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<document id="8166FD6775AA341F52666EA2FD99D3E1" ID-CLB-Dataset="298900" ID-DOI="10.1643/ci-18-055" ID-GBIF-Dataset="4919bfd6-3d35-48b9-9421-c852fa35abab" ID-Zenodo-Dep="12536069" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="julia" IM.metadata_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1719340789171" checkinUser="julia" docAuthor="Ma, Ka Yan &amp; Craig, Matthew T." docDate="2018" docId="03FEC024E12EDF39FCB83C1B6CE1FF13" docLanguage="en" docName="copeia.106.3.443-456.pdf" docOrigin="Copeia 106 (3)" docSource="http://dx.doi.org/10.1643/ci-18-055" docTitle="Plectropomus Oken 1817" docType="treatment" docVersion="5" lastPageNumber="452" masterDocId="FFC7B85CE126DF33FFE33B236D19FFA4" masterDocTitle="An Inconvenient Monophyly: An Update on the Taxonomy of the Groupers (Epinephelidae)" masterLastPageNumber="456" masterPageNumber="443" pageNumber="451" updateTime="1719854615312" updateUser="julia">
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<mods:title id="A86A68C5C6F277400AA6EDF3F761C535">An Inconvenient Monophyly: An Update on the Taxonomy of the Groupers (Epinephelidae)</mods:title>
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<mods:name id="9C7F8DA62FC663A576658C243F35AB92" type="personal">
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<mods:namePart id="5E9CB46E39C380CB18F20631AB71C641">Ma, Ka Yan</mods:namePart>
<mods:affiliation id="F4337370DC3A66B7872C2FC206CB5641">Simon F. S. Li Marine Science Laboratory, School of Life Sciences, The Chinese University of Hong Kong, Sha Tin, 999077, Hong Kong SAR.</mods:affiliation>
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<mods:namePart id="421DCD3D8CB067D4E7466669071085F9">Craig, Matthew T.</mods:namePart>
<mods:affiliation id="071321261CAF11C5B7B63ABA93F6A8A2">2 National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Southwest Fisheries Science Center, 8901 La Jolla Shores Drive, La Jolla, California 92037</mods:affiliation>
<mods:nameIdentifier id="36B1903CBE8BEA873F2452389C338178" type="email">matthew.craig@noaa.gov.</mods:nameIdentifier>
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<treatment id="03FEC024E12EDF39FCB83C1B6CE1FF13" ID-DOI="http://doi.org/10.5281/zenodo.12536324" ID-GBIF-Taxon="233767945" ID-Zenodo-Dep="12536324" LSID="urn:lsid:plazi:treatment:03FEC024E12EDF39FCB83C1B6CE1FF13" httpUri="http://treatment.plazi.org/id/03FEC024E12EDF39FCB83C1B6CE1FF13" lastPageId="10" lastPageNumber="452" pageId="8" pageNumber="451">
<subSubSection id="C34D22B9E12EDF3BFCB83C1B6986F8EB" box="[859,1183,1848,1871]" pageId="8" pageNumber="451" type="nomenclature">
<paragraph id="8BE87132E12EDF3BFCB83C1B6986F8EB" blockId="8.[859,1530,1848,2024]" box="[859,1183,1848,1871]" pageId="8" pageNumber="451">
<taxonomicName id="4C570AB1E12EDF3BFCB83C1B6EECF8EB" ID-CoL="6SD4" authority="Oken, 1817" authorityName="Oken" authorityYear="1817" box="[859,1013,1848,1871]" family="Epinephelidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="456" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12EDF3BFCB83C1B6EECF8EB" box="[859,1013,1848,1871]" pageId="8" pageNumber="451">Plectropomus</emphasis>
</taxonomicName>
<emphasis id="B923AD20E12EDF3BFBE73C1B6986F8EB" box="[1028,1183,1848,1871]" pageId="8" pageNumber="451">and Its Allies.</emphasis>
</paragraph>
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<subSubSection id="C34D22B9E12EDF39FB7C3C1B6CE1FF13" lastPageId="10" lastPageNumber="453" pageId="8" pageNumber="451" type="discussion">
<paragraph id="8BE87132E12EDF3AFB7C3C1B6CAAF829" blockId="8.[859,1530,1848,2024]" lastBlockId="9.[103,773,1522,2024]" lastPageId="9" lastPageNumber="452" pageId="8" pageNumber="451">
—The genus
