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<document id="738281493E52071462DBB7DB0FA5BF33" ID-CLB-Dataset="306537" ID-DOI="10.1093/zoolinnean/zlae114" ID-GBIF-Dataset="ad4a3e13-83d6-4cee-b8b5-3a2749f9ebfe" ID-ISSN="0024-4082" ID-Zenodo-Dep="14502811" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1734031824042" checkinUser="plazi" docAuthor="Schweiger, Susan, Rödel, Mark-Oliver, Hammel, Jörg. U. &amp; Müller, Hendrik" docDate="2024" docId="7C754916FF8FFFA8FF4BEC02D7A9FA50" docLanguage="en" docName="zlae114.pdf" docOrigin="Zoological Journal of the Linnean Society 202" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Nyctibates corrugatus Boulenger 1904" docType="treatment" docVersion="3" lastPageNumber="9" masterDocId="804C316EFF89FFA1FF89EB56D203FFBE" masterDocTitle="Don’t go with the flow: cranial adaptations of stream tadpoles in the Afrobatrachian family Arthroleptidae" masterLastPageNumber="17" masterPageNumber="1" pageNumber="7" updateTime="1734376722614" updateUser="ExternalLinkService">
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<mods:title id="48FA5BE2C8A21B85F4BEE2E832AFA859">Don’t go with the flow: cranial adaptations of stream tadpoles in the Afrobatrachian family Arthroleptidae</mods:title>
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<mods:namePart id="CB0CB601913CA83959EB211A9BE1C41C">Schweiger, Susan</mods:namePart>
<mods:affiliation id="CEF0D912480BC6AEE7C744D7CB268A1A">Institute of Zoology and Evolutionary Research, Friedrich-Schiller-University Jena, Jena, Germany</mods:affiliation>
<mods:nameIdentifier id="978E1BB4AFA4D9E4BD38B513786AA9BF" type="email">susan.schweiger@uni-jena.de</mods:nameIdentifier>
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<mods:namePart id="745ED6A7324E4A4376B4E33D6857F129">Rödel, Mark-Oliver</mods:namePart>
<mods:affiliation id="8D071E9A37B0EB10A04A3DB769007A89">Museum für Naturkunde - Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany</mods:affiliation>
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<mods:namePart id="5BA48707C9EC220C5A8F3B87C239D445">Hammel, Jörg. U.</mods:namePart>
<mods:affiliation id="E38BD07D90BD1AE4E72E1FEE1CABC2F9">Helmholtz-Zentrum Hereon, Insitute of Materials Physics, Geesthacht, Germany</mods:affiliation>
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<mods:namePart id="1CD94BA5C228FD1B15B1C49006AF86C0">Müller, Hendrik</mods:namePart>
<mods:affiliation id="AF04FDB5D82D30264A072AAED8F8840D">Zentralmagazin Naturwissenschaftlicher Sammlungen, Martin-Luther-Universität Halle-Wittenberg, Halle (Saale), Germany</mods:affiliation>
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<mods:title id="9B48D254A5CFD9F0543A06768842566D">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="62CD9B81EC4FF8A97D9D8CE751CF4C0F">2024</mods:date>
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<treatment id="7C754916FF8FFFA8FF4BEC02D7A9FA50" ID-DOI="http://doi.org/10.5281/zenodo.14502817" ID-Zenodo-Dep="14502817" LSID="urn:lsid:plazi:treatment:7C754916FF8FFFA8FF4BEC02D7A9FA50" httpUri="http://treatment.plazi.org/id/7C754916FF8FFFA8FF4BEC02D7A9FA50" lastPageId="9" lastPageNumber="9" pageId="6" pageNumber="7" scope_family="Arthroleptidae">
