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<document id="49D2CA9E553FEBAE744B67DE685EF377" ID-CLB-Dataset="306267" ID-DOI="10.3897/zookeys.1220.123541" ID-GBIF-Dataset="d9472aa9-c5c2-482b-bbbd-6cb337e2e9ca" ID-ZooBank="D8BC336A-6DA7-4819-9217-536078DA6755" ID-publisher-id="123541" URI-arpha="CBDCB3CC-CBED-5D3C-971A-BA79CB338801" URI-zoobank="http://zoobank.org/D8BC336A-6DA7-4819-9217-536078DA6755" XM.bibliography_approvedBy="admin" XM.materialsCitations_approvedBy="admin" XM.treatmentCitations_approvedBy="admin" article-type="research-article" checkinTime="1733757363100" checkinUser="pensoft" docAuthor="Su, Yo &amp; Ho, Hsuan-Ching" docDate="2024" docId="13BF485C3ACE583CB90203D6804ACF4B" docLanguage="en" docName="ZooKeys 1220: 165-174" docOrigin="ZooKeys 1220" docSource="https://zookeys.pensoft.net/article/123541/download/xml/" docStyle="DocumentStyle:PensoftTaxPub.0000.journal_article.generic" docStyleName="PensoftTaxPub.0000.journal_article.generic" docTitle="Opistognathus cryos Su &amp; Ho, 2024, sp. nov." docType="treatment" docVersion="7" dtd-version="3.0" lastPageNumber="174" masterDocId="CBDCB3CCCBED5D3C971ABA79CB338801" masterDocTitle="A new species of the jawfish genus Opistognathus from Taiwan, northwestern Pacific Ocean (Perciformes, Opistognathidae)" masterLastPageNumber="174" masterPageNumber="165" pageNumber="165" updateTime="1733779277351" updateUser="ExternalLinkService">
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<mods:title id="F6EB996867DD00F7D8CC6C12A5508A38">A new species of the jawfish genus Opistognathus from Taiwan, northwestern Pacific Ocean (Perciformes, Opistognathidae)</mods:title>
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<mods:namePart id="7349CC23F3A11F3995D117B7F360A62F">Su, Yo</mods:namePart>
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<mods:affiliation id="075E9861FA8C93DADB22592292E15281">Department and Graduate Institute of Aquaculture, National Kaohsiung University of Science Technology, Kaohsiung, Taiwan</mods:affiliation>
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<mods:namePart id="8609E571916F7B3B607BDB834BB36D28">Ho, Hsuan-Ching</mods:namePart>
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<mods:affiliation id="676B4264F272B65772C8AE2D6DDCA958">Department and Graduate Institute of Aquaculture, National Kaohsiung University of Science Technology, Kaohsiung, Taiwan &amp; Institute of Marine Biology, National Dong Hwa University, Pingtung, Taiwan &amp; Australian Museum, Sydney, Australia</mods:affiliation>
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<mods:date id="1892BF9E8E8090ECCB27F4D564E54AC1">2024</mods:date>
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<mods:number id="019C5FA6F8308B9AD4702DB46B6F8E8A">2024-12-09</mods:number>
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<mods:number id="2A5B5C05E5FF3038A7346751BF8D2986">1220</mods:number>
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<treatment id="13BF485C3ACE583CB90203D6804ACF4B" ID-DOI="http://doi.org/10.5281/zenodo.14343485" ID-GBIF-Taxon="248824518" ID-Zenodo-Dep="14343485" ID-arpha="13BF485C-3ACE-583C-B902-03D6804ACF4B" ID-zoobank="https://zoobank.org/8F808D49-8B17-4E9C-A4B8-E273BAC2742C" LSID="urn:lsid:plazi:treatment:13BF485C3ACE583CB90203D6804ACF4B" LSID-ZBK="urn:lsid:zoobank.org:act:8F808D49-8B17-4E9C-A4B8-E273BAC2742C" httpUri="http://treatment.plazi.org/id/13BF485C3ACE583CB90203D6804ACF4B" scope_family="Opistognathidae" scope_order="Perciformes">
<subSubSection id="1CF6CFDB8F18E57C48005CB3EE97C817" type="nomenclature">
<paragraph id="2C09C2CE1545C025B9A63812F71FCC5B">
<taxonomicName id="30337A36DDD262F31C2FDFA0CA18E0A3" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos" status="sp. nov.">
<emphasis id="1A6501C7B0E7940825824A3E7DEA8ED7" italics="true">Opistognathus cryos</emphasis>
</taxonomicName>
<taxonomicNameLabel id="8562A94860B76D73C24F82679DD512FF" rank="species">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection id="9C3592329C7A0950FEDA6A9D9BC5DB9A" type="description">
<paragraph id="CE66A4D3AA90E15EFD0D5DE42A652640">
<figureCitation id="D29DDABD3BD8D079E9B7A9F3236603CA" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Preserved specimen of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL A on black background B on white background C dorsal view of nape. Photos by Y. - C. Hsu." figureDoi="10.3897/zookeys.1220.123541.figure1" httpUri="https://binary.pensoft.net/fig/1192524">Figs 1</figureCitation>
,
<figureCitation id="9AF0D8BB584FBA8568DE89C294161C7A" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Head pores and adjacent area of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL, with scaled area shaded in gray A dorsal view B lateral view. Abbreviations: AN = anterior nostril; PN = posterior nostril. Dotted lines indicate dorsal- and pectoral-fin bases. Anterior to left. Figure not to scale." figureDoi="10.3897/zookeys.1220.123541.figure2" httpUri="https://binary.pensoft.net/fig/1192525">2</figureCitation>
,
<figureCitation id="5D23D9F5B48EE186931F3A10D57B7945" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. X-radiograph of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL." figureDoi="10.3897/zookeys.1220.123541.figure3" httpUri="https://binary.pensoft.net/fig/1192526">3</figureCitation>
,
<tableCitation id="FBD5E6CB24A0A724DD6E9AC3BA99EA62" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Meristic characters of Opistognathus cryos sp. nov. and similar species. Paired characters are presented as left / right whenever available. Data of other species were retrieved from Smith-Vaniz (2023)." httpUri="http://table.plazi.org/id/35CB17D5C0691D99A665ABB9F92676F6" tableUuid="35CB17D5C0691D99A665ABB9F92676F6">Tables 1</tableCitation>
,
<tableCitation id="18EDB5EB170B8CF1C67B03FC58A7B9DE" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Morphometric characters of Opistognathus cryos sp. nov. and similar species. Data of other species were retrieved from Smith-Vaniz (2023). Abbreviations: A = anal-fin; C = caudal-fin; D = dorsal-fin; HL = head length; SL = standard length; V = pelvic-fin." httpUri="http://table.plazi.org/id/48A203DFB18FA58BE55BDBF4397F5ECB" tableUuid="48A203DFB18FA58BE55BDBF4397F5ECB">2 New English name: Frozen jawfish New Chinese name: 冷鋒後頷鱚</tableCitation>
</paragraph>
</subSubSection>
<subSubSection id="5C8588C161EFD3B6C51171AB80DEF8B9" type="reference_group">
<paragraph id="9A0449D7279C5AEBC93B33163BC7C897">
<treatmentCitationGroup id="323F44A2F09C76CA99140D8B32E25130">
<taxonomicName id="D96602BC1E4B290C69AB8E50F5FB2345" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="undefined-1">
<emphasis id="AD9A79A58D50E18CC86A78557F39132A" italics="true">Opistognathus</emphasis>
<comment id="8AF7FC0F770DC18D3C60ED81C93E6824">
sp. 1:
<treatmentCitation id="57F119F6D579E70EF6FF6D61419A4485" author="Smith-Vaniz WF" page="72" year="2023">
<bibRefCitation id="E51B14FC16ED373DF36E6800174F0CA3" DOI="10.11646/zootaxa.5252.1.1" author="Smith-Vaniz WF" firstAuthor="Smith-Vaniz" issue="1" journalOrPublisher="Zootaxa" page="72" pagination="1-180" refId="B21" refString="Smith-Vaniz WF (2023) Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species. Zootaxa 5252 (1): 1180. https://doi.org/10.11646/zootaxa.5252.1.1" title="Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species." volume="5252" year="2023">Smith-Vaniz 2023: 72</bibRefCitation>
</treatmentCitation>
, fig. 103.
</comment>
</taxonomicName>
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="SECID0ERHAC" type="type locality">
<paragraph id="B9A89DB351689319C73AD763643D4F30">
<heading id="95D587881F17D73C5E657961DEA403B3" reason="title">Type locality.</heading>
</paragraph>
<paragraph id="03B7B268B405D475ECDC6D131906FEA4">
<collectingCountry id="947A29C59B9C52420B8FD4699879B8B4" name="Taiwan">Taiwan</collectingCountry>
,
<collectingCounty id="C256F835F6CD1A962F32586BA6B60453">Peng-hu Islands</collectingCounty>
,
<location id="5091D3B82C3F1AB91FE971F009148984" LSID="urn:lsid:plazi:treatment:13BF485C3ACE583CB90203D6804ACF4B:5091D3B82C3F1AB91FE971F009148984" country="Taiwan" county="Peng-hu Islands" latitude="23.67" longLatPrecision="21" longitude="119.602776" name="Chih-kan beach">Chih-kan beach</location>
, ca
<named-content content-type="dwc:verbatimCoordinates" id="NCID0E1HAC" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[119.602778,23.670000]}">
<geoCoordinate id="953090814867E2897BDF744A8BC2D3B9" degrees="23" direction="north" minutes="40" orientation="latitude" precision="15" seconds="12" value="23.67">23°40'12&quot;N</geoCoordinate>
,
<geoCoordinate id="27858162625D60EACD0C398FF3A39B3E" degrees="119" direction="east" minutes="36" orientation="longitude" precision="15" seconds="10" value="119.602776">119°36'10&quot;E</geoCoordinate>
</named-content>
,
<date id="4858C43A8C9E162670EA7A3834AA05FB" value="2022-02-25">
<collectingDate id="DCE54F00CEB58E5BACAADEDABB23E817" value="2022-02-25">25 February 2022</collectingDate>
</date>
,
<collectorName id="A6D10482FF355029CADE37E9F1E81EAF">H.-C. Ho</collectorName>
leg.
