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<mods:title id="9B18A902EDB1153F25C67DF905F32E2D">A new classification of the family Ariidae (Osteichthyes: Ostariophysi: Siluriformes) based on combined analyses of morphological and molecular data</mods:title>
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<mods:namePart id="CF59E1310D5C90DD7F41FFB0627EAC4E">Marceniuk, Alexandre Pires</mods:namePart>
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<mods:namePart id="10060E7D2B1AF26F935ED4DA64D5E645">Oliveira, Claudio</mods:namePart>
<mods:affiliation id="A6F8C3A4DCABFEEB59B0F05713B22B98">Departamento de Biologia Estrutural e Funcional, Instituto de Biociências, Universidade Estadual Paulista Júlio de Mesquita Filho (UNESP), Botucatu, SP 18618 - 970, Brazil</mods:affiliation>
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<paragraph id="8BE59A61FFC7FF810E83FD89FABD4867" blockId="32.[967,1302,607,634]" box="[967,1302,607,634]" pageId="32" pageNumber="458">
<heading id="D0AD2D0DFFC7FF810E83FD89FABD4867" bold="true" box="[967,1302,607,634]" centered="true" fontSize="11" level="2" pageId="32" pageNumber="458" reason="2">
<taxonomicName id="4C5AE1E2FFC7FF810E83FD89FABD4867" ID-CoL="4WZ5" ID-ENA="591301" authority="Bleeker, 1858" authorityName="Bleeker" authorityYear="1858" box="[967,1302,607,634]" family="Ariidae" genus="Hexanematichthys" kingdom="Animalia" order="Siluriformes" pageId="32" pageNumber="458" phylum="Chordata" rank="genus">
<emphasis id="B92E4673FFC7FF810E83FD89FABD4867" bold="true" box="[967,1302,607,634]" pageId="32" pageNumber="458">
<emphasis id="B92E4673FFC7FF810E83FD89FB2A4864" bold="true" box="[967,1153,607,634]" italics="true" pageId="32" pageNumber="458">Hexanematichthys</emphasis>
<bibRefCitation id="EFCBE790FFC7FF8109C1FDB6FABD4867" author="Bleeker P" box="[1157,1302,607,634]" pageId="32" pageNumber="458" pagination="1 - 370" refId="ref29792" refString="Bleeker P. De Visschen van den Indischen Archipel. Beschreven en toegelicht. Siluri. Acta Societatis Regiae Scientiarum Indo-Neerlandicae 1858; 4: 1 - 370." type="journal article" year="1858">Bleeker, 1858</bibRefCitation>
</emphasis>
</taxonomicName>
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<subSubSection id="C340C9EAFFC7FF80094CFD58FF6E4886" lastPageId="33" lastPageNumber="459" pageId="32" pageNumber="458" type="description">
<paragraph id="8BE59A61FFC7FF81094CFD58FB7E48B6" blockId="32.[1032,1237,654,681]" box="[1032,1237,654,681]" pageId="32" pageNumber="458">
<emphasis id="B92E4673FFC7FF81094CFD58FB7E48B6" bold="true" box="[1032,1237,654,681]" pageId="32" pageNumber="458">
(
<figureCitation id="136186E4FFC7FF810956FD59FBC748B7" box="[1042,1132,655,681]" captionStart-0="Figure 1" captionStart-1="Figure 2" captionStart-2="Figure 3" captionStartId-0="3.[129,194,1805,1829]" captionStartId-1="3.[825,890,1803,1827]" captionStartId-2="4.[113,178,1813,1837]" captionTargetBox-1="[153,1434,148,1768]" captionTargetBox-2="[151,708,146,1775]" captionTargetId-2="figure-643@4.[149,717,144,1776]" captionTargetPageId-1="3" captionTargetPageId-2="4" captionText-0="Figure 1. MP analysis.Subfamilies (SF) Galeichthyinae (pink), Bagreinae (bright blue), and Ariinae (dark blue). Tribe (T) Ariini (blue), Cathopsini (orange), Genidentini tribe nov. (brown), and Sciadeini tribe nov. (light green).Subtribe (ST) Ariina (light blue) and Doiichthyina (green). Incertae sedis taxa (IS) (red). Numbers over branches correspond to the bootstrap support." captionText-1="Figure 2. ML analysis. Subfamilies (SF) Galeichthyinae (pink), Bagreinae (bright blue), and Ariinae (dark blue). Tribe (T) Ariini (blue), Cathopsini (orange), Genidentini tribe nov. (brown), and Sciadeini tribe nov. (light green). Subtribe (ST) Ariina (light blue) and Doiichthyina (green). Incertae sedis taxa (IS) (red). Numbers over branches correspond to the bootstrap support." captionText-2="Figure 3. BI analysis.Subfamilies (SF) Galeichthyinae (pink), Bagreinae (bright blue), and Ariinae (dark blue). Tribe (T) Ariini (blue), Cathopsini (orange), Genidentini tribe nov. (brown), and Sciadeini tribe nov. (light green).Subtribe (ST) Ariina (light blue) and Doiichthyina (green). Incertae sedis taxa (IS) (red). Numbers over branches correspond to the bootstrap support." figureDoi-1="http://doi.org/10.5281/zenodo.11240122" figureDoi-2="http://doi.org/10.5281/zenodo.11240126" httpUri-1="https://zenodo.org/record/11240122/files/figure.png" httpUri-2="https://zenodo.org/record/11240126/files/figure.png" pageId="32" pageNumber="458">Figs 13</figureCitation>