<subSubSection id="BCC6AB8BFF8FFFA7FF4BEC02D0CBF8D1" box="[194,712,1876,1903]" pageId="6" pageNumber="7" type="nomenclature">
<paragraph id="F463F800FF8FFFA7FF4BEC02D0CBF8D1" blockId="6.[129,778,1876,1971]" box="[194,712,1876,1903]" pageId="6" pageNumber="7">
<emphasis id="C6A82412FF8FFFA7FF4BEC02D0CBF8D1" bold="true" box="[194,712,1876,1903]" pageId="6" pageNumber="7">
<taxonomicName id="33DC8383FF8FFFA7FF4BEC02D399F8D1" ID-CoL="485FQ" authorityName="Boulenger" authorityYear="1904" box="[194,410,1876,1903]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8FFFA7FF4BEC02D399F8D1" bold="true" box="[194,410,1876,1903]" italics="true" pageId="6" pageNumber="7">Nyctibates corrugatus</emphasis>
</taxonomicName>
and
<taxonomicName id="33DC8383FF8FFFA7FE42EC02D0CBF8D1" authority="Boulenger, 1900" authorityName="Boulenger" authorityYear="1900" box="[459,712,1876,1903]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8FFFA7FE42EC02D0CBF8D1" bold="true" box="[459,712,1876,1903]" italics="true" pageId="6" pageNumber="7">Trichobatrachus robustus</emphasis>
</taxonomicName>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="BCC6AB8BFF8FFFA8FF08EC2DD7A9FA50" lastPageId="9" lastPageNumber="10" pageId="6" pageNumber="7" type="discussion">
<paragraph id="F463F800FF8FFFA7FF08EC2DD62CF906" blockId="6.[129,778,1876,1971]" lastBlockId="6.[825,1476,1539,1971]" pageId="6" pageNumber="7">
Both taxa had more muscular tails than the previously described two taxa but were otherwise similar to them in having low dorsal and ventral fins (Fig. 1). The oral disc of
<taxonomicName id="33DC8383FF8FFFA7FB50ED52D75FF9A5" authorityName="Boulenger" authorityYear="1904" box="[1241,1372,1540,1563]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8FFFA7FB50ED52D75FF9A5" box="[1241,1372,1540,1563]" italics="true" pageId="6" pageNumber="7">N. corrugatus</emphasis>
</taxonomicName>
consisted of semicircular, wide, and flattened lobes that were surrounded by marginal papillae (Fig. 3C, also see
<bibRefCitation id="904D85F1FF8FFFA7FB40ED14D7C1F9E4" author="Griesbaum F &amp; Hirschfeld M &amp; Barej MF" box="[1225,1474,1602,1626]" pageId="6" pageNumber="7" pagination="133 - 60" refId="ref9642" refString="Griesbaum F, Hirschfeld M, Barej MF et al. Tadpoles of three western African frog genera: Astylosternus Werner, 1898, Nyctibates Boulenger, 1904, and Scotobleps Boulenger, 1900 (Amphibia, Anura, Arthroleptidae). Zoosystematics and Evolution 2019; 95: 133 - 60. https: // doi. org / 10. 3897 / zse. 95. 32793" type="journal article" year="2019">
Griesbaum
<emphasis id="C6A82412FF8FFFA7FACBED15D771F9E4" box="[1346,1394,1602,1626]" italics="true" pageId="6" pageNumber="7">et al.</emphasis>
(2019)
</bibRefCitation>
for a detailed description). The oral apparatus of
<taxonomicName id="33DC8383FF8FFFA7FAA2ED34D794F9C7" box="[1323,1431,1633,1657]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8FFFA7FAA2ED34D794F9C7" box="[1323,1431,1633,1657]" italics="true" pageId="6" pageNumber="7">T. robustus</emphasis>
</taxonomicName>
was highly modified into a large oral sucker with multiple rows of small papillae (Fig. 3D).