<taxonomicName id="4C570AB1E12EDF3BFAD83C1B68DBF8EA" authorityName="Oken" authorityYear="1817" box="[1339,1474,1848,1870]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="451" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12EDF3BFAD83C1B68DBF8EA" box="[1339,1474,1848,1870]" italics="true" pageId="8" pageNumber="451">Plectropomus</emphasis>
</taxonomicName>
was recovered as monophyletic; however, both the monotypic
<taxonomicName id="4C570AB1E12EDF3BFCB83C566EFEF82F" box="[859,999,1909,1931]" family="Serranidae" genus="Gonioplectrus" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="451" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12EDF3BFCB83C566EFEF82F" box="[859,999,1909,1931]" italics="true" pageId="8" pageNumber="451">Gonioplectrus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E12EDF3BFBC23C56696AF82F" authorityName="Smith" authorityYear="1964" box="[1057,1139,1909,1931]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="8" pageNumber="451" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12EDF3BFBC23C56696AF82F" box="[1057,1139,1909,1931]" italics="true" pageId="8" pageNumber="451">Saloptia</emphasis>
</taxonomicName>
are included in a larger monophyletic clade. All three genera share a relatively low dorsal-fin spine count (78 vs. 911 in the remaining grouper genera). Smith (1964), Leis (1986), and Heemstra and Randall (1993) hypothesized a close relationship between
<taxonomicName id="4C570AB1E12FDF3AFD953ED16FD3F9AC" authorityName="Smith" authorityYear="1964" box="[630,714,1522,1544]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFD953ED16FD3F9AC" box="[630,714,1522,1544]" italics="true" pageId="9" pageNumber="452">Saloptia</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E12FDF3AFF843D336DF7F982" authorityName="Oken" authorityYear="1817" box="[103,238,1552,1574]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFF843D336DF7F982" box="[103,238,1552,1574]" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
. Species in these genera are the only groupers with a single supraneural bone, low dorsal-fin spine count, three large antrorse spines on the lower edge of the preopercle, and only 13 branched caudal-fin rays.
<taxonomicName id="4C570AB1E12FDF3AFD513D496E1DF924" authorityName="Smith" authorityYear="1964" box="[690,772,1642,1664]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFD513D496E1DF924" box="[690,772,1642,1664]" italics="true" pageId="9" pageNumber="452">Saloptia</emphasis>
</taxonomicName>
has been treated as a monotypic genus on the basis of the relative strength of the median fin spines (strong in
<taxonomicName id="4C570AB1E12FDF3AFF843DE76DA0F97E" authorityName="Smith" authorityYear="1964" box="[103,185,1732,1754]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFF843DE76DA0F97E" box="[103,185,1732,1754]" italics="true" pageId="9" pageNumber="452">Saloptia</emphasis>
</taxonomicName>
and weak in
<taxonomicName id="4C570AB1E12FDF3AFE8C3DE76CEFF97E" authorityName="Oken" authorityYear="1817" box="[367,502,1732,1754]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFE8C3DE76CEFF97E" box="[367,502,1732,1754]" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
; Heemstra and Randall, 1993) and an absence of enlarged canine teeth on the lower jaw. Given the numerous synapomorphies shared by these genera, the argument for retaining
<taxonomicName id="4C570AB1E12FDF3AFE0C3C3D6F48F897" authorityName="Smith" authorityYear="1964" box="[495,593,1821,1843]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="species" species="powelli">
<emphasis id="B923AD20E12FDF3AFE0C3C3D6F48F897" box="[495,593,1821,1843]" italics="true" pageId="9" pageNumber="452">S. powelli</emphasis>
</taxonomicName>
in a monotypic genus is relatively weak. The loss of canine teeth appears to be an autapomorphic character and is insufficient to exclude the species from
