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<document id="CD83BD627D679BCB38D2727325EE26C7" 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="03FEC024E124DF36FF6D3D116836FA2F" docLanguage="en" docName="copeia.106.3.443-456.pdf" docOrigin="Copeia 106 (3)" docSource="http://dx.doi.org/10.1643/ci-18-055" docTitle="Cephalopholis Bloch &amp; Schneider 1801" docType="treatment" docVersion="5" lastPageNumber="448" masterDocId="FFC7B85CE126DF33FFE33B236D19FFA4" masterDocTitle="An Inconvenient Monophyly: An Update on the Taxonomy of the Groupers (Epinephelidae)" masterLastPageNumber="456" masterPageNumber="443" pageNumber="445" updateTime="1719854615312" updateUser="julia">
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<mods:title id="649B65067A8E8EA6D28DB1C5AA061B1B">An Inconvenient Monophyly: An Update on the Taxonomy of the Groupers (Epinephelidae)</mods:title>
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<mods:namePart id="7E3A1F95E36D4A61B65A7A02C57084F8">Ma, Ka Yan</mods:namePart>
<mods:affiliation id="9184868E73DC9BA0AF919B61C1DF1BB2">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:affiliation id="3F21C61E8FD56B30617BAC9753909B6A">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>
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<treatment id="03FEC024E124DF36FF6D3D116836FA2F" ID-DOI="http://doi.org/10.5281/zenodo.12536312" ID-GBIF-Taxon="233767946" ID-Zenodo-Dep="12536312" LSID="urn:lsid:plazi:treatment:03FEC024E124DF36FF6D3D116836FA2F" httpUri="http://treatment.plazi.org/id/03FEC024E124DF36FF6D3D116836FA2F" lastPageId="5" lastPageNumber="448" pageId="2" pageNumber="445">
<emphasis id="B923AD20E124DF31FF6D3D116C58F9ED" bold="true" box="[142,321,1586,1609]" italics="true" pageId="2" pageNumber="445">
<subSubSection id="C34D22B9E124DF31FF6D3D116C33F9ED" box="[142,298,1586,1609]" pageId="2" pageNumber="445" type="nomenclature">
<paragraph id="8BE87132E124DF31FF6D3D116C33F9ED" blockId="2.[142,812,1586,2024]" box="[142,298,1586,1609]" pageId="2" pageNumber="445">
<taxonomicName id="4C570AB1E124DF31FF6D3D116C3FF9ED" ID-CoL="3KG8" authority="Bloch &amp; Schneider, 1801" authorityName="Bloch &amp; Schneider" authorityYear="1801" box="[142,294,1586,1609]" family="Epinephelidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">Cephalopholis</taxonomicName>
.
</paragraph>
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<subSubSection id="C34D22B9E124DF36FEC93D116836FA2F" lastPageId="5" lastPageNumber="448" pageId="2" pageNumber="445" type="discussion">
<paragraph id="8BE87132E124DF31FEC93D116CC4F8D5" blockId="2.[142,812,1586,2024]" pageId="2" pageNumber="445">
—As with all previous molecular phylogenies where they have been sampled,
<emphasis id="B923AD20E124DF31FE1C3D736E3CF9C2" box="[511,805,1616,1638]" italics="true" pageId="2" pageNumber="445">
<taxonomicName id="4C570AB1E124DF31FE1C3D736F9FF9C2" authorityName=", Randall" authorityYear="1964" box="[511,646,1616,1638]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">Aethaloperca</taxonomicName>
,
<taxonomicName id="4C570AB1E124DF31FD773D736E3CF9C2" authorityName=", Randall" authorityYear="1964" box="[660,805,1616,1638]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">Cephalopholis</taxonomicName>
</emphasis>
,
<taxonomicName id="4C570AB1E124DF31FF6D3D4E6DC0F927" authorityName="Randall" authorityYear="1964" box="[142,217,1645,1667]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FF6D3D4E6DC0F927" box="[142,217,1645,1667]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C570AB1E124DF31FEF93D4E6C90F927" box="[282,393,1645,1667]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FEF93D4E6C90F927" box="[282,393,1645,1667]" italics="true" pageId="2" pageNumber="445">Paranthias</emphasis>
</taxonomicName>
form a clade and morphological data suggest that they are close allies (e.g., Randall, 1964; Smith-Vaniz et al., 1988). The basis for the exclusion of