</paragraph>
<paragraph id="F463F800FF8FFFA9FCDCED96D03DF87B" blockId="6.[825,1476,1539,1971]" lastBlockId="8.[129,779,1683,1989]" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7">
In
<taxonomicName id="33DC8383FF8FFFA7FCFAED96D1FAF969" authorityName="Boulenger" authorityYear="1904" box="[883,1017,1728,1751]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8FFFA7FCFAED96D1FAF969" box="[883,1017,1728,1751]" italics="true" pageId="6" pageNumber="7">N. corrugatus</emphasis>
</taxonomicName>
, the cornua trabeculae were fused together and formed a broad, trapezoidal plate, which was very different from the other three tadpoles examined (Fig. 4C). Laterally, the cornua trabeculae articulated with the cartilago labialis superior. In
<taxonomicName id="33DC8383FF8FFFA7FC17EC6BD60AF8EB" box="[926,1033,1853,1877]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8FFFA7FC17EC6BD60AF8EB" box="[926,1033,1853,1877]" italics="true" pageId="6" pageNumber="7">T. robustus</emphasis>
</taxonomicName>
, the cornua trabeculae were also fused, but each trabecular horn curved ventrad at its anterior end (Fig. 4D). The processus muscularis of the palatoquadrate was much wider in
<taxonomicName id="33DC8383FF8FFFA7FC18ECCDD1F8F80D" box="[913,1019,1947,1971]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8FFFA7FC18ECCDD1F8F80D" box="[913,1019,1947,1971]" italics="true" pageId="6" pageNumber="7">T. robustus</emphasis>
</taxonomicName>
compared to
<taxonomicName id="33DC8383FF8FFFA7FB0AECCDD706F80C" authorityName="Boulenger" authorityYear="1904" box="[1155,1285,1947,1970]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8FFFA7FB0AECCDD706F80C" box="[1155,1285,1947,1970]" italics="true" pageId="6" pageNumber="7">N. corrugatus</emphasis>
</taxonomicName>
(Fig. 5C, D). In
<taxonomicName id="33DC8383FF8FFFA6FA22ECCDD2D4F97F" authorityName="Boulenger" authorityYear="1904" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" lastPageId="7" lastPageNumber="8" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8FFFA6FA22ECCDD2D4F97F" italics="true" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">N. corrugatus</emphasis>
</taxonomicName>
, the commissura quadratoorbitalis formed a curved, cartilaginous strip that connected the processus muscularis of the palatoquadrate to the trabecula cranii, whereas it was broad and massive in
<taxonomicName id="33DC8383FF8EFFA6FE86EC5ED378F89E" box="[271,379,1800,1824]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8EFFA6FE86EC5ED378F89E" box="[271,379,1800,1824]" italics="true" pageId="7" pageNumber="8">T. robustus</emphasis>
</taxonomicName>
(Fig. 5C, D). In
<taxonomicName id="33DC8383FF8EFFA6FDA3EC5ED0ACF8A1" authorityName="Boulenger" authorityYear="1904" box="[554,687,1800,1823]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8EFFA6FDA3EC5ED0ACF8A1" box="[554,687,1800,1823]" italics="true" pageId="7" pageNumber="8">N. corrugatus</emphasis>
</taxonomicName>
, a long processus hyoquadratis extended from the medial part of the palatoquadrate (Fig. 5C). The processus hyoquadratis articulated widely with the hypobranchial apparatus via the lateral part of the ceratohyale. There was a wide gap between this process and the processus muscularis. The ceratohyal fitted into this gap, articulating with the palatoquadrate. This structure was not seen in any of the three other tadpoles examined in this study. In
<taxonomicName id="33DC8383FF8EFFA6FA12EDBFD193F8A1" authorityName="Boulenger" authorityYear="1904" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8EFFA6FA12EDBFD193F8A1" italics="true" pageId="7" pageNumber="8">N. corrugatus</emphasis>
</taxonomicName>
and
<taxonomicName id="33DC8383FF8EFFA6FC42EC5ED638F89E" box="[971,1083,1800,1824]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8EFFA6FC42EC5ED638F89E" box="[971,1083,1800,1824]" italics="true" pageId="7" pageNumber="8">T. robustus</emphasis>
</taxonomicName>
, the palatoquadrate was connected anteroventrally to the neurocranium by a broad commissura quadratocranialis (Fig. 6C, D). Laterally, the palatoquadrate of
