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<document ID-DOI="10.26107/RBZ-2022-0028" ID-GBIF-Dataset="71e44b33-6313-4f48-8226-7b8bf4b2804d" ID-ISSN="2345-7600" ID-Zenodo-Dep="7399882" ID-ZooBank="3622B8D7-9786-424C-97B8-B7F962B91B94" checkinTime="1670242845483" checkinUser="felipe" docAuthor="Bohlen, Jörg, Kottelat, Maurice &amp; Šlechtová, Vendula" docDate="2022" docId="722E87D4AF71A448FF5BF8CAFD2CFB35" docLanguage="en" docName="RafflesBZool.70.511518.pdf" docOrigin="Raffles Bulletin of Zoology 70" docStyle="DocumentStyle:AF09A0581917EACD12E7F0991C3E7C26.5:RafflesBZool.2019-.journal_article" docStyleId="AF09A0581917EACD12E7F0991C3E7C26" docStyleName="RafflesBZool.2019-.journal_article" docStyleVersion="5" docTitle="Nemacheilus cacao Bohlen &amp; Kottelat &amp; Šlechtová 2022, new species" docType="treatment" docVersion="4" lastPageNumber="515" masterDocId="8E17FFACAF70A44CFFA1FFA4FF9DFFD8" masterDocTitle="Nemacheilus cacao, a new species of loach (Teleostei: Nemacheilidae) from the middle Mekong basin in Laos" masterLastPageNumber="518" masterPageNumber="511" pageNumber="512" updateTime="1672774617846" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Nemacheilus cacao, a new species of loach (Teleostei: Nemacheilidae) from the middle Mekong basin in Laos</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bohlen, Jörg</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kottelat, Maurice</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Šlechtová, Vendula</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Raffles Bulletin of Zoology</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-11-16</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>70</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>511</mods:start>
<mods:end>518</mods:end>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.26107/RBZ-2022-0028</mods:identifier>
<mods:identifier type="GBIF-Dataset">71e44b33-6313-4f48-8226-7b8bf4b2804d</mods:identifier>
<mods:identifier type="ISSN">2345-7600</mods:identifier>
<mods:identifier type="Zenodo-Dep">7399882</mods:identifier>
<mods:identifier type="ZooBank">3622B8D7-9786-424C-97B8-B7F962B91B94</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.7502277" ID-Zenodo-Dep="7502277" LSID="urn:lsid:plazi:treatment:722E87D4AF71A448FF5BF8CAFD2CFB35" httpUri="http://treatment.plazi.org/id/722E87D4AF71A448FF5BF8CAFD2CFB35" lastPageId="4" lastPageNumber="515" pageId="1" pageNumber="512">
<subSubSection box="[250,615,1902,1927]" pageId="1" pageNumber="512" type="nomenclature">
<paragraph blockId="1.[250,615,1902,1959]" box="[250,615,1902,1927]" pageId="1" pageNumber="512">
<heading bold="true" box="[250,615,1902,1927]" centered="true" fontSize="10" level="2" pageId="1" pageNumber="512" reason="2">
<emphasis bold="true" box="[250,615,1902,1927]" pageId="1" pageNumber="512">
<taxonomicName authority="Bohlen &amp; Kottelat &amp; Šlechtová, 2022" authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[250,467,1902,1926]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="cacao" status="new species">
<emphasis bold="true" box="[250,467,1902,1926]" italics="true" pageId="1" pageNumber="512">Nemacheilus cacao</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[482,615,1902,1927]" pageId="1" pageNumber="512" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection box="[324,541,1934,1959]" pageId="1" pageNumber="512" type="description">
<paragraph blockId="1.[250,615,1902,1959]" box="[324,541,1934,1959]" pageId="1" pageNumber="512">
(
<figureCitation box="[332,436,1934,1959]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStart-4="Fig" captionStartId-0="2.[135,170,656,678]" captionStartId-1="3.[113,148,2064,2086]" captionStartId-2="4.[135,170,683,705]" captionStartId-3="4.[835,870,683,705]" captionStartId-4="5.[113,148,1791,1813]" captionTargetBox-0="[135,1473,189,641]" captionTargetBox-1="[180,1380,221,2031]" captionTargetBox-2="[135,783,189,668]" captionTargetBox-3="[826,1474,189,668]" captionTargetBox-4="[113,1451,433,1775]" captionTargetId-0="figure-8@2.[132,1476,189,641]" captionTargetId-1="figure-61@3.[151,1414,189,2049]" captionTargetId-2="figure-8@4.[135,775,186,668]" captionTargetId-3="figure-9@4.[835,1474,189,668]" captionTargetId-4="figure-14@5.[113,1451,433,1775]" captionTargetPageId-0="2" captionTargetPageId-1="3" captionTargetPageId-2="4" captionTargetPageId-3="4" captionTargetPageId-4="5" captionText-0="Fig. 1. Live specimen of Nemacheilus cacao, new species, ZRC 62554, paratype, male, 61.5 mm SL; Laos: Khamouane province: Thakkhet district: small tributary of Nam Thorn at cave Tham Nang Eng (Photo: J. Kühne)." captionText-1="Fig. 2. Nemacheilus cacao, new species, Laos: Khammouane prov: Thaket district: small tributary of Nam Thorn at cave Tham Nang Eng. a, ZRC 62553, holotype, 56.9 mm SL, adult male; b, ZRC 62558, paratype, 50.4 mm SL, adult female; c, ZRC 62556, 27.7 mm SL, juvenile." captionText-2="Fig. 3. Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL; mouth in ventral view." captionText-3="Fig. 4. Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL; left suborbital flap in lateral view." captionText-4="Fig. 5. Tubercles on body side and pectoral fin of large male Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL. A, red outline depicts field of tubercles on body side; BC, area of main tuberculation enlarged; D, dorsal view on right pectoral fin; E, tubercles on pectoral fin enlarged." figureDoi-0="http://doi.org/10.5281/zenodo.7399886" figureDoi-1="http://doi.org/10.5281/zenodo.7399888" figureDoi-2="http://doi.org/10.5281/zenodo.7399890" figureDoi-3="http://doi.org/10.5281/zenodo.7502275" figureDoi-4="http://doi.org/10.5281/zenodo.7399892" httpUri-0="https://zenodo.org/record/7399886/files/figure.png" httpUri-1="https://zenodo.org/record/7399888/files/figure.png" httpUri-2="https://zenodo.org/record/7399890/files/figure.png" httpUri-3="https://zenodo.org/record/7502275/files/figure.png" httpUri-4="https://zenodo.org/record/7399892/files/figure.png" pageId="1" pageNumber="512">Figs. 15</figureCitation>
,
<tableCitation box="[449,531,1935,1959]" captionStart="Table 1" captionStartId="1.[113,168,189,211]" captionTargetPageId="1" captionText="Table 1. Morphometric data of holotype and eight paratypes (ZRC 62553 to 62558) of Nemacheilus cacao, new species. Range and mean include holotype data." httpUri="http://table.plazi.org/id/AEF8664AAF71A44DFFD0FF19FED6FF28" pageId="1" pageNumber="512" tableUuid="AEF8664AAF71A44DFFD0FF19FED6FF28">Table 1</tableCitation>
)
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="512" type="materials_examined">
<paragraph blockId="1.[113,751,1998,2119]" pageId="1" pageNumber="512">
<materialsCitation accessionNumber="ZRC 62553" collectingDate="2013-06-09" collectingDateMax="2013-10" collectingDateMin="2013-06-09" collectionCode="ZRC" country="Laos" county="Thakhek district" latitude="17.444166" location="cave Tham Nang Eng" longLatPrecision="21" longitude="104.94861" municipality="Nam Thorn" pageId="1" pageNumber="512" specimenCount="1" stateProvince="Khammouan" typeStatus="holotype">
<emphasis bold="true" box="[113,226,1999,2023]" pageId="1" pageNumber="512">
<typeStatus box="[113,220,1999,2023]" pageId="1" pageNumber="512">Holotype</typeStatus>
.
</emphasis>
<accessionNumber box="[236,368,1998,2023]" httpUri="http://www.ncbi.nlm.nih.gov/protein/ZRC62553" pageId="1" pageNumber="512">
<collectionCode box="[236,288,1999,2023]" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="512">ZRC</collectionCode>
62553
</accessionNumber>
,
<quantity box="[383,482,1998,2022]" metricMagnitude="-2" metricUnit="m" metricValue="5.6899999999999995" pageId="1" pageNumber="512" unit="mm" value="56.9">56.9 mm</quantity>
SL;
<collectingCountry box="[542,599,1999,2022]" name="Laos" pageId="1" pageNumber="512">Laos</collectingCountry>
:
<collectingRegion country="Laos" name="Khammouan" pageId="1" pageNumber="512">Khammouan province</collectingRegion>
:
<collectingCounty box="[221,399,2031,2055]" pageId="1" pageNumber="512">Thakhek district</collectingCounty>
: small tributary of
<collectingMunicipality box="[602,725,2031,2055]" pageId="1" pageNumber="512">Nam Thorn</collectingMunicipality>
at
<location LSID="urn:lsid:plazi:treatment:722E87D4AF71A448FF5BF8CAFD2CFB35:FF586019AF71A44DFFD0F7ABFEFFF7FF" box="[113,354,2063,2087]" country="Laos" county="Thakhek district" latitude="17.444166" longLatPrecision="21" longitude="104.94861" municipality="Nam Thorn" name="cave Tham Nang Eng" pageId="1" pageNumber="512" stateProvince="Khammouan">cave Tham Nang Eng</location>
,
<geoCoordinate box="[366,494,2061,2086]" degrees="17" direction="north" minutes="26" orientation="latitude" pageId="1" pageNumber="512" precision="15" seconds="39" value="17.444166">17°2639″N</geoCoordinate>
<geoCoordinate box="[502,643,2061,2086]" degrees="104" direction="east" minutes="56" orientation="longitude" pageId="1" pageNumber="512" precision="15" seconds="55" value="104.94861">104°5655″E</geoCoordinate>
; collected
<date box="[113,262,2095,2119]" pageId="1" pageNumber="512" value="2013-06-09">
<collectingDate box="[113,262,2095,2119]" pageId="1" pageNumber="512" value="2013-06-09">09 June 2013</collectingDate>
</date>
, reared until
<date box="[408,555,2095,2119]" pageId="1" pageNumber="512" value="2013-10">
<collectingDate box="[408,555,2095,2119]" pageId="1" pageNumber="512" value="2013-10">
October
<quantity box="[502,555,2095,2119]" metricMagnitude="1" metricUnit="m" metricValue="5.11302" pageId="1" pageNumber="512" unit="in" value="2013.0">2013</quantity>
</collectingDate>
</date>
in aquaria.
</materialsCitation>
</paragraph>
<paragraph blockId="1.[813,1452,1326,1639]" pageId="1" pageNumber="512">
<materialsCitation accessionNumber="ZRC 62554" collectionCode="ZRC" country="Laos" location="Khammouan province" pageId="1" pageNumber="512" specimenCode="1" specimenCount="2" stateProvince="Khammouan" typeStatus="paratype">
<emphasis bold="true" box="[813,930,1327,1351]" pageId="1" pageNumber="512">
<typeStatus box="[813,930,1327,1351]" pageId="1" pageNumber="512">Paratypes</typeStatus>
</emphasis>
.
