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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.606.9310" ID-GBIF-Dataset="7b02d07d-db82-4d57-aafb-11a3d0604281" ID-PMC="PMC4978008" ID-Pensoft-Pub="1313-2970-606-11" ID-PubMed="27551219" ID-ZBK="EB98FE3B665B4CF28E8B740D167BA2BB" ModsDocAuthor="" ModsDocDate="2016" ModsDocID="1313-2970-606-11" ModsDocOrigin="ZooKeys 606" ModsDocTitle="A new solitary free-living species of the genus Sphenopus (Cnidaria, Anthozoa, Zoantharia, Sphenopidae) from Okinawa-jima Island, Japan" checkinTime="1469161625121" checkinUser="pensoft" docAuthor="Fujii, Takuma &amp; Reimer, James Davis" docDate="2016" docId="8B59E29DC1F988689BEED295A21975DF" docLanguage="en" docName="ZooKeys 606: 11-24" docOrigin="ZooKeys 606" docSource="http://dx.doi.org/10.3897/zookeys.606.9310" docTitle="Sphenopus exilis Fujii &amp; Reimer, 2016, sp. n." docType="treatment" docUuid="30C107C6-8104-4EC9-9DC9-CF2DEE4A9638" docUuidSource="ZooBank" docVersion="5" lastPageNumber="19" masterDocId="FFBDFF8BB575A578EB71FF9EFFA0FFF7" masterDocTitle="A new solitary free-living species of the genus Sphenopus (Cnidaria, Anthozoa, Zoantharia, Sphenopidae) from Okinawa-jima Island, Japan" masterLastPageNumber="24" masterPageNumber="11" pageNumber="13" updateTime="1668163344316" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>A new solitary free-living species of the genus Sphenopus (Cnidaria, Anthozoa, Zoantharia, Sphenopidae) from Okinawa-jima Island, Japan</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fujii, Takuma</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Reimer, James Davis</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2016</mods:date>
<mods:detail type="volume">
<mods:number>606</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>11</mods:start>
<mods:end>24</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.606.9310</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.606.9310</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-606-11</mods:identifier>
<mods:identifier type="ZBK">EB98FE3B665B4CF28E8B740D167BA2BB</mods:identifier>
<mods:identifier type="ZooBank">EB98FE3B665B4CF28E8B740D167BA2BB</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="127861685" LSID="urn:lsid:zoobank.org:act:30C107C6-8104-4EC9-9DC9-CF2DEE4A9638" httpUri="http://treatment.plazi.org/id/8B59E29DC1F988689BEED295A21975DF" lastPageId="8" lastPageNumber="19" pageId="2" pageNumber="13">
<subSubSection pageId="2" pageNumber="13" type="multiple">
<paragraph pageId="2" pageNumber="13">Taxon classification Animalia Zoantharia Sphenopidae</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="13" type="nomenclature">
<paragraph pageId="2" pageNumber="13">
<taxonomicName LSID="http://zoobank.org/30C107C6-8104-4EC9-9DC9-CF2DEE4A9638" class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="2" pageNumber="13" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
<taxonomicNameLabel pageId="2" pageNumber="13">sp. n.</taxonomicNameLabel>
Figures 2, 3
</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="14" pageId="2" pageNumber="13" type="holotype">
<paragraph pageId="2" pageNumber="13">Holotype.</paragraph>
<paragraph lastPageId="3" lastPageNumber="14" pageId="2" pageNumber="13">
Specimen number NSMT-Co1576 (MISE-TF-107): Kin Bay, Uruma, Okinawa-jima Island, Japan (
<geoCoordinate direction="north" orientation="latitude" precision="15" value="26.373611">26°22'25&quot;N</geoCoordinate>
,
<geoCoordinate direction="east" orientation="longitude" precision="15" value="127.89166">127°53'30&quot;E</geoCoordinate>
), 15 m depth, collected by
<pageBreakToken pageId="3" pageNumber="14" start="start">Takuma</pageBreakToken>
Fujii, 29 October 2011, fixed in 5-10% SW formalin, deposited in National Museum of Nature and Science, Tokyo, Japan (NSMT). Polyp length 2.4 cm, maximum width 0.8 cm, minimum width 0.3 cm. Figure 2B.