<taxonomicName id="33DC8383FF8EFFA6FA16EC11D17FF8C0" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8EFFA6FA16EC11D17FF8C0" italics="true" pageId="7" pageNumber="8">T. robustus</emphasis>
</taxonomicName>
bore an extremely elongated processus ventralis, which was much shorter in
<taxonomicName id="33DC8383FF8EFFA6FC72ECD0D67EF823" authorityName="Boulenger" authorityYear="1904" box="[1019,1149,1926,1949]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8EFFA6FC72ECD0D67EF823" box="[1019,1149,1926,1949]" italics="true" pageId="7" pageNumber="8">N. corrugatus</emphasis>
</taxonomicName>
(Fig. 5C, D). In both tadpoles, the cartilago labialis superior formed a massive, single element of the upper jaw (Fig. 6C, D). Compared to
<taxonomicName id="33DC8383FF81FFA9FDEDEDE5D0E8F974" authorityName="Boulenger" authorityYear="1904" box="[612,747,1715,1738]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FDEDEDE5D0E8F974" box="[612,747,1715,1738]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
, it appeared much shorter and more compressed in
<taxonomicName id="33DC8383FF81FFA9FD1DED85D107F954" box="[660,772,1746,1770]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FD1DED85D107F954" box="[660,772,1746,1770]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
. The cartilago labialis inferior consisted of two separated elements; it was of a more triangular shape in
<taxonomicName id="33DC8383FF81FFA9FDA7EC47D098F897" box="[558,667,1809,1833]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FDA7EC47D098F897" box="[558,667,1809,1833]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
(Fig. 6D). The cartilago meckeli was transversally oriented and L-shaped in frontal view in
<taxonomicName id="33DC8383FF81FFA9FECDEC06D3B0F8D9" box="[324,435,1871,1895]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FECDEC06D3B0F8D9" box="[324,435,1871,1895]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
, while it appeared thicker in
<taxonomicName id="33DC8383FF81FFA9FD7BEC06D2E4F838" authorityName="Boulenger" authorityYear="1904" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FD7BEC06D2E4F838" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
(Fig. 6C, D). For a detailed description of the complete cranial skeleton see Supporting Information (Fig. S3 for
<taxonomicName id="33DC8383FF81FFA9FD7BECD9D2E4F87B" authorityName="Boulenger" authorityYear="1904" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FD7BECD9D2E4F87B" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
, Fig. S4 for
<taxonomicName id="33DC8383FF81FFA9FED6ECF8D3C9F87B" box="[351,458,1965,1989]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FED6ECF8D3C9F87B" box="[351,458,1965,1989]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
;
<tableCitation id="B95ECDBBFF81FFA9FE5FECFBD02DF87B" box="[470,558,1965,1989]" captionStart="Table 2" captionStartId="10.[112,167,612,637]" captionTargetId="graphics-12@10.[112,1943,644,1539]" captionTargetPageId="10" captionText="Table 2. Comparison of cranial adaptations in the chondrocranium of different tadpole species in relation to their habitat (see Table 3)" httpUri="http://table.plazi.org/id/A0A3A888FF83FFABFFF9E932D75CFDC3" pageId="8" pageNumber="9" tableUuid="A0A3A888FF83FFABFFF9E932D75CFDC3">Table S2</tableCitation>
).
</paragraph>
<caption id="A0A3A888FF8EFFA6FFF8EEA0D079F9DC" ID-DOI="http://doi.org/10.5281/zenodo.14502827" ID-Zenodo-Dep="14502827" httpUri="https://zenodo.org/record/14502827/files/figure.png" pageId="7" pageNumber="8" startId="7.[113,178,1526,1550]" targetBox="[115,1456,145,1496]" targetPageId="7" targetType="figure">
<paragraph id="F463F800FF8EFFA6FFF8EEA0D079F9DC" blockId="7.[113,1454,1526,1634]" pageId="7" pageNumber="8">
<emphasis id="C6A82412FF8EFFA6FFF8EEA0D2CBF9B0" bold="true" box="[113,200,1526,1550]" pageId="7" pageNumber="8">Figure 6.</emphasis>
3D reconstruction of the chondrocranium, ventral view. A, slow-flowing stream tadpole of
<taxonomicName id="33DC8383FF8EFFA6FBA9EEA0D6C2F9B0" box="[1056,1217,1526,1550]" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="C6A82412FF8EFFA6FBA9EEA0D6C2F9B0" box="[1056,1217,1526,1550]" italics="true" pageId="7" pageNumber="8">Leptopelis parkeri</emphasis>
</taxonomicName>
