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<document id="32C54F9BA7D494C324EEA64E6B061554" ID-DOI="10.1093/zoolinnean/zlad078" ID-ISSN="0024-4082" ID-Zenodo-Dep="11240120" ID-ZooBank="5D6C7EBF-E568-4100-9364-2DD357003878" IM.bibliography_approvedBy="guilherme" IM.illustrations_approvedBy="guilherme" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="guilherme" IM.treatments_approvedBy="guilherme" checkinTime="1716361325736" checkinUser="plazi" docAuthor="Marceniuk, Alexandre Pires, Oliveira, Claudio &amp; Ferraris Jr, Carl J." docDate="2024" docId="03F32B77FFC2FF860950FD49FDCC4FD2" docLanguage="en" docName="zlad078.pdf" docOrigin="Zoological Journal of the Linnean Society 200 (2)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad078" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.5:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="5" docTitle="Bleekeriella Marceniuk &amp; Oliveira &amp; Ferraris Jr 2024, gen. nov." docType="treatment" docVersion="3" lastPageNumber="463" masterDocId="FFCA530FFFE7FFA10D44FFD6FFAB4A1E" masterDocTitle="A new classification of the family Ariidae (Osteichthyes: Ostariophysi: Siluriformes) based on combined analyses of morphological and molecular data" masterLastPageNumber="476" masterPageNumber="426" pageNumber="463" updateTime="1716560620312" updateUser="guilherme">
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<mods:title id="F545E6F51F13935E28F7DEBF6C594061">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="D9A13029BDD1DE6B1E3B795F6B1A0BB5">Marceniuk, Alexandre Pires</mods:namePart>
<mods:affiliation id="40A8C53735061AF423E3BB759E308AF2">Programa de Pós-Graduação em Ciências Biológicas, Universidade Federal da Paraíba, conjunto Presidente Castelo Branco III, João Pessoa, PB 58050 - 000, Brazil &amp; Programa de Pós-Graduação em Ecologia e Conservação, Universidade Estadual da Paraíba, Campina Grande, PB 58429 - 500, Brazil &amp; 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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<mods:namePart id="D2AF650F19DD18CF86B7D787ED6AEC51">Oliveira, Claudio</mods:namePart>
<mods:affiliation id="2A16339F4C035F125D1411E759E5733F">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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<mods:namePart id="898648DE2992F710CA80495A030ED06F">Ferraris Jr, Carl J.</mods:namePart>
<mods:affiliation id="5530B8A79C81C0AD7D4E062E74A38515">545 NE Floral Pl., Portland, OR 97232, USA</mods:affiliation>
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<mods:date id="0AC35B130AB9081C61E064AD986F4D1B">2024</mods:date>
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<treatment id="03F32B77FFC2FF860950FD49FDCC4FD2" LSID="urn:lsid:plazi:treatment:03F32B77FFC2FF860950FD49FDCC4FD2" httpUri="http://treatment.plazi.org/id/03F32B77FFC2FF860950FD49FDCC4FD2" lastPageId="39" lastPageNumber="463" pageId="37" pageNumber="463">
<subSubSection id="C340C9EAFFC2FF840950FD49FB4C48A7" box="[1044,1255,671,698]" pageId="37" pageNumber="463" type="nomenclature">
<paragraph id="8BE59A61FFC2FF840950FD49FB4C48A7" blockId="37.[1044,1255,671,698]" box="[1044,1255,671,698]" pageId="37" pageNumber="463">