<taxonomicName id="4C570AB1E12FDF3AFEC63C546CB5F829" authorityName="Oken" authorityYear="1817" box="[293,428,1911,1933]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFEC63C546CB5F829" box="[293,428,1911,1933]" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="DF2821BAE12FDF3AFB903E40684CFA08" ID-DOI="http://doi.org/10.5281/zenodo.12536102" ID-Zenodo-Dep="12536102" httpUri="https://zenodo.org/record/12536102/files/figure.png" pageId="9" pageNumber="452" startId="9.[1139,1167,1379,1399]" targetBox="[103,1090,131,1446]" targetPageId="9" targetType="figure">
<paragraph id="8BE87132E12FDF3AFB903E40684CFA08" blockId="9.[1139,1490,1378,1452]" pageId="9" pageNumber="452">
<emphasis id="B923AD20E12FDF3AFB903E4069ABFA08" bold="true" pageId="9" pageNumber="452">
FIG. 7.
<taxonomicName id="4C570AB1E12FDF3AFB5C3E416898FAD2" baseAuthorityName="Valenciennes" baseAuthorityYear="1828" box="[1215,1409,1378,1398]" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="species" species="altivelis">
<emphasis id="B923AD20E12FDF3AFB5C3E416898FAD2" bold="true" box="[1215,1409,1378,1398]" italics="true" pageId="9" pageNumber="452">Epinephelus altivelis</emphasis>
</taxonomicName>
juvenile (top) and adult (bottom). Photographs
</emphasis>
©
<taxonomicName id="4C570AB1E12FDF3AFB323EBB684CFA08" authority="Myers." authorityName="Myers." box="[1233,1365,1432,1452]" class="Insecta" family="Coccinellidae" genus="Robert" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="452" phylum="Arthropoda" rank="genus">Robert Myers.</taxonomicName>
</paragraph>
</caption>
<paragraph id="8BE87132E12FDF3AFF9C3CB56912F8A3" blockId="9.[103,773,1522,2024]" lastBlockId="9.[820,1491,1522,2024]" pageId="9" pageNumber="452">
The monotypic genus
<taxonomicName id="4C570AB1E12FDF3AFE9D3CB56F13F808" box="[382,522,1942,1964]" family="Serranidae" genus="Gonioplectrus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFE9D3CB56F13F808" box="[382,522,1942,1964]" italics="true" pageId="9" pageNumber="452">Gonioplectrus</emphasis>
</taxonomicName>
is also present in this clade (clade A,
<figureCitation id="136C6DB7E12FDF3AFEEC3C976C4CF86D" box="[271,341,1972,1994]" captionStart="FIG" captionStartId="3.[103,131,1851,1871]" captionTargetBox="[217,1376,130,1821]" captionTargetId="figure-1@3.[217,1376,130,1821]" captionTargetPageId="3" captionText="FIG. 1. Majority rule consensus cladogram of maximum likelihood analysis, reconstructed using RAxML with 1000 bootstrap replicates following Ma et al. (2016). Numbers at nodes represent bootstrap support values (%)." figureDoi="http://doi.org/10.5281/zenodo.12536076" httpUri="https://zenodo.org/record/12536076/files/figure.png" pageId="9" pageNumber="452">Figs. 1</figureCitation>
,
<figureCitation id="136C6DB7E12FDF3AFE873C906C6DF86E" box="[356,372,1971,1994]" captionStart="FIG" captionStartId="4.[142,170,1851,1871]" captionTargetBox="[213,1458,130,1821]" captionTargetId="figure-0@4.[213,1458,130,1821]" captionTargetPageId="4" captionText="FIG. 2. Bayesian inference phylogram reconstructed using MrBayes following Ma et al. (2016). Numbers on branches represent posterior probabilities." figureDoi="http://doi.org/10.5281/zenodo.12536081" httpUri="https://zenodo.org/record/12536081/files/figure.png" pageId="9" pageNumber="452">2</figureCitation>
). This is not entirely surprising as it shares the low number of dorsal-fin spines (8) with species of
<emphasis id="B923AD20E12FDF3AFCD73ED16947F9AC" box="[820,1118,1522,1544]" italics="true" pageId="9" pageNumber="452">
<taxonomicName id="4C570AB1E12FDF3AFCD73ED16EA6F9AC" authorityName="Oken" authorityYear="1817" box="[820,959,1522,1544]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">Plectropomus</taxonomicName>
.