</paragraph>
<caption pageId="3" pageNumber="14">
<paragraph pageId="3" pageNumber="14">
Figure 2. Polyps of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="3" pageNumber="14" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. A In situ image of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="3" pageNumber="14" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n., polyp with no black patterns, from the type locality in Kin Bay, Okinawa, Japan on 29 October 2011 B Polyps of NSMT-Co1576 &amp; NSMT-Co1577 from Kin Bay, Okinawa-jima Island, Japan. The white circle points to the holotype C In situ image of NSMT-Co1578 from Oura Bay, Okinawa-jima Island, Japan, on 13 November 2012. Faint black patterns and bands appear on the oral disc and the tentacles D Polyps of lot number NSMT-Co1578 showing phenotypic variation with black stripes on the upper part of the polyps. Scale bars: 1 cm.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="3" pageNumber="14" type="paratypes">
<paragraph pageId="3" pageNumber="14">Paratypes.</paragraph>
<paragraph pageId="3" pageNumber="14">
Specimen number NSMT-Co1577 (MISE-TF-107), a lot of total 11 polyps collected on the same dive, collection data same as holotype, five polyps fixed in 5-10% formalin, six polyps fixed in 99% EtOH, polyp length 1.3 to 2.2 cm (average 1.7
<normalizedToken originalValue="±">+/-</normalizedToken>
0.3 cm), maximum width 0.4 to 1.0 cm (average 0.5
<normalizedToken originalValue="±">+/-</normalizedToken>
0.2 cm), minimum width 0.2 cm, deposited in NSMT. GenBank accession numbers: COI, KX400760-KX400768; mt 16S rDNA, KX400756-KX400759; ITS-rDNA, KX400769-KX400772. Figure 2A and B; Specimen number RMNH Coel. 42121 (MISE-TF-144): a lot of total 16 polyps collected on the same dive, Kin Bay, Uruma, Okinawa-jima Island, Japan (
<geoCoordinate direction="north" orientation="latitude" precision="15" value="26.373611">26°22'25&quot;N</geoCoordinate>
,
<geoCoordinate direction="east" orientation="longitude" precision="15" value="127.89166">127°53'30&quot;E</geoCoordinate>
), 15 m depth, collected by Takuma Fujii, 24 May 2012, 11 polyps fixed in 5-10% formalin, five polyps fixed in 99% EtOH, polyp length 1.1 to 2.2 cm (average 1.7
<normalizedToken originalValue="±">+/-</normalizedToken>
0.4 cm), maximum width 0.4 to 0.5 cm (average 0.5
<normalizedToken originalValue="±">+/-</normalizedToken>
0.1 cm), minimum width 0.1 to 0.3 cm (average 0.2
<normalizedToken originalValue="±">+/-</normalizedToken>
0.1), deposited in Naturalis Biodiversity Center, Leiden, Netherlands (RMNH); Specimen number NSMT-Co1578 (MISE-TF-151), a lot of total six polyps collected on the same dive, Oura Bay, Nago, Okinawa-jima Island, Japan (
<geoCoordinate direction="north" orientation="latitude" precision="15" value="26.541388">26°32'29&quot;N</geoCoordinate>
,
<geoCoordinate direction="east" orientation="longitude" precision="15" value="128.05444">128°3'16&quot;E</geoCoordinate>
), 17 m depth, collected by Takuma Fujii, 13 November 2012, five polyps fixed in 5-10% formalin, 1 polyp fixed in 99% EtOH, polyp length 1.0 to 1.9 cm (average 1.5
<normalizedToken originalValue="±">+/-</normalizedToken>
0.4 cm), maximum width 0.3 to 1.1 cm (average 0.7
<normalizedToken originalValue="±">+/-</normalizedToken>
0.3 cm), minimum width 0.1 to 0.3 cm (average 0.2
<normalizedToken originalValue="±">+/-</normalizedToken>
0.1 cm), deposited in NSMT. Figure 2C and D.