<taxonomicName id="4C570AB1E124DF31FD423D8B6E3CF91A" authorityName=", Randall" authorityYear="1964" box="[673,805,1704,1726]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FD423D8B6E3CF91A" box="[673,805,1704,1726]" italics="true" pageId="2" pageNumber="445">Aethaloperca</emphasis>
</taxonomicName>
,
<taxonomicName id="4C570AB1E124DF31FF6D3DE56DC0F978" authorityName="Randall" authorityYear="1964" box="[142,217,1734,1756]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FF6D3DE56DC0F978" box="[142,217,1734,1756]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C570AB1E124DF31FEFB3DE56C9EF978" box="[280,391,1734,1756]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FEFB3DE56C9EF978" box="[280,391,1734,1756]" italics="true" pageId="2" pageNumber="445">Paranthias</emphasis>
</taxonomicName>
from
<taxonomicName id="4C570AB1E124DF31FE283DE56F45F978" authorityName=", Randall" authorityYear="1964" box="[459,604,1734,1756]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FE283DE56F45F978" box="[459,604,1734,1756]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
in past treatments (e.g., Heemstra and Randall, 1993) was solely based on apomorphic characters. However, given that they are deeply nested within
<taxonomicName id="4C570AB1E124DF31FED63C3C6CDFF891" authorityName=", Randall" authorityYear="1964" box="[309,454,1823,1845]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FED63C3C6CDFF891" box="[309,454,1823,1845]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
, these characters become autapomorphic, and their usefulness to exclude them from
<taxonomicName id="4C570AB1E124DF31FF6D3C796C06F8D4" authorityName=", Randall" authorityYear="1964" box="[142,287,1882,1904]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FF6D3C796C06F8D4" box="[142,287,1882,1904]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
is rendered null.
</paragraph>
<paragraph id="8BE87132E124DF31FF453C5A69ADFD4F" blockId="2.[142,812,1586,2024]" lastBlockId="2.[859,1530,131,2024]" pageId="2" pageNumber="445">
Smith (1954) elevated the sub-genus
<taxonomicName id="4C570AB1E124DF31FDB33C5B6FCDF82A" authorityName=", Randall" authorityYear="1964" box="[592,724,1912,1934]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FDB33C5B6FCDF82A" box="[592,724,1912,1934]" italics="true" pageId="2" pageNumber="445">Aethaloperca</emphasis>
</taxonomicName>
for the species
<taxonomicName id="4C570AB1E124DF31FF013CB56F02F808" baseAuthorityName="Fowler &amp; Bean" baseAuthorityYear="1930" box="[226,539,1942,1964]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="albomarginatus">
<emphasis id="B923AD20E124DF31FF013CB56F02F808" box="[226,539,1942,1964]" italics="true" pageId="2" pageNumber="445">Cephalopholis albomarginatus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E124DF31FDB73CB46FD3F808" authorityName="Forsskal" authorityYear="1775" box="[596,714,1943,1964]" family="Percidae" genus="Perca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="rogaa">
<emphasis id="B923AD20E124DF31FDB73CB46FD3F808" box="[596,714,1943,1964]" italics="true" pageId="2" pageNumber="445">Perca rogaa</emphasis>
</taxonomicName>
. Randall (1964) removed
<taxonomicName id="4C570AB1E124DF31FEB73C976F02F86E" baseAuthorityName="Fowler &amp; Bean" baseAuthorityYear="1930" box="[340,539,1972,1994]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="albomarginatus">
<emphasis id="B923AD20E124DF31FEB73C976F02F86E" box="[340,539,1972,1994]" italics="true" pageId="2" pageNumber="445">C. albomarginatus</emphasis>
</taxonomicName>
from
<taxonomicName id="4C570AB1E124DF31FD8C3C976FEAF86E" authorityName=", Randall" authorityYear="1964" box="[623,755,1972,1994]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FD8C3C976FEAF86E" box="[623,755,1972,1994]" italics="true" pageId="2" pageNumber="445">Aethaloperca</emphasis>
</taxonomicName>
and placed it into the new genus
<taxonomicName id="4C570AB1E124DF31FDE13CF26F54F843" authorityName="Randall" authorityYear="1964" box="[514,589,2001,2023]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FDE13CF26F54F843" box="[514,589,2001,2023]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
(discussed below).
