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<document id="5D1F10EF94FBFF213C927EB66F1C601B" ID-DOI="10.1163/18759866-BJA10015" ID-ISSN="1875-9866" ID-Zenodo-Dep="8343299" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="juliana" checkinTime="1694663820210" checkinUser="felipe" docAuthor="Yap, Nicholas Wei Liang, Quek, Zheng Bin Randolph, Tan, Ria, Nugroho, Dharma Arif, Lee, Jen Nie, Lee Berumen, Michael, Tan, Koh Siang &amp; Huang, Danwei" docDate="2021" docId="48500508B374FFF8EBDCB198FC25FB57" docLanguage="en" docName="ContribZool.90.2.155-182.pdf" docOrigin="Contributions to Zoology 90 (2)" docSource="http://dx.doi.org/10.1163/18759866-bja10015" docStyle="DocumentStyle:E1C3C5FD6CDDA08D62BFE928526826AF.2:ContribZool.2019-.journal_article" docStyleId="E1C3C5FD6CDDA08D62BFE928526826AF" docStyleName="ContribZool.2019-.journal_article" docStyleVersion="2" docTitle="Heteranthus Klunzinger 1877" docType="treatment" docVersion="1" lastPageNumber="163" masterDocId="B4697D70B373FFF0EB5DB72BFFD8FFD8" masterDocTitle="Carlgrens hesitation allayed: redescription and systematics of Heteranthus verruculatus Klunzinger, 1877 (Cnidaria, Actiniaria), with a redefinition of Heteranthidae Carlgren, 1900" masterLastPageNumber="182" masterPageNumber="155" pageNumber="162" updateTime="1695054941782" updateUser="juliana" zenodo-license-document="CC-BY-4.0">
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<mods:title id="1AF38130B448CF4E7AFCB9A449ED2863">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="C046B41EB374FFF7EBDCB198FDC9F915" blockId="7.[98,610,207,1741]" box="[129,529,1715,1741]" pageId="7" pageNumber="162">
Genus
<taxonomicName id="07F9CF9DB374FFF7EB93B198FDC9F915" ID-CoL="7P5TY" ID-ENA="1326988" authority="Klunzinger, 1877" authorityName="Klunzinger" authorityYear="1877" box="[206,529,1715,1741]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="genus">
<emphasis id="F28D680CB374FFF7EB93B198FE8EF915" box="[206,342,1715,1741]" italics="true" pageId="7" pageNumber="162">Heteranthus</emphasis>
<bibRefCitation id="A468C9EFB374FFF7EA01B198FDC9F915" author="Klunzinger, C. B." box="[348,529,1715,1741]" 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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<subSubSection id="88E3E795B374FFF7E9F9B780FB5CFBB3" pageId="7" pageNumber="162" type="diagnosis">
<paragraph id="C046B41EB374FFF7E9F9B780FBF5FEEA" blockId="7.[645,1157,171,1742]" pageId="7" pageNumber="162">
Diagnosis modified after
<bibRefCitation id="A468C9EFB374FFF7E88FB780FB5CFF1D" author="Carlgren, O." box="[978,1156,171,197]" pageId="7" pageNumber="162" pagination="1 - 121" refId="ref13118" refString="Carlgren, O. (1949) A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria. K. Svenska Vet. Akad. Handl., 1, 1 - 121." type="journal article" year="1949">Carlgren (1949)</bibRefCitation>
and
<bibRefCitation id="A468C9EFB374FFF7E99EB7E4FC29FF32" author="Fautin, D. G. &amp; Crowther, A. L. &amp; Wallace, C. C." box="[707,1009,207,234]" pageId="7" pageNumber="162" pagination="35 - 64" refId="ref13721" refString="Fautin, D. G., Crowther, A. L. &amp; Wallace, C. C. (2008) Sea anemones (Cnidaria: Anthozoa: Actiniaria) of Moreton Bay. Mem. Queensl. Mus. - Nature, 54, 35 - 64." type="journal article" year="2008">Fautin et al. (2008: 52)</bibRefCitation>
. Substantial changes from our research are indicated in
<emphasis id="F28D680CB374FFF7E9D8B63CFD61FEE9" bold="true" box="[645,697,279,305]" pageId="7" pageNumber="162">bold</emphasis>
; all minor additions are in
<emphasis id="F28D680CB374FFF7E8BBB63CFBFFFEE9" box="[998,1063,279,305]" italics="true" pageId="7" pageNumber="162">italics</emphasis>
.
