treatments-xml/data/03/EC/87/03EC87C9FF92FFA1FE9FFBFCFB20F241.xml

<document id="4FB2C35C572DED62BCF655D9FB31237D" ID-DOI="10.5281/zenodo.4504614" ID-ISSN="2345-7600" ID-Zenodo-Dep="4504614" ID-ZooBank="DF8D43A5-742E-4571-9838-DC0935576F45" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1612466763449" checkinUser="carolina" docAuthor="Hui, Tan Heok &amp; Ahmad, Amirrudin Bin" docDate="2018" docId="03EC87C9FF92FFA1FE9FFBFCFB20F241" docLanguage="en" docName="RafflesBZool.66.402-407.pdf" docOrigin="Raffles Bulletin of Zoology 66" docStyle="DocumentStyle:F2D34FF25DBF9998BB8DF6F24A5B097A.5:RafflesBZool.2014-2018.journal_article" docStyleId="F2D34FF25DBF9998BB8DF6F24A5B097A" docStyleName="RafflesBZool.2014-2018.journal_article" docStyleVersion="5" docTitle="Betta omega Hui &amp; Ahmad 2018, new species" docType="treatment" docVersion="4" lastPageNumber="406" masterDocId="FFD5FFB1FF93FFA5FFA8FFBCFF99F51B" masterDocTitle="Betta omega, a new species of black water fighting fish (Teleostei: Osphronemidae) from Malaysia" masterLastPageNumber="407" masterPageNumber="402" pageNumber="403" updateTime="1698866299707" updateUser="plazi" zenodo-license-document="CC-BY-4.0">
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<mods:title id="F72532AEADA72817FCDEA00310D7EAF7">Betta omega, a new species of black water fighting fish (Teleostei: Osphronemidae) from Malaysia</mods:title>
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<mods:namePart id="01DBB2F2156574259927F503356C0019">Hui, Tan Heok</mods:namePart>
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<mods:namePart id="5BE28BF66AD44F8234F0CA140EF8F51D">Ahmad, Amirrudin Bin</mods:namePart>
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<mods:title id="1CA51CF95D832C5ADE4360CA913E856A">Raffles Bulletin of Zoology</mods:title>
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<mods:date id="39DC8356B4A0C97EF249B246AF12948F">2018</mods:date>
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<heading id="D0B281B3FF92FFA4FE9FFBFCFDCEF143" bold="true" box="[311,599,1088,1112]" centered="true" fontSize="10" level="2" pageId="1" pageNumber="403" reason="2">
<emphasis id="B931EACDFF92FFA4FE9FFBFCFDCEF143" bold="true" box="[311,599,1088,1112]" pageId="1" pageNumber="403">
<taxonomicName id="4C454D5CFF92FFA4FE9FFBFCFE5AF14C" authority="Hui &amp; Ahmad, 2018" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[311,451,1088,1111]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="omega" status="sp. nov.">
<emphasis id="B931EACDFF92FFA4FE9FFBFCFE5AF14C" bold="true" box="[311,451,1088,1111]" italics="true" pageId="1" pageNumber="403">Betta omega</emphasis>
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, 
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<paragraph id="8BFA36DFFF92FFA4FE24FBDCFD9AF163" blockId="1.[311,599,1088,1144]" box="[396,515,1120,1144]" pageId="1" pageNumber="403">
(