</paragraph>
</subSubSection>
<subSubSection id="SECID0E6HAC" type="Type specimen">
<paragraph id="13D1675A0D0DE07AC21E99F161F5C054">
<heading id="561DE27F917C9960C59F59A519299F17" reason="title">Type specimen.</heading>
</paragraph>
<paragraph id="65982BB63AC6BB28E7CE5DD5A2D02CDC">
<materialsCitation id="AD474F8980C215D33F5D57554A501AD9" ID-GBIF-Occurrence="4989283301" collectingDate="2022-02-25" collectorName="H.-C. Ho" country="Taiwan" county="Peng-hu Islands" latitude="23.67" location="Chih-kan beach" longLatPrecision="21" longitude="119.602776" specimenCode="NMMB-P 36179" specimenCount="1" typeStatus="holotype">
<typeStatus id="9E1CD25C70EDE35CA3D9A3B94BE57464" type="holotype">Holotype</typeStatus>
:
<specimenCode id="868E06F9EF7BCEA3177E0695A07EC8FA" collectionCode="NMMB-P">
<named-content id="0BA9694A53532EB49343849E3F09BCBC" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-marine-biology-and-aquarium" xlink_title="National Museum of Marine Biology and Aquarium, Pingtung, Taiwan">NMMB-P</named-content>
36179
</specimenCode>
,
<quantity id="E118BC3DCBA6A4F85AF0063468942D4E" metricMagnitude="-2" metricUnit="m" metricValue="6.51" unit="mm" value="65.1">65.1 mm</quantity>
<abbrev id="ABBRID0EKIAC" xlink_title="standard length">SL</abbrev>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection id="SECID0EOIAC" type="diagnosis">
<paragraph id="E58BBF07FCC3172611ACB69A87CA795F">
<heading id="A37BEF75BC003843C9C789A85BFB47BF" reason="title">Diagnosis.</heading>
</paragraph>
<paragraph id="AA72CFC0938C6116141A04C740E6A14F">
A species of
<taxonomicName id="D6333D2376AD4BA3940B33641CC23ABD" authorityName="Cuvier" authorityYear="1816" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="genus">
<emphasis id="507778060D69527E5FC3B4DE6A0C7F16" italics="true">Opistognathus</emphasis>
</taxonomicName>
differing from its congeners in having the following combination of characters: upper jaw rigid, without a distinct flexible lamina posteriorly; dorsal-fin rays XI, 11; anal-fin rays II, 10; gill rakers 1011 + 1 + 1922 = 3133; scale rows in lateral series 55; vertebral formula 10 + 16 = 26; dorsal-fin interdigitation pattern S / S / 1 / 1 + 1 / 1 /; lateral-line ends at base of fourth segmented dorsal-fin ray (15
<superScript id="BC445376257F0E18153E74F738536658">th</superScript>
in total rays); head, nape, dorsal-fin base above lateral line, throat, chest, and pectoral-fin base scaleless; upper two preopercular and fifth mandibular pores bipored; vomer edentate; head mottled with small, dark blotches; dorsal fin with eight blotches along its base; distal portion of membrane between dorsal-fin spines white; body with five horizontal, dark stripes; nape with two dark blotches in front of dorsal-fin origin; and caudal fin with five narrow, dark bands.
</paragraph>
</subSubSection>
<subSubSection id="SECID0E4IAC" type="description">
<paragraph id="C43DD4099B80BE89D644E3C79BFE4330">
<heading id="A9C5878D657AB21BABA2577F1735DF90" reason="title">Description.</heading>
</paragraph>
<paragraph id="1E6B0D1B0F1B3897F365822821FEBDE8">
Meristic and morphometric data are provided in Tables
<tableCitation id="729297C59BC8BB34788D66225EFFC08F" captionStart="Table 1" captionStartId="T1" captionText="Table 1. Meristic characters of Opistognathus cryos sp. nov. and similar species. Paired characters are presented as left / right whenever available. Data of other species were retrieved from Smith-Vaniz (2023)." httpUri="http://table.plazi.org/id/35CB17D5C0691D99A665ABB9F92676F6" tableUuid="35CB17D5C0691D99A665ABB9F92676F6">1</tableCitation>
,
<tableCitation id="3692C9B92EB63ED471EF9D52C73EC3C8" captionStart="Table 2" captionStartId="T2" captionText="Table 2. Morphometric characters of Opistognathus cryos sp. nov. and similar species. Data of other species were retrieved from Smith-Vaniz (2023). Abbreviations: A = anal-fin; C = caudal-fin; D = dorsal-fin; HL = head length; SL = standard length; V = pelvic-fin." httpUri="http://table.plazi.org/id/48A203DFB18FA58BE55BDBF4397F5ECB" tableUuid="48A203DFB18FA58BE55BDBF4397F5ECB">2</tableCitation>
. Paired characters are presented as left / right whenever available.
</paragraph>
<caption id="35CB17D5C0691D99A665ABB9F92676F6" ID-Table-UUID="35CB17D5C0691D99A665ABB9F92676F6" httpUri="http://table.plazi.org/id/35CB17D5C0691D99A665ABB9F92676F6" startId="T1" targetIsTable="true">
<paragraph id="8A0160B0128A0F808C495DE8B04A4C7C">
<label id="6F288E55DF52544F7BCBFB4EB78CD2F9">Table 1.</label>
</paragraph>
<paragraph id="D85D6C0F94189E30E0C76108A605BD02">
Meristic characters of
<taxonomicName id="889DA9686C43531DD83578A94942C17F" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos" status="sp. nov.">
<emphasis id="B4527AC72834A52519A966FAD5F4A86D" italics="true">Opistognathus cryos</emphasis>
</taxonomicName>
<taxonomicNameLabel id="F3F19E057D1FF3B8943E03A785E81D12" rank="species">sp. nov.</taxonomicNameLabel>
and similar species. Paired characters are presented as left / right whenever available. Data of other species were retrieved from
<bibRefCitation id="E6C5B75AF9089B5E3092A1913761D396" DOI="10.11646/zootaxa.5252.1.1" author="Smith-Vaniz WF" firstAuthor="Smith-Vaniz" issue="1" journalOrPublisher="Zootaxa" pagination="1-180" refId="B21" refString="Smith-Vaniz WF (2023) Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species. Zootaxa 5252 (1): 1180. https://doi.org/10.11646/zootaxa.5252.1.1" title="Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species." volume="5252" year="2023">Smith-Vaniz (2023)</bibRefCitation>
.
</paragraph>
</caption>
<paragraph id="586C3A9AEE48A8DEF4C3E9FC79741DE1">
<table id="TID0ERXAE" rules="all">
<tbody id="7B6FAD6DA2987DEA74B110B959936499">
<tr id="414CBCFC2E98C15EA1004AA88FCF4644">
<th id="C0380B83237700859AAD1BF85F043259" colspan="1" isEmpty="true" rowspan="3">-</th>
<th id="6122C637CEC8682681B6CAE934F9FDAA" colspan="1" rowspan="1">
<taxonomicName id="4624FBE639A09210B0B3557C2CB04BB8" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos" status="sp. nov.">
<emphasis id="69B525EBE38B34451650B61EE2FED120" italics="true">O. cryos</emphasis>
</taxonomicName>
<taxonomicNameLabel id="8C2C48697116AA9EBD4FBEA768506127" rank="species">sp. nov.</taxonomicNameLabel>
</th>
<emphasis id="676303A729250A46E4793000023EB448" italics="true">
<th id="6DAB2BDE58FECEDE4A298580371FF019" colspan="1" rowspan="1">
<taxonomicName id="D3AB2926FF4DF75592A96F8CEECD455D" authorityName="Smith-Vaniz" authorityYear="2023" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="asper">O. asper</taxonomicName>
</th>
<th id="6F4BB475F5B11561C749998566FC763A" colspan="1" rowspan="1">
<taxonomicName id="209C7AC09C1EBCB0FCC2A9033D00046F" authorityName="Smith-Vaniz &amp; Yoshino" authorityYear="1985" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="liturus">O. liturus</taxonomicName>
</th>
</emphasis>
</tr>
<tr id="6D68B549C2AC352CE41D774599F2797B">
<th id="803BEF465FDBBE16D3D9B17DEC6D3D86" colspan="1" rowspan="1">This study</th>
<th id="32F71EBBEB663B613C5BF6684EDAFF7D" colspan="1" rowspan="1">
<bibRefCitation id="5159644CE492AB616B326A114310A4C8" DOI="10.11646/zootaxa.5252.1.1" author="Smith-Vaniz WF" firstAuthor="Smith-Vaniz" issue="1" journalOrPublisher="Zootaxa" pagination="1-180" refId="B21" refString="Smith-Vaniz WF (2023) Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species. Zootaxa 5252 (1): 1180. https://doi.org/10.11646/zootaxa.5252.1.1" title="Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species." volume="5252" year="2023">Smith-Vaniz (2023)</bibRefCitation>
</th>
<th id="BB5B62460F0ACFAB0B5ED3DE9F5B9346" colspan="1" rowspan="1">
<bibRefCitation id="74C9653DA6B28F9D202321DC2F3C6892" DOI="10.11646/zootaxa.5252.1.1" author="Smith-Vaniz WF" firstAuthor="Smith-Vaniz" issue="1" journalOrPublisher="Zootaxa" pagination="1-180" refId="B21" refString="Smith-Vaniz WF (2023) Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species. Zootaxa 5252 (1): 1180. https://doi.org/10.11646/zootaxa.5252.1.1" title="Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species." volume="5252" year="2023">Smith-Vaniz (2023)</bibRefCitation>