<collectingCountry box="[947,1004,1327,1350]" name="Laos" pageId="1" pageNumber="512">Laos</collectingCountry>
:
<collectingRegion box="[1018,1269,1327,1351]" country="Laos" name="Khammouan" pageId="1" pageNumber="512">Khammouan province</collectingRegion>
:
<accessionNumber box="[1283,1417,1326,1351]" httpUri="http://www.ncbi.nlm.nih.gov/protein/ZRC62554" pageId="1" pageNumber="512">
<collectionCode box="[1283,1335,1327,1351]" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="512">ZRC</collectionCode>
62554
</accessionNumber>
,
<specimenCode box="[1431,1444,1327,1351]" collectionCode="ZRC" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="512">1</specimenCode>
,
<quantity box="[813,910,1358,1382]" metricMagnitude="-2" metricUnit="m" metricValue="6.15" pageId="1" pageNumber="512" unit="mm" value="61.5">61.5 mm</quantity>
SL, collected and reared together with holotype
</materialsCitation>
;
<materialsCitation accessionNumber="ZRC 62555, CMK 19403" collectingDate="2018-03" collectingDateMax="2013-10" collectingDateMin="2013-06-09" collectionCode="ZRC" country="Laos" county="Thakhek district" latitude="17.444166" location="cave Tham Nang Eng" longLatPrecision="21" longitude="104.94861" municipality="Nam Thorn" pageId="1" pageNumber="512" specimenCode="ZRC 62555-62558, 7" specimenCount="1" stateProvince="Khammouan province" typeStatus="paratype">
<specimenCode box="[813,1023,1390,1415]" collectionCode="ZRC" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="512">
<accessionNumber box="[813,941,1390,1415]" httpUri="http://www.ncbi.nlm.nih.gov/protein/ZRC62555" pageId="1" pageNumber="512">ZRC 62555</accessionNumber>
62558
</specimenCode>
,
<specimenCode box="[1035,1050,1391,1414]" collectionCode="ZRC" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="512">7</specimenCode>
,
<quantity box="[1064,1220,1390,1414]" metricMagnitude="-2" metricUnit="m" metricValue="3.905" metricValueMax="5.04" metricValueMin="2.77" pageId="1" pageNumber="512" unit="mm" value="39.05" valueMax="50.4" valueMin="27.7">27.750.4 mm</quantity>
SL; same locality as holotype,
<date box="[925,1064,1423,1447]" pageId="1" pageNumber="512" value="2018-03">
<collectingDate box="[925,1064,1423,1447]" pageId="1" pageNumber="512" value="2018-03">March 2018</collectingDate>
</date>
. —
<accessionNumber box="[1115,1256,1423,1447]" httpUri="http://www.ncbi.nlm.nih.gov/protein/CMK19403" pageId="1" pageNumber="512">CMK 19403</accessionNumber>
</materialsCitation>
,
<materialsCitation accessionNumber="CMK 23170" collectingDate="2006-05-07" collectorName="M. Kottelat" country="Laos" county="Southwest of Gnommalat" latitude="17.541666" location="Phou Phathoung" longLatPrecision="21" longitude="105.166115" municipality="Tham Pong cave" pageId="1" pageNumber="512" specimenCount="1" stateProvince="Khammouan province" typeStatus="paratype">
1,
<quantity box="[1302,1402,1423,1447]" metricMagnitude="-2" metricUnit="m" metricValue="3.22" pageId="1" pageNumber="512" unit="mm" value="32.2">32.2 mm</quantity>
SL;
<collectingCounty box="[813,1108,1455,1479]" pageId="1" pageNumber="512">Southwest of Gnommalat</collectingCounty>
,
<collectingMunicipality box="[1123,1321,1455,1479]" pageId="1" pageNumber="512">Tham Pong cave</collectingMunicipality>
, near
<location LSID="urn:lsid:plazi:treatment:722E87D4AF71A448FF5BF8CAFD2CFB35:FF586019AF71A44DFAD2FA0BFC3DFA3F" country="Laos" county="Southwest of Gnommalat" latitude="17.541666" longLatPrecision="21" longitude="105.166115" municipality="Tham Pong cave" name="Phou Phathoung" pageId="1" pageNumber="512" stateProvince="Khammouan province">Phou Phathoung</location>
quarry, inside cave,
<geoCoordinate box="[1167,1296,1485,1510]" degrees="17" direction="north" minutes="32" orientation="latitude" pageId="1" pageNumber="512" precision="15" seconds="30" value="17.541666">17°3230″N</geoCoordinate>
<geoCoordinate box="[1306,1448,1485,1510]" degrees="105" direction="east" minutes="09" orientation="longitude" pageId="1" pageNumber="512" precision="15" seconds="58" value="105.166115">105°0958″E</geoCoordinate>
;
<collectorName box="[813,936,1519,1543]" pageId="1" pageNumber="512">M. Kottelat</collectorName>
et al.,
<date box="[1007,1140,1519,1543]" pageId="1" pageNumber="512" value="2006-05-07">
<collectingDate box="[1007,1140,1519,1543]" pageId="1" pageNumber="512" value="2006-05-07">7 May 2006</collectingDate>
</date>
. —
<accessionNumber box="[1183,1319,1519,1543]" httpUri="http://www.ncbi.nlm.nih.gov/protein/CMK23170" pageId="1" pageNumber="512">CMK 23170</accessionNumber>
</materialsCitation>
,
<materialsCitation collectingDate="2012-05-22" collectorName="M. Kottelat" country="Laos" county="Xe Bangfai" latitude="17.34889" location="Nam Oula" longLatPrecision="21" longitude="105.33528" municipality="Houay Kalo" pageId="1" pageNumber="512" specimenCount="1" stateProvince="Khammouan province" typeStatus="paratype">
1,
<quantity box="[1357,1451,1519,1543]" metricMagnitude="-2" metricUnit="m" metricValue="3.69" pageId="1" pageNumber="512" unit="mm" value="36.9">36.9 mm</quantity>