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<emphasis id="F28D680CB374FFF7E9F9B617FCCEFE8D" box="[676,790,316,341]" italics="true" pageId="7" pageNumber="162">Diagnosis.</emphasis>
Pedal disc well-developed. Column, apart from most proximal part, with verrucae, which are smaller and more numerous at the margin, and overhang the fosse,
<emphasis id="F28D680CB374FFF7E85BB6E1FC3DFE3C" box="[774,997,458,484]" italics="true" pageId="7" pageNumber="162">resembling papillae</emphasis>
. Endodermal marginal sphincter muscle,
<emphasis id="F28D680CB374FFF7E8B4B6C5FB5DFDD1" bold="true" box="[1001,1157,494,521]" pageId="7" pageNumber="162">restricted to</emphasis>
circumscribed
<emphasis id="F28D680CB374FFF7E864B538FC4CFDF4" box="[825,916,530,556]" italics="true" pageId="7" pageNumber="162">in form</emphasis>
,
<emphasis id="F28D680CB374FFF7E8F3B539FBF1FDF4" bold="true" box="[942,1065,530,556]" pageId="7" pageNumber="162">may form</emphasis>
a few folds.
<emphasis id="F28D680CB374FFF7E98AB51DFD36FDAC" bold="true" pageId="7" pageNumber="162">Tentacles as marginal and discal tentacles</emphasis>
. Marginal tentacles conical, simple,
<emphasis id="F28D680CB374FFF7E9B0B555FBA7FD40" bold="true" box="[749,1151,638,664]" pageId="7" pageNumber="162">without ramified protuberances</emphasis>
, arranged in cycles,
<emphasis id="F28D680CB374FFF7E833B589FCE3FD38" bold="true" pageId="7" pageNumber="162">innermost cycle longer than outermost</emphasis>
; discal tentacles shorter and papilliform,
<emphasis id="F28D680CB374FFF7E846B5C1FC72FCDC" box="[795,938,746,772]" italics="true" pageId="7" pageNumber="162">arranged as</emphasis>
<emphasis id="F28D680CB374FFF7E8E1B5C1FB5CFCDC" bold="true" box="[956,1156,746,772]" pageId="7" pageNumber="162">dense or sparse</emphasis>
radial rows.
<emphasis id="F28D680CB374FFF7E876B425FBA7FCF0" box="[811,1151,782,808]" italics="true" pageId="7" pageNumber="162">Siphonoglyphs well-developed</emphasis>
,
<emphasis id="F28D680CB374FFF7E9D8B419FCFBFC94" bold="true" box="[645,803,818,844]" pageId="7" pageNumber="162">when present</emphasis>
.
<emphasis id="F28D680CB374FFF7E872B419FB96FC93" box="[815,1102,818,844]" italics="true" pageId="7" pageNumber="162">Number of directives pairs</emphasis>
,
<emphasis id="F28D680CB374FFF7EF07B419FD65FCB7" italics="true" pageId="7" pageNumber="162">and pairs</emphasis>
of
<emphasis id="F28D680CB374FFF7E9BEB47EFC2EFCB7" box="[739,1014,853,879]" italics="true" pageId="7" pageNumber="162">complete and incomplete</emphasis>
mesenteries
<emphasis id="F28D680CB374FFF7E9D8B452FD3BFC6F" bold="true" pageId="7" pageNumber="162">variable, gametogenic tissue may be present on both</emphasis>
; retractor muscles diffuse
<emphasis id="F28D680CB374FFF7EF48B4B6FCC1FC03" bold="true" pageId="7" pageNumber="162">to diffuse circumscript</emphasis>
, well developed. Parietobasilar muscles weak to fairly strong.