</th>
</tr>
<tr id="EA431F6EDF7AABD528E6205E01A78862">
<th id="1F6C45FA4A188A0A871C4B261F3D93A6" colspan="1" rowspan="1">Holotype</th>
<th id="85E9E78CD5D849444C6931C351B7C450" colspan="1" rowspan="1">
All types (
<emphasis id="170704087A92511F969C077C04CC7AB1" italics="true">n</emphasis>
= 3)
</th>
<th id="983BCD06C86993BCAF6835180A40AA66" colspan="1" rowspan="1">Holotype</th>
</tr>
<tr id="84C59DAE756908FB43AD339D027C10EF">
<td id="C17D52978E60E92F7B0689E00A315582" colspan="1" rowspan="1">Dorsal-fin rays</td>
<td id="0165EA48C7B31ACD7BB095EA41222960" colspan="1" rowspan="1">XI, 11</td>
<td id="A1805E95991ACD0316F7228BD3302F02" colspan="1" rowspan="1">XI, 1112</td>
<td id="AF830B01ECE6197AAEE6D765576D7A15" colspan="1" rowspan="1">XI, 11</td>
</tr>
<tr id="3E33E3706D9251F7BEB04DB84CC0EBAC">
<td id="C33D27878DC12CC6C6FF5B10B7F9D9E8" colspan="1" rowspan="1">Pectoral-fin rays</td>
<td id="77EAB945BEE82A36DB741D38D2BC2E19" colspan="1" rowspan="1">19 / 19</td>
<td id="B9C9B0CC2BCA865A54F207CC9515F846" colspan="1" rowspan="1">19</td>
<td id="51C9AF25F95CA65C39CEF961D8396777" colspan="1" rowspan="1">19</td>
</tr>
<tr id="0D7B8AEDEC9A21870A867E2C80845B07">
<td id="6197AD8E1DEAF512AF9EF885F5CA426F" colspan="1" rowspan="1">Anal-fin rays</td>
<td id="F65C0D3E21839C3EB840F3AEB682B762" colspan="1" rowspan="1">II, 10</td>
<td id="28BD24735E9A199489EDF15555A410E6" colspan="1" rowspan="1">II, 10</td>
<td id="A14C759B51FE98B1006085AF312ED4D1" colspan="1" rowspan="1">II, 10</td>
</tr>
<tr id="0EB82B5DC5B24B376FA903D03E628193">
<td id="5F759FF308880379AB29180F691705F6" colspan="1" rowspan="1">Gill rakers</td>
<td id="795B352E2EDF0D169057912196716694" colspan="1" rowspan="1">11 + 1 + 19 = 31 / 10 + 1 + 22 = 33</td>
<td id="382EC333E79E68D21CD515966A42F718" colspan="1" rowspan="1">1011 + 2224 = 3234</td>
<td id="04DAF6CB1AC9E55513AE56CD30D8955B" colspan="1" rowspan="1">9 + 1 + 21 = 31</td>
</tr>
<tr id="3B766093B1DBB8B79680FD99897ED0C9">
<td id="3C8FAD30F14C9F69D50D4A15C52A09DF" colspan="1" rowspan="1">Scale rows in lateral series</td>
<td id="E9F6D54A513BA6B9D455299CD389E8DA" colspan="1" rowspan="1">55 / 55</td>
<td id="D4165BD66E252FD6A49B9243B1543CAE" colspan="1" rowspan="1">4244</td>
<td id="3B6D915972CF1119FE47854EEB933A5C" colspan="1" rowspan="1">44</td>
</tr>
<tr id="6BFB7D30EDCAAEA56571BE786058F63E">
<td id="B2868FE6712BB8B7678C8321934A90A8" colspan="1" rowspan="1">Vertebrae</td>
<td id="50A4D9E4B266B413BB0A894349452ABC" colspan="1" rowspan="1">10 + 16 = 26</td>
<td id="AACABB3CE033A904FE039534B5786228" colspan="1" rowspan="1">10 + 16 = 26</td>
<td id="8236EF8A02F6912B37CB1958B5897A5D" colspan="1" rowspan="1">10 + 16 = 26</td>
</tr>
<tr id="0F018193E5CF17D655E58178F5714E29">
<td id="D5F071C8E76CBB6AB9CED3455A5212AA" colspan="1" rowspan="1">Dorsal-fin interdigitation anterior patterns</td>
<td id="D8030A14E37913A3A7CB9112505F8EDB" colspan="1" rowspan="1">S / S / 1 / 1 + 1 / 1 /</td>
<td id="FBAB9B149B5AC20FD52D2DCC66B52BE9" colspan="1" rowspan="1">S / S / 1 / 1 + 1 / 1 /</td>
<td id="B989ABD387B77C2CA89100BFE26CECD8" colspan="1" rowspan="1">/ / 1 / 1 + 1 / 1 /</td>
</tr>
<tr id="37CBA799EFE0D0AD491EAE8398DE14D5">
<td id="34F49DCFB4460C7C0E317AA1C112830C" colspan="1" rowspan="1">Lateral-line terminus total dorsal-fin rays</td>
<td id="1A9ACB78D5370A4A27A639357711580E" colspan="1" rowspan="1">15 / 15</td>
<td id="9FD97C35702FC757C785A8885D0B2A11" colspan="1" rowspan="1">1115</td>
<td id="EF07F6D93D7135DDE6176924AEC49A71" colspan="1" rowspan="1">15</td>
</tr>
</tbody>
</table>
</paragraph>
<paragraph id="FF18EBD165AD4C26CDEFF211896CB4D5">Dorsal-fin rays XI, 11. Anal-fin rays II, 10. Pectoral-fin rays 19 / 19. Pelvic-fin rays I, 5 / I, 5. Principal caudal-fin rays 8 + 8, with innermost 6 + 6 rays branched; procurrent caudal-fin rays 4 on both upper and lower lobes. Gill rakers 11 + 1 + 19 = 31 / 10 + 1 + 22 = 33. Scale rows in lateral series 55 / 55. Vertebrae 10 + 16 = 26. Dorsal-fin interdigitation pattern S / S / 1 / 1 + 1 / 1 /; anal-fin interdigitation pattern / 1 / 1 / 1 / 1 / 1 /.</paragraph>
<paragraph id="E091C54E246D61C8FEDDF5D5C899A13F">
Body slender, depth at anal-fin origin
<quantity id="1BEDDDE96F033343E070C1BD45B53AEC" metricMagnitude="-1" metricUnit="m" metricValue="1.0668" unit="in" value="4.2">4.2 in</quantity>
<abbrev id="ABBRID0EFAAE" xlink_title="standard length">SL</abbrev>
; both dorsal and ventral profiles of body flat. Head large, length
<quantity id="26DFDB9A8D323E709461FFB46CF4365A" metricMagnitude="-2" metricUnit="m" metricValue="6.35" unit="in" value="2.5">2.5 in</quantity>
<abbrev id="ABBRID0EJAAE" xlink_title="standard length">SL</abbrev>
; anterior profile of head rounded, gently curved to dorsal-fin origin. Eyes large, eye diameter 3.0 in
<abbrev id="ABBRID0ENAAE" xlink_title="head length">HL</abbrev>
. Two nostrils, anterior one a short tube with small flap; posterior one without flap, situated immediately in front of eye. Preoprecle and opercle covered by skin, their posterior margins without spines; single, small flap present on upper end of opercle.
</paragraph>
<paragraph id="E93115623878925299189C9351D909BA">
Mouth lower in position, slightly oblique, forming ca 10 ° angle with body axis. Jaws terminal, with lower jaw slightly included. Upper-jaw length
<quantity id="D0687795BAA718E78EA702D7713B1BB6" metricMagnitude="-2" metricUnit="m" metricValue="3.81" unit="in" value="1.5">1.5 in</quantity>
<abbrev id="ABBRID0ETAAE" xlink_title="head length">HL</abbrev>
, its end exceeding
<quantity id="E64194C37B6098B8875A476FB09B807E" metricMagnitude="-2" metricUnit="m" metricValue="2.032" unit="in" value="0.8">0.8 in</quantity>
eye diameter behind posterior margin of orbit; posterior margin of maxilla truncated, without distinct flexible lamina; its anterior margin without crenulae. Supramaxilla present, greatest width about one-third of maxilla. Premaxilla with two or three rows of small canine teeth anteriorly; posterior portion forming single row of conical teeth, their size decreasing posteriorly. Dentary with small canine teeth forming two or three rows anteriorly and single row posteriorly. Vomer and palatine without teeth.
</paragraph>
<paragraph id="EA90425F1E28A27E0E58D2ACA2ACF836">
Body scales cycloid. Scales absent on head, nape, dorsal-fin base above lateral line, throat, chest, and pectoral-fin base (Fig.
<figureCitation id="EB135A06BFAC8DFA830D9F9581E02165" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Head pores and adjacent area of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL, with scaled area shaded in gray A dorsal view B lateral view. Abbreviations: AN = anterior nostril; PN = posterior nostril. Dotted lines indicate dorsal- and pectoral-fin bases. Anterior to left. Figure not to scale." figureDoi="10.3897/zookeys.1220.123541.figure2" httpUri="https://binary.pensoft.net/fig/1192525">2</figureCitation>
). Lateral-line terminus below 4
<superScript id="1783CA9F51A866559F39E47002BD3986">th</superScript>
/ 4
<superScript id="5F3B1C1127CB519E38C2EB4165E5F397">th</superScript>
or 15
<superScript id="3C80DA65CB0DCF61240C2872E07E8AAC">th</superScript>
/ 15
<superScript id="7891E6260C36B904661BC2FCE3BBB298">th</superScript>
in soft ray or total elements of dorsal fin, respectively; lateral-line pores arranged in irregular series along embedded lateral-line canals. Cephalic sensory pores rather sparsely scattered (Fig.
<figureCitation id="2303AA9668ABF7971D73373C7628A9F6" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Head pores and adjacent area of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL, with scaled area shaded in gray A dorsal view B lateral view. Abbreviations: AN = anterior nostril; PN = posterior nostril. Dotted lines indicate dorsal- and pectoral-fin bases. Anterior to left. Figure not to scale." figureDoi="10.3897/zookeys.1220.123541.figure2" httpUri="https://binary.pensoft.net/fig/1192525">2</figureCitation>
); upper-two preopercular pores bipored, others simple; fifth mandibular pore bipored, others simple.