<heading id="D0AD2D0DFFC2FF840950FD49FB4C48A7" bold="true" box="[1044,1255,671,698]" centered="true" fontSize="11" level="2" pageId="37" pageNumber="463" reason="2">
<emphasis id="B92E4673FFC2FF840950FD49FB4C48A7" bold="true" box="[1044,1255,671,698]" pageId="37" pageNumber="463">
<taxonomicName id="4C5AE1E2FFC2FF840950FD49FB2348A4" authority="Marceniuk &amp; Oliveira &amp; Ferraris Jr, 2024" authorityName="Marceniuk &amp; Oliveira &amp; Ferraris Jr" authorityYear="2024" box="[1044,1160,671,698]" family="Ariidae" genus="Bleekeriella" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B92E4673FFC2FF840950FD49FB2348A4" bold="true" box="[1044,1160,671,698]" italics="true" pageId="37" pageNumber="463">Bleekeriella</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21DFB08FFC2FF8409C8FD49FB4C48A7" box="[1164,1255,671,697]" pageId="37" pageNumber="463" rank="genus">gen. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C340C9EAFFC2FF84094DFD08FB0A498A" pageId="37" pageNumber="463" type="description">
<paragraph id="8BE59A61FFC2FF84094DFD08FB5948E6" blockId="37.[1033,1266,734,760]" box="[1033,1266,734,760]" pageId="37" pageNumber="463">
<heading id="D0AD2D0DFFC2FF84094DFD08FB5948E6" allCaps="true" bold="true" box="[1033,1266,734,760]" centered="true" fontSize="11" level="1" pageId="37" pageNumber="463" reason="1">
<emphasis id="B92E4673FFC2FF84094DFD08FB5948E6" bold="true" box="[1033,1266,734,760]" pageId="37" pageNumber="463">MP 100, BI 1, ML 100</emphasis>
</heading>
</paragraph>
<paragraph id="8BE59A61FFC2FF840977FCCAFB634928" blockId="37.[1075,1224,796,823]" box="[1075,1224,796,823]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840977FCCAFB634928" bold="true" box="[1075,1224,796,823]" pageId="37" pageNumber="463">
(
<figureCitation id="136186E4FFC2FF84097AFCCAFB3C4928" box="[1086,1175,796,822]" 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-0="[152,776,143,1771]" captionTargetBox-1="[843,1439,143,1771]" captionTargetBox-2="[151,708,146,1775]" captionTargetId-0="figure-8@3.[152,744,149,1755]" captionTargetId-1="figure-61@3.[851,1439,143,1771]" 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="37" pageNumber="463">Figs 13</figureCitation>
,
<figureCitation id="136186E4FFC2FF8409E6FCCAFB154928" box="[1186,1214,796,822]" captionStart="Figure 40" captionStartId="39.[129,194,1267,1291]" captionTargetBox="[129,1473,687,1239]" captionTargetId="figure-271@39.[129,1473,687,1239]" captionTargetPageId="39" captionText="Figure 40. Bleekeriella leptaspis, northern Australia and New Guinea, photograph by Ricardo Betancur-R., type species of the genus." figureDoi="http://doi.org/10.5281/zenodo.11240201" httpUri="https://zenodo.org/record/11240201/files/figure.png" pageId="37" pageNumber="463">40</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC2FF840E7DFC8BFB0A498A" blockId="37.[825,1474,861,916]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840E7DFC8BFB8B496B" box="[825,1056,861,885]" italics="true" pageId="37" pageNumber="463">ZooBank registration:</emphasis>
<uri id="FFCB9663FFC2FF84097EFC8BFB30498A" pageId="37" pageNumber="463">
urn:lsid:zoobank.org:act:
<uuid id="FFFCA0B4FFC2FF84087DFC8BFB30498A" pageId="37" pageNumber="463">8CB66BBD-FB43-495E-B45A-EBF42667194F</uuid>
</uri>
.