<figureCitation id="137E2A5AFF92FFA4FE3BFBDCFE63F163" box="[403,506,1120,1144]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStartId-0="1.[136,171,947,969]" captionStartId-1="2.[114,149,2058,2080]" captionStartId-2="3.[137,173,1028,1050]" captionTargetBox-0="[133,1477,189,932]" captionTargetBox-1="[114,1452,1458,2052]" captionTargetBox-2="[136,775,189,1012]" captionTargetId-0="figure-218@1.[133,1477,189,932]" captionTargetId-1="figure-12@2.[114,1452,1458,2052]" captionTargetId-2="figure-736@3.[136,775,189,1012]" captionTargetPageId-0="1" captionTargetPageId-1="2" captionTargetPageId-2="3" captionText-0="Fig. 1. Betta omega, new species – ZRC 59663, 70.0 mm SL holotype, life colouration (photograph by Z. Zakaria)." captionText-1="Fig. 2. Betta omega, new species – ZRC 59663, 70.0 mm SL holotype, preserved specimen." captionText-2="Fig. 3. Betta omega, new species – ZRC 59664, 74.0 mm SL paratype, front view of head showing lower jaw and throat pattern (photograph by Z. Zakaria)." figureDoi-0="http://doi.org/10.5281/zenodo.4504616" figureDoi-1="http://doi.org/10.5281/zenodo.4504618" figureDoi-2="http://doi.org/10.5281/zenodo.4504620" httpUri-0="https://zenodo.org/record/4504616/files/figure.png" httpUri-1="https://zenodo.org/record/4504618/files/figure.png" httpUri-2="https://zenodo.org/record/4504620/files/figure.png" pageId="1" pageNumber="403">Figs. 1–3</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C35F6554FF92FFA4FF20FB23FDC8F1CA" pageId="1" pageNumber="403" type="reference_group">
<paragraph id="8BFA36DFFF92FFA4FF20FB23FDC8F1CA" blockId="1.[136,774,1182,1233]" pageId="1" pageNumber="403">
<treatmentCitationGroup id="AB5511F1FF92FFA4FF20FB23FDC8F1CA" pageId="1" pageNumber="403">
<taxonomicName id="4C454D5CFF92FFA4FF20FB23FEC6F1A8" ID-CoL="5WJYP" authorityName="Regan" authorityYear="1910" box="[136,351,1182,1204]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="macrophthalma">
<emphasis id="B931EACDFF92FFA4FF20FB23FEC6F1A8" box="[136,351,1182,1204]" italics="true" pageId="1" pageNumber="403">Betta macrophthalma</emphasis>
</taxonomicName>
(non-Regan, 1910) – 
<treatmentCitation id="0AE410CEFF92FFA4FDE9FB22FC9FF1AF" author="Schmidt J" box="[577,774,1182,1204]" page="341" pageId="1" pageNumber="403" year="1988">
<bibRefCitation id="EFD44B2EFF92FFA4FDE9FB22FC9FF1AF" author="Schmidt J" box="[577,774,1182,1204]" pageId="1" pageNumber="403" pagination="341 - 344" refId="ref3345" refString="Schmidt J (1988) Wasers grosser maulbrutender Kampffisch Betta macrophthalma Regan, 1910. Aquarien und Terrarien, 41: 341 - 344." type="journal article" year="1988">Schmidt, 1988: 341</bibRefCitation>
</treatmentCitation>
(illustrated in 
<bibRefCitation id="EFD44B2EFF92FFA4FE90FB07FE66F1CB" author="Kottelat M &amp; Whitten AJ &amp; Kartikasari SN &amp; Wirjoatmodjo S" box="[312,511,1211,1233]" pageId="1" pageNumber="403" refId="ref3222" refString="Kottelat M, Whitten AJ, Kartikasari SN &amp; Wirjoatmodjo S (1993) Freshwater Fishes of Western Indonesia and Sulawesi. Periplus Editions, Hong Kong, 259 pp., 84 pls." type="book" year="1993">Kottelat et al., 1993</bibRefCitation>
: pl. 77).