</paragraph>
<paragraph id="F933E797761F94BF0F273501E28D5930">
Dorsal fin with long base, originated at vertical through upper end of gill slit; its distal nearly straight, with slight elevation on soft rays; no distinct notch between spines and soft rays. Pectoral-fin tip pointed; its origin at same horizontal through lower margin of eye; its tip reaching near vertical through anal-fin origin. Pelvic fin elongated, its origin below second dorsal-fin spine and in advance of pectoral fin; its tip reaching anal-fin origin when adpressed; outermost segmented ray not bounded to adjacent ray, with incision on interradial membrane. Anal-fin base moderately long, its posterior end slightly anterior to that of dorsal fin; its origin below first and second dorsal-fin soft rays (between 12
<superScript id="130F4E87E461DE904E714B0F5C91EDC4">th</superScript>
and 13
<superScript id="96A53A376666E341B7F52F0E272678C3">th</superScript>
total dorsal-fin rays); fin rays gradually longer posteriorly; its distal margin nearly straight. Caudal fin rounded, slightly pointed; its length
<quantity id="CF8C7FC93199579E80AB879FB4E04053" metricMagnitude="-2" metricUnit="m" metricValue="3.5559999999999996" unit="in" value="1.4">1.4 in</quantity>
<abbrev id="ABBRID0EPBAE" xlink_title="head length">HL</abbrev>
. Caudal peduncle broad, depth
<quantity id="ECECF2D1ABAC72D35D3A1BE8489D09A8" metricMagnitude="-2" metricUnit="m" metricValue="7.111999999999999" unit="in" value="2.8">2.8 in</quantity>
<abbrev id="ABBRID0ETBAE" xlink_title="head length">HL</abbrev>
.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EXBAE" type="Coloration">
<paragraph id="BFC80C86FE8D4F031927EAAF4F2ADF96">
<heading id="F50A7685512C7CE820ACCD836B3CC1CB" reason="title">Coloration.</heading>
</paragraph>
<paragraph id="3B0531AFE27D15F59C6BDBB27758AA8B">
When preserved (Fig.
<figureCitation id="4F1C71C9C79DEACD7B40C811DEC0A212" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Preserved specimen of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL A on black background B on white background C dorsal view of nape. Photos by Y. - C. Hsu." figureDoi="10.3897/zookeys.1220.123541.figure1" httpUri="https://binary.pensoft.net/fig/1192524">1 A, B</figureCitation>
), body pale, with one horizontal dark line near dorsal-fin base, and four rows of thinner dark lines on lateral side of body above anal-fin base. Head pale, scattered with irregularly sized dark blotches on operculum, upper and lower jaws, and infraorbital; their sizes smaller than pupil. Nape with two dark blotches in front of dorsal-fin origin (Fig.
<figureCitation id="5936B639E0CB10F16E21B3A9FAAB55B9" captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Preserved specimen of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL A on black background B on white background C dorsal view of nape. Photos by Y. - C. Hsu." figureDoi="10.3897/zookeys.1220.123541.figure1" httpUri="https://binary.pensoft.net/fig/1192524">1 C</figureCitation>
). Dorsal fin dusky, with eight dark blotches along its base, slightly extending onto dorsum; distal end of fin spines white; soft rays with two horizontal, dark stripes. Caudal fin dusky, with five narrow, dark bands. Anal, pectoral, and pelvic fins dusky. Oral cavity, including underside of tongue pale.
</paragraph>
<caption id="7EE1C14CEA12191CD1D6C13E33429DCE" ID-DOI="10.3897/zookeys.1220.123541.figure1" ID-arpha="21A1323E-4C4E-57C1-9513-3063E09F96D2" httpUri="https://binary.pensoft.net/fig/1192524" startId="F1">
<paragraph id="A757EF83CB1795897041A47F0C89FA68">
<label id="BCCB365FEE6FE1A02951DD6234D7AFEC">Figure 1.</label>
</paragraph>
<paragraph id="38C3FA6F6C3050F5EA6F15AA01EFA9DA">
Preserved specimen of
<taxonomicName id="0F753E76E696D2BD285753209ED20C9B" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos" status="sp. nov.">
<emphasis id="C8561D030E3CFAFDCE78CE9A3A911A77" italics="true">Opistognathus cryos</emphasis>
</taxonomicName>
<taxonomicNameLabel id="B9437315E79DD759C431D4BAC51B26F8" rank="species">sp. nov.</taxonomicNameLabel>
, holotype,
<named-content id="1A94EB5D06D093BC836BE6CA54333CC2" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-marine-biology-and-aquarium" xlink_title="National Museum of Marine Biology and Aquarium, Pingtung, Taiwan">NMMB-P</named-content>
36179, 65.1 mm
<abbrev id="ABBRID0E4CAE" xlink_title="standard length">SL</abbrev>
<emphasis id="38A835CE8E55B8F0687B8F04373060D3" bold="true">A</emphasis>
on black background
<emphasis id="BD725D78F085D257481F64C6AB247F3F" bold="true">B</emphasis>
on white background
<emphasis id="56ED1387AA312B98E002E4DCBD4DDBB1" bold="true">C</emphasis>
dorsal view of nape. Photos by Y. - C. Hsu.
</paragraph>
</caption>
<caption id="16860E268B568EFEBEBDDBDC3EDF4D38" ID-DOI="10.3897/zookeys.1220.123541.figure2" ID-arpha="D5F648A3-6FBF-5A99-8A17-FFB35F60EDE3" httpUri="https://binary.pensoft.net/fig/1192525" startId="F2">
<paragraph id="25F213B53D39D2D8986C71EDAE7C57A4">
<label id="39FAD9223E8A9A241453E24A9D132926">Figure 2.</label>
</paragraph>
<paragraph id="F82757E2914570AACAE5EE056D5ABF6D">
Head pores and adjacent area of
<taxonomicName id="67A8AB2A3754492C7556830E5556E70E" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos" status="sp. nov.">
<emphasis id="6A92D2038AF2778F49DEDFB7CFB4E068" italics="true">Opistognathus cryos</emphasis>
</taxonomicName>
<taxonomicNameLabel id="264D629D64DD042DF1768ECA1DF37A12" rank="species">sp. nov.</taxonomicNameLabel>
, holotype,
<named-content id="5EF64FA4D71BFE31E4744F3CFD1F2AA1" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-marine-biology-and-aquarium" xlink_title="National Museum of Marine Biology and Aquarium, Pingtung, Taiwan">NMMB-P</named-content>
36179, 65.1 mm
<abbrev id="ABBRID0EEEAE" xlink_title="standard length">SL</abbrev>
, with scaled area shaded in gray
<emphasis id="FC00B1A9F725458D1387FCD1A9BACDDF" bold="true">A</emphasis>
dorsal view
<emphasis id="98C94112969B7077706A30F5D512F656" bold="true">B</emphasis>
lateral view. Abbreviations: AN = anterior nostril; PN = posterior nostril. Dotted lines indicate dorsal- and pectoral-fin bases. Anterior to left. Figure not to scale.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0EREAE" type="Osteology">
<paragraph id="5822FEFF03651E4AE835CA58566BE341">
<heading id="7C83931E1FAF81A518D9A5EBC5C737FC" reason="title">Osteology.</heading>
</paragraph>
<paragraph id="EEC46BD9371BBCC97F6707D5B7C59B4B">
Pleural ribs present on fourth to tenth vertebra; epineurals present and epipleurals absent. Caudal skeleton consists of four plates, including three hypurals and one parhypural; hypural 1 and 2 fused; hypural 3 and 4 fused; hypural 5 present (Fig.
<figureCitation id="C376A9A09B72577F01D9325F3C67B9DF" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. X-radiograph of Opistognathus cryos sp. nov., holotype, NMMB-P 36179, 65.1 mm SL." figureDoi="10.3897/zookeys.1220.123541.figure3" httpUri="https://binary.pensoft.net/fig/1192526">3</figureCitation>
).
</paragraph>
<caption id="5673A2CB35215976B9EAD73B86B335F5" ID-DOI="10.3897/zookeys.1220.123541.figure3" ID-arpha="E6F2DCB5-0C93-50F3-BA60-EB6BF29FAA48" httpUri="https://binary.pensoft.net/fig/1192526" startId="F3">
<paragraph id="40679F322E81617D055DB304D1374A5A">
<label id="325A3C9EB9F01E3AE3A2621A82691B7B">Figure 3.</label>
</paragraph>
<paragraph id="25AFFB95B73A26B4797A89C4BDC631D5">
X-radiograph of
<taxonomicName id="7289461CC160645900B033BB98F1DE77" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos" status="sp. nov.">
<emphasis id="E945DBFFB7382E11484081B65A0AB38D" italics="true">Opistognathus cryos</emphasis>
</taxonomicName>
<taxonomicNameLabel id="6F78A6C830C0326CD83EC59FABC811AA" rank="species">sp. nov.</taxonomicNameLabel>
, holotype,
<named-content id="D8D9839A6F6F220F8E38F06C98893F4A" content-type="dwc:institutional_code" xlink_href="http://grbio.org/institution/national-museum-marine-biology-and-aquarium" xlink_title="National Museum of Marine Biology and Aquarium, Pingtung, Taiwan">NMMB-P</named-content>
36179, 65.1 mm
<abbrev id="ABBRID0ETFAE" xlink_title="standard length">SL</abbrev>
.
</paragraph>
</caption>
</subSubSection>
<subSubSection id="SECID0E3FAE" type="etymology">
<paragraph id="2DF8BF32B84E29C2EBB6E5B1EA393DD3">
<heading id="0B4B83EE5FEC1DD475B4A0B9C60EBCDF" reason="title">Etymology.</heading>
</paragraph>
<paragraph id="5BB1E036CB95941A7FB37B2956BE63C4">
The specific name
<taxonomicName id="677C29716F42C12AEAE2CF78B84359AB" authorityName="Su &amp; Ho" authorityYear="2024" family="Opistognathidae" genus="Opistognathus" kingdom="Animalia" order="Perciformes" phylum="Chordata" rank="species" species="cryos">
<emphasis id="B6B791B28366F0351225C6B23826C08C" italics="true">cryos</emphasis>
</taxonomicName>
, is from the Greek “ κρύος ” meaning cold or chilled, indicating that the
<typeStatus id="1CF6FF689129CCC03CC588DDFC695AA8" type="holotype">holotype</typeStatus>
was collected during a cold snap in 2022. The common name “ frozen jawfish ” is also a reference to the fantasy film “ Frozen ” produced by Walt Disney Animation Studios.