SL;
<collectingCounty box="[859,982,1551,1575]" pageId="1" pageNumber="512">Xe Bangfai</collectingCounty>
drainage:
<collectingMunicipality box="[1098,1231,1551,1575]" pageId="1" pageNumber="512">Houay Kalo</collectingMunicipality>
, a small tributary of
<location LSID="urn:lsid:plazi:treatment:722E87D4AF71A448FF5BF8CAFD2CFB35:FF586019AF71A44DFC8CF98BFC02F99F" box="[813,927,1583,1607]" country="Laos" county="Xe Bangfai" latitude="17.34889" longLatPrecision="21" longitude="105.33528" municipality="Houay Kalo" name="Nam Oula" pageId="1" pageNumber="512" stateProvince="Khammouan province">Nam Oula</location>
near
<location LSID="urn:lsid:plazi:treatment:722E87D4AF71A448FF5BF8CAFD2CFB35:FF586019AF71A44DFC40F98BFB1FF99F" box="[993,1154,1583,1607]" country="Laos" county="Xe Bangfai" latitude="17.34889" longLatPrecision="21" longitude="105.33528" municipality="Houay Kalo" name="Ban Phonexay" pageId="1" pageNumber="512" stateProvince="Khammouan province">Ban Phonexay</location>
,
<geoCoordinate box="[1168,1296,1581,1606]" degrees="17" direction="north" minutes="20" orientation="latitude" pageId="1" pageNumber="512" precision="15" seconds="56" value="17.34889">17°2056″N</geoCoordinate>
<geoCoordinate box="[1306,1447,1581,1606]" degrees="105" direction="east" minutes="20" orientation="longitude" pageId="1" pageNumber="512" precision="15" seconds="07" value="105.33528">105°2007″E</geoCoordinate>
;
<collectorName box="[813,938,1615,1639]" pageId="1" pageNumber="512">M. Kottelat</collectorName>
et al.,
<date box="[1015,1165,1615,1639]" pageId="1" pageNumber="512" value="2012-05-22">
<collectingDate box="[1015,1165,1615,1639]" pageId="1" pageNumber="512" value="2012-05-22">22 May 2012</collectingDate>
</date>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="1" pageNumber="512" type="diagnosis">
<paragraph blockId="1.[813,1453,1678,2119]" pageId="1" pageNumber="512">
<emphasis bold="true" box="[813,933,1678,1702]" pageId="1" pageNumber="512">Diagnosis.</emphasis>
<taxonomicName authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[944,1161,1679,1703]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="cacao">
<emphasis box="[944,1161,1679,1703]" italics="true" pageId="1" pageNumber="512">Nemacheilus cacao</emphasis>
</taxonomicName>
is distinguished from all other species of the genus except
<taxonomicName authorityName="Kottelat" authorityYear="1990" box="[1181,1308,1711,1735]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="platiceps">
<emphasis box="[1181,1197,1711,1734]" italics="true" pageId="1" pageNumber="512">N</emphasis>
.
<emphasis box="[1212,1308,1711,1735]" italics="true" pageId="1" pageNumber="512">platiceps</emphasis>
</taxonomicName>
in having an incomplete lateral line reaching between verticals of pelvicfin origin and of anus, with 3349 pores (vs. complete).
<taxonomicName authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[813,1029,1807,1831]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="cacao">
<emphasis box="[813,1029,1807,1831]" italics="true" pageId="1" pageNumber="512">Nemacheilus cacao</emphasis>
</taxonomicName>
is most easily distinguished from
<taxonomicName authorityName="Kottelat" authorityYear="1990" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="platiceps">
<emphasis italics="true" pageId="1" pageNumber="512">N. platiceps</emphasis>
</taxonomicName>
by its colour pattern, with a uniform dark brown body in adult males. In contrast,
<taxonomicName authorityName="Kottelat" authorityYear="1990" box="[1192,1327,1871,1895]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="platiceps">
<emphasis box="[1192,1210,1871,1894]" italics="true" pageId="1" pageNumber="512">N</emphasis>
.
<emphasis box="[1228,1327,1871,1895]" italics="true" pageId="1" pageNumber="512">platiceps</emphasis>
</taxonomicName>
has 1216 narrow bars on the flank, clearly distinct at all sizes and both sexes. In addition,
<taxonomicName authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[1079,1177,1935,1958]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="cacao">
<emphasis box="[1079,1097,1935,1958]" italics="true" pageId="1" pageNumber="512">N</emphasis>
.
<emphasis box="[1113,1177,1935,1958]" italics="true" pageId="1" pageNumber="512">cacao</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName authorityName="Kottelat" authorityYear="1990" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="1" pageNumber="512" phylum="Chordata" rank="species" species="platiceps">
<emphasis box="[1427,1445,1935,1958]" italics="true" pageId="1" pageNumber="512">N</emphasis>
.
<emphasis box="[813,909,1967,1991]" italics="true" pageId="1" pageNumber="512">platiceps</emphasis>
</taxonomicName>
by males having: a conspicuous suborbital flap (vs. poorly developed); dorsal surface of first 37 pectoral-fin rays covered by densely-set small tubercles; flank with a patch of scales each with a small tubercle (vs. absence of tubercles on pectoral fin and flanks).