<taxonomicName id="4C570AB1E124DF31FCB83BA06906FF3D" baseAuthorityName="Forsskal" baseAuthorityYear="1775" box="[859,1055,131,153]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="rogaa">
<emphasis id="B923AD20E124DF31FCB83BA06906FF3D" box="[859,1055,131,153]" italics="true" pageId="2" pageNumber="445">Aethaloperca rogaa</emphasis>
</taxonomicName>
was considered to be appropriately placed in a monotypic genus based on the shape of its cranial crests, its deeper body, the steeper profile of its head, the shape of its dorsal fin, and the presence of asymmetrical pectoral fins (Randall, 1964; Smith-Vaniz et al., 1988; Heemstra and Randall, 1993). While these characters may seem to indicate a generic difference, they are not all unique and some are shared with other species of
<taxonomicName id="4C570AB1E124DF31FB7E3A716837FECC" authorityName=", Randall" authorityYear="1964" box="[1181,1326,338,360]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FB7E3A716837FECC" box="[1181,1326,338,360]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
. For example, the presence of a steep head profile is present in
<taxonomicName id="4C570AB1E124DF31FAB63A5268E0FE22" authorityName="Katayama" authorityYear="1957" box="[1365,1529,368,390]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="igarashiensis">
<emphasis id="B923AD20E124DF31FAB63A5268E0FE22" box="[1365,1529,368,390]" italics="true" pageId="2" pageNumber="445">C. igarashiensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E124DF31FC723AAD690BFE00" baseAuthorityName="Valenciennes" baseAuthorityYear="1828" box="[913,1042,398,420]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="sonnerati">
<emphasis id="B923AD20E124DF31FC723AAD690BFE00" box="[913,1042,398,420]" italics="true" pageId="2" pageNumber="445">C. sonnerati</emphasis>
</taxonomicName>
(
<figureCitation id="136C6DB7E124DF31FBC63AAD6971FE01" box="[1061,1128,398,421]" captionStart="FIG" captionStartId="5.[1139,1170,1248,1268]" captionTargetBox="[103,1089,131,1316]" captionTargetId="figure-381@5.[103,1089,131,1316]" captionTargetPageId="5" captionText="FIG. 3. Cephalopholis sonnerati (top) and C. rogaa (bottom). Photographs © Robert Myers." figureDoi="http://doi.org/10.5281/zenodo.12536083" httpUri="https://zenodo.org/record/12536083/files/figure.png" pageId="2" pageNumber="445">Fig. 3</figureCitation>
). Its body depth (2.12.4 times in standard length [SL]; Heemstra and Randall, 1993) is well within the range of other species of
<taxonomicName id="4C570AB1E124DF31FAE63AEA688FFE7B" authorityName=", Randall" authorityYear="1964" box="[1285,1430,457,479]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FAE63AEA688FFE7B" box="[1285,1430,457,479]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
(2.03.2 times in SL; Heemstra and Randall, 1993; Craig et al., 2011). The remaining external characters (shape of dorsal and pectoral fins) are the only obvious morphological differences between this monotypic genus and
<taxonomicName id="4C570AB1E124DF31FB0E39636867FDF2" authorityName=", Randall" authorityYear="1964" box="[1261,1406,576,598]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FB0E39636867FDF2" box="[1261,1406,576,598]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
. As will be discussed below, there are several synapomorphic characters that unite
<taxonomicName id="4C570AB1E124DF31FC3639586940FD35" authorityName=", Randall" authorityYear="1964" box="[981,1113,635,657]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FC3639586940FD35" box="[981,1113,635,657]" italics="true" pageId="2" pageNumber="445">Aethaloperca</emphasis>
</taxonomicName>
with
<taxonomicName id="4C570AB1E124DF31FB7D39586836FD35" authorityName=", Randall" authorityYear="1964" box="[1182,1327,635,657]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FB7D39586836FD35" box="[1182,1327,635,657]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
and there is thus little reason to exclude
<taxonomicName id="4C570AB1E124DF31FBBE39BA69F8FD0B" authorityName=", Randall" authorityYear="1964" box="[1117,1249,665,687]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FBBE39BA69F8FD0B" box="[1117,1249,665,687]" italics="true" pageId="2" pageNumber="445">Aethaloperca</emphasis>
</taxonomicName>
from
<taxonomicName id="4C570AB1E124DF31FAC739BA68ACFD0B" authorityName=", Randall" authorityYear="1964" box="[1316,1461,665,687]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FAC739BA68ACFD0B" box="[1316,1461,665,687]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
given its nested position within the latter based on these autapomorphic specializations.