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C35F6554FF92FFA4FF20FB45FC98F389" pageId="1" pageNumber="403" type="materials_examined">
<paragraph id="8BFA36DFFF92FFA4FF20FB45FD38F04A" blockId="1.[136,774,1273,1362]" pageId="1" pageNumber="403">
<emphasis id="B931EACDFF92FFA4FF20FB45FEE8F00A" bold="true" box="[136,369,1273,1297]" pageId="1" pageNumber="403">Material examined.</emphasis>
– 
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<typeStatus id="54FE887DFF92FFA4FE3CFB45FE66F00A" box="[404,511,1273,1297]" pageId="1" pageNumber="403" type="holotype">Holotype</typeStatus>
: 
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59663, 70.0 mm SL; Peninsular 
<collectingCountry id="F352764FFF92FFA4FE98FAA5FE0FF02A" box="[304,406,1305,1329]" name="Malaysia" pageId="1" pageNumber="403">Malaysia</collectingCountry>
: 
<collectingRegion id="4981F83DFF92FFA4FE09FAA5FE79F02A" box="[417,480,1305,1329]" country="Malaysia" name="Johor" pageId="1" pageNumber="403">Johor</collectingRegion>
: 
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, Sungai Burong near Jeram Batu; 
<collectorName id="26B05309FF92FFA4FE0CFA86FD9AF04A" box="[420,515,1338,1361]" pageId="1" pageNumber="403">J. J. Foo</collectorName>
, 
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<collectingDate id="EFBFE9F7FF92FFA4FDB8FA86FD04F04A" box="[528,669,1337,1362]" pageId="1" pageNumber="403" value="2016-02-19">19 Feb 2016</collectingDate>
</date>
.
</materialsCitation>
</paragraph>
<paragraph id="8BFA36DFFF92FFA4FF20FAC6FD80F0E9" blockId="1.[136,774,1401,1522]" pageId="1" pageNumber="403">
<materialsCitation id="3B2D3C82FF92FFA4FF20FAC6FEA6F0AA" accessionNumber="ZRC 59664" collectionCode="ZRC" pageId="1" pageNumber="403" specimenCount="1" typeStatus="paratype">
<typeStatus id="54FE887DFF92FFA4FF20FAC6FF6DF089" box="[136,244,1402,1426]" pageId="1" pageNumber="403" type="paratype">Paratypes</typeStatus>
: 
<collectionCode id="ED54AE1AFF92FFA4FEA9FAC6FEACF089" box="[257,309,1402,1426]" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="403">ZRC</collectionCode>
59664, 
<specimenCount id="9D43FD56FF92FFA4FE3AFAC6FE52F08A" box="[402,459,1402,1426]" pageId="1" pageNumber="403" type="generic">1 ex.</specimenCount>
, 74.0 mm SL; same locality data as 
<typeStatus id="54FE887DFF92FFA4FF76FA25FEA6F0AA" box="[222,319,1433,1457]" pageId="1" pageNumber="403" type="holotype">holotype</typeStatus>
</materialsCitation>
. – 
<materialsCitation id="3B2D3C82FF92FFA4FECBFA26FD8FF0E9" accessionNumber="ZRC 59665" collectionCode="ZRC" collectorName="Allan &amp; Barbara Brown" country="Malaysia" location="Pekan Nanas area" pageId="1" pageNumber="403" specimenCount="3" stateProvince="Johor" typeStatus="paratype">
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59665, 
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, 
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SL; Peninsular 
<collectingCountry id="F352764FFF92FFA4FEAFFA05FEE9F0CA" box="[263,368,1465,1489]" name="Malaysia" pageId="1" pageNumber="403">Malaysia</collectingCountry>
: 
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: 
<location id="8E9A6004FF92FFA4FE65FA05FD03F0CA" LSID="urn:lsid:plazi:treatment:03EC87C9FF92FFA1FE9FFBFCFB20F241:8E9A6004FF92FFA4FE65FA05FD03F0CA" box="[461,666,1465,1489]" country="Malaysia" name="Pekan Nanas area" pageId="1" pageNumber="403" stateProvince="Johor">Pekan Nanas area</location>
; 
<collectorName id="26B05309FF92FFA4FD01FA05FD7FF0CA" box="[681,742,1465,1489]" pageId="1" pageNumber="403">Allan</collectorName>
&amp; 
<collectorName id="26B05309FF92FFA4FF20FA65FEA9F0EA" box="[136,304,1497,1521]" pageId="1" pageNumber="403">Barbara Brown</collectorName>
, (preserved in) 1984
</materialsCitation>
.