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7399886" ID-Zenodo-Dep="7399886" httpUri="https://zenodo.org/record/7399886/files/figure.png" pageId="2" pageNumber="513" startId="2.[135,170,656,678]" targetBox="[135,1473,189,641]" targetPageId="2">
<paragraph blockId="2.[135,1474,656,707]" pageId="2" pageNumber="513">
Fig. 1. Live specimen of
<taxonomicName authority="Bohlen &amp; Kottelat &amp; Šlechtová, 2022" authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[374,561,656,678]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="2" pageNumber="513" phylum="Chordata" rank="species" species="cacao" status="new species">
<emphasis box="[374,561,656,678]" italics="true" pageId="2" pageNumber="513">Nemacheilus cacao</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[573,690,656,678]" pageId="2" pageNumber="513" rank="species">new species</taxonomicNameLabel>
, ZRC 62554, paratype, male, 61.5 mm SL; Laos: Khamouane province: Thakkhet district: small tributary of Nam Thorn at cave Tham Nang Eng (Photo: J. Kühne).
</paragraph>
</caption>
<subSubSection lastPageId="4" lastPageNumber="515" pageId="2" pageNumber="513" type="description">
<paragraph blockId="2.[134,777,762,1234]" pageId="2" pageNumber="513">
<emphasis bold="true" box="[136,282,762,786]" pageId="2" pageNumber="513">Description.</emphasis>
See
<figureCitation box="[344,424,762,786]" captionStart="Fig" captionStartId="2.[135,170,656,678]" captionTargetBox="[135,1473,189,641]" captionTargetId="figure-8@2.[132,1476,189,641]" captionTargetPageId="2" captionText="Fig. 1. Live specimen of Nemacheilus cacao, new species, ZRC 62554, paratype, male, 61.5 mm SL; Laos: Khamouane province: Thakkhet district: small tributary of Nam Thorn at cave Tham Nang Eng (Photo: J. Kühne)." figureDoi="http://doi.org/10.5281/zenodo.7399886" httpUri="https://zenodo.org/record/7399886/files/figure.png" pageId="2" pageNumber="513">Figs. 1</figureCitation>
and
<figureCitation box="[487,500,762,786]" captionStart="Fig" captionStartId="3.[113,148,2064,2086]" captionTargetBox="[180,1380,221,2031]" captionTargetId="figure-61@3.[151,1414,189,2049]" captionTargetPageId="3" captionText="Fig. 2. Nemacheilus cacao, new species, Laos: Khammouane prov: Thaket district: small tributary of Nam Thorn at cave Tham Nang Eng. a, ZRC 62553, holotype, 56.9 mm SL, adult male; b, ZRC 62558, paratype, 50.4 mm SL, adult female; c, ZRC 62556, 27.7 mm SL, juvenile." figureDoi="http://doi.org/10.5281/zenodo.7399888" httpUri="https://zenodo.org/record/7399888/files/figure.png" pageId="2" pageNumber="513">2</figureCitation>
for general appearance and
<tableCitation box="[185,269,794,818]" captionStart="Table 1" captionStartId="1.[113,168,189,211]" captionTargetPageId="1" captionText="Table 1. Morphometric data of holotype and eight paratypes (ZRC 62553 to 62558) of Nemacheilus cacao, new species. Range and mean include holotype data." httpUri="http://table.plazi.org/id/AEF8664AAF71A44DFFD0FF19FED6FF28" pageId="2" pageNumber="513" tableUuid="AEF8664AAF71A44DFFD0FF19FED6FF28">Table 1</tableCitation>
for morphometric data of
<typeStatus box="[570,664,794,818]" pageId="2" pageNumber="513">holotype</typeStatus>
and
<specimenCount pageId="2" pageNumber="513" type="generic" typeStatus="paratypes">eight paratypes</specimenCount>
. A moderately elongated nemacheilid loach with body depth about 57 times in SL. Body depth gradually increasing until dorsal-fin origin, then decreasing until posterior extremity of anal-fin base, uniform from there to caudal-fin base. Dorsal outline between nape and dorsalfin origin slightly convex. Head slightly depressed; body moderately compressed behind head, increasingly compressed until caudal-fin base. Interorbital area flat or nearly flat. In lateral view, upper margin of eye flush with dorsal profile of head. Cheeks not swollen; snout rounded. Depth of caudal peduncle 1.21.5 times in its length. Very short and low ventral crest on posterior end of caudal peduncle, absent or nearly absent dorsal crest. Largest recorded size
<quantity box="[641,733,1210,1234]" metricMagnitude="-2" metricUnit="m" metricValue="6.15" pageId="2" pageNumber="513" unit="mm" value="61.5">61.5 mm</quantity>
SL.
</paragraph>
<paragraph blockId="2.[135,776,1273,1682]" pageId="2" pageNumber="513">
Dorsal fin with 4 unbranched and 9½ branched rays, distal margin straight to slightly convex; second branched ray longest. Anal fin with 3 unbranched and 5½ branched rays, distal margin straight, second branched ray longest. Pectoral fin with 1 unbranched and 1112 branched rays, reaching halfway to pelvic-fin base; no axillary lobe. Pelvic fin with 1 unbranched and 7 (
<quantity box="[339,380,1466,1490]" metricMagnitude="-1" metricUnit="m" metricValue="1.524" pageId="2" pageNumber="513" unit="in" value="6.0">6 in</quantity>
<specimenCount box="[386,508,1466,1490]" pageId="2" pageNumber="513" type="generic">1 specimen</specimenCount>
) branched rays, reaching half-way to anal-fin base, not or just reaching anus, which is located 1½2 eye diameter in front of anal-fin origin: axillary lobe small and on whole length attached to body. Caudal fin with 9+8 (8+
<quantity box="[412,456,1594,1618]" metricMagnitude="-1" metricUnit="m" metricValue="2.032" pageId="2" pageNumber="513" unit="in" value="8.0">8 in</quantity>
<specimenCount box="[467,595,1594,1618]" pageId="2" pageNumber="513" type="generic">1 specimen</specimenCount>
) branched rays, slightly forked, lobes rounded, both lobes same length or upper lobe up to 1.1 times length of lower lobe.