</paragraph>
<paragraph id="8BE87132E124DF31FC9039D1699AFB95" blockId="2.[859,1530,131,2024]" pageId="2" pageNumber="445">
Following Smiths (1954) inclusion of
<taxonomicName id="4C570AB1E124DF31FACA39D16EC8FC82" baseAuthorityName="Fowler &amp; Bean" baseAuthorityYear="1930" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="albomarginatus">
<emphasis id="B923AD20E124DF31FACA39D16EC8FC82" italics="true" pageId="2" pageNumber="445">Cephalopholis albomarginatus</emphasis>
</taxonomicName>
in
<taxonomicName id="4C570AB1E124DF31FBEA3833684AFC83" authority=", Randall (1964)" authorityName=", Randall" authorityYear="1964" box="[1033,1363,784,807]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FBEA38336996FC82" box="[1033,1167,784,806]" italics="true" pageId="2" pageNumber="445">Aethaloperca</emphasis>
, Randall (1964)
</taxonomicName>
removed the species and erected the new genus
<taxonomicName id="4C570AB1E124DF31FAE3380D6852FCE0" authorityName="Randall" authorityYear="1964" box="[1280,1355,814,836]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FAE3380D6852FCE0" box="[1280,1355,814,836]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
. While noting that the species showed affinities to
<taxonomicName id="4C570AB1E124DF31FAE038686EBCFC24" authority=", Randall (1964)" authorityName=", Randall" authorityYear="1964" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FAE03868688DFCC5" box="[1283,1428,843,865]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
, Randall (1964)
</taxonomicName>
highlighted differences in external morphology including a shorter head and an emarginate caudal fin. He also noted that
<taxonomicName id="4C570AB1E124DF31FBFA3887697DFC1E" authorityName="Randall" authorityYear="1964" box="[1049,1124,932,954]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FBFA3887697DFC1E" box="[1049,1124,932,954]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
displayed a different mode of life (semi-pelagic) from other species of
<taxonomicName id="4C570AB1E124DF31FAE738E1688CFC7C" authorityName=", Randall" authorityYear="1964" box="[1284,1429,962,984]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FAE738E1688CFC7C" box="[1284,1429,962,984]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
(
<figureCitation id="136C6DB7E124DF31FA4B38E168F5FC7D" box="[1448,1516,962,985]" captionStart="FIG" captionStartId="6.[142,170,944,964]" captionTargetBox="[142,1530,130,914]" captionTargetId="figure-590@6.[142,1530,130,914]" captionTargetPageId="6" captionText="FIG. 4. Cephalopholis argus (top left), C. albomarginata (bottom left), C. polleni (top right photo © Luiz Rocha; bottom right photo © Bob Fenner, www.wetwebmedia.com)." figureDoi="http://doi.org/10.5281/zenodo.12536088" httpUri="https://zenodo.org/record/12536088/files/figure.png" pageId="2" pageNumber="445">Fig. 4</figureCitation>
). Katayama (1974) described and placed a second species within
<taxonomicName id="4C570AB1E124DF31FC5738DE6EE6FBB7" authorityName="Randall" authorityYear="1964" box="[948,1023,1021,1043]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FC5738DE6EE6FBB7" box="[948,1023,1021,1043]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
,
<taxonomicName id="4C570AB1E124DF31FBF638DD6987FBB7" authorityName="Katayama" authorityYear="1974" box="[1045,1182,1021,1043]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="okinawae">
<emphasis id="B923AD20E124DF31FBF638DD6987FBB7" box="[1045,1182,1021,1043]" italics="true" pageId="2" pageNumber="445">G. okinawae</emphasis>
</taxonomicName>
(
<emphasis id="B923AD20E124DF31FB5038DF69DCFBB7" bold="true" box="[1203,1221,1020,1043]" pageId="2" pageNumber="445">¼</emphasis>
<taxonomicName id="4C570AB1E124DF31FB3138DD6824FBB7" baseAuthorityName="Bleeker" baseAuthorityYear="1868" box="[1234,1341,1021,1043]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="polleni">
<emphasis id="B923AD20E124DF31FB3138DD6824FBB7" box="[1234,1341,1021,1043]" italics="true" pageId="2" pageNumber="445">C. polleni</emphasis>
</taxonomicName>
), based on the presence of a truncate tail.