<emphasis id="F28D680CB374FFF7EF5BB4CEFBA6FBFB" bold="true" pageId="7" pageNumber="162">Frequently propagates by longitudinal fission</emphasis>
. Cnidom: spirocysts, basitrichs,
<emphasis id="F28D680CB374FFF7EF58B306FCE5FBB3" bold="true" pageId="7" pageNumber="162">microbasic amastigophores</emphasis>
, microbasic
<emphasis id="F28D680CB374FFF7E894B379FC0FFBB3" box="[969,983,1106,1131]" italics="true" pageId="7" pageNumber="162">p</emphasis>
-mastigophores
</paragraph>
</subSubSection>
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<paragraph id="C046B41EB374FFF7E9F9B35EFCA9FB57" blockId="7.[645,1157,171,1742]" box="[676,881,1141,1167]" pageId="7" pageNumber="162">
<emphasis id="F28D680CB374FFF7E9F9B35EFD20FB57" box="[676,760,1141,1167]" italics="true" pageId="7" pageNumber="162">Gender.</emphasis>
Masculine
</paragraph>
</subSubSection>
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<emphasis id="F28D680CB374FFF7E9F9B3B2FC9AFB6A" box="[676,834,1177,1202]" italics="true" pageId="7" pageNumber="162">
<typeStatus id="1F420ABCB374FFF7E9F9B3B2FD0FFB6A" box="[676,727,1177,1202]" pageId="7" pageNumber="162">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="07F9CF9DB374FFF7E803B3B2FC9AFB0F" authority="Klunzinger, 1877" authorityName="Klunzinger" authorityYear="1877" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="7" pageNumber="162" phylum="Cnidaria" rank="species" species="verruculatus">
<emphasis id="F28D680CB374FFF7E803B3B2FB5DFB6B" box="[862,1157,1177,1203]" italics="true" pageId="7" pageNumber="162">Heteranthus verruculatus</emphasis>
<bibRefCitation id="A468C9EFB374FFF7E9D8B397FC9AFB0F" author="Klunzinger, C. B." box="[645,834,1212,1239]" 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>
</taxonomicName>
, by monotypy (see
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)
</paragraph>
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<emphasis id="F28D680CB374FFF7E9F9B22FFCD4FAC6" box="[676,780,1284,1310]" italics="true" pageId="7" pageNumber="162">Remarks.</emphasis>
<bibRefCitation id="A468C9EFB374FFF7E852B22FFC07FAC7" author="Carlgren, O." box="[783,991,1284,1311]" pageId="7" pageNumber="162" pagination="1 - 121" refId="ref13118" refString="Carlgren, O. (1949) A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria. K. Svenska Vet. Akad. Handl., 1, 1 - 121." type="journal article" year="1949">Carlgren (1949: 75)</bibRefCitation>
stated that the column is “provided with large verrucae,” but we found this character to be variable in size and visibility, among preserved specimens. We adhere to the rewording by
<bibRefCitation id="A468C9EFB374FFF7EF4AB2BFFD2DFA0A" author="Fautin, D. G. &amp; Crowther, A. L. &amp; Wallace, C. C." pageId="7" pageNumber="162" pagination="35 - 64" refId="ref13721" refString="Fautin, D. G., Crowther, A. L. &amp; Wallace, C. C. (2008) Sea anemones (Cnidaria: Anthozoa: Actiniaria) of Moreton Bay. Mem. Queensl. Mus. - Nature, 54, 35 - 64." type="journal article" year="2008">Fautin et al. (2008)</bibRefCitation>
in not specifying size of verrucae to allow for this variation. On marginal tentacles,
<bibRefCitation id="A468C9EFB374FFF7E85FB12BFC0CF9C2" author="Klunzinger, C. B." box="[770,980,1536,1562]" 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>
reported that these were of equal length, but