. B, moderate-flowing stream tadpole of
<taxonomicName id="33DC8383FF8EFFA6FEECED44D04AF994" box="[357,585,1554,1578]" class="Amphibia" family="Arthroleptidae" genus="Astylosternus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="occidentalis">
<emphasis id="C6A82412FF8EFFA6FEECED44D04AF994" box="[357,585,1554,1578]" italics="true" pageId="7" pageNumber="8">Astylosternus occidentalis</emphasis>
</taxonomicName>
. C, fast-flowing stream tadpole of
<taxonomicName id="33DC8383FF8EFFA6FC00ED44D648F994" authorityName="Boulenger" authorityYear="1904" box="[905,1099,1554,1578]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8EFFA6FC00ED44D648F994" box="[905,1099,1554,1578]" italics="true" pageId="7" pageNumber="8">Nyctibates corrugatus</emphasis>
</taxonomicName>
. D, fast-flowing stream tadpole of
<taxonomicName id="33DC8383FF8EFFA6FFF8ED78D356F9F8" box="[113,341,1582,1606]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8EFFA6FFF8ED78D356F9F8" box="[113,341,1582,1606]" italics="true" pageId="7" pageNumber="8">Trichobatrachus robustus</emphasis>
</taxonomicName>
. The cartilago labialis superior is fused and the ceratohyalia are enlarged in
<taxonomicName id="33DC8383FF8EFFA6FB9CED78D695F9F8" box="[1045,1174,1582,1606]" class="Amphibia" family="Arthroleptidae" genus="Astylosternus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="occidentalis">
<emphasis id="C6A82412FF8EFFA6FB9CED78D695F9F8" box="[1045,1174,1582,1606]" italics="true" pageId="7" pageNumber="8">A. occidentalis</emphasis>
</taxonomicName>
,
<taxonomicName id="33DC8383FF8EFFA6FB16ED79D71AF9F8" authorityName="Boulenger" authorityYear="1904" box="[1183,1305,1583,1606]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF8EFFA6FB16ED79D71AF9F8" box="[1183,1305,1583,1606]" italics="true" pageId="7" pageNumber="8">N. corrugatus</emphasis>
</taxonomicName>
, and
<taxonomicName id="33DC8383FF8EFFA6FAC2ED79D7ADF9F8" box="[1355,1454,1582,1606]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF8EFFA6FAC2ED79D7ADF9F8" box="[1355,1454,1582,1606]" italics="true" pageId="7" pageNumber="8">T. robustus</emphasis>
</taxonomicName>
compared to the more generalized tadpole of
<taxonomicName id="33DC8383FF8EFFA6FD95ED1DD076F9DC" box="[540,629,1610,1634]" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="C6A82412FF8EFFA6FD95ED1DD076F9DC" box="[540,629,1610,1634]" italics="true" pageId="7" pageNumber="8">L. parkeri</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<caption id="A0A3A888FF81FFA9FF08EEA1D00EF9DA" ID-DOI="http://doi.org/10.5281/zenodo.14502829" ID-Zenodo-Dep="14502829" httpUri="https://zenodo.org/record/14502829/files/figure.png" pageId="8" pageNumber="9" startId="8.[129,194,1527,1551]" targetBox="[130,1471,145,1497]" targetPageId="8" targetType="figure">
<paragraph id="F463F800FF81FFA9FF08EEA1D00EF9DA" blockId="8.[129,1463,1527,1636]" pageId="8" pageNumber="9">
<emphasis id="C6A82412FF81FFA9FF08EEA1D2DBF9AE" bold="true" box="[129,216,1527,1552]" pageId="8" pageNumber="9">Figure 7.</emphasis>
3D reconstruction of the hyobranchial skeleton. A, slow-flowing stream tadpole of
<taxonomicName id="33DC8383FF81FFA9FC6CEEA1D686F9B1" box="[997,1157,1527,1551]" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="C6A82412FF81FFA9FC6CEEA1D686F9B1" box="[997,1157,1527,1551]" italics="true" pageId="8" pageNumber="9">Leptopelis parkeri</emphasis>
</taxonomicName>
. B, moderate-flowing stream tadpole of
<taxonomicName id="33DC8383FF81FFA9FF6CED45D3CBF995" box="[229,456,1555,1579]" class="Amphibia" family="Arthroleptidae" genus="Astylosternus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="occidentalis">
<emphasis id="C6A82412FF81FFA9FF6CED45D3CBF995" box="[229,456,1555,1579]" italics="true" pageId="8" pageNumber="9">Astylosternus occidentalis</emphasis>
</taxonomicName>
. C, fast-flowing stream tadpole of
<taxonomicName id="33DC8383FF81FFA9FC81ED45D1C9F995" authorityName="Boulenger" authorityYear="1904" box="[776,970,1555,1579]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FC81ED45D1C9F995" box="[776,970,1555,1579]" italics="true" pageId="8" pageNumber="9">Nyctibates corrugatus</emphasis>
</taxonomicName>