</paragraph>
<paragraph id="8BE87132E124DF31FC903F1A69F9F9A8" blockId="2.[859,1530,131,2024]" pageId="2" pageNumber="445">
The head length of
<taxonomicName id="4C570AB1E124DF31FBBA3F1A69BDFBEB" authorityName="Randall" authorityYear="1964" box="[1113,1188,1081,1103]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FBBA3F1A69BDFBEB" box="[1113,1188,1081,1103]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
(2.93.2 in SL; Heemstra and Randall, 1993; Craig et al., 2011) is well within the range of other species of
<taxonomicName id="4C570AB1E124DF31FBF53F5769BEFB2E" authorityName=", Randall" authorityYear="1964" box="[1046,1191,1140,1162]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FBF53F5769BEFB2E" box="[1046,1191,1140,1162]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
(2.23.1 in SL; Heemstra and Randall, 1993; Craig et al., 2011). The emarginated caudal fin of
<taxonomicName id="4C570AB1E124DF31FC993F936929FB62" baseAuthorityName="Fowler &amp; Bean" baseAuthorityYear="1930" box="[890,1072,1200,1222]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="albomarginata">
<emphasis id="B923AD20E124DF31FC993F936929FB62" box="[890,1072,1200,1222]" italics="true" pageId="2" pageNumber="445">G. albomarginata</emphasis>
</taxonomicName>
is emarginate only in juveniles, while in adults it is only weakly emarginated and is closer to truncate. A nearly emarginate caudal fin is also present in
<taxonomicName id="4C570AB1E124DF31FA023FCF6EC6FABB" authorityName="Katayama" authorityYear="1957" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="igarashiensis">
<emphasis id="B923AD20E124DF31FA023FCF6EC6FABB" italics="true" pageId="2" pageNumber="445">C. igarashiensis</emphasis>
</taxonomicName>
, and a truncate caudal fin is present in
<taxonomicName id="4C570AB1E124DF31FA753E2A68E1FABB" baseAuthorityName="Bleeker" baseAuthorityYear="1868" box="[1430,1528,1289,1311]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="polleni">
<emphasis id="B923AD20E124DF31FA753E2A68E1FABB" box="[1430,1528,1289,1311]" italics="true" pageId="2" pageNumber="445">C. polleni</emphasis>
</taxonomicName>
(although in some specimens the caudal is rounded;
<figureCitation id="136C6DB7E124DF31FA4E3E0468F5FA99" box="[1453,1516,1318,1341]" captionStart="FIG" captionStartId="6.[142,170,944,964]" captionTargetBox="[142,1530,130,914]" captionTargetId="figure-590@6.[142,1530,130,914]" captionTargetPageId="6" captionText="FIG. 4. Cephalopholis argus (top left), C. albomarginata (bottom left), C. polleni (top right photo © Luiz Rocha; bottom right photo © Bob Fenner, www.wetwebmedia.com)." figureDoi="http://doi.org/10.5281/zenodo.12536088" httpUri="https://zenodo.org/record/12536088/files/figure.png" pageId="2" pageNumber="445">Fig. 4</figureCitation>
). These three species also form a clade (
<figureCitation id="136C6DB7E124DF31FAEE3E67684CFAFE" box="[1293,1365,1348,1370]" 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="2" pageNumber="445">Figs. 1</figureCitation>
,
<figureCitation id="136C6DB7E124DF31FA863E67686CFAFF" box="[1381,1397,1348,1371]" 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="2" pageNumber="445">2</figureCitation>
). The semi-pelagic behavior of this species is more-or-less restricted to juveniles; adults display a roving behavior but are still closely associated with the substrate. Given the sum-total of the synapomorphies which are present in this clade, both morphological (see below) and molecular, excluding
<taxonomicName id="4C570AB1E124DF31FA4D3EFB68E0FA4A" authorityName="Randall" authorityYear="1964" box="[1454,1529,1496,1518]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FA4D3EFB68E0FA4A" box="[1454,1529,1496,1518]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
from
<taxonomicName id="4C570AB1E124DF31FC7A3ED56933F9A8" authorityName=", Randall" authorityYear="1964" box="[921,1066,1526,1548]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FC7A3ED56933F9A8" box="[921,1066,1526,1548]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
is unwarranted.
</paragraph>
<paragraph id="8BE87132E124DF31FC903D376858F84C" blockId="2.[859,1530,131,2024]" pageId="2" pageNumber="445">
The most surprising of the species included in the monophyletic
<taxonomicName id="4C570AB1E124DF31FBE03D11698DF9EC" authorityName=", Randall" authorityYear="1964" box="[1027,1172,1586,1608]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FBE03D11698DF9EC" box="[1027,1172,1586,1608]" italics="true" pageId="2" pageNumber="445">Cephalopholis</emphasis>
</taxonomicName>
are those of the genus