<bibRefCitation id="A468C9EFB374FFF7EF4AB10FFD2FF9BA" author="Fautin, D. G. &amp; Crowther, A. L. &amp; Wallace, C. C." pageId="7" pageNumber="162" pagination="35 - 64" refId="ref13721" refString="Fautin, D. G., Crowther, A. L. &amp; Wallace, C. C. (2008) Sea anemones (Cnidaria: Anthozoa: Actiniaria) of Moreton Bay. Mem. Queensl. Mus. - Nature, 54, 35 - 64." type="journal article" year="2008">Fautin et al. (2008)</bibRefCitation>
found otherwise in studying live specimens. Likewise, we observed that the innermost cycle of marginal tentacles was longer than those at the outermost in living individuals. Note that in many preserved specimens, this feature may be less-pronounced and tentacles may contract. Specification of the number of siphonoglyphs present was removed as this varied among individuals or be absent entirely. Similarly, the number of directive pairs, and pairs of complete and incomplete mesenteries also varies among individuals. Like both
<bibRefCitation id="A468C9EFB37BFFF8E940B22FFF1FFA9B" author="Carlgren, O." pageId="8" pageNumber="163" pagination="1 - 121" refId="ref13118" refString="Carlgren, O. (1949) A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria. K. Svenska Vet. Akad. Handl., 1, 1 - 121." type="journal article" year="1949">Carlgren (1949)</bibRefCitation>
and
<bibRefCitation id="A468C9EFB37BFFF8EA5DB203FE07FA9B" author="Fautin, D. G. &amp; Crowther, A. L. &amp; Wallace, C. C." box="[256,479,1320,1347]" pageId="8" pageNumber="163" pagination="35 - 64" refId="ref13721" refString="Fautin, D. G., Crowther, A. L. &amp; Wallace, C. C. (2008) Sea anemones (Cnidaria: Anthozoa: Actiniaria) of Moreton Bay. Mem. Queensl. Mus. - Nature, 54, 35 - 64." type="journal article" year="2008">Fautin et al. (2008)</bibRefCitation>
, we too failed to find any gametogenic tissues in all but those specimens from
<collectingRegion id="023D7AFCB37BFFF8EAFEB25AFDA4FA52" box="[419,636,1392,1419]" country="Australia" name="Western Australia" pageId="8" pageNumber="163">Western Australia</collectingRegion>
. Moreover, we observed that living polyps readily undergo longitudinal fission. The diagnosis is modified to indicate the presence of gametogenic tissue in some individuals and its reproductive mode.
</paragraph>
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<paragraph id="C046B41EB37BFFF8EBDCB5E9FD3AFC7D" blockId="8.[129,1174,706,934]" pageId="8" pageNumber="163">
FIGURE 2 Name-bearing type materials of
<taxonomicName id="07F9CF9DB37BFFF8E97DB5E9FCDDFD0F" authorityName="Klunzinger" authorityYear="1877" box="[544,773,706,727]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="8" pageNumber="163" phylum="Cnidaria" rank="species" species="verruculatus">
<emphasis id="F28D680CB37BFFF8E97DB5E9FCDDFD0F" box="[544,773,706,727]" italics="true" pageId="8" pageNumber="163">Heteranthus verruculatus</emphasis>
</taxonomicName>
and
<taxonomicName id="07F9CF9DB37BFFF8E86FB5E9FC32FD00" authorityName="Carlgren" authorityYear="1943" box="[818,1002,706,728]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="8" pageNumber="163" phylum="Cnidaria" rank="species" species="insignis">
<emphasis id="F28D680CB37BFFF8E86FB5E9FC32FD00" box="[818,1002,706,728]" italics="true" pageId="8" pageNumber="163">Heteranthus insignis</emphasis>
</taxonomicName>
. A. dried lectotype of
<taxonomicName id="07F9CF9DB37BFFF8EA4CB5CBFDDFFD2E" authority="(ZMB 1852)" baseAuthorityName="ZMB" baseAuthorityYear="1852" box="[273,519,736,758]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="8" pageNumber="163" phylum="Cnidaria" rank="species" species="verruculatus">