<taxonomicName id="4C570AB1E12FDF3AFC313ED16947F9AC" box="[978,1118,1522,1544]" family="Serranidae" genus="Gonioplectrus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">Gonioplectrus</taxonomicName>
</emphasis>
has two well-developed supraneurals (versus only one in
<taxonomicName id="4C570AB1E12FDF3AFB9C3D31681FF98C" authorityName="Oken" authorityYear="1817" box="[1151,1286,1554,1576]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFB9C3D31681FF98C" box="[1151,1286,1554,1576]" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E12FDF3AFAAA3D316882F98C" authorityName="Smith" authorityYear="1964" box="[1353,1435,1554,1576]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFAAA3D316882F98C" box="[1353,1435,1554,1576]" italics="true" pageId="9" pageNumber="452">Saloptia</emphasis>
</taxonomicName>
), 15 branched caudal-fin rays (versus 13 in
<taxonomicName id="4C570AB1E12FDF3AFAF23D116881F9EC" authorityName="Oken" authorityYear="1817" box="[1297,1432,1586,1608]" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFAF23D116881F9EC" box="[1297,1432,1586,1608]" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E12FDF3AFCD73D716E9FF9CC" authorityName="Smith" authorityYear="1964" box="[820,902,1618,1640]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFCD73D716E9FF9CC" box="[820,902,1618,1640]" italics="true" pageId="9" pageNumber="452">Saloptia</emphasis>
</taxonomicName>
), a prominent knob on the lower corner of the maxilla (absent in
<taxonomicName id="4C570AB1E12FDF3AFBE93D51682AF92C" authority=", Plectropomus" authorityName="Plectropomus" box="[1034,1331,1650,1672]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFBE93D516982F92C" box="[1034,1179,1650,1672]" italics="true" pageId="9" pageNumber="452">Cephalopholis</emphasis>
,
<emphasis id="B923AD20E12FDF3AFB4F3D51682AF92C" box="[1196,1331,1650,1672]" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C570AB1E12FDF3AFA943D5168D0F92C" authorityName="Smith" authorityYear="1964" box="[1399,1481,1650,1672]" family="Serranidae" genus="Saloptia" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFA943D5168D0F92C" box="[1399,1481,1650,1672]" italics="true" pageId="9" pageNumber="452">Saloptia</emphasis>
</taxonomicName>
) and unique cranial morphology (Heemstra and Randall, 1993). Given these differences,
<taxonomicName id="4C570AB1E12FDF3AFB763D926838F963" box="[1173,1313,1713,1735]" family="Serranidae" genus="Gonioplectrus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFB763D926838F963" box="[1173,1313,1713,1735]" italics="true" pageId="9" pageNumber="452">Gonioplectrus</emphasis>
</taxonomicName>
appears to be a related, but distinct genus and is retained for the single species
<taxonomicName id="4C570AB1E12FDF3AFC6B3DD1691CF8A3" baseAuthorityName="Cuvier" baseAuthorityYear="1828" box="[904,1029,1777,1799]" family="Serranidae" genus="Gonioplectrus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="species" species="hispanus">
<emphasis id="B923AD20E12FDF3AFC6B3DD1691CF8A3" box="[904,1029,1777,1799]" italics="true" pageId="9" pageNumber="452">G. hispanus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BE87132E12FDF39FCAF3C316CE1FF13" blockId="9.[820,1491,1522,2024]" lastBlockId="10.[142,812,131,183]" lastPageId="10" lastPageNumber="453" pageId="9" pageNumber="452">