</paragraph>
</subSubSection>
<subSubSection id="C340C9EAFFC2FF840E7DFC60FB7649D0" box="[825,1245,950,974]" pageId="37" pageNumber="463" type="type_taxon">
<paragraph id="8BE59A61FFC2FF840E7DFC60FB7649D0" blockId="37.[825,1245,950,974]" box="[825,1245,950,974]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840E7DFC60FC1E49D3" box="[825,949,950,973]" italics="true" pageId="37" pageNumber="463">
<typeStatus id="54E124C3FFC2FF840E7DFC60FCC249D3" box="[825,873,950,973]" pageId="37" pageNumber="463">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="4C5AE1E2FFC2FF840EFEFC60FB7249D0" authority="Bleeker, 1862" authorityName="Bleeker" authorityYear="1862" box="[954,1241,950,974]" family="Ariidae" genus="Arius" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="species" species="leptaspis">
<emphasis id="B92E4673FFC2FF840EFEFC60FBEC49D0" box="[954,1095,950,974]" italics="true" pageId="37" pageNumber="463">Arius leptaspis</emphasis>
<bibRefCitation id="EFCBE790FFC2FF840908FC60FB7249D0" author="Bleeker P" box="[1100,1241,950,974]" pageId="37" pageNumber="463" refId="ref29821" refString="Bleeker P. Atlas ichthyologique des Indes Orientales Neerlandaises, publie sous les auspices du Gouvernement colonial neerlandais. Tome II. Siluroides, Chacoides et Heterobranchoides. Amsterdam, 1862." type="book" year="1862">Bleeker, 1862</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C340C9EAFFC2FF840EFDFC38FB7D4FDD" pageId="37" pageNumber="463" type="diagnosis">
<paragraph id="8BE59A61FFC2FF840EFDFC38FAE94E16" blockId="37.[825,1475,1005,1475]" box="[953,1346,1005,1032]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840EFDFC38FAE94E16" box="[953,1346,1005,1032]" italics="true" pageId="37" pageNumber="463">Diagnosis (all of ambiguous optimization)</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC2FF840E7DFBC2FB7D4FDD" blockId="37.[825,1475,1005,1475]" pageId="37" pageNumber="463">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); medial groove of cranium absent (30, 0&gt; 1); temporal fossa very reduced (39, 1&gt; 0); posterior process of exoccipital sutured to Müllerian ramus (93, 0&gt; 1); autopalatine posterior portion conspicuously compressed (106, 0&gt; 1); articulation of autopalatine with lateral ethmoid posteromesially 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); one-half or less of interopercle posterior part contacting ventral margin of opercle (131, 1&gt; 0); sesamoid bone I very long and subtriangular (145, 0&gt; 1); Müllerian ramus bone blade evident only basally (206, 2&gt; 1); Müllerian ramus distal third markedly curved (208, 1&gt; 2).</paragraph>
</subSubSection>
<subSubSection id="C340C9EAFFC2FF84090FFA34FA194C7E" pageId="37" pageNumber="463" type="etymology">
<paragraph id="8BE59A61FFC2FF84090FFA34FB1A4FE2" blockId="37.[1099,1201,1506,1532]" box="[1099,1201,1506,1532]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF84090FFA34FB1A4FE2" box="[1099,1201,1506,1532]" italics="true" pageId="37" pageNumber="463">Etymology</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC2FF840E7DF9DFFA194C7E" blockId="37.[825,1475,1545,1632]" pageId="37" pageNumber="463">Named for the Dutch ichthyologist Pieter Bleeker (18191878) who contributed greatly to our understanding of the taxonomy of ariids from the IndoMalaysian archipelago. Gender: feminine.</paragraph>
</subSubSection>
<subSubSection id="C340C9EAFFC2FF840971F9A9FB814CC2" pageId="37" pageNumber="463" type="taxon_list">