<emphasis id="F28D680CB37BFFF8EA4CB5CBFE43FD2D" box="[273,411,736,757]" italics="true" pageId="8" pageNumber="163">H. verruculatus</emphasis>
(ZMB 1852)
</taxonomicName>
, top view. B. close-up of ZMB 1852 in two pieces, showing details of distalmost end, and mesenteries, top view. Note that a slice of the distalmost end is missing. C. the piece of distalmost end from lectotype ZMB 1852, now at Naturhistoriska Riksmuseet (NRS 4861), Sweden, side view. Note details of conspicuous verrucae and marginal projections with papillae- like bumps. D. holotype of
<taxonomicName id="07F9CF9DB37BFFF8EAB3B47DFD93FCB3" authorityName="Carlgren" authorityYear="1943" box="[494,587,854,875]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="8" pageNumber="163" phylum="Cnidaria" rank="species" species="insignis">
<emphasis id="F28D680CB37BFFF8EAB3B47DFD93FCB3" box="[494,587,854,875]" italics="true" pageId="8" pageNumber="163">H. insignis</emphasis>
</taxonomicName>
, an entire specimen (NRS 4076), side view. Abbreviations: a, actinopharynx; m, mesenteries; mp, marginal projection; mt, marginal tentacles; pd, pedal disc; v, verrucae. Scale bar = 1 mm. Photographs by NWL Yap.
</paragraph>
</caption>
<paragraph id="C046B41EB37BFFF8EBC2B163FC25FB57" blockId="8.[129,640,997,1742]" lastBlockId="8.[676,1187,997,1742]" pageId="8" pageNumber="163">
We confirm the presence of microbasic amastigophores in live materials examined for cnidae; this cnidae
<typeStatus id="1F420ABCB37BFFF8EA0BB1BBFE5FF972" box="[342,391,1680,1706]" pageId="8" pageNumber="163">type</typeStatus>
has not been reported in
<taxonomicName id="07F9CF9DB37BFFF8EBFFB198FEF2F915" authorityName="Klunzinger" authorityYear="1877" box="[162,298,1715,1741]" class="Anthozoa" family="Heteranthidae" genus="Heteranthus" kingdom="Animalia" order="Actiniaria" pageId="8" pageNumber="163" phylum="Cnidaria" rank="genus">
<emphasis id="F28D680CB37BFFF8EBFFB198FEF2F915" box="[162,298,1715,1741]" italics="true" pageId="8" pageNumber="163">Heteranthus</emphasis>
</taxonomicName>
. Because unfired capsules of microbasic amastigophores strongly resemble those of microbasic
<emphasis id="F28D680CB37BFFF8E8CDB321FC7EFBFB" box="[912,934,1034,1059]" italics="true" pageId="8" pageNumber="163">p-</emphasis>
mastigophores in preserved specimens, they may have been mistaken for the latter and were not reported (see
<bibRefCitation id="A468C9EFB37BFFF8E9F9B35BFC9BFB57" author="Ostman, C." box="[676,835,1136,1167]" pageId="8" pageNumber="163" pagination="31 - 46" refId="ref14298" refString="Ostman, C. (2000) A guideline to nematocyst nomenclature and classification, and some notes on the systematic value of nematocysts. Sci. Mar., 64, 31 - 46." type="journal article" year="2000">Östman, 2000</bibRefCitation>
;
<bibRefCitation id="A468C9EFB37BFFF8E811B35EFC29FB57" author="Yap, N. W. L. &amp; Fautin, D. G. &amp; Ramos, D. A. &amp; Tan, R." box="[844,1009,1141,1167]" pageId="8" pageNumber="163" pagination="439 - 454" refId="ref14872" refString="Yap, N. W. L., Fautin, D. G., Ramos, D. A. &amp; Tan, R. (2014) Sea anemones of Singapore: Synpeachia temasek new genus, new species, and redescription of Metapeachia tropica (Cnidaria: Actiniaria: Haloclavidae). Proc. Biol. Soc. Wash., 127, 439 - 454." type="journal article" year="2014">Yap et al., 2014</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>