,
<figureCitation id="136186E4FFC7FF810932FD59FB0E48B7" box="[1142,1189,655,681]" captionStart="Figure 24" captionStartId="27.[129,194,1851,1875]" captionTargetBox="[259,1343,151,1822]" captionTargetId="figure-8@27.[257,1345,145,1823]" captionTargetPageId="27" captionText="Figure 24. Representative skulls, in dorsal view, and distribution of the subtribe Ariina.A, Arius maculatus, AUM 44875. B, Cephalocassis melanochir, CAS 49426. C, Cryptarius truncatus, AUM 50243. D, Hexanematichthys sagor, AUM 50242. E, Kyataphisa nenga, AUM 46280. F, Osteogeneiosus militaris, AUM 44880. G, Pseudosciades sona, AUM 44861. * marine, ** brackish water, *** freshwater." figureDoi="http://doi.org/10.5281/zenodo.11240169" httpUri="https://zenodo.org/record/11240169/files/figure.png" pageId="32" pageNumber="458">24D</figureCitation>
,
<figureCitation id="136186E4FFC7FF8109EBFD59FB6048B7" box="[1199,1227,655,681]" captionStart="Figure 32" captionStartId="33.[129,194,1270,1294]" captionTargetBox="[129,1473,770,1243]" captionTargetId="figure-214@33.[129,1473,770,1243]" captionTargetPageId="33" captionText="Figure 32. Hexanematichthys sagor (senior synonym of Bagrus sondaicus), Indo-West Pacific, photograph by Ricardo Betancur-R., type species of the genus." figureDoi="http://doi.org/10.5281/zenodo.11240185" httpUri="https://zenodo.org/record/11240185/files/figure.png" pageId="32" pageNumber="458">32</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC7FF810E6EFD06FA8848F9" blockId="32.[810,1315,719,743]" box="[810,1315,719,743]" pageId="32" pageNumber="458">
<emphasis id="B92E4673FFC7FF810E6EFD06FC0D48F9" box="[810,934,720,743]" italics="true" pageId="32" pageNumber="458">
<typeStatus id="54E124C3FFC7FF810E6EFD06FCF148F9" box="[810,858,720,743]" pageId="32" pageNumber="458">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="4C5AE1E2FFC7FF810EEFFD06FAB548F9" authority="Valenciennes, 1840" authorityName="Valenciennes" authorityYear="1840" box="[939,1310,719,743]" family="Bagridae" genus="Bagrus" kingdom="Animalia" order="Siluriformes" pageId="32" pageNumber="458" phylum="Chordata" rank="species" species="sondaicus">
<emphasis id="B92E4673FFC7FF810EEFFD06FBF948F9" box="[939,1106,719,743]" italics="true" pageId="32" pageNumber="458">Bagrus sondaicus</emphasis>
Valenciennes, 1840
</taxonomicName>
.
</paragraph>
<paragraph id="8BE59A61FFC7FF810904FCDEFB36493F" blockId="32.[810,1459,776,1245]" box="[1088,1181,776,801]" pageId="32" pageNumber="458">
<emphasis id="B92E4673FFC7FF810904FCDEFB36493F" box="[1088,1181,776,801]" italics="true" pageId="32" pageNumber="458">Diagnosis</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC7FF810E6EFCFBFB304EC3" blockId="32.[810,1459,776,1245]" pageId="32" pageNumber="458">Posterior branches of mesethmoid moderately long, delimiting between one-fourth and one-half of length of anterior cranial fontanel (8, 0&gt; 1); posterior branch of lateral ethmoid depressed (13, 0&gt; 1); posterior cranial fontanel absent (26, 1&gt; 0); epiphyseal bar indistinct (28, 0&gt; 1); temporal fossa absent (38, 0&gt; 1); accessory tooth plates small, oval to rounded (60, 2&gt; 1); lateral margins of orbitosphenoid progressively diverging anteriorly (63, 0&gt; 1); autopalatine posterior portion conspicuously compressed (106, 0&gt; 1); articulation of autopalatine with lateral ethmoid mesoposteriorly oriented (110, 0&gt; 1); lateral and mesial portions of premaxilla with different sizes (121, 0&gt; 1); anteroventral portion of opercle subtrapezoidal, very long (127, 1&gt; 0); Müllerian ramus distal third markedly curved (208, 1&gt; 2).</paragraph>
<paragraph id="8BE59A61FFC7FF800E6EFAD1FEA04AD9" blockId="32.[810,1459,1287,1373]" lastBlockId="33.[129,777,144,199]" lastPageId="33" lastPageNumber="459" pageId="32" pageNumber="458">
<emphasis id="B92E4673FFC7FF810E6EFAD1FB924F00" box="[810,1081,1287,1311]" italics="true" pageId="32" pageNumber="458">Ambiguous optimization:</emphasis>
Exoccipital bony crest perpendicular to vertebral column and ventrolaterally directed (92, 1&gt; 0); 15 or more ribs (213, 0&gt; 1); second dorsal cleithral process posteriorly directed and parallel to posterior process (226, 1&gt; 0).