</paragraph>
<paragraph blockId="2.[136,776,1722,1906]" pageId="2" pageNumber="513">Body entirely scaled. Lateral line incomplete, ending usually below or shortly behind dorsal-fin base, occasionally reaching level of anal-fin origin; visible pores (3349) only until vertical of anus. Cephalic lateral line system with 5 supraorbital, 4+1011 infraorbital, 910 preoperculomandibular and 3 supratemporal pores.</paragraph>
<paragraph blockId="2.[136,776,1946,2098]" lastBlockId="2.[835,1476,762,882]" pageId="2" pageNumber="513">
Anterior nare pierced at tip of an obliquely truncate tube. Posterior nare adjacent to anterior one. Mouth moderately arched, gape about twice wider than long (
<figureCitation box="[615,684,2010,2034]" captionStart="Fig" captionStartId="4.[135,170,683,705]" captionTargetBox="[135,783,189,668]" captionTargetId="figure-8@4.[135,775,186,668]" captionTargetPageId="4" captionText="Fig. 3. Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL; mouth in ventral view." figureDoi="http://doi.org/10.5281/zenodo.7399890" httpUri="https://zenodo.org/record/7399890/files/figure.png" pageId="2" pageNumber="513">Fig. 3</figureCitation>
). Upper lip moderately thick, without median notch, with numerous wrinkles. Processus dentiformis present. Lower lip with narrow median interruption; median part with very weakly marked sulci; thinner towards rictus. Barbels long; inner rostral barbel reaching beyond vertical through middle of eye, outer one and mandibular barbels reaching beyond eye.
</paragraph>
<paragraph blockId="2.[835,1474,922,1138]" pageId="2" pageNumber="513">
<emphasis bold="true" box="[835,1080,922,946]" pageId="2" pageNumber="513">Sexual dimorphism.</emphasis>
Adult males with suborbital flap (
<figureCitation box="[843,914,954,978]" captionStart="Fig" captionStartId="4.[835,870,683,705]" captionTargetBox="[826,1474,189,668]" captionTargetId="figure-9@4.[835,1474,189,668]" captionTargetPageId="4" captionText="Fig. 4. Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL; left suborbital flap in lateral view." figureDoi="http://doi.org/10.5281/zenodo.7502275" httpUri="https://zenodo.org/record/7502275/files/figure.png" pageId="2" pageNumber="513">Fig. 4</figureCitation>
) and with uniform brown flank (vs. females and juveniles with narrow brown bars). Adult males (= males with suborbital flap;
<quantity box="[1059,1174,1018,1042]" metricMagnitude="-2" metricUnit="m" metricValue="5.5" metricValueMax="6.2" metricValueMin="4.8" pageId="2" pageNumber="513" unit="mm" value="55.0" valueMax="62.0" valueMin="48.0">4862 mm</quantity>
SL) with patch of tubercles on flank under dorsal-fin base, on upper surface of pectoralfin rays, and scattered tubercles on postorbital area of head and nape (
<figureCitation box="[949,1016,1114,1138]" captionStart="Fig" captionStartId="5.[113,148,1791,1813]" captionTargetBox="[113,1451,433,1775]" captionTargetId="figure-14@5.[113,1451,433,1775]" captionTargetPageId="5" captionText="Fig. 5. Tubercles on body side and pectoral fin of large male Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL. A, red outline depicts field of tubercles on body side; BC, area of main tuberculation enlarged; D, dorsal view on right pectoral fin; E, tubercles on pectoral fin enlarged." figureDoi="http://doi.org/10.5281/zenodo.7399892" httpUri="https://zenodo.org/record/7399892/files/figure.png" pageId="2" pageNumber="513">Fig. 5</figureCitation>
).
</paragraph>
<paragraph blockId="2.[835,1477,1178,1714]" pageId="2" pageNumber="513">
Extent of patches of tubercles on flank variable, reaching longitudinally from tip of adpressed pectoral fin to end of anal-fin base, single tubercles even until caudal-fin base. Shape of patch irregular, usually elongated oval (
<figureCitation box="[1381,1462,1274,1298]" captionStart="Fig" captionStartId="5.[113,148,1791,1813]" captionTargetBox="[113,1451,433,1775]" captionTargetId="figure-14@5.[113,1451,433,1775]" captionTargetPageId="5" captionText="Fig. 5. Tubercles on body side and pectoral fin of large male Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL. A, red outline depicts field of tubercles on body side; BC, area of main tuberculation enlarged; D, dorsal view on right pectoral fin; E, tubercles on pectoral fin enlarged." figureDoi="http://doi.org/10.5281/zenodo.7399892" httpUri="https://zenodo.org/record/7399892/files/figure.png" pageId="2" pageNumber="513">Fig. 5a</figureCitation>
). In dorsoventral direction, patch stretching across 5 rows of scales above lateral line and up to 10 rows below, highest under dorsal fin and shortly behind. In this patch, each scale with a single large, white and triangular tubercle (
<figureCitation captionStart="Fig" captionStartId="5.[113,148,1791,1813]" captionTargetBox="[113,1451,433,1775]" captionTargetId="figure-14@5.[113,1451,433,1775]" captionTargetPageId="5" captionText="Fig. 5. Tubercles on body side and pectoral fin of large male Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL. A, red outline depicts field of tubercles on body side; BC, area of main tuberculation enlarged; D, dorsal view on right pectoral fin; E, tubercles on pectoral fin enlarged." figureDoi="http://doi.org/10.5281/zenodo.7399892" httpUri="https://zenodo.org/record/7399892/files/figure.png" pageId="2" pageNumber="513">Fig. 5b, c</figureCitation>
). Tubercles on pectoral fin much smaller than those on body, white, conical, in 15 rows on dorsal side of rays and on unculiferous pad behind; stretching to tip of ray on unbranched and first branched ray, with decreasing length on following branched rays until only on base of third-last ray (
<figureCitation box="[888,996,1594,1618]" captionStart="Fig" captionStartId="5.[113,148,1791,1813]" captionTargetBox="[113,1451,433,1775]" captionTargetId="figure-14@5.[113,1451,433,1775]" captionTargetPageId="5" captionText="Fig. 5. Tubercles on body side and pectoral fin of large male Nemacheilus cacao, new species, ZRC 62553, holotype, 56.9 mm SL. A, red outline depicts field of tubercles on body side; BC, area of main tuberculation enlarged; D, dorsal view on right pectoral fin; E, tubercles on pectoral fin enlarged." figureDoi="http://doi.org/10.5281/zenodo.7399892" httpUri="https://zenodo.org/record/7399892/files/figure.png" pageId="2" pageNumber="513">Fig. 5d, e</figureCitation>
). Last two branched rays without tubercles. Tubercles on sides of head small, white, conical, only tip pointing out of skin. Tubercles on body, pectoral fin, and head either all present or all absent.