<taxonomicName id="4C570AB1E124DF31FA573D106E97F9C1" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FA573D106E97F9C1" italics="true" pageId="2" pageNumber="445">Paranthias</emphasis>
</taxonomicName>
. While rare among groupers, their semi-pelagic behavior is shared with
<taxonomicName id="4C570AB1E124DF31FC1B3D4D6932F927" authorityName="Randall, Johnson &amp; Lowe" authorityYear="1989" box="[1016,1067,1646,1667]" family="Serranidae" genus="Triso" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FC1B3D4D6932F927" box="[1016,1067,1646,1667]" italics="true" pageId="2" pageNumber="445">Triso</emphasis>
</taxonomicName>
and juveniles of
<taxonomicName id="4C570AB1E124DF31FB063D4E6829F927" authorityName="Randall" authorityYear="1964" box="[1253,1328,1645,1667]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FB063D4E6829F927" box="[1253,1328,1645,1667]" italics="true" pageId="2" pageNumber="445">Gracila</emphasis>
</taxonomicName>
, and their distinct morphological features such as a deeply forked tail, high gill raker counts, and small teeth, appear to be associated with a transition to the water column (
<figureCitation id="136C6DB7E124DF31FB5D3DE569E5F979" box="[1214,1276,1734,1757]" captionStart="FIG" captionStartId="7.[1139,1167,1701,1721]" captionTargetBox="[103,1060,131,1822]" captionTargetId="figure-77@7.[103,1060,131,1822]" captionTargetPageId="7" captionText="FIG. 5. Cephalopholis fulva (top), C. furcifer (middle), and hybrid C. fulva x C. furcifer (bottom). Top and middle photographs © Robert Myers, bottom photograph © Luiz Rocha." figureDoi="http://doi.org/10.5281/zenodo.12536091" httpUri="https://zenodo.org/record/12536091/files/figure.png" pageId="2" pageNumber="445">Fig. 5</figureCitation>
). Species of
<taxonomicName id="4C570AB1E124DF31FA603DE568EBF978" box="[1411,1522,1734,1756]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FA603DE568EBF978" box="[1411,1522,1734,1756]" italics="true" pageId="2" pageNumber="445">Paranthias</emphasis>
</taxonomicName>
, while clearly semi-pelagic, behave very much like other groupers in that when provided with a negative stimulus, they retreat into reef crevices rather than swim away, clearly reflecting their preference for the reef. It should also be noted that
<taxonomicName id="4C570AB1E124DF31FC6D3C786940F8D5" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[910,1113,1883,1905]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="fulva">
<emphasis id="B923AD20E124DF31FC6D3C786940F8D5" box="[910,1113,1883,1905]" italics="true" pageId="2" pageNumber="445">Cephalopholis fulva</emphasis>
</taxonomicName>
and
<taxonomicName id="4C570AB1E124DF31FB6C3C786854F8D5" baseAuthorityName="Valenciennes" baseAuthorityYear="1828" box="[1167,1357,1883,1905]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="furcifer">
<emphasis id="B923AD20E124DF31FB6C3C786854F8D5" box="[1167,1357,1883,1905]" italics="true" pageId="2" pageNumber="445">Paranthias furcifer</emphasis>
</taxonomicName>
are well known to hybridize in the wild (the hybrid being originally described as the monotypic
<taxonomicName id="4C570AB1E124DF31FB463CB56884F808" authorityName="Poey" authorityYear="1875" box="[1189,1437,1942,1964]" family="Labridae" genus="Menephorus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="punctiferus">
<emphasis id="B923AD20E124DF31FB463CB56884F808" box="[1189,1437,1942,1964]" italics="true" pageId="2" pageNumber="445">Menephorus punctiferus</emphasis>
</taxonomicName>
; Smith, 1966; Bostrom et al., 2002), and the two species of
<taxonomicName id="4C570AB1E124DF31FA6A3C9768E1F86E" box="[1417,1528,1972,1994]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E124DF31FA6A3C9768E1F86E" box="[1417,1528,1972,1994]" italics="true" pageId="2" pageNumber="445">Paranthias</emphasis>
</taxonomicName>
form the sister group to
<taxonomicName id="4C570AB1E124DF31FB8F3CF169A6F843" baseAuthorityName="Linnaeus" baseAuthorityYear="1758" box="[1132,1215,2001,2023]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="445" phylum="Chordata" rank="species" species="fulva">
<emphasis id="B923AD20E124DF31FB8F3CF169A6F843" box="[1132,1215,2001,2023]" italics="true" pageId="2" pageNumber="445">C. fulva</emphasis>
</taxonomicName>
(
<figureCitation id="136C6DB7E124DF31FB2C3CF1680FF843" box="[1231,1302,2002,2024]" 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="2" pageNumber="445">Figs. 1</figureCitation>
,
<figureCitation id="136C6DB7E124DF31FAC63CF2682CF84C" box="[1317,1333,2001,2024]" 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="2" pageNumber="445">2</figureCitation>
).