<paragraph id="8BE59A61FFC2FF840971F9A9FB6D4C87" blockId="37.[1077,1222,1663,1689]" box="[1077,1222,1663,1689]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840971F9A9FB6D4C87" box="[1077,1222,1663,1689]" italics="true" pageId="37" pageNumber="463">Included species</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC2FF840E7DF970FB3B4CA0" blockId="37.[825,1168,1702,1757]" box="[825,1168,1702,1726]" pageId="37" pageNumber="463">
<taxonomicName id="4C5AE1E2FFC2FF840E7DF970FB3B4CA0" authority="Bleeker, 1862" authorityName="Bleeker" authorityYear="1862" box="[825,1168,1702,1726]" family="Ariidae" genus="Bleekeriella" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="species" species="leptaspis">
<emphasis id="B92E4673FFC2FF840E7DF970FBAB4CA0" box="[825,1024,1702,1726]" italics="true" pageId="37" pageNumber="463">Bleekeriella leptaspis</emphasis>
<bibRefCitation id="EFCBE790FFC2FF840942F970FB3B4CA0" author="Bleeker P" box="[1030,1168,1702,1726]" pageId="37" pageNumber="463" refId="ref29821" refString="Bleeker P. Atlas ichthyologique des Indes Orientales Neerlandaises, publie sous les auspices du Gouvernement colonial neerlandais. Tome II. Siluroides, Chacoides et Heterobranchoides. Amsterdam, 1862." type="book" year="1862">Bleeker, 1862</bibRefCitation>
</taxonomicName>
</paragraph>
<paragraph id="8BE59A61FFC2FF840E7DF913FB814CC2" blockId="37.[825,1168,1702,1757]" box="[825,1066,1733,1757]" pageId="37" pageNumber="463">
<taxonomicName id="4C5AE1E2FFC2FF840E7DF913FB8F4CC3" authorityName="Bleeker" authorityYear="1862" box="[825,1060,1733,1757]" family="Ariidae" genus="Bleekeriella" isUncertain="true" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="species" species="leptaspis">
<emphasis id="B92E4673FFC2FF840E7DF913FC034CC3" box="[825,936,1733,1757]" italics="true" pageId="37" pageNumber="463">Bleekeriella</emphasis>
aff.
<emphasis id="B92E4673FFC2FF840E95F913FB8F4CC3" box="[977,1060,1733,1757]" italics="true" pageId="37" pageNumber="463">leptaspis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C340C9EAFFC2FF840E7DF928FAC64D28" pageId="37" pageNumber="463" type="distribution">
<paragraph id="8BE59A61FFC2FF840E7DF928FAC64D28" blockId="37.[825,1474,1790,1846]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840E7DF928FBE64D08" box="[825,1101,1790,1814]" italics="true" pageId="37" pageNumber="463">Habitat and distribution:</emphasis>
Predominantly brackish waters, southern New
<collectingCountry id="F34DDAF1FFC2FF840E8BF8C8FBB14D28" box="[975,1050,1822,1846]" name="Guinea-Bissau" pageId="37" pageNumber="463">Guinea</collectingCountry>
and northern
<collectingCountry id="F34DDAF1FFC2FF8409E8F8C8FAAC4D28" box="[1196,1287,1822,1846]" name="Australia" pageId="37" pageNumber="463">Australia</collectingCountry>
(
<figureCitation id="136186E4FFC2FF840853F8C8FAF74D28" box="[1303,1372,1822,1846]" captionStart="Figure 38" captionStartId="38.[117,182,1915,1939]" captionTargetBox="[247,1333,150,1886]" captionTargetId="figure-7@38.[245,1333,144,1888]" captionTargetPageId="38" captionText="Figure 38. Representative skulls, in dorsal view, and distribution of the subtribe Doiichthyina.A, Brustiarius nox, AUM 47488. B, Paracinetodus carinatus, AUM 47550.C, Cochlefelis spatula, AUM 50296. D, Nedystoma dayi, AUM 47500. E, Aceroichthys dioctes, AUM 47507. F, Pachyula crassilabris, AUM 47509. G, Pararius proximus, AUM 47486.H, Potamosilurus velutinus, AUM 47489.* marine, ** brackish waters, *** freshwater." figureDoi="http://doi.org/10.5281/zenodo.11240197" httpUri="https://zenodo.org/record/11240197/files/figure.png" pageId="37" pageNumber="463">Fig. 38</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C340C9EAFFC2FF860917F883FDCC4FD2" lastPageId="39" lastPageNumber="465" pageId="37" pageNumber="463" type="discussion">
