,1167]" italics="true" pageId="7" pageNumber="162">Remarks.</emphasis>
Based on morphological similarity,
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synonymised
<taxonomicName id="07F9CF9DB374FFF7EAE2B3B2FDBAFB6B" authorityName="Carlgren" authorityYear="1900" box="[447,610,1177,1203]" class="Anthozoa" family="Heteranthidae" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="family">Heteranthidae</taxonomicName>
and
<taxonomicName id="07F9CF9DB374FFF7EBCAB396FEE5FB0F" authorityName="Andres" authorityYear="1883" box="[151,317,1213,1239]" class="Anthozoa" family="Phymanthidae" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="family">Phymanthidae</taxonomicName>
, and classed
<taxonomicName id="07F9CF9DB374FFF7EA84B396FDB9FB0F" authorityName="Klunzinger" authorityYear="1877" box="[473,609,1213,1239]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="genus">
<emphasis id="F28D680CB374FFF7EA84B396FDB9FB0F" box="[473,609,1213,1239]" italics="true" pageId="7" pageNumber="162">Heteranthus</emphasis>
</taxonomicName>
in the latter. We resurrect
<taxonomicName id="07F9CF9DB374FFF7EAE2B3CBFDBAFB22" authorityName="Carlgren" authorityYear="1900" box="[447,610,1248,1274]" class="Anthozoa" family="Heteranthidae" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="family">Heteranthidae</taxonomicName>
for
<taxonomicName id="07F9CF9DB374FFF7EBCCB22FFEC1FAC6" authorityName="Klunzinger" authorityYear="1877" box="[145,281,1284,1310]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="genus">
<emphasis id="F28D680CB374FFF7EBCCB22FFEC1FAC6" box="[145,281,1284,1310]" italics="true" pageId="7" pageNumber="162">Heteranthus</emphasis>
</taxonomicName>
as molecular phylogenetic results from this and previous studies (see
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;
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) do not support
<bibRefCitation id="A468C9EFB374FFF7EAFCB25BFDB9FA52" author="Carlgren, O." box="[417,609,1392,1418]" pageId="7" pageNumber="162" pagination="1 - 43" refId="ref13090" refString="Carlgren, O. (1943) East-Asiatic Corallimorpharia and Actiniaria. K. Svenska Vet. Akad. Handl., 20, 1 - 43." type="journal article" year="1943">Carlgrens (1943)</bibRefCitation>
conclusion. Following this line of evidence, we also revised the mention of higher-level classification in the original diagnosis. Among characters absent from
<bibRefCitation id="A468C9EFB374FFF7EACAB12BFDBAF9C2" author="Carlgren, O." box="[407,610,1536,1562]" pageId="7" pageNumber="162" pagination="21 - 144" refId="ref13017" refString="Carlgren, O. (1900) Ostafrikanische Actinien. Gesammelt von Herrn Dr. F. Stuhlmann 1888 und 1889. Mitt. Naturh. Museum, 17, 21 - 144." type="journal article" year="1900">Carlgrens (1900)</bibRefCitation>
diagnosis are cnidom and arrangement of the internal musculature. We included these here as they are a necessary part of contemporary sea anemone taxonomy.
</paragraph>
</subSubSection>
</treatment>
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