. D, fast-flowing stream tadpole of
<taxonomicName id="33DC8383FF81FFA9FA82ED45D2CEF9F9" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FA82ED45D2CEF9F9" italics="true" pageId="8" pageNumber="9">Trichobatrachus robustus</emphasis>
</taxonomicName>
. Note the large ceratohyalia and the comparatively small branchial baskets in
<taxonomicName id="33DC8383FF81FFA9FC2BED66D621F9F9" box="[930,1058,1583,1607]" class="Amphibia" family="Arthroleptidae" genus="Astylosternus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="occidentalis">
<emphasis id="C6A82412FF81FFA9FC2BED66D621F9F9" box="[930,1058,1583,1607]" italics="true" pageId="8" pageNumber="9">A. occidentalis</emphasis>
</taxonomicName>
,
<taxonomicName id="33DC8383FF81FFA9FBA5ED66D6A5F9F9" authorityName="Boulenger" authorityYear="1904" box="[1068,1190,1584,1607]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FBA5ED66D6A5F9F9" box="[1068,1190,1584,1607]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
, and
<taxonomicName id="33DC8383FF81FFA9FB51ED66D738F9F9" box="[1240,1339,1583,1607]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FB51ED66D738F9F9" box="[1240,1339,1583,1607]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
compared to the more generalized tadpole of
<taxonomicName id="33DC8383FF81FFA9FE26ED1AD00BF9DD" box="[431,520,1611,1635]" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="parkeri">
<emphasis id="C6A82412FF81FFA9FE26ED1AD00BF9DD" box="[431,520,1611,1635]" italics="true" pageId="8" pageNumber="9">L. parkeri</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="F463F800FF81FFA9FCDCEDC5D79BF839" blockId="8.[825,1475,1683,1989]" pageId="8" pageNumber="9">
The hyobranchial skeleton in
<taxonomicName id="33DC8383FF81FFA9FBFDEDC2D6D9F915" box="[1140,1242,1683,1707]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FBFDEDC2D6D9F915" box="[1140,1242,1683,1707]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
was transversally orientated.Incontrastitwasmoreobliquein
<taxonomicName id="33DC8383FF81FFA9FB20EDE5D720F974" authorityName="Boulenger" authorityYear="1904" box="[1193,1315,1715,1738]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FB20EDE5D720F974" box="[1193,1315,1715,1738]" italics="true" pageId="8" pageNumber="9">N.corrugatus</emphasis>
</taxonomicName>
.Theceratohyale of
<taxonomicName id="33DC8383FF81FFA9FCDAED85D1D2F954" authorityName="Boulenger" authorityYear="1904" box="[851,977,1747,1770]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FCDAED85D1D2F954" box="[851,977,1747,1770]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
and
<taxonomicName id="33DC8383FF81FFA9FB88ED85D664F954" box="[1025,1127,1746,1770]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FB88ED85D664F954" box="[1025,1127,1746,1770]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
were even more massive compared to
<taxonomicName id="33DC8383FF81FFA9FCDDEDA4D1D9F8B7" box="[852,986,1777,1801]" class="Amphibia" family="Arthroleptidae" genus="Astylosternus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="occidentalis">
<emphasis id="C6A82412FF81FFA9FCDDEDA4D1D9F8B7" box="[852,986,1777,1801]" italics="true" pageId="8" pageNumber="9">A. occidentalis</emphasis>
</taxonomicName>
(Fig. 7). The ceratohyale of
<taxonomicName id="33DC8383FF81FFA9FB7CEDA4D75FF8B7" box="[1269,1372,1777,1801]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FB7CEDA4D75FF8B7" box="[1269,1372,1777,1801]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
was more stout and transversely oriented in comparison to
<taxonomicName id="33DC8383FF81FFA9FA9BEC47D793F896" authorityName="Boulenger" authorityYear="1904" box="[1298,1424,1809,1832]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FA9BEC47D793F896" box="[1298,1424,1809,1832]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
(Fig. 7C, D). A basihyale was only present in
<taxonomicName id="33DC8383FF81FFA9FB5FEC67D754F8F6" authorityName="Boulenger" authorityYear="1904" box="[1238,1367,1841,1864]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FB5FEC67D754F8F6" box="[1238,1367,1841,1864]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
(Fig. 7C).