</paragraph>
<caption id="DF25CAE9FFC7FF810D37F850FD374DA5" ID-DOI="http://doi.org/10.5281/zenodo.11240183" ID-Zenodo-Dep="11240183" httpUri="https://zenodo.org/record/11240183/files/figure.png" pageId="32" pageNumber="458" startId="32.[115,180,1926,1950]" targetBox="[114,1457,1490,1898]" targetPageId="32" targetType="figure">
<paragraph id="8BE59A61FFC7FF810D37F850FD374DA5" blockId="32.[113,1410,1926,1979]" pageId="32" pageNumber="458">
<emphasis id="B92E4673FFC7FF810D37F850FF7C4D81" bold="true" box="[115,215,1926,1951]" pageId="32" pageNumber="458">Figure 31.</emphasis>
<taxonomicName id="4C5AE1E2FFC7FF810D98F851FDD24D81" authority="Bleeker, 1858" authorityName="Bleeker" authorityYear="1858" box="[220,633,1927,1951]" class="Actinopterygii" family="Ariidae" genus="Hemipimelodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Siluriformes" pageId="32" pageNumber="458" phylum="Chordata" rank="species" species="macrocephalus">
<emphasis id="B92E4673FFC7FF810D98F851FE584D81" box="[220,499,1927,1951]" italics="true" pageId="32" pageNumber="458">Hemipimelodus macrocephalus</emphasis>
<bibRefCitation id="EFCBE790FFC7FF810CB3F851FDD24D81" author="Bleeker P" box="[503,633,1927,1951]" pageId="32" pageNumber="458" pagination="1 - 370" refId="ref29792" refString="Bleeker P. De Visschen van den Indischen Archipel. Beschreven en toegelicht. Siluri. Acta Societatis Regiae Scientiarum Indo-Neerlandicae 1858; 4: 1 - 370." type="journal article" year="1858">Bleeker, 1858</bibRefCitation>
</taxonomicName>
, junior synonym of
<taxonomicName id="4C5AE1E2FFC7FF810E71F851FB014D81" authority="Bleeker, 1851" authorityName="Bleeker" authorityYear="1851" box="[821,1194,1927,1951]" class="Actinopterygii" family="Ariidae" genus="Hemipimelodus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Siluriformes" pageId="32" pageNumber="458" phylum="Chordata" rank="species" species="borneensis">
<emphasis id="B92E4673FFC7FF810E71F851FB884D81" box="[821,1059,1927,1951]" italics="true" pageId="32" pageNumber="458">Hemipimelodus borneensis</emphasis>
Bleeker, 1851
</taxonomicName>
, BMNH 1863.12.4.78, Sundaland, Malaysia to Vietnam, type species of the genus.
</paragraph>
</caption>
<paragraph id="8BE59A61FFC6FF800C39FF31FDA64B1F" blockId="33.[129,539,231,293]" box="[381,525,231,257]" pageId="33" pageNumber="459">
<emphasis id="B92E4673FFC6FF800C39FF31FDA64B1F" box="[381,525,231,257]" italics="true" pageId="33" pageNumber="459">Included species</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC6FF800DC5FEDBFDB04B3B" blockId="33.[129,539,231,293]" box="[129,539,269,293]" pageId="33" pageNumber="459">
<taxonomicName id="4C5AE1E2FFC6FF800DC5FEDBFDBC4B3B" authority="Hamilton, 1822" authorityName="Hamilton" authorityYear="1822" box="[129,535,269,293]" family="Ariidae" genus="Hexanematichthys" kingdom="Animalia" order="Siluriformes" pageId="33" pageNumber="459" phylum="Chordata" rank="species" species="sagor">
<emphasis id="B92E4673FFC6FF800DC5FEDBFEC64B3B" box="[129,365,269,293]" italics="true" pageId="33" pageNumber="459">Hexanematichthys sagor</emphasis>
Hamilton, 1822
</taxonomicName>
.