<paragraph id="8BE59A61FFC2FF840917F883FB0C4D71" blockId="37.[1107,1191,1877,1903]" box="[1107,1191,1877,1903]" pageId="37" pageNumber="463">
<emphasis id="B92E4673FFC2FF840917F883FB0C4D71" box="[1107,1191,1877,1903]" italics="true" pageId="37" pageNumber="463">Remarks</emphasis>
</paragraph>
<paragraph id="8BE59A61FFC2FF860E7DF8AAFDCC4FD2" blockId="37.[825,1474,1916,1971]" lastBlockId="39.[129,778,1366,1484]" lastPageId="39" lastPageNumber="465" pageId="37" pageNumber="463">
<taxonomicName id="4C5AE1E2FFC2FF840E7DF8AAFC6E4D8A" ID-CoL="GQWD" authorityName="Bleeker" authorityYear="1862" box="[825,965,1916,1940]" family="Ariidae" genus="Arius" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="species" species="leptaspis">
<emphasis id="B92E4673FFC2FF840E7DF8AAFC6E4D8A" box="[825,965,1916,1940]" italics="true" pageId="37" pageNumber="463">Arius leptaspis</emphasis>
</taxonomicName>
was previously included in
<taxonomicName id="4C5AE1E2FFC2FF840991F8AAFA034D8A" ID-CoL="8V54V" authority="Kailola 2004" authorityName="Kailola" authorityYear="2004" box="[1237,1448,1916,1940]" family="Ariidae" genus="Ariopsis" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="genus">
<emphasis id="B92E4673FFC2FF840991F8AAFA8F4D8D" box="[1237,1316,1916,1939]" italics="true" pageId="37" pageNumber="463">Ariopsis</emphasis>
<bibRefCitation id="EFCBE790FFC2FF840863F8AAFA034D8A" author="Kailola PJ" box="[1319,1448,1916,1940]" pageId="37" pageNumber="463" pagination="87 - 166" refId="ref30669" refString="Kailola PJ. A phylogenetic exploration of the catfish family Ariidae (Otophysi: Siluriformes). The Beagle, Records of Museum Art Galleries of North Territory 2004; 20: 87 - 166." type="journal article" year="2004">Kailola 2004</bibRefCitation>
</taxonomicName>
or
<taxonomicName id="4C5AE1E2FFC2FF840E7DF84DFC2B4DAD" ID-CoL="63RGQ" authorityName="Muller &amp; Troschel" authorityYear="1849" box="[825,896,1947,1971]" family="Ariidae" genus="Sciades" kingdom="Animalia" order="Siluriformes" pageId="37" pageNumber="463" phylum="Chordata" rank="genus">
<emphasis id="B92E4673FFC2FF840E7DF84DFC2B4DAD" box="[825,896,1947,1971]" italics="true" pageId="37" pageNumber="463">Sciades</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFCBE790FFC2FF840ED5F84DFB6C4DAD" author="Marceniuk AP &amp; Menezes NA" box="[913,1223,1947,1971]" pageId="37" pageNumber="463" pagination="1 - 126" refId="ref30826" refString="Marceniuk AP, Menezes NA. Systematics of the family Ariidae (Ostariophysi, Siluriformes), with a redefinition of the genera. Zootaxa 2007; 1416: 1 - 126." type="journal article" year="2007">Marceniuk and Menezes 2007</bibRefCitation>
,
<bibRefCitation id="EFCBE790FFC2FF840997F84DFA1A4DAD" author="Marceniuk AP &amp; Menezes NA &amp; Britto MR" box="[1235,1457,1947,1971]" pageId="37" pageNumber="463" 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="B92E4673FFC2FF84080CF84AFAD34DAD" box="[1352,1400,1947,1971]" italics="true" pageId="37" pageNumber="463">et al.</emphasis>
2012
</bibRefCitation>
), but results of the total-evidence analysis indicate that it is not closely related to species of either genus, but is related closely to an unnamed species from
<collectingCountry id="F34DDAF1FFC0FF860CCDFA42FE5A4FB2" box="[393,497,1428,1452]" name="Indonesia" pageId="39" pageNumber="465">Indonesia</collectingCountry>
. These findings corroborate a previous molecular study (Betancur-R. 2009).
</paragraph>
</subSubSection>
</treatment>
</document>