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="14" type="diagnosis">
<paragraph pageId="3" pageNumber="14">Diagnosis: external morphology.</paragraph>
<paragraph pageId="3" pageNumber="14">
Solitary, cylindrical polyp. Length of polyps 1.0 to 2.4 cm (average 1.7
<normalizedToken originalValue="±">+/-</normalizedToken>
0.3 cm), maximum width 0.3 to 1.1 cm (average 0.6
<normalizedToken originalValue="±">+/-</normalizedToken>
0.2 cm), minimum width 0.1 to 0.3 cm (average 0.2
<normalizedToken originalValue="±">+/-</normalizedToken>
0.1 cm) (n=34). Tentacles longer than half diameter of the expanded oral disc (Figure 2A). Oral disc gently hollowing into mouth, with stellate grooves as many as tentacles (Figure 2A, C). Capitular ridges present but not strongly pronounced when polyps closed (Figure 1). The upper part of the polyp between capitulum and the column slightly constricted (the width of the most constricted region approximately 0.1 cm to 0.4 cm thinner than the width of contracted capitulum) when polyp contracted (Figures 1, 2B, D). Upper part of the column generally thick and oval (Figures 1, 2B, D). Aboral narrow bottom portion of column extended (=foot), thinner than upper portion of column, like a cone (Figures 1, 2B, D), with the distal portion round and thicker than the extended foot (=physa) (Figures 1, 2B, D). Column smooth, with encrusted fine dense sand particles. Occasionally broken piece(s) of bivalve shells attached to the aboral end (Figure 2B).
</paragraph>
</subSubSection>
<subSubSection pageId="3" pageNumber="14" type="diagnosis">
<paragraph pageId="3" pageNumber="14">Diagnosis: internal morphology.</paragraph>
<paragraph pageId="3" pageNumber="14">
Fine sand particles heavily encrusted into ectoderm and mesoglea. Mesenteries in brachycnemic arrangement. Mesentery number 36, complete 18, incomplete 18 (Figure 3A; n=6 polyps). Single siphonoglyph apparent. Mesogleal sphincter muscle well developed, visible under dissecting microscope (Figure 3B). Endosymbiotic
<taxonomicName class="Cryptophyceae" family="Zooxanthellaceae" genus="Symbiodinium" higherTaxonomySource="GBIF" kingdom="Chromista" lsidName="Symbiodinium" order="Cryptomonadales" pageId="3" pageNumber="14" phylum="Cryptophyta" rank="genus">Symbiodinium</taxonomicName>
spp. (zooxanthellae) absent (=azooxanthellate).
</paragraph>
<caption pageId="3" pageNumber="14">
<paragraph pageId="3" pageNumber="14">
Figure 3. Morphological features of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="3" pageNumber="14" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. A Cross section of holotype NSMT-Co1576 through the actinopharynx showing the mesenterial arrangement and dense sand encrustations B Well-developed mesogleal sphincter muscles visible on a hand-cut longitudinal section of the holotype NSMT-Co1576 C Comparison of polyp shape between
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="3" pageNumber="14" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. NSMT-Co1577 and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="3" pageNumber="14" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
(from Brunei, refer to
<bibRefCitation author="Reimer, JD" journalOrPublisher="Contributions to Zoology" pageId="11" pageNumber="22" pagination="43 - 54" title="The phylogenetic position of the solitary zoanthid genus Sphenopus (Cnidaria: Hexacorallia)." volume="81" year="2012">Reimer et al. 2012</bibRefCitation>
).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="3" pageNumber="14" type="diagnosis">
<paragraph pageId="3" pageNumber="14">Diagnosis: cnidae.</paragraph>
<paragraph pageId="3" pageNumber="14">Basitrichs and spirocysts in tentacles and actinopharynx. Basitrichs, holotrichs, microbasic p-mastigophores and basitrichs in mesenterial filaments. Holotrichs in column (Table 1).</paragraph>
<caption pageId="3" pageNumber="14">
<paragraph pageId="3" pageNumber="14">
Table 1. Cnidae types and sizes in different tissue sections of the holotype of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="3" pageNumber="14" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n.