<taxonomicName id="33DC8383FF81FFA9FCB0EC19D602F8D9" authorityName="Boulenger" authorityYear="1904" box="[825,1025,1871,1895]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FCB0EC19D602F8D9" box="[825,1025,1871,1895]" italics="true" pageId="8" pageNumber="9">Nyctibates corrugatus</emphasis>
</taxonomicName>
had four separated ceratobranchialia, whereas ceratobranchialia I and II were fused in
<taxonomicName id="33DC8383FF81FFA9FB3AEC39D71BF838" box="[1203,1304,1902,1926]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FB3AEC39D71BF838" box="[1203,1304,1902,1926]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
(Fig. 7C, D).
</paragraph>
<paragraph id="F463F800FF81FFA8FCDCECD8D7A9FA50" blockId="8.[825,1475,1683,1989]" lastBlockId="9.[810,1462,1462,1518]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
The m. orbitohyoideus in
<taxonomicName id="33DC8383FF81FFA9FBECECD9D6E8F818" authorityName="Boulenger" authorityYear="1904" box="[1125,1259,1935,1958]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FBECECD9D6E8F818" box="[1125,1259,1935,1958]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
was more elongated in comparison to
<taxonomicName id="33DC8383FF81FFA9FC7FECF8D666F87B" box="[1014,1125,1965,1989]" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF81FFA9FC7FECF8D666F87B" box="[1014,1125,1965,1989]" italics="true" pageId="8" pageNumber="9">T. robustus</emphasis>
</taxonomicName>
. In
<taxonomicName id="33DC8383FF81FFA9FB18ECF8D71BF87B" authorityName="Boulenger" authorityYear="1904" box="[1169,1304,1966,1989]" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF81FFA9FB18ECF8D71BF87B" box="[1169,1304,1966,1989]" italics="true" pageId="8" pageNumber="9">N. corrugatus</emphasis>
</taxonomicName>
, it originated at the lateral part of the processus muscularis and the processus ventralis on the palatoquadrate and inserted along the lateral part of the ceratohyale and on the processus hyoquadratis (Fig. 8C). For a complete description of the cranial musculature of
<taxonomicName id="33DC8383FF80FFA8FD6AED43D2D4F9F5" authorityName="Boulenger" authorityYear="1904" class="Amphibia" family="Arthroleptidae" genus="Nyctibates" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="corrugatus">
<emphasis id="C6A82412FF80FFA8FD6AED43D2D4F9F5" italics="true" pageId="9" pageNumber="10">N. corrugatus</emphasis>
</taxonomicName>
see Supporting Information (Fig. S7;
<tableCitation id="B95ECDBBFF80FFA8FDECED62D0B1F9F2" box="[613,690,1588,1612]" captionStart="Table 2" captionStartId="10.[112,167,612,637]" captionTargetId="graphics-12@10.[112,1943,644,1539]" captionTargetPageId="10" captionText="Table 2. Comparison of cranial adaptations in the chondrocranium of different tadpole species in relation to their habitat (see Table 3)" httpUri="http://table.plazi.org/id/A0A3A888FF83FFABFFF9E932D75CFDC3" pageId="9" pageNumber="10" tableUuid="A0A3A888FF83FFABFFF9E932D75CFDC3">Table 2</tableCitation>
). In
<taxonomicName id="33DC8383FF80FFA8FD6EED62D2C0F9D5" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF80FFA8FD6EED62D2C0F9D5" italics="true" pageId="9" pageNumber="10">T. robustus</emphasis>
</taxonomicName>
, the m. orbitohyoideus was very thick and consisted of three layers of muscle, a feature which we did not observe in any of the three other tadpoles (