</paragraph>
</subSubSection>
<subSubSection id="SECID0EMGAE" type="distribution">
<paragraph id="EA47E7C3FC0AAEACECAF1C4A2870E173">
<heading id="8E24E6907BF20A3DD85BB1823479B17E" reason="title">Distribution.</heading>
</paragraph>
<paragraph id="69314CE657F1677E09FF6654C9896917">
The
<typeStatus id="EA25BAFC6A283FE94311F50A9BFE667E" type="holotype">holotype</typeStatus>
was collected ashore from the northern portion of the Peng-hu Islands, western Pacific Ocean. Another possible record from
<collectingCountry id="314B4FC9B45F25273CA28400DB85803B" name="Japan">Japan</collectingCountry>
(
<bibRefCitation id="E55185C660E4ECE8D37A821CD5910798" DOI="10.11646/zootaxa.5252.1.1" author="Smith-Vaniz" firstAuthor="Smith-Vaniz" issue="1" journalOrPublisher="Zootaxa" pagination="1-180" refId="B21" refString="Smith-Vaniz WF (2023) Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species. Zootaxa 5252 (1): 1180. https://doi.org/10.11646/zootaxa.5252.1.1" title="Review of Indo-West Pacific jawfishes (Opistognathus: Opistognathidae), with descriptions of 18 new species." volume="5252" year="2023">Smith-Vaniz 2023</bibRefCitation>
) may suggest a wide distribution in the northwestern Pacific Ocean.
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title id="3FE5062807111A3A098406BBC3AD35D7">Stranger things: on the novel buccopharyngeal anatomy and functional morphology of sand-eating Malagasy tadpoles (Anura: Mantellidae: Mantidactylus)</mods:title>
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<heading id="CB1781ECFFA0FFF6FC499475FB3CF9D7" box="[1004,1264,1548,1575]" centered="true" fontSize="9" level="2" pageId="1" pageNumber="2" reason="2">
<taxonomicName id="57E04D03FFA0FFF6FC499475FB3CF9D7" ID-CoL="3XWCQ" authority="Mocquard, 1895" authorityName="Mocquard" authorityYear="1895" box="[1004,1264,1548,1575]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="ambreensis">
<emphasis id="A294EA92FFA0FFF6FC499475FB3CF9D7" bold="true" box="[1004,1264,1548,1575]" italics="true" pageId="1" pageNumber="2">Mantidactylus ambreensis</emphasis>
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<emphasis id="A294EA92FFA0FFF6FC8F9434FC47F995" box="[810,907,1613,1637]" italics="true" pageId="1" pageNumber="2">Oral disc:</emphasis>
Tadpoles of this species are not formally described, and only a few features of the oral disc, mostly from the upper lip, are evident in SEM micrographs. The upper lip is bordered by a single row of conical, alternate marginal papillae, and the inner surface is covered with small, conical, submarginal papillae. The labial tooth row formula (LTRF) could not be assessed, but two rows of labial teeth are present in the upper lip, a long row A1 and A2 as two short sections separated by a wide gap. Labial teeth are short, with four to eight long, distal cusps (
<figureCitation id="08DB2A05FFA0FFF6FB4D953EFAFEF8AF" box="[1256,1330,1863,1887]" captionStart="Figure 2" captionStartId="2.[129,194,1723,1747]" captionTargetBox="[129,1473,1087,1695]" captionTargetId="figure-77@2.[129,1473,1087,1695]" captionTargetPageId="2" captionText="Figure 2. Oral disc and buccopharyngeal morphology of Mantidactylus ambreensis (ZSM 755/2004). A, detail of the upper lip, showing keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infralabial and lingual papillae.C, buccal floor; note the unique crest-like structures delimiting the buccal floor arena, formed of transverse ruffled ridges composed of cauliflower-like papillae.D, buccal roof; note the large narial vacuities with ornamented anterior and posterior margins, and the buccal roof arena papillae tightly aligned in two longitudinal crests. Abbreviations: bfa, buccal floor arena; bra buccal roof arena; brap, buccal roof arena papillae; dv, dorsal velum, gl, glottis; ilp, infralabial papillae; in, internal nare; ljs, lower jaw sheath; lp, lingual papillae; mr, median ridge; pnr, prenarial ridge; rr, ruffled ridges; vv, ventral velum.Scale bars: 100 µm (A, B),200 µm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.14343377" httpUri="https://zenodo.org/record/14343377/files/figure.png" pageId="1" pageNumber="2">Fig. 2A</figureCitation>
). The upper jaw sheath was not visible in micrographs, and the lower sheath is V-shaped and marginally serrated (Fig. 2B).
</paragraph>
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<paragraph id="905F3680FFA3FFF5FF2491DDFD63FBE0" blockId="2.[129,1445,932,1040]" pageId="2" pageNumber="3">
<emphasis id="A294EA92FFA3FFF5FF2491DDFF14FC4C" bold="true" box="[129,216,932,956]" pageId="2" pageNumber="3">Figure 1.</emphasis>
Oral discs of
<taxonomicName id="57E04D03FFA3FFF5FEFD91DDFE16FC4C" authorityName="Boulenger" authorityYear="1985" box="[344,474,932,956]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis id="A294EA92FFA3FFF5FEFD91DDFE16FC4C" box="[344,474,932,956]" italics="true" pageId="2" pageNumber="3">Mantidactylus</emphasis>
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tadpoles examined. Adapted from
<bibRefCitation id="F4714B71FFA3FFF5FC8791DDFBE7FC4C" author="Randrianiaina RD &amp; Strauss A &amp; Glos J" box="[802,1067,932,956]" etAl="et al." firstAuthor="Randrianiaina" pageId="2" pageNumber="3" refId="ref9748" refString="Randrianiaina RD, Strauss A, Glos J et al. Diversity, external morphology and ' reverse taxonomy' in the specialized tadpoles of Malagasy river bank frogs of the subgenus Ochthomantis (genus Mantidactylus %. Contributions in Zoology 2011 a; 80: 17 - S 11. hups: ** doi. org * 10.1163 * 18759866 - 08001002" type="journal volume" year="2011" yearSuffix="a">
Randrianiaina
<emphasis id="A294EA92FFA3FFF5FC0891DCFC15FC4C" box="[941,985,932,956]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
(2011a)
</bibRefCitation>
. Note the extraordinary diversity of oral discs, from configurations similar to pond-type tadpoles in
<taxonomicName id="57E04D03FFA3FFF5FD0991B8FCD2FC28" authorityName="Methuen" authorityYear="1920" box="[684,798,961,984]" class="Amphibia" family="Mantellidae" genus="Maitsomantis" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="argenteus">
<emphasis id="A294EA92FFA3FFF5FD0991B8FCD2FC28" box="[684,798,961,984]" italics="true" pageId="2" pageNumber="3">M. argenteus</emphasis>
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(with keratinized jaw sheaths and labial tooth ridges) to the highly derived morphologies in
<taxonomicName id="57E04D03FFA3FFF5FEC891A4FE20FC04" authorityName="Angel" authorityYear="1929" box="[365,492,988,1012]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="mocquardi">
<emphasis id="A294EA92FFA3FFF5FEC891A4FE20FC04" box="[365,492,988,1012]" italics="true" pageId="2" pageNumber="3">M. mocquardi</emphasis>
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and
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<emphasis id="A294EA92FFA3FFF5FDBC91A4FDBFFC04" box="[537,627,989,1012]" italics="true" pageId="2" pageNumber="3">M. majori</emphasis>
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, with minimum keratinization and extensive papillation. Scale bars: 1 mm. Reprinted with permission of the Journal &quot;Contributions to Zoology&quot;.
</paragraph>
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<paragraph id="905F3680FFA3FFF5FF2494C2FAD6F88C" blockId="2.[129,1427,1723,1916]" pageId="2" pageNumber="3">
<emphasis id="A294EA92FFA3FFF5FF2494C2FF14F924" bold="true" box="[129,216,1723,1748]" pageId="2" pageNumber="3">Figure 2.</emphasis>
Oral disc and buccopharyngeal morphology of
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<emphasis id="A294EA92FFA3FFF5FD3F94C5FC4EF924" box="[666,898,1724,1748]" italics="true" pageId="2" pageNumber="3">Mantidactylus ambreensis</emphasis>
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(ZSM 755/2004). A, detail of the upper lip, showing keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infralabial and lingual papillae. C, buccal floor; note the unique crest-like structures delimiting the buccal floor arena, formed of transverse ruffled ridges composed of cauliflower-like papillae. D, buccal roof; note the large narial vacuities with ornamented anterior and posterior margins, and the buccal roof arena papillae tightly aligned in two longitudinal crests. Abbreviations: bfa, buccal floor arena; bra buccal roof arena; brap, buccal roof arena papillae; dv, dorsal velum, gl, glottis; ilp, infralabial papillae; in, internal nare; ljs, lower jaw sheath; lp, lingual papillae; mr, median ridge; pnr, prenarial ridge; rr, ruffled ridges; vv, ventral velum. Scale bars: 100 µm (A, B), 200 µm (C, D).
</paragraph>
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<paragraph id="905F3680FFA2FFF4FFD492E9FE87FD2C" blockId="3.[113,763,144,732]" pageId="3" pageNumber="4">
<emphasis id="A294EA92FFA2FFF4FFD492E9FF2FFF58" box="[113,227,144,168]" italics="true" pageId="3" pageNumber="4">Buccal floor</emphasis>
(Fig. 2B, C): Buccal floor triangular, caudally wider. Three pairs of tall, conical infralabial papillae overlapping each other. Tongue anlage wide, elliptical, with bulging lateral regions; medially, one pair of conical, tall, lingual papillae with bifid tips. Buccal floor arena delimited by two long, thick crests formed of approximately 15 transverse ruffled ridges closely spaced (100 350 µm long, 95140 µm wide); these ridges are apparently formed of densely packed papillae, distally wider, with a pustulated surface giving them a cauliflower-like appearance, more elaborate rostral than caudally; crests get closer caudally but do not meet medially. Caudal half of the central area of buccal floor arena covered by rounded, short pustulations densely arranged. Buccal pockets slit shaped, deep, oriented obliquely. Ventral velum very short medially, arch-shaped, with long marginal projections, deep median notch, and secretory pits on the margin; spicular support evident. Branchial basket triangular, shallow, with three filter cavities. Branchial plates with a mesh of filter rows visible under the caudal margin of the ventral velum. Glottis fully exposed.