</paragraph>
<paragraph id="8BE59A61FFC6FF800DC5FE91FED64B60" blockId="33.[129,777,327,382]" pageId="33" pageNumber="459">
<emphasis id="B92E4673FFC6FF800DC5FE91FEDD4B41" box="[129,374,327,351]" italics="true" pageId="33" pageNumber="459">Habitat and distribution:</emphasis>
Brackish and marine waters, South and Southeast Asia (
<figureCitation id="136186E4FFC6FF800C63FEB0FEC74B60" box="[295,364,358,382]" captionStart="Figure 24" captionStartId="27.[129,194,1851,1875]" captionTargetBox="[259,1343,151,1822]" captionTargetId="figure-8@27.[257,1345,145,1823]" captionTargetPageId="27" captionText="Figure 24. Representative skulls, in dorsal view, and distribution of the subtribe Ariina.A, Arius maculatus, AUM 44875. B, Cephalocassis melanochir, CAS 49426. C, Cryptarius truncatus, AUM 50243. D, Hexanematichthys sagor, AUM 50242. E, Kyataphisa nenga, AUM 46280. F, Osteogeneiosus militaris, AUM 44880. G, Pseudosciades sona, AUM 44861. * marine, ** brackish water, *** freshwater." figureDoi="http://doi.org/10.5281/zenodo.11240169" httpUri="https://zenodo.org/record/11240169/files/figure.png" pageId="33" pageNumber="459">Fig. 24</figureCitation>
).
</paragraph>
<paragraph id="8BE59A61FFC6FF800CDFFE4BFE444BA9" blockId="33.[411,495,413,439]" box="[411,495,413,439]" pageId="33" pageNumber="459">
<emphasis id="B92E4673FFC6FF800CDFFE4BFE444BA9" box="[411,495,413,439]" italics="true" pageId="33" pageNumber="459">Remarks</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC6FF800DC5FE12FF6E4886" blockId="33.[129,778,452,664]" pageId="33" pageNumber="459">
The total-evidence analysis supports the validity of
<taxonomicName id="4C5AE1E2FFC6FF800DC5FE35FE9E4BE5" authorityName="Bleeker" authorityYear="1858" box="[129,309,483,507]" family="Ariidae" genus="Hexanematichthys" kingdom="Animalia" order="Siluriformes" pageId="33" pageNumber="459" phylum="Chordata" rank="genus">
<emphasis id="B92E4673FFC6FF800DC5FE35FE9E4BE5" box="[129,309,483,507]" italics="true" pageId="33" pageNumber="459">Hexanematichthys</emphasis>
</taxonomicName>
, corroborating the result of a previous molecular study (Betancur-R. 2009) that reveals a strong morphological convergence between
<taxonomicName id="4C5AE1E2FFC6FF800C9DFDF4FD264824" authorityName="Bleeker" authorityYear="1858" box="[473,653,546,570]" family="Ariidae" genus="Hexanematichthys" kingdom="Animalia" order="Siluriformes" pageId="33" pageNumber="459" phylum="Chordata" rank="genus">
<emphasis id="B92E4673FFC6FF800C9DFDF4FD264824" box="[473,653,546,570]" italics="true" pageId="33" pageNumber="459">Hexanematichthys</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5AE1E2FFC6FF800F86FDF4FCA24824" authorityName="Muller &amp; Troschel" authorityYear="1849" box="[706,777,546,570]" family="Ariidae" genus="Sciades" kingdom="Animalia" order="Siluriformes" pageId="33" pageNumber="459" phylum="Chordata" rank="genus">
<emphasis id="B92E4673FFC6FF800F86FDF4FCA24824" box="[706,777,546,570]" italics="true" pageId="33" pageNumber="459">Sciades</emphasis>
</taxonomicName>
(characters 26, 28, 93, 106, and 110), which had resulted in them having previously been treated as synonyms (
<bibRefCitation id="EFCBE790FFC6FF800F25FDB7FF1E4886" author="Marceniuk AP &amp; Menezes NA &amp; Britto MR" pageId="33" pageNumber="459" pagination="534 - 669" refId="ref30859" refString="Marceniuk AP, Menezes NA, Britto MR. Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera. Zoological Journal of the Linnean Society 2012; 165: 534 - 669. https: // doi. org / 10.1111 / j. 1096 - 3642.2012.00822. x" type="journal article" year="2012">
Marceniuk
<emphasis id="B92E4673FFC6FF800F9CFDB7FCA14867" box="[728,778,609,633]" italics="true" pageId="33" pageNumber="459">et al.</emphasis>
2012
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).
</paragraph>
</subSubSection>
</treatment>
</document>