</paragraph>
<paragraph blockId="2.[835,1476,1754,2098]" lastBlockId="4.[135,776,789,1069]" lastPageId="4" lastPageNumber="515" pageId="2" pageNumber="513">
<emphasis bold="true" box="[835,980,1754,1778]" pageId="2" pageNumber="513">Colouration.</emphasis>
<taxonomicName authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[990,1202,1754,1778]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="2" pageNumber="513" phylum="Chordata" rank="species" species="cacao">
<emphasis box="[990,1202,1754,1778]" italics="true" pageId="2" pageNumber="513">Nemacheilus cacao</emphasis>
</taxonomicName>
undergoes a remarkable ontogenetic and sexual change of colouration. A juvenile of
<quantity box="[867,964,1818,1842]" metricMagnitude="-2" metricUnit="m" metricValue="2.77" pageId="2" pageNumber="513" unit="mm" value="27.7">27.7 mm</quantity>
SL had 22 dark brown bars on body meeting their contralaterals on back, their width at least twice width of interspaces, some vertically split. In females, most bars split and fade with increasing size, resulting in a pattern of numerous narrow brown bars along body and broader dorsal saddles. In adult males, bars and saddles absent and whole fish with uniform chocolate brown body and head, only ventral side of head and body white. A faint dark stripe from snout to eye in most specimens, no black bar on caudal-fin base.
<emphasis bold="true" box="[136,391,789,813]" pageId="4" pageNumber="515">Phylogenetic position.</emphasis>
The topologies of the single gene trees were congruent; therefore, the data have been combined.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7399888" ID-Zenodo-Dep="7399888" httpUri="https://zenodo.org/record/7399888/files/figure.png" pageId="3" pageNumber="514" startId="3.[113,148,2064,2086]" targetBox="[180,1380,221,2031]" targetPageId="3">
<paragraph blockId="3.[113,1453,2064,2115]" pageId="3" pageNumber="514">
Fig. 2.
<taxonomicName authority="Bohlen &amp; Kottelat &amp; Šlechtová, 2022" authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[180,366,2064,2086]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="514" phylum="Chordata" rank="species" species="cacao" status="new species">
<emphasis box="[180,366,2064,2086]" italics="true" pageId="3" pageNumber="514">Nemacheilus cacao</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[377,495,2064,2086]" pageId="3" pageNumber="514" rank="species">new species</taxonomicNameLabel>
, Laos: Khammouane prov: Thaket district: small tributary of Nam Thorn at cave Tham Nang Eng. a, ZRC 62553, holotype, 56.9 mm SL, adult male; b, ZRC 62558, paratype, 50.4 mm SL, adult female; c, ZRC 62556, 27.7 mm SL, juvenile.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7399890" ID-Zenodo-Dep="7399890" httpUri="https://zenodo.org/record/7399890/files/figure.png" pageId="4" pageNumber="515" startId="4.[135,170,683,705]" targetBox="[135,783,189,668]" targetPageId="4">
<paragraph blockId="4.[135,776,683,734]" pageId="4" pageNumber="515">
Fig. 3.
<taxonomicName authority="Bohlen &amp; Kottelat &amp; Šlechtová, 2022" authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[210,403,683,705]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="515" phylum="Chordata" rank="species" species="cacao" status="new species">
<emphasis box="[210,403,683,705]" italics="true" pageId="4" pageNumber="515">Nemacheilus cacao</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[418,540,683,705]" pageId="4" pageNumber="515" rank="species">new species</taxonomicNameLabel>
, ZRC 62553, holotype, 56.9 mm SL; mouth in ventral view.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.7502275" ID-Zenodo-Dep="7502275" httpUri="https://zenodo.org/record/7502275/files/figure.png" pageId="4" pageNumber="515" startId="4.[835,870,683,705]" targetBox="[826,1474,189,668]" targetPageId="4">
<paragraph blockId="4.[835,1474,683,734]" pageId="4" pageNumber="515">
Fig. 4.