</paragraph>
<caption id="DF2821BAE125DF30FF843C186E07F8CD" ID-DOI="http://doi.org/10.5281/zenodo.12536076" ID-Zenodo-Dep="12536076" httpUri="https://zenodo.org/record/12536076/files/figure.png" pageId="3" pageNumber="446" startId="3.[103,131,1851,1871]" targetBox="[217,1376,130,1821]" targetPageId="3" targetType="figure">
<paragraph id="8BE87132E125DF30FF843C186E07F8CD" blockId="3.[103,1490,1851,1897]" pageId="3" pageNumber="446">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 (%).</paragraph>
</caption>
<caption id="DF2821BAE122DF37FF6D3C186C11F8CD" ID-DOI="http://doi.org/10.5281/zenodo.12536081" ID-Zenodo-Dep="12536081" httpUri="https://zenodo.org/record/12536081/files/figure.png" pageId="4" pageNumber="447" startId="4.[142,170,1851,1871]" targetBox="[213,1458,130,1821]" targetPageId="4" targetType="figure">
<paragraph id="8BE87132E122DF37FF6D3C186C11F8CD" blockId="4.[142,1529,1851,1897]" pageId="4" pageNumber="447">FIG. 2. Bayesian inference phylogram reconstructed using MrBayes following Ma et al. (2016). Numbers on branches represent posterior probabilities.</paragraph>
</caption>
<caption id="DF2821BAE123DF36FB903FC3684CFA8D" ID-DOI="http://doi.org/10.5281/zenodo.12536083" ID-Zenodo-Dep="12536083" httpUri="https://zenodo.org/record/12536083/files/figure.png" pageId="5" pageNumber="448" startId="5.[1139,1170,1248,1268]" targetBox="[103,1089,131,1316]" targetPageId="5" targetType="figure">
<paragraph id="8BE87132E123DF36FB903FC3684CFA8D" blockId="5.[1139,1490,1247,1321]" pageId="5" pageNumber="448">
<emphasis id="B923AD20E123DF36FB903FC369ABFA8D" bold="true" pageId="5" pageNumber="448">
FIG. 3.
<taxonomicName id="4C570AB1E123DF36FB2F3FC368CBFB57" baseAuthorityName="Valenciennes" baseAuthorityYear="1828" box="[1228,1490,1247,1268]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="species" species="sonnerati">
<emphasis id="B923AD20E123DF36FB2F3FC368CBFB57" bold="true" box="[1228,1490,1247,1268]" italics="true" pageId="5" pageNumber="448">Cephalopholis sonnerati</emphasis>
</taxonomicName>
(top) and
<taxonomicName id="4C570AB1E123DF36FB353FD96830FAAA" baseAuthorityName="Forsskal" baseAuthorityYear="1775" box="[1238,1321,1274,1294]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="species" species="rogaa">
<emphasis id="B923AD20E123DF36FB353FD96830FAAA" bold="true" box="[1238,1321,1274,1294]" italics="true" pageId="5" pageNumber="448">C. rogaa</emphasis>
</taxonomicName>
(bottom). Photographs
</emphasis>
©
<taxonomicName id="4C570AB1E123DF36FB323E36684CFA8D" authority="Myers." authorityName="Myers." box="[1233,1365,1301,1321]" class="Insecta" family="Coccinellidae" genus="Robert" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="448" phylum="Arthropoda" rank="genus">Robert Myers.</taxonomicName>
</paragraph>
</caption>
<paragraph id="8BE87132E123DF36FF9C3E766C56F973" blockId="5.[103,773,1365,2024]" pageId="5" pageNumber="448">
Evolutionary transitions involving radical changes to morphology within a clade are not unique to groupers. For example, the fusiliers (Caesioninae) are deeply nested within the larger snapper family (
<taxonomicName id="4C570AB1E123DF36FE6E3E936F1DFA63" authorityName="Gill" authorityYear="1861" box="[397,516,1456,1479]" family="Lutjanidae" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="family">Lutjanidae</taxonomicName>
) despite having clearly different external adaptations to mid-water habitat (Miller and Cribb, 2007). The semi-pelagic boga (
<taxonomicName id="4C570AB1E123DF36FDD03ECF6FE5F9A6" baseAuthorityName="Poey" baseAuthorityYear="1860" box="[563,764,1516,1538]" family="Haemulidae" genus="Haemulon" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="species" species="vittatum">
<emphasis id="B923AD20E123DF36FDD03ECF6FE5F9A6" box="[563,764,1516,1538]" italics="true" pageId="5" pageNumber="448">Haemulon vittatum</emphasis>
</taxonomicName>
) was long regarded as belonging to the monotypic genus
<taxonomicName id="4C570AB1E123DF36FF843D096DAEF99B" box="[103,183,1578,1599]" family="Haemulidae" genus="Inermia" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E123DF36FF843D096DAEF99B" box="[103,183,1578,1599]" italics="true" pageId="5" pageNumber="448">Inermia</emphasis>
</taxonomicName>
; however, molecular evidence clearly indicates its placement within
<taxonomicName id="4C570AB1E123DF36FEA03D646FB6F9FA" authority="(Rocha et al., 2008)" baseAuthorityName="Rocha" baseAuthorityYear="2008" box="[323,687,1607,1630]" family="Haemulidae" genus="Haemulon" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E123DF36FEA03D646CB7F9F9" box="[323,430,1607,1629]" italics="true" pageId="5" pageNumber="448">Haemulon</emphasis>
(Rocha et al., 2008)
</taxonomicName>
. While certainly unexpected, the placement of the two species of
<taxonomicName id="4C570AB1E123DF36FF843DA76DCFF93E" box="[103,214,1668,1690]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E123DF36FF843DA76DCFF93E" box="[103,214,1668,1690]" italics="true" pageId="5" pageNumber="448">Paranthias</emphasis>
</taxonomicName>
nested within
<taxonomicName id="4C570AB1E123DF36FE693DA76F02F93E" authorityName=", Randall" authorityYear="1964" box="[394,539,1668,1690]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E123DF36FE693DA76F02F93E" box="[394,539,1668,1690]" italics="true" pageId="5" pageNumber="448">Cephalopholis</emphasis>
</taxonomicName>
is clearly warranted given the morphological and molecular synapomorphies that unite this clade.