</paragraph>
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Non-type material: 
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<collectionCode id="ED54AE1AFF92FFA4FEF4F9A6FE09F329" box="[348,400,1562,1586]" country="Singapore" httpUri="http://grbio.org/cool/16z4-1tbx" name="Zoological Reference Collection, National University of Singapore" pageId="1" pageNumber="403">ZRC</collectionCode>
59666, 
<specimenCount id="9D43FD56FF92FFA4FE41F9A6FDB7F32A" box="[489,558,1562,1586]" pageId="1" pageNumber="403" type="generic">14 ex.</specimenCount>
, 
<quantity id="4CBD9B3AFF92FFA4FD93F9A5FD4EF32A" box="[571,727,1561,1585]" metricMagnitude="-2" metricUnit="m" metricValue="3.2600000000000002" metricValueMax="3.5100000000000002" metricValueMin="3.0100000000000002" pageId="1" pageNumber="403" unit="mm" value="32.6" valueMax="35.1" valueMin="30.1">30.1–35.1 mm</quantity>
SL
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; 
<materialsCitation id="3B2D3C82FF92FFA4FF22F986FD64F389" accessionNumber="ZRC 59667" collectionCode="ZRC" collectorName="Allan &amp; Barbara Brown" country="Malaysia" location="Pekan Nanas area" pageId="1" pageNumber="403" specimenCount="9" stateProvince="Johor">
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59667, 
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, 
<quantity id="4CBD9B3AFF92FFA4FEFFF985FE6BF34A" box="[343,498,1593,1617]" metricMagnitude="-2" metricUnit="m" metricValue="7.435" metricValueMax="7.92" metricValueMin="6.95" pageId="1" pageNumber="403" unit="mm" value="74.35" valueMax="79.2" valueMin="69.5">69.5–79.2 mm</quantity>
SL; Peninsular 
<collectingCountry id="F352764FFF92FFA4FD36F985FC9DF34A" box="[670,772,1593,1617]" name="Malaysia" pageId="1" pageNumber="403">Malaysia</collectingCountry>
: 
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: 
<location id="8E9A6004FF92FFA4FF7DF9E5FE05F36A" LSID="urn:lsid:plazi:treatment:03EC87C9FF92FFA1FE9FFBFCFB20F241:8E9A6004FF92FFA4FF7DF9E5FE05F36A" box="[213,412,1625,1649]" country="Malaysia" name="Pekan Nanas area" pageId="1" pageNumber="403" stateProvince="Johor">Pekan Nanas area</location>
; F1 generation from wild caught individuals; 
<collectorName id="26B05309FF92FFA4FEA7F9C5FED3F38A" box="[271,330,1657,1681]" pageId="1" pageNumber="403">Allan</collectorName>
&amp; 
<collectorName id="26B05309FF92FFA4FED8F9C5FD81F38A" box="[368,536,1657,1681]" pageId="1" pageNumber="403">Barbara Brown</collectorName>
, (preserved in) 1988
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.
</paragraph>
</subSubSection>
<subSubSection id="C35F6554FF92FFA4FF20F905FA3CF1A3" pageId="1" pageNumber="403" type="diagnosis">
<paragraph id="8BFA36DFFF92FFA4FF20F905FD71F2EA" blockId="1.[134,775,1721,2033]" pageId="1" pageNumber="403">
<emphasis id="B931EACDFF92FFA4FF20F905FF66F3CA" bold="true" box="[136,255,1721,1745]" pageId="1" pageNumber="403">Diagnosis.</emphasis>
<taxonomicName id="4C454D5CFF92FFA4FEA1F906FE0CF3CA" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[265,405,1722,1745]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF92FFA4FEA1F906FE0CF3CA" box="[265,405,1722,1745]" italics="true" pageId="1" pageNumber="403">Betta omega</emphasis>
</taxonomicName>
can be distinguished from other members of the 
<taxonomicName id="4C454D5CFF92FFA4FEE9F966FE32F3EA" authorityName="Krummenacher" authorityYear="1986" box="[321,427,1754,1777]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="waseri">
<emphasis id="B931EACDFF92FFA4FEE9F966FE32F3EA" box="[321,427,1754,1777]" italics="true" pageId="1" pageNumber="403">B. waseri</emphasis>
</taxonomicName>
group in having the following combination of characters: black markings on lower jaw continuous with two black downward curved bars on throat, ending with rounded bulging ends near edge of lower jaw (vertical line projection downwards from lower jaw posterior end; see 
<figureCitation id="137E2A5AFF92FFA4FF45F8C5FEA3F28A" box="[237,314,1913,1937]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="1" pageNumber="403">Fig. 4i</figureCitation>
); opercle uniform brown with light dark brown mottling, operculum without lower distal margin black; black transverse bars on the dorsal and caudal fin interradial membranes; absence of a dark distal border on anal fin.