<taxonomicName authority="Bohlen &amp; Kottelat &amp; Šlechtová, 2022" authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[910,1102,683,705]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="515" phylum="Chordata" rank="species" species="cacao" status="new species">
<emphasis box="[910,1102,683,705]" italics="true" pageId="4" pageNumber="515">Nemacheilus cacao</emphasis>
</taxonomicName>
,
<taxonomicNameLabel box="[1117,1238,683,705]" pageId="4" pageNumber="515" rank="species">new species</taxonomicNameLabel>
, ZRC 62553, holotype, 56.9 mm SL; left suborbital flap in lateral view.
</paragraph>
</caption>
<paragraph blockId="4.[135,776,789,1069]" pageId="4" pageNumber="515">
The analysis of the concatenated dataset estimated the phylogenetic position of
<taxonomicName authorityName="Bohlen &amp; Kottelat &amp; Šlechtová" authorityYear="2022" box="[402,497,885,909]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="515" phylum="Chordata" rank="species" species="cacao">
<emphasis box="[402,420,886,909]" italics="true" pageId="4" pageNumber="515">N</emphasis>
.
<emphasis box="[433,497,886,909]" italics="true" pageId="4" pageNumber="515">cacao</emphasis>
</taxonomicName>
as a member of the genus
<taxonomicName box="[136,274,917,941]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="515" phylum="Chordata" rank="genus">
<emphasis box="[136,274,917,941]" italics="true" pageId="4" pageNumber="515">Nemacheilus</emphasis>
</taxonomicName>
, member of the Selangoricus clade and sister species to
<taxonomicName authorityName="Kottelat" authorityYear="1990" box="[246,373,949,973]" class="Actinopterygii" family="Nemacheilidae" genus="Nemacheilus" kingdom="Animalia" order="Cypriniformes" pageId="4" pageNumber="515" phylum="Chordata" rank="species" species="platiceps">
<emphasis box="[246,264,950,973]" italics="true" pageId="4" pageNumber="515">N</emphasis>
.
<emphasis box="[277,373,949,973]" italics="true" pageId="4" pageNumber="515">platiceps</emphasis>
</taxonomicName>
(
<figureCitation box="[387,453,949,973]" captionStart="Fig" captionStartId="6.[136,171,1660,1682]" captionTargetBox="[136,1476,547,1645]" captionTargetId="figure-7@6.[136,1476,547,1645]" captionTargetPageId="6" captionText="Fig. 6. Bayesian tree of the genus Nemacheilus based on the concatenated datasets of mitochondrial cytochrome b and nuclear RAG-1 marker. Datasets from Šlechtová et al. (2021) and Sember et al. (2015) plus two specimens of Nemacheilus cacao, new species. The tree shows N. cacao to be the sister species to N. platiceps and both to be located at the base of the Selangoricus clade within Nemacheilus. The outgroup contains 25 nemacheilid species from genera other than Nemacheilus and one species from the family Cobitidae." figureDoi="http://doi.org/10.5281/zenodo.7399894" httpUri="https://zenodo.org/record/7399894/files/figure.png" pageId="4" pageNumber="515">Fig. 6</figureCitation>
). Genbank accession numbers for the sequences of the
<typeStatus box="[392,485,981,1005]" pageId="4" pageNumber="515">holotype</typeStatus>
(
<accessionNumber box="[498,625,981,1005]" httpUri="http://www.ncbi.nlm.nih.gov/protein/ZRC62553" pageId="4" pageNumber="515">ZRC 62553</accessionNumber>
) and
<typeStatus box="[683,775,981,1005]" pageId="4" pageNumber="515">paratype</typeStatus>
(
<accessionNumber box="[143,269,1013,1037]" httpUri="http://www.ncbi.nlm.nih.gov/protein/ZRC62554" pageId="4" pageNumber="515">ZRC 62554</accessionNumber>
) are:
<accessionNumber box="[326,444,1013,1037]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/ON720270" pageId="4" pageNumber="515">ON720270</accessionNumber>
and
<accessionNumber box="[492,610,1013,1037]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/ON720269" pageId="4" pageNumber="515">ON720269</accessionNumber>
for cytochrome
<emphasis box="[136,149,1045,1069]" italics="true" pageId="4" pageNumber="515">b</emphasis>
,
<accessionNumber box="[165,283,1045,1069]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/ON720272" pageId="4" pageNumber="515">ON720272</accessionNumber>
and
<accessionNumber box="[339,458,1045,1069]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/ON720271" pageId="4" pageNumber="515">ON720271</accessionNumber>
for RAG-1.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="515" type="etymology">
<paragraph blockId="4.[135,775,1109,1261]" pageId="4" pageNumber="515">
<emphasis bold="true" box="[136,266,1109,1133]" pageId="4" pageNumber="515">Etymology.</emphasis>
From
<taxonomicName box="[340,535,1109,1133]" class="Magnoliopsida" family="Malvaceae" genus="Theobroma" kingdom="Plantae" order="Malvales" pageId="4" pageNumber="515" phylum="Tracheophyta" rank="species" species="cacao">
<emphasis box="[340,535,1109,1133]" italics="true" pageId="4" pageNumber="515">Theobroma cacao</emphasis>
</taxonomicName>
, the cacao tree whose seeds are used to produce chocolate; itself derived from kakawa in some ancient Mesoamerican language (
<bibRefCitation author="Kaufman T &amp; Justeson J" pageId="4" pageNumber="515" pagination="193 - 237" refId="ref3869" refString="Kaufman T &amp; Justeson J (2007) The history of the word for Cacao in ancient Meso-America. Ancient Mesoamerica, 18 (2): 193 - 237." type="journal article" year="2007">Kaufman &amp; Justeson, 2007</bibRefCitation>
). A reference to the chocolate brown colour of large males. A noun in apposition, indeclinable.
</paragraph>
</subSubSection>
</treatment>
</document>