</paragraph>
<paragraph id="905F3680FFA2FFF4FFD49084FDB7FADA" blockId="3.[113,763,765,1322]" pageId="3" pageNumber="4">
<emphasis id="A294EA92FFA2FFF4FFD49084FF12FCE5" box="[113,222,765,789]" italics="true" pageId="3" pageNumber="4">Buccal roof</emphasis>
(
<figureCitation id="08DB2A05FFA2FFF4FF559084FEF0FCE5" box="[240,316,765,789]" captionStart="Figure 2" captionStartId="2.[129,194,1723,1747]" captionTargetBox="[129,1473,1087,1695]" captionTargetId="figure-77@2.[129,1473,1087,1695]" captionTargetPageId="2" captionText="Figure 2. Oral disc and buccopharyngeal morphology of Mantidactylus ambreensis (ZSM 755/2004). A, detail of the upper lip, showing keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infralabial and lingual papillae.C, buccal floor; note the unique crest-like structures delimiting the buccal floor arena, formed of transverse ruffled ridges composed of cauliflower-like papillae.D, buccal roof; note the large narial vacuities with ornamented anterior and posterior margins, and the buccal roof arena papillae tightly aligned in two longitudinal crests. Abbreviations: bfa, buccal floor arena; bra buccal roof arena; brap, buccal roof arena papillae; dv, dorsal velum, gl, glottis; ilp, infralabial papillae; in, internal nare; ljs, lower jaw sheath; lp, lingual papillae; mr, median ridge; pnr, prenarial ridge; rr, ruffled ridges; vv, ventral velum.Scale bars: 100 µm (A, B),200 µm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.14343377" httpUri="https://zenodo.org/record/14343377/files/figure.png" pageId="3" pageNumber="4">Fig. 2D</figureCitation>
): Buccal roof triangular, longer than wide. Prenarial arena triangular, with a wide inverted-V crest with pustulated margin and scattered small pustulations. Internal nares large and subcircular, forming shallow vacuities covered with cilia; margins thick and mostly smooth, with a short, finger-like papilla at the anteromedial corner, and a long, flap-like, pustulated papilla at the anterolateral edge; posterior margin mostly concealed by a tall, wide, triangular postnarial arena papilla anteriorly directed. Median ridge triangular, wide, and with crenulated free margin. Lateral ridge papillae absent. Buccal roof arena delimited by two parallel crests of
<emphasis id="A294EA92FFA2FFF4FE66964EFE00FBBE" box="[451,460,1079,1102]" italics="true" pageId="3" pageNumber="4">c</emphasis>
. 25 packed individual papillae distally pustulated, longer and more elaborated at the middle portion, and shorter and simpler anterior and caudally. Central area of buccal roof arena covered by short, rounded pustulation, denser caudally. Glandular zone with secretory pits arranged in a band anterior to the dorsal velum. Dorsal velum long, medially discontinuous, with medial margins and caudal surface covered by small pustulations and short, finger-like papillae.
</paragraph>
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<mods:title id="C4B7DED9C4ED7602E37A3C5D9E865913">Stranger things: on the novel buccopharyngeal anatomy and functional morphology of sand-eating Malagasy tadpoles (Anura: Mantellidae: Mantidactylus)</mods:title>
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<mods:date id="6CDDB8B44B173B9A97A2BE1DF2C756BD">2024</mods:date>
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<taxonomicName id="57E04D03FFA2FFF4FBA796FAFB10FB6D" ID-CoL="736MP" authority="Boulenger, 1896" authorityName="Boulenger" authorityYear="1896" box="[1026,1244,1155,1182]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="majori">
<emphasis id="A294EA92FFA2FFF4FBA796FAFB10FB6D" bold="true" box="[1026,1244,1155,1182]" italics="true" pageId="3" pageNumber="4">Mantidactylus majori</emphasis>
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<emphasis id="A294EA92FFA2FFF4FC8F96BDFC42FB2C" box="[810,910,1220,1244]" italics="true" pageId="3" pageNumber="4">Oral disc:</emphasis>
Oral configuration is described and illustrated in
<bibRefCitation id="F4714B71FFA2FFF4FC8F969AFB84FB0B" author="Randrianiaina RD &amp; Strauss A &amp; Glos J" box="[810,1096,1251,1275]" etAl="et al." firstAuthor="Randrianiaina" pageId="3" pageNumber="4" refId="ref9748" refString="Randrianiaina RD, Strauss A, Glos J et al. Diversity, external morphology and ' reverse taxonomy' in the specialized tadpoles of Malagasy river bank frogs of the subgenus Ochthomantis (genus Mantidactylus %. Contributions in Zoology 2011 a; 80: 17 - S 11. hups: ** doi. org * 10.1163 * 18759866 - 08001002" type="journal volume" year="2011" yearSuffix="a">
Randrianiaina
<emphasis id="A294EA92FFA2FFF4FC65969DFC23FB0B" box="[960,1007,1251,1275]" italics="true" pageId="3" pageNumber="4">et al.</emphasis>
(2011a)
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and reproduced here (
<figureCitation id="08DB2A05FFA2FFF4FA94969AFAA5FB0B" box="[1329,1385,1251,1275]" captionStart="Figure 1" captionStartId="2.[129,194,932,956]" captionTargetBox="[135,1467,150,898]" captionTargetId="figure-6@2.[129,1473,144,904]" captionTargetPageId="2" captionText="Figure 1. Oral discs of Mantidactylus tadpoles examined. Adapted from Randrianiaina et al.(2011a). Note the extraordinary diversity of oral discs, from configurations similar to pond-type tadpoles in M.argenteus (with keratinized jaw sheaths and labial tooth ridges) to the highly derived morphologies in M. mocquardi and M.majori, with minimum keratinization and extensive papillation. Scale bars: 1 mm. Reprinted with permission of the Journal &quot;Contributions to Zoology&quot;." figureDoi="http://doi.org/10.5281/zenodo.14343375" httpUri="https://zenodo.org/record/14343375/files/figure.png" pageId="3" pageNumber="4">Fig. 1</figureCitation>
). SEM micrographs show slight flaking of a keratinized epithelium on the surface of the three, large, thorn-like papillae in the normal position of the upper jaw sheath (
<figureCitation id="08DB2A05FFA2FFF4FB2C9738FB22FAA9" box="[1161,1262,1345,1369]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="3" pageNumber="4">Fig. 5A, B</figureCitation>
); lower jaw sheath U-shaped, slightly keratinized, and marginally smooth (
<figureCitation id="08DB2A05FFA2FFF4FAFD9718FA6FFA88" box="[1368,1443,1376,1401]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="3" pageNumber="4">Fig. 5D</figureCitation>
).
</paragraph>
<paragraph id="905F3680FFA2FFF3FC8F97E3FD66FB47" blockId="3.[810,1460,1434,1521]" lastBlockId="4.[129,779,964,1207]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
<emphasis id="A294EA92FFA2FFF4FC8F97E3FC59FA42" box="[810,917,1434,1458]" italics="true" pageId="3" pageNumber="4">B uccalfloor</emphasis>
(
<figureCitation id="08DB2A05FFA2FFF4FC0697E3FC33FA41" box="[931,1023,1434,1458]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="3" pageNumber="4">Fig.5D, E</figureCitation>
): Fivepairsoftall, conicalinfra-labialpapillae aligned on the posterior margin of Meckels cartilage. Tongue anlage with one pair of tall, flat, lingual papillae with pustulated tips. Buccal floor arena delimited by two long, thick crests formed of about 20 transverse ruffled ridges closely spaced (365375 µm long, 40200 µm wide; detail in
<figureCitation id="08DB2A05FFA5FFF3FEE4967BFE4BFBEA" box="[321,391,1026,1050]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="4" pageNumber="5">Fig. 5E</figureCitation>
); crests meet medially thus forming a horse-shoe intricate margin surrounding the whole buccal floor arena. Central region of the arena smooth, except a medial, posterior area where several scattered short, pustulated papillae appear. Buccal pockets oriented transversely. Ventral velum without marginal projections and median notch. Glottis fully covered.
</paragraph>
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<emphasis id="A294EA92FFA2FFF4FFD49509FF04F878" bold="true" box="[113,200,1904,1928]" pageId="3" pageNumber="4">Figure 3.</emphasis>
Oral disc and buccopharyngeal morphology of
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<emphasis id="A294EA92FFA2FFF4FD2F9509FC95F878" box="[650,857,1904,1928]" italics="true" pageId="3" pageNumber="4">Mantidactylus ambony</emphasis>
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(ZSM 762/2004 Stage 25). Unfortunately, only a limited area of the oral cavity was well preserved and suitable for illustration. A, detail of the upper jaw sheath. B, detail of the ruffled ridges of the buccal floor. C, detail of the buccal roof arena papillae. Scale bars: 100 µm (A), 200 µm (B, C).
</paragraph>
</caption>
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<emphasis id="A294EA92FFA5FFF3FF269175FF16FCD4" bold="true" box="[131,218,780,804]" pageId="4" pageNumber="5">Figure 4.</emphasis>
Oral disc and buccopharyngeal morphology of
<taxonomicName id="57E04D03FFA5FFF3FD399175FCBFFCD4" baseAuthorityName="Boulenger" baseAuthorityYear="1882" box="[668,883,780,804]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="femoralis">
<emphasis id="A294EA92FFA5FFF3FD399175FCBFFCD4" box="[668,883,780,804]" italics="true" pageId="4" pageNumber="5">Mantidactylus femoralis</emphasis>
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(ZSM 1928/2007 Stage 26 for oral disc, and ZSM 1781/2007 the remaining). A, detail of the keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infra-labial and lingual papillae. C, buccal floor; note the thick ruffled ridges closely spaced along the longitudinal crests. D, buccal roof; note the large, ornamented narial vacuities and the longitudinal crests with tightly aligned buccal roof arena papillae. Scale bars: 100 µm (A), 200 µm (BD).