Also included in the monophyletic clade A (
<figureCitation id="136C6DB7E12FDF3AFAB93C316882F88D" box="[1370,1435,1810,1833]" captionStart="FIG" captionStartId="3.[103,131,1851,1871]" captionTargetBox="[217,1376,130,1821]" captionTargetId="figure-1@3.[217,1376,130,1821]" captionTargetPageId="3" captionText="FIG. 1. Majority rule consensus cladogram of maximum likelihood analysis, reconstructed using RAxML with 1000 bootstrap replicates following Ma et al. (2016). Numbers at nodes represent bootstrap support values (%)." figureDoi="http://doi.org/10.5281/zenodo.12536076" httpUri="https://zenodo.org/record/12536076/files/figure.png" pageId="9" pageNumber="452">Fig. 1</figureCitation>
) are members of the genus
<taxonomicName id="4C570AB1E12FDF3AFBD83C11699CF8EC" authorityName="Swainson" authorityYear="1839" box="[1083,1157,1842,1864]" family="Serranidae" genus="Variola" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFBD83C11699CF8EC" box="[1083,1157,1842,1864]" italics="true" pageId="9" pageNumber="452">Variola</emphasis>
</taxonomicName>
. With nine dorsal-fin spines, this genus would seem to be closely related to
<taxonomicName id="4C570AB1E12FDF3AFAD93C7168D2F8CC" box="[1338,1483,1874,1896]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFAD93C7168D2F8CC" box="[1338,1483,1874,1896]" italics="true" pageId="9" pageNumber="452">Cephalopholis</emphasis>
</taxonomicName>
; however, it is clear from the analyses that this is not the case. Species of
<taxonomicName id="4C570AB1E12FDF3AFC583CB1691FF80C" authorityName="Swainson" authorityYear="1839" box="[955,1030,1938,1960]" family="Serranidae" genus="Variola" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFC583CB1691FF80C" box="[955,1030,1938,1960]" italics="true" pageId="9" pageNumber="452">Variola</emphasis>
</taxonomicName>
are the only grouper species with a pronounced, lunate tail. While closely allied with
<taxonomicName id="4C570AB1E12FDF3AFA8D3C926E46F843" authorityName="Oken" authorityYear="1817" family="Serranidae" genus="Plectropomus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFA8D3C926E46F843" italics="true" pageId="9" pageNumber="452">Plectropomus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E12FDF3AFC473CF26929F843" box="[932,1072,2001,2023]" family="Serranidae" genus="Gonioplectrus" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFC473CF26929F843" box="[932,1072,2001,2023]" italics="true" pageId="9" pageNumber="452">Gonioplectrus</emphasis>
</taxonomicName>
, the genus
<taxonomicName id="4C570AB1E12FDF3AFB203CF26814F843" authorityName="Swainson" authorityYear="1839" box="[1219,1293,2001,2023]" family="Serranidae" genus="Variola" kingdom="Animalia" order="Perciformes" pageId="9" pageNumber="452" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E12FDF3AFB203CF26814F843" box="[1219,1293,2001,2023]" italics="true" pageId="9" pageNumber="452">Variola</emphasis>
</taxonomicName>
appears to be a distinct group among the groupers with low dorsal-fin spine counts and is therefore retained.
</paragraph>
</subSubSection>
</treatment>
</document>