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="14">
<table pageId="3" pageNumber="14">
<tr pageId="3" pageNumber="14">
<th colspan="1" pageId="3" pageNumber="14" rowspan="1">
Image (Scale bars: 50
<normalizedToken originalValue="μm">μm</normalizedToken>
)
</th>
<th colspan="1" pageId="3" pageNumber="14" rowspan="1">Length*</th>
<th colspan="1" pageId="3" pageNumber="14" rowspan="1">Width*</th>
<th colspan="1" pageId="3" pageNumber="14" rowspan="1">Frequency**</th>
</tr>
<tr pageId="3" pageNumber="14">
<td colspan="1" pageId="3" pageNumber="14" rowspan="1">p</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="15" type="habitat">
<paragraph pageId="4" pageNumber="15">
<pageBreakToken pageId="4" pageNumber="15" start="start">Habitat</pageBreakToken>
.
</paragraph>
<paragraph pageId="4" pageNumber="15">Specimens were found at approximately 10 to 20 m depths on the slopes of silty seafloors in enclosed bays. Most polyps semi-burrowed in silt, with only the open oral disc visible and protruding out from the seafloor.</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="15" type="colour">
<paragraph pageId="4" pageNumber="15">Colour.</paragraph>
<paragraph pageId="4" pageNumber="15">Tentacles and oral disc whitish and translucent in life. Faint black narrow horizontal bands appear on tentacles, and similar faint patterns on the oral disc of a few polyps (Figure 2C). Column colour of encrusted sand particles, a few polyps with 2 to 6 faint black vertical stripes approximately 15 mm wide on the upper part of the column, reaching from oral end to aboral end (Figure 2C, D).</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="16" type="etymology">
<paragraph pageId="5" pageNumber="16">
<pageBreakToken pageId="5" pageNumber="16" start="start">Etymology</pageBreakToken>
.
</paragraph>
<paragraph pageId="5" pageNumber="16">
Named from latin '
<taxonomicName genus="Brachycnemina" lsidName="Brachycnemina exilis" pageId="5" pageNumber="16" rank="species" species="exilis">exilis</taxonomicName>
' meaning
<normalizedToken originalValue="‘slender’">'slender'</normalizedToken>
or
<normalizedToken originalValue="‘small’">'small'</normalizedToken>
, as polyps have an elongate and narrow foot more slender than other known species in this genus to the exception of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus pedunculatus" order="Zoantharia" pageId="5" pageNumber="16" phylum="Cnidaria" rank="species" species="pedunculatus">Sphenopus pedunculatus</taxonomicName>
. Polyps of this species are also much smaller than those of all three other species in the genus.
</paragraph>
</subSubSection>
<subSubSection pageId="5" pageNumber="16" type="common name">
<paragraph pageId="5" pageNumber="16">Common name.</paragraph>
<paragraph pageId="5" pageNumber="16">Hime-daruma-sunaginchaku (new Japanese name)</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="17" pageId="5" pageNumber="16" type="molecular phylogeny">
<paragraph pageId="5" pageNumber="16">Molecular phylogeny.</paragraph>
<paragraph pageId="5" pageNumber="16">
The results of the phylogenetic analyses of both mitochondrial cytochrome oxidase subunit I(COI) and 16S rDNA showed very few differences between sequences of our specimens and those of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="5" pageNumber="16" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
, as well as compared with those of various
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa" order="Zoantharia" pageId="5" pageNumber="16" phylum="Cnidaria" rank="genus">Palythoa</taxonomicName>
species. These results are not incongruous with previous studies on the molecular phylogeny of family
<taxonomicName family="Sphenopidae" lsidName="" pageId="5" pageNumber="16" rank="family">Sphenopidae</taxonomicName>
, where intra-family variation levels of mitochondrial DNA sequences were relatively low (
<bibRefCitation author="Reimer, JD" journalOrPublisher="Zoological Science" pageId="11" pageNumber="22" pagination="87 - 94" title="Molecular evidence suggesting species in the zoanthid genera Palythoa and Protopalythoa (Anthozoa: Hexacorallia) are congeneric." url="10.2108/zsj.23.87" volume="23" year="2006">Reimer et al. 2006</bibRefCitation>
,
<bibRefCitation author="Reimer, JD" journalOrPublisher="Contributions to Zoology" pageId="11" pageNumber="22" pagination="43 - 54" title="The phylogenetic position of the solitary zoanthid genus Sphenopus (Cnidaria: Hexacorallia)." volume="81" year="2012">2012</bibRefCitation>
).