</paragraph>
<paragraph id="8BE87132E123DF36FF9C3DFC6836FA2F" blockId="5.[103,773,1365,2024]" lastBlockId="5.[820,1490,1365,1419]" pageId="5" pageNumber="448">
Despite their seemingly different morphologies, these four genera share a number of characters, including the presence of nine dorsal fin spines (shared only with
<taxonomicName id="4C570AB1E123DF36FD613C3F6DFDF8F4" baseAuthorityName="Gilbert" baseAuthorityYear="1892" family="Serranidae" genus="Hyporthodus" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="species" species="acanthistius">
<emphasis id="B923AD20E123DF36FD613C3F6DFDF8F4" italics="true" pageId="5" pageNumber="448">Hyporthodus acanthistius</emphasis>
</taxonomicName>
), the ninth of which is formed indirectly from a soft ray (Kendall, 1979; Leis, 1986), and the presence of trisegmental pterygiophores in the dorsal fin (bisegmental in remaining epinephelines; Heemstra and Randall, 1993).
<emphasis id="B923AD20E123DF36FF843C906C4EF86D" box="[103,343,1971,1993]" italics="true" pageId="5" pageNumber="448">
<taxonomicName id="4C570AB1E123DF36FF843C906DE2F86D" authorityName=", Randall" authorityYear="1964" box="[103,251,1971,1993]" family="Serranidae" genus="Cephalopholis" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">Cephalopholis</taxonomicName>
,
<taxonomicName id="4C570AB1E123DF36FEEF3C906C4EF86D" authorityName="Randall" authorityYear="1964" box="[268,343,1971,1993]" family="Serranidae" genus="Gracila" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">Gracila</taxonomicName>
</emphasis>
, and
<taxonomicName id="4C570AB1E123DF36FE413C906F08F86D" box="[418,529,1971,1993]" family="Serranidae" genus="Paranthias" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E123DF36FE413C906F08F86D" box="[418,529,1971,1993]" italics="true" pageId="5" pageNumber="448">Paranthias</emphasis>
</taxonomicName>
also share epineural ribs on vertebrae 19 (
<emphasis id="B923AD20E123DF36FE843CF26F65F843" box="[359,636,2001,2023]" italics="true" pageId="5" pageNumber="448">
<taxonomicName id="4C570AB1E123DF36FE843CF26CF7F843" authorityName=", Randall" authorityYear="1964" box="[359,494,2001,2023]" family="Serranidae" genus="Aethaloperca" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">Aethaloperca</taxonomicName>
,
<taxonomicName id="4C570AB1E123DF36FE183CF26F61F843" authorityName="Bloch" authorityYear="1793" box="[507,632,2001,2023]" family="Serranidae" genus="Epinephelus" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">Epinephelus</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C570AB1E123DF36FD5B3CF16E6CFACF" authorityName="Gill" authorityYear="1862" box="[696,885,1366,2023]" family="Serranidae" genus="Mycteroperca" kingdom="Animalia" order="Perciformes" pageId="5" pageNumber="448" phylum="Chordata" rank="genus">
<emphasis id="B923AD20E123DF36FD5B3CF16E6CFACF" box="[696,885,1366,2023]" italics="true" pageId="5" pageNumber="448">Mycteroperca</emphasis>
</taxonomicName>
have epineurals on vertebrae 110; Baldwin and Johnson, 1993; Heemstra and Randall, 1993).
</paragraph>
</subSubSection>
</emphasis>
</treatment>
</document>