</paragraph>
</caption>
<paragraph id="905F3680FFA5FFF3FF2496A1FCC6F8F5" blockId="4.[128,779,1240,1797]" pageId="4" pageNumber="5">
<emphasis id="A294EA92FFA5FFF3FF2496A1FF27FB01" box="[129,235,1240,1265]" italics="true" pageId="4" pageNumber="5">Buccal roof</emphasis>
(
<figureCitation id="08DB2A05FFA5FFF3FF5896A1FEAEFB01" box="[253,354,1240,1265]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="4" pageNumber="5">Fig. 5F, G</figureCitation>
): Prenarial arena with a wide inverted-V crest with smooth margin. Internal nares large and slit-like, transversely oriented; margins thick and mostly smooth, with a long, conical papilla at the anterolateral edge, projecting posteriorly. Median ridge rectangular, taller than wide, and with crenulated free margin. Buccal roof arena entirely ornamented with
<emphasis id="A294EA92FFA5FFF3FD27970FFD47FA7D" box="[642,651,1398,1421]" italics="true" pageId="4" pageNumber="5">c</emphasis>
. 22 bilateral, transverse, parallel ruffled ridges of densely packed, short papillae; posteriorly to the median ridge, these ridges meet medially with the contralateral one and papillae appear less packed; some larger papillae are scattered along the midline; laterally, each ridge ends in a hand-like flap papilla that recurves medially (detail in
<figureCitation id="08DB2A05FFA5FFF3FD08946BFD3BF9DA" box="[685,759,1554,1578]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="4" pageNumber="5">Fig. 5G</figureCitation>
); these flaps are large in the middle section, and gradually decrease in size rostral and caudally. Overall, ridges are 400830 µm long and 90160 µm wide, and cover about 45% of the buccal roof. Threedimensional reconstruction of the buccal cavity (
<figureCitation id="08DB2A05FFA5FFF3FDD594F6FD76F957" box="[624,698,1679,1703]" captionStart="Figure 5" captionStartId="5.[113,178,1260,1284]" captionTargetBox="[114,1458,144,1232]" captionTargetId="figure-180@5.[114,1458,144,1232]" captionTargetPageId="5" captionText="Figure 5. Oral disc and buccopharyngeal morphology of Mantidactylus majori (ZSM 042/2007 for oral disc and ZSM 1684/2004 Stage 29 the remaining).A, upper lip, showing three large papillae in the place of upper jaw sheath. B, detail of the central papillae, showing a likely slight keratinization of the surface, as suggested by tissue flaking.C, three-dimensional reconstruction of a CT scanned specimen, showing the arrangement of dorsal (yellow, central region) and ventral (cream, lateral regions) ruffled ridges regarding the chondrocranium:ventral view of the chondrocranium, transverse section at the level of subocular fenestrae, and sagittal section at midline; in the latter image, the hyobranchial apparatus was not removed to show the tight space between buccal roof and floor that crowded ruffled ridges occupy; arrows indicate anterior. D, buccal floor; note the extraordinary development of the ruffled ridges, and the bilateral crests meeting medially forming a horse-shoe structure. E, detail of ventral ruffled ridges from the anterior section of the right crest; note the rows of tall, cauliflower-like papillae packed together forming each ridge.F, buccal roof; note the transverse, bilateral ruffled ridges occupying the entire buccal roof arena. G, detail of dorsal ruffled ridges from the mid portion of the left crest; note the lateral edge of each ridge retaining a papilla- or flap-like aspect and the medial portion proliferating with a cauliflower-like appearance as in the buccal floor. Scale bars: 500 µm (A, D, F), 100 µm (B, E, G)." figureDoi="http://doi.org/10.5281/zenodo.14343387" httpUri="https://zenodo.org/record/14343387/files/figure.png" pageId="4" pageNumber="5">Fig. 5C</figureCitation>
) shows how buccal roof and floor structures align relative to each other, with dorsal crests fitting exactly within the space defined by ventral crests. Unfortunately, the dorsal velum was broken in SEM images.
</paragraph>
</subSubSection>
</treatment>
</document>

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@ -0,0 +1,139 @@
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<heading id="CB1781ECFFA2FFF4FC5393E4FB2BFE48" box="[1014,1255,413,440]" centered="true" fontSize="9" level="2" pageId="3" pageNumber="4" reason="2">
<taxonomicName id="57E04D03FFA2FFF4FC5393E4FB2BFE48" ID-CoL="3XWDP" authority="(Boulenger, 1882)" baseAuthorityName="Boulenger" baseAuthorityYear="1882" box="[1014,1255,413,440]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="femoralis">
<emphasis id="A294EA92FFA2FFF4FC5393E4FB2BFE48" bold="true" box="[1014,1255,413,440]" italics="true" pageId="3" pageNumber="4">Mantidactylus femoralis</emphasis>
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<emphasis id="A294EA92FFA2FFF4FC8F93A7FC42FE06" box="[810,910,478,502]" italics="true" pageId="3" pageNumber="4">Oral disc:</emphasis>
Oral configuration is described and illustrated in
<bibRefCitation id="F4714B71FFA2FFF4FC8F9384FB84FDE5" author="Randrianiaina RD &amp; Strauss A &amp; Glos J" box="[810,1096,509,533]" etAl="et al." firstAuthor="Randrianiaina" pageId="3" pageNumber="4" refId="ref9748" refString="Randrianiaina RD, Strauss A, Glos J et al. Diversity, external morphology and ' reverse taxonomy' in the specialized tadpoles of Malagasy river bank frogs of the subgenus Ochthomantis (genus Mantidactylus %. Contributions in Zoology 2011 a; 80: 17 - S 11. hups: ** doi. org * 10.1163 * 18759866 - 08001002" type="journal volume" year="2011" yearSuffix="a">
Randrianiaina
<emphasis id="A294EA92FFA2FFF4FC659387FC23FDE5" box="[960,1007,509,533]" italics="true" pageId="3" pageNumber="4">et al.</emphasis>
(2011a)
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and reproduced here (
<figureCitation id="08DB2A05FFA2FFF4FA949384FAA5FDE5" box="[1329,1385,509,533]" captionStart="Figure 1" captionStartId="2.[129,194,932,956]" captionTargetBox="[135,1467,150,898]" captionTargetId="figure-6@2.[129,1473,144,904]" captionTargetPageId="2" captionText="Figure 1. Oral discs of Mantidactylus tadpoles examined. Adapted from Randrianiaina et al.(2011a). Note the extraordinary diversity of oral discs, from configurations similar to pond-type tadpoles in M.argenteus (with keratinized jaw sheaths and labial tooth ridges) to the highly derived morphologies in M. mocquardi and M.majori, with minimum keratinization and extensive papillation. Scale bars: 1 mm. Reprinted with permission of the Journal &quot;Contributions to Zoology&quot;." figureDoi="http://doi.org/10.5281/zenodo.14343375" httpUri="https://zenodo.org/record/14343375/files/figure.png" pageId="3" pageNumber="4">Fig. 1</figureCitation>
). SEM micrographs show labial teeth short, with wide sheath and head with four to eight long, distal cusps (
<figureCitation id="08DB2A05FFA2FFF4FB0D9045FB3EFDA4" box="[1192,1266,572,596]" captionStart="Figure 4" captionStartId="4.[131,196,780,804]" captionTargetBox="[129,1473,144,752]" captionTargetId="figure-584@4.[129,1473,144,752]" captionTargetPageId="4" captionText="Figure 4. Oral disc and buccopharyngeal morphology of Mantidactylus femoralis (ZSM 1928/2007 Stage 26 for oral disc, and ZSM 1781/2007 the remaining). A, detail of the keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infra-labial and lingual papillae.C, buccal floor; note the thick ruffled ridges closely spaced along the longitudinal crests. D, buccal roof; note the large, ornamented narial vacuities and the longitudinal crests with tightly aligned buccal roof arena papillae. Scale bars: 100 µm (A), 200 µm (BD)." figureDoi="http://doi.org/10.5281/zenodo.14343383" httpUri="https://zenodo.org/record/14343383/files/figure.png" pageId="3" pageNumber="4">Fig. 4A</figureCitation>
). Upper jaw finely serrated and lower jaw sheath V-shaped, poorly keratinized, and marginally serrated (
<figureCitation id="08DB2A05FFA2FFF4FC589002FB8FFD63" box="[1021,1091,635,659]" captionStart="Figure 4" captionStartId="4.[131,196,780,804]" captionTargetBox="[129,1473,144,752]" captionTargetId="figure-584@4.[129,1473,144,752]" captionTargetPageId="4" captionText="Figure 4. Oral disc and buccopharyngeal morphology of Mantidactylus femoralis (ZSM 1928/2007 Stage 26 for oral disc, and ZSM 1781/2007 the remaining). A, detail of the keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infra-labial and lingual papillae.C, buccal floor; note the thick ruffled ridges closely spaced along the longitudinal crests. D, buccal roof; note the large, ornamented narial vacuities and the longitudinal crests with tightly aligned buccal roof arena papillae. Scale bars: 100 µm (A), 200 µm (BD)." figureDoi="http://doi.org/10.5281/zenodo.14343383" httpUri="https://zenodo.org/record/14343383/files/figure.png" pageId="3" pageNumber="4">Fig. 4B</figureCitation>
).
</paragraph>
<paragraph id="905F3680FFA2FFF4FC8F90CDFA78FCDA" blockId="3.[810,1460,692,810]" pageId="3" pageNumber="4">
<emphasis id="A294EA92FFA2FFF4FC8F90CDFC50FD3C" box="[810,924,692,716]" italics="true" pageId="3" pageNumber="4">Buccal floor</emphasis>
(
<figureCitation id="08DB2A05FFA2FFF4FC0990CDFBDDFD3C" box="[940,1041,692,716]" captionStart="Figure 4" captionStartId="4.[131,196,780,804]" captionTargetBox="[129,1473,144,752]" captionTargetId="figure-584@4.[129,1473,144,752]" captionTargetPageId="4" captionText="Figure 4. Oral disc and buccopharyngeal morphology of Mantidactylus femoralis (ZSM 1928/2007 Stage 26 for oral disc, and ZSM 1781/2007 the remaining). A, detail of the keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infra-labial and lingual papillae.C, buccal floor; note the thick ruffled ridges closely spaced along the longitudinal crests. D, buccal roof; note the large, ornamented narial vacuities and the longitudinal crests with tightly aligned buccal roof arena papillae. Scale bars: 100 µm (A), 200 µm (BD)." figureDoi="http://doi.org/10.5281/zenodo.14343383" httpUri="https://zenodo.org/record/14343383/files/figure.png" pageId="3" pageNumber="4">Fig. 4B, C</figureCitation>
): Four pairs of short, conical infra-labial papillae not overlapping each other. Buccal floor arena crests formed of approximately 15 transverse ruffled ridges closely spaced (45140 µm long, 2050 µm wide). Glottis fully covered.