</paragraph>
<paragraph pageId="6" pageNumber="17">
<pageBreakToken pageId="6" pageNumber="17" start="start">The</pageBreakToken>
results of the phylogenetic analyses of nuclear internal transcribed spacer rDNA region showed
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. forming a well-supported clade in the maximum likelihood and Bayesian analyses (Figure 4; ML=94%, Bayes=0.91). As well, together with sequences of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
,
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. formed a strongly supported
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
clade (Figure 4; ML=99%, Bayes=1.00). In comparing the ITS-rDNA sequences between
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
, there were 12 to 27 b.p. differences over a total 470 b.p. (=2.5~5.7% difference).
</paragraph>
<caption pageId="6" pageNumber="17">
<paragraph pageId="6" pageNumber="17">
Figure 4. Maximum likelihood tree of nuclear internal transcribed spacer of ribosomal DNA (ITS-rDNA) region for newly obtained sequences from
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. in this study along with previously published GenBank sequences of family
<taxonomicName family="Sphenopidae" lsidName="" pageId="6" pageNumber="17" rank="family">Sphenopidae</taxonomicName>
. Bootstrap values of ML&gt;60% are shown at respective nodes. Nodes supported by Bayesian posterior probabilities&gt;0.90 are marked with asterisks. Species
<normalizedToken originalValue="names’">names'</normalizedToken>
of sequences obtained from GenBank follow with accession numbers. The subtree shown in b) shows only the clade formed by genus
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
,
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa mizigama" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="mizigama">Palythoa mizigama</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa umbrosa" order="Zoantharia" pageId="6" pageNumber="17" phylum="Cnidaria" rank="species" species="umbrosa">Palythoa umbrosa</taxonomicName>
, delineated by the gray square in a).
</paragraph>
</caption>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="19" pageId="7" pageNumber="18" type="remarks">
<paragraph pageId="7" pageNumber="18">
<pageBreakToken pageId="7" pageNumber="18" start="start">Remarks</pageBreakToken>
.
</paragraph>
<paragraph pageId="7" pageNumber="18">
Until now three species have been considered valid within
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
;
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
(Gmelin, 1791),
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus arenaceus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="arenaceus">Sphenopus arenaceus</taxonomicName>
Hertwig, 1882, and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus pedunculatus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="pedunculatus">Sphenopus pedunculatus</taxonomicName>
Hertwig, 1888.
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. is easily distinguished from these other species by its small polyp size (length of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. &lt;2.5 cm and width &lt;1 cm), and by the shape of its elongated foot and physa. Polyps of both
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus arenaceus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="arenaceus">Sphenopus arenaceus</taxonomicName>
are round on the aboral end, and not elongated as in
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. (Figure 3C).