<figureCitation id="6CE7E485FF80FFA8FE02EDC4D3DAF914" box="[395,473,1682,1706]" captionStart="Figure 8" captionStartId="9.[114,179,1278,1302]" captionTargetBox="[115,1456,145,1249]" captionTargetId="figure-315@9.[114,1458,144,1251]" captionTargetPageId="9" captionText="Figure 8. 3D reconstruction of the chondrocranium and the m. orbitohyoideus, lateral view. A, slow-flowing stream tadpole of Leptopelis parkeri. B, moderate-flowing stream tadpole of Astylosternus occidentalis. C, fast-flowing stream tadpole of Nyctibates corrugatus. D, fast-flowing stream tadpole of Trichobatrachus robustus. The processus ventralis functions as an extended origin and therefore expands the surface of the large m. orbitohyoideus in A. occidentalis, N.corrugatus, and T.robustus. Note that the m. orbitohyoideus has separate deep and superficial muscle layers in T. robustus." figureDoi="http://doi.org/10.5281/zenodo.14502831" httpUri="https://zenodo.org/record/14502831/files/figure.png" pageId="9" pageNumber="10">Figs 8D</figureCitation>
,
<figureCitation id="6CE7E485FF80FFA8FE6BEDC4D3ECF914" box="[482,495,1682,1706]" captionStart="Figure 9" captionStartId="11.[113,178,1294,1318]" captionTargetBox="[145,1426,146,1264]" captionTargetId="figure-362@11.[144,1428,144,1266]" captionTargetPageId="11" captionText="Figure 9. 3D reconstruction of the chondrocranium and associated musculature of Trichobatrachus robustus. Left, dorsal view. Right, lateral view. Shown separately are the three layers of the m. orbitohyoideus: (1) superficial layer, (2) medial layer, (3) deeper layer." figureDoi="http://doi.org/10.5281/zenodo.14502833" httpUri="https://zenodo.org/record/14502833/files/figure.png" pageId="9" pageNumber="10">9</figureCitation>
). The fibres of the deepest layer originated dorsolaterally on the processus muscularis and the inner side of the processus ventralis and inserted at the anterior region of the ceratohyale (Fig. 9). The middle layer originated anterolaterally on the processus muscularis and along the processus ventralis and inserted medioventrally and mediodorsally on the ceratohyale. The outer layer originated dorsolaterally on the processus muscularis and also on along the processus ventralis, and inserted distally on the ceratohyale (Fig. 9). For a complete description of the cranial musculature of
<taxonomicName id="33DC8383FF80FFA8FA16EEE1D17FFA50" class="Amphibia" family="Arthroleptidae" genus="Trichobatrachus" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="robustus">
<emphasis id="C6A82412FF80FFA8FA16EEE1D17FFA50" italics="true" pageId="9" pageNumber="10">T. robustus</emphasis>
</taxonomicName>
see Supporting Information (File S1, Fig. S8;
<tableCitation id="B95ECDBBFF80FFA8FAC7EE80D79AFA50" box="[1358,1433,1494,1518]" captionStart="Table 2" captionStartId="10.[112,167,612,637]" captionTargetId="graphics-12@10.[112,1943,644,1539]" captionTargetPageId="10" captionText="Table 2. Comparison of cranial adaptations in the chondrocranium of different tadpole species in relation to their habitat (see Table 3)" httpUri="http://table.plazi.org/id/A0A3A888FF83FFABFFF9E932D75CFDC3" pageId="9" pageNumber="10" tableUuid="A0A3A888FF83FFABFFF9E932D75CFDC3">Table 2</tableCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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