</paragraph>
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<emphasis id="A294EA92FFA2FFF4FC8F9132FC55FC93" box="[810,921,843,867]" italics="true" pageId="3" pageNumber="4">Buccal roof</emphasis>
(
<figureCitation id="08DB2A05FFA2FFF4FC099132FC37FC93" box="[940,1019,843,867]" captionStart="Figure 4" captionStartId="4.[131,196,780,804]" captionTargetBox="[129,1473,144,752]" captionTargetId="figure-584@4.[129,1473,144,752]" captionTargetPageId="4" captionText="Figure 4. Oral disc and buccopharyngeal morphology of Mantidactylus femoralis (ZSM 1928/2007 Stage 26 for oral disc, and ZSM 1781/2007 the remaining). A, detail of the keratinized teeth of row A1. B, detail of the anterior region of the buccal floor, showing the keratinized and serrated lower jaw sheath, and infra-labial and lingual papillae.C, buccal floor; note the thick ruffled ridges closely spaced along the longitudinal crests. D, buccal roof; note the large, ornamented narial vacuities and the longitudinal crests with tightly aligned buccal roof arena papillae. Scale bars: 100 µm (A), 200 µm (BD)." figureDoi="http://doi.org/10.5281/zenodo.14343383" httpUri="https://zenodo.org/record/14343383/files/figure.png" pageId="3" pageNumber="4">Fig. 4D</figureCitation>
): Prenarial arena with a wide inverted-V crest with a mostly smooth margin. Internal nares with a long, conical papilla at the anterolateral edge; posterior margin partially concealed by a short, wide, triangular postnarial arena papilla anteriorly directed. Median ridge very short, triangular, and with smooth free margin. Buccal roof arena delimited by two slightly convergent crests formed of
<emphasis id="A294EA92FFA2FFF4FB329671FB6CFBEF" box="[1175,1184,1032,1055]" italics="true" pageId="3" pageNumber="4">c</emphasis>
. 30 closely spaced papillae. Central area of buccal roof arena smooth. Glandular zone not evident in micrographs.
</paragraph>
</subSubSection>
</treatment>
</document>

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<mods:title id="F8DB037CD0478F423F61B507DE32E067">Stranger things: on the novel buccopharyngeal anatomy and functional morphology of sand-eating Malagasy tadpoles (Anura: Mantellidae: Mantidactylus)</mods:title>
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<heading id="CB1781ECFFA2FFF4FEE79730FDE6FA94" box="[322,554,1353,1380]" centered="true" fontSize="9" level="2" pageId="3" pageNumber="4" reason="2">
<taxonomicName id="57E04D03FFA2FFF4FEE79730FDE6FA94" authority="Scherz, Rasolonjatovo, Köhler, Rancilhac, Rakotoarison, Raselimanana, Ohler, Preick, Hofreiter, Glaw, and Vences, 2020" authorityName="Scherz, Rasolonjatovo, Köhler, Rancilhac, Rakotoarison, Raselimanana, Ohler, Preick, Hofreiter, Glaw, and Vences" authorityYear="2020" box="[322,554,1353,1380]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ambony">
<emphasis id="A294EA92FFA2FFF4FEE79730FDE6FA94" bold="true" box="[322,554,1353,1380]" italics="true" pageId="3" pageNumber="4">Mantidactylus ambony</emphasis>
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<paragraph id="905F3680FFA2FFF4FFD49702FB24FF17" blockId="3.[113,762,1403,1521]" lastBlockId="3.[810,1459,144,231]" pageId="3" pageNumber="4">
<emphasis id="A294EA92FFA2FFF4FFD49702FF1AFA63" box="[113,214,1403,1427]" italics="true" pageId="3" pageNumber="4">Oral disc:</emphasis>
Oral configuration is described and illustrated in
<bibRefCitation id="F4714B71FFA2FFF4FFD497E3FE45FA42" author="Randrianiaina" box="[113,393,1434,1458]" etAl="et al." firstAuthor="Randrianiaina" pageId="3" pageNumber="4" refId="ref9748" refString="Randrianiaina RD, Strauss A, Glos J et al. Diversity, external morphology and ' reverse taxonomy' in the specialized tadpoles of Malagasy river bank frogs of the subgenus Ochthomantis (genus Mantidactylus %. Contributions in Zoology 2011 a; 80: 17 - S 11. hups: ** doi. org * 10.1163 * 18759866 - 08001002" type="journal volume" year="2011" yearSuffix="a">
Randrianiaina
<emphasis id="A294EA92FFA2FFF4FEAC97E2FEF5FA42" box="[265,313,1434,1458]" italics="true" pageId="3" pageNumber="4">et al.</emphasis>
(2011a
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; as
<taxonomicName id="57E04D03FFA2FFF4FE1697E2FDF3FA42" authorityName="Mocquard" authorityYear="1895" box="[435,575,1434,1458]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ambreensis">
<emphasis id="A294EA92FFA2FFF4FE1697E2FDF3FA42" box="[435,575,1434,1458]" italics="true" pageId="3" pageNumber="4">M. ambreensis</emphasis>
</taxonomicName>
), and reproduced here (
<figureCitation id="08DB2A05FFA2FFF4FF0B97C0FF2AFA21" box="[174,230,1465,1489]" captionStart="Figure 1" captionStartId="2.[129,194,932,956]" captionTargetBox="[135,1467,150,898]" captionTargetId="figure-6@2.[129,1473,144,904]" captionTargetPageId="2" captionText="Figure 1. Oral discs of Mantidactylus tadpoles examined. Adapted from Randrianiaina et al.(2011a). Note the extraordinary diversity of oral discs, from configurations similar to pond-type tadpoles in M.argenteus (with keratinized jaw sheaths and labial tooth ridges) to the highly derived morphologies in M. mocquardi and M.majori, with minimum keratinization and extensive papillation. Scale bars: 1 mm. Reprinted with permission of the Journal &quot;Contributions to Zoology&quot;." figureDoi="http://doi.org/10.5281/zenodo.14343375" httpUri="https://zenodo.org/record/14343375/files/figure.png" pageId="3" pageNumber="4">Fig. 1</figureCitation>
). SEM micrographs show the strong serration of the upper jaw sheath (
<figureCitation id="08DB2A05FFA2FFF4FE8897A0FEBAFA01" box="[301,374,1497,1521]" captionStart="Figure 3" captionStartId="3.[113,178,1904,1928]" captionTargetBox="[114,1458,1569,1876]" captionTargetId="figure-770@3.[114,1458,1569,1876]" captionTargetPageId="3" captionText="Figure 3. Oral disc and buccopharyngeal morphology of Mantidactylus ambony (ZSM 762/2004 Stage 25). Unfortunately, only a limited area the oral cavity was well preserved and suitable for illustration. A, detail of the upper jaw sheath. B, detail of the ruffled ridges of the buccal. C, detail of the buccal roof arena papillae.Scale bars: 100 µm (A), 200 µm (B, C)." figureDoi="http://doi.org/10.5281/zenodo.14343379" httpUri="https://zenodo.org/record/14343379/files/figure.png" pageId="3" pageNumber="4">Fig. 3A</figureCitation>
).
<emphasis id="A294EA92FFA2FFF4FC8F92E9FC53FF58" box="[810,927,144,168]" italics="true" pageId="3" pageNumber="4">Buccal floor</emphasis>
(Fig. 3B): Almost identical to
<emphasis id="A294EA92FFA2FFF4FB4A92E8FAB1FF58" box="[1263,1405,144,168]" italics="true" pageId="3" pageNumber="4">M. ambreensis</emphasis>
con- figuration; crests of the buccal floor with slightly smaller ruffled ridges (100155 µm long, 2376 µm wide).
</paragraph>
<paragraph id="905F3680FFA2FFF4FC8F9371FBBBFE8E" blockId="3.[810,1459,264,382]" pageId="3" pageNumber="4">
<emphasis id="A294EA92FFA2FFF4FC8F9371FC59FED0" box="[810,917,264,288]" italics="true" pageId="3" pageNumber="4">Buccal roof</emphasis>
(
<figureCitation id="08DB2A05FFA2FFF4FC019371FC21FED0" box="[932,1005,264,288]" captionStart="Figure 3" captionStartId="3.[113,178,1904,1928]" captionTargetBox="[114,1458,1569,1876]" captionTargetId="figure-770@3.[114,1458,1569,1876]" captionTargetPageId="3" captionText="Figure 3. Oral disc and buccopharyngeal morphology of Mantidactylus ambony (ZSM 762/2004 Stage 25). Unfortunately, only a limited area the oral cavity was well preserved and suitable for illustration. A, detail of the upper jaw sheath. B, detail of the ruffled ridges of the buccal. C, detail of the buccal roof arena papillae.Scale bars: 100 µm (A), 200 µm (B, C)." figureDoi="http://doi.org/10.5281/zenodo.14343379" httpUri="https://zenodo.org/record/14343379/files/figure.png" pageId="3" pageNumber="4">Fig. 3C</figureCitation>
): Features of the prenarial, narial, and dorsal velum regions unfortunately could not be observed. Other features as in
<taxonomicName id="57E04D03FFA2FFF4FC31933EFBD2FEAF" authorityName="Mocquard" authorityYear="1895" box="[916,1054,327,351]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ambreensis">
<emphasis id="A294EA92FFA2FFF4FC31933EFBD2FEAF" box="[916,1054,327,351]" italics="true" pageId="3" pageNumber="4">M. ambreensis</emphasis>
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, except less numerous buccal roof arena papillae (about 20 on each side).
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