<bibRefCitation author="Soong, K" journalOrPublisher="Zoological Studies" pageId="12" pageNumber="23" pagination="333 - 343" title="Morphological and life history divergence of the zoanthid, Sphenopusmarsupialis off the Taiwanese coast." volume="38" year="1999">Soong et al. (1999)</bibRefCitation>
examined various sized
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
collected from around Taiwan including small polyps without any narrow elongated foot (length &lt;2 cm). Additionally,
<bibRefCitation author="Reimer, JD" journalOrPublisher="Fauna Ryukyuana" pageId="11" pageNumber="22" pagination="37 - 40" title="First record of the genus Sphenopus (Anthozoa: Hexacorallia: Zoantharia) from Japan." volume="29" year="2016">Reimer et al. (2016)</bibRefCitation>
recently reported on a
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
specimen of the typical rounded shape and large size (~9 cm in height) from Okinawa-jima Island. No polyps with intermediate morphology between
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. have ever been found. Thus, the specimens collected in this study cannot be considered to be immature polyps of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
. The morphologically most similar species to
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. is
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus pedunculatus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="pedunculatus">Sphenopus pedunculatus</taxonomicName>
as it also has a narrow foot, but
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus pedunculatus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="pedunculatus">Sphenopus pedunculatus</taxonomicName>
is much larger than
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n., with polyp lengths of 2.4 to 3.2 cm and widths of 2 to 2.4 cm, and with approximately 60 mesenteries. As well, the aboral end of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus pedunculatus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="pedunculatus">Sphenopus pedunculatus</taxonomicName>
is shaped like a clasping disc, different from that of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. with a narrow rounded shape (
<bibRefCitation author="Hertwig, R" journalOrPublisher="Reports on the Scientific Results of the Exploring Voyage of H. M. S. Challenger during the years 1873 - 1876, Zoology" pageId="10" pageNumber="21" pagination="1 - 56" title="Report on the Actiniaria dredged by H. M. S. Challenger during the years 1873 - 1876, Supplement." volume="26" year="1888">Hertwig 1888</bibRefCitation>
,
<bibRefCitation author="Reimer, JD" journalOrPublisher="ZooKeys" pageId="11" pageNumber="22" pagination="1 - 57" title="Shallow-water zoantharians (Cnidaria, Hexacorallia) from the Central Indo-Pacific." url="10.3897/zookeys.444.7537" volume="444" year="2014">Reimer et al. 2014</bibRefCitation>
).
</paragraph>
<paragraph lastPageId="8" lastPageNumber="19" pageId="7" pageNumber="18">
In contrast to the morphological differentiation from other
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
species, only a few differences were found in molecular analyses. The COI sequences of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. were identical to those of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
,
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa tuberculosa" order="Zoantharia" pageId="7" pageNumber="18" phylum="Cnidaria" rank="species" species="tuberculosa">Palythoa tuberculosa</taxonomicName>
(Esper, 1805), and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa umbrosa" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="umbrosa">
<pageBreakToken pageId="8" pageNumber="19" start="start">Palythoa</pageBreakToken>
umbrosa
</taxonomicName>
Irei, Singer &amp; Reimer, 2015. However, it is known that the evolutionary rate of mitochondrial DNA markers is quite slow in most
<taxonomicName class="Anthozoa" lsidName="Brachycnemina" pageId="8" pageNumber="19" rank="class">Anthozoa</taxonomicName>
(Shearer et al. 2004;
<bibRefCitation author="Huang, D" journalOrPublisher="Journal of Molecular Evolution" pageId="10" pageNumber="21" pagination="164 - 174" title="Slow mitochondrial COI sequence evolution at the base of the Metazoan tree and its implications for DNA barcording." url="10.1007/s00239-008-9069-5" volume="66" year="2008">Huang et al. 2008</bibRefCitation>
;
<bibRefCitation author="Stampar, SN" journalOrPublisher="PLoS ONE" pageId="12" pageNumber="23" title="Fast-evolving mitochondrial DNA in Ceriantharia: A refletion of Hexacorallia paraphyly?" url="10.1371/journal.pone.0086612" volume="9" year="2014">Stampar et al. 2014</bibRefCitation>
), and the nuclear ITS-rDNA region is currently the fastest evolving DNA marker that has been utilized for species-level analyses of suborder
<taxonomicName lsidName="Brachycnemina" pageId="8" pageNumber="19" rank="suborder" suborder="Brachycnemina">Brachycnemina</taxonomicName>
(
<bibRefCitation author="Reimer, JD" journalOrPublisher="Coral Reefs" pageId="11" pageNumber="22" pagination="399 - 410" title="Diversity and evolution in the zoanthid genus Palythoa (Cnidaria: Hexacorallia) utilizing nuclear ITS-rDNA." url="10.1007/s00338-007-0210-5" volume="26" year="2007">Reimer et al. 2007</bibRefCitation>
). Although there are only relatively few differences between the ITS-rDNA sequences of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
(2.5~5.7% sequence divergence), the formation of a supported monophyletic clade confirms the results of our morphological analyses that the specimens collected in this study belong to a species different from
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
(Figure 4). Moreover, these results suggest the possibility of the presence of multiple, cryptic species within
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
as previously mentioned by
<bibRefCitation author="Soong, K" journalOrPublisher="Zoological Studies" pageId="12" pageNumber="23" pagination="333 - 343" title="Morphological and life history divergence of the zoanthid, Sphenopusmarsupialis off the Taiwanese coast." volume="38" year="1999">Soong et al. (1999)</bibRefCitation>
.
</paragraph>
<paragraph pageId="8" pageNumber="19">
Currently, very little is known about the ecology and species diversity of the genus
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
, demonstrated by the fact that there have been no or few records of both
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus arenaceus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="arenaceus">Sphenopus arenaceus</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus pedunculatus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="pedunculatus">Sphenopus pedunculatus</taxonomicName>
within the last 100 years. Thus, morphological and molecular analyses of newly obtained specimens from type localities followed by reviewing each
<normalizedToken originalValue="species’">species'</normalizedToken>
description carefully are required to clearly understand the species distinction of
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
species. As mentioned in previous studies, the phylogenetic results of this study indicate a need to re-examine the validity of the genus
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
as it is positioned within the genus
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Palythoa</taxonomicName>
, and by extension the definition of genera within the family
<taxonomicName family="Sphenopidae" lsidName="" pageId="8" pageNumber="19" rank="family">Sphenopidae</taxonomicName>
should be reconsidered (
<bibRefCitation author="Reimer, JD" journalOrPublisher="Contributions to Zoology" pageId="11" pageNumber="22" pagination="43 - 54" title="The phylogenetic position of the solitary zoanthid genus Sphenopus (Cnidaria: Hexacorallia)." volume="81" year="2012">Reimer et al. 2012</bibRefCitation>
,
<bibRefCitation author="Irei, Y" journalOrPublisher="Zookeys" pageId="11" pageNumber="22" pagination="1 - 26" title="Descriptions of two azooxanthellate Palythoa species (Subclass Hexacorallia, Order Zoantharia) from the Ryukyu Archipelago, southern Japan." url="10.3897/zookeys.478.8512" volume="478" year="2015">Irei et al. 2015</bibRefCitation>
).
</paragraph>
<paragraph pageId="8" pageNumber="19">
In the ITS-rDNA molecular phylogeny, it is notable that two recently described azooxanthellate
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Palythoa</taxonomicName>
species from caves,
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa umbrosa" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="umbrosa">Palythoa umbrosa</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa mizigama" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="mizigama">Palythoa mizigama</taxonomicName>
, form a well-supported subclade with
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus exilis" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="exilis">Sphenopus exilis</taxonomicName>
sp. n. and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus marsupialis" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="species" species="marsupialis">Sphenopus marsupialis</taxonomicName>
. As the phylogenetic relationship between
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Palythoa</taxonomicName>
is not yet clear, and likely does not reflect the traditional taxonomy (
<bibRefCitation author="Reimer, JD" journalOrPublisher="Contributions to Zoology" pageId="11" pageNumber="22" pagination="43 - 54" title="The phylogenetic position of the solitary zoanthid genus Sphenopus (Cnidaria: Hexacorallia)." volume="81" year="2012">Reimer et al. 2012</bibRefCitation>
), construction of a large ITS-rDNA phylogeny with additional sequences from other
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Palythoa" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Palythoa" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Palythoa</taxonomicName>
and
<taxonomicName class="Anthozoa" family="Sphenopidae" genus="Sphenopus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphenopus" order="Zoantharia" pageId="8" pageNumber="19" phylum="Cnidaria" rank="genus">Sphenopus</taxonomicName>
species is needed. At the same time, investigation with additional DNA markers asides from the mt DNA and ITS-rDNA currently utilized in zoantharian phylogeny may be helpful.
</paragraph>
</subSubSection>
</treatment>
</document>
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