</paragraph>
<paragraph id="8BFA36DFFF92FFA4FCEBFBBDFA3CF1A3" blockId="1.[834,1475,1024,1208]" pageId="1" pageNumber="403">
<taxonomicName id="4C454D5CFF92FFA4FCEBFBBDFC48F103" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[835,977,1025,1048]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF92FFA4FCEBFBBDFC48F103" box="[835,977,1025,1048]" italics="true" pageId="1" pageNumber="403">Betta omega</emphasis>
</taxonomicName>
differs from its putative closest relative 
<taxonomicName id="4C454D5CFF92FFA4FA03FBBDFC54F123" authorityName="P.K.L.Ng &amp; Kottelat" authorityYear="1994" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="hipposideros">
<emphasis id="B931EACDFF92FFA4FA03FBBDFC54F123" italics="true" pageId="1" pageNumber="403">B. hipposideros</emphasis>
</taxonomicName>
in having an omega (Ω) pattern on throat (vs. downward slender black bars, resembling a horseshoe shape), lateral scale 15 below the dorsal-fin origin (vs. mode 16), lateral scale 6 above the anal-fin origin (vs. mode 7) and postdorsal scales 10–12, mode 11 (vs. 9–10, mode 9½).
</paragraph>
</subSubSection>
<subSubSection id="C35F6554FF92FFA6FCEBFB5CFA5DF46E" lastPageId="3" lastPageNumber="405" pageId="1" pageNumber="403" type="description">
<paragraph id="8BFA36DFFF92FFA4FCEBFB5CFB73F383" blockId="1.[835,1476,1248,1689]" pageId="1" pageNumber="403">
<emphasis id="B931EACDFF92FFA4FCEBFB5CFC57F1E3" bold="true" box="[835,974,1248,1272]" pageId="1" pageNumber="403">Description.</emphasis>
General body form as in 
<figureCitation id="137E2A5AFF92FFA4FB59FB5CFAA4F1E3" box="[1265,1341,1248,1272]" captionStart="Fig" captionStartId="1.[136,171,947,969]" captionTargetBox="[133,1477,189,932]" captionTargetId="figure-218@1.[133,1477,189,932]" captionTargetPageId="1" captionText="Fig. 1. Betta omega, new species – ZRC 59663, 70.0 mm SL holotype, life colouration (photograph by Z. Zakaria)." figureDoi="http://doi.org/10.5281/zenodo.4504616" httpUri="https://zenodo.org/record/4504616/files/figure.png" pageId="1" pageNumber="403">Figs. 1</figureCitation>
, 
<figureCitation id="137E2A5AFF92FFA4FAE6FB5CFAC7F1E3" box="[1358,1374,1248,1272]" captionStart="Fig" captionStartId="2.[114,149,2058,2080]" captionTargetBox="[114,1452,1458,2052]" captionTargetId="figure-12@2.[114,1452,1458,2052]" captionTargetPageId="2" captionText="Fig. 2. Betta omega, new species – ZRC 59663, 70.0 mm SL holotype, preserved specimen." figureDoi="http://doi.org/10.5281/zenodo.4504618" httpUri="https://zenodo.org/record/4504618/files/figure.png" pageId="1" pageNumber="403">2</figureCitation>
; meristic and morphometric information listed in 
<tableCitation id="C6C70364FF92FFA4FAB3FABCFAECF003" box="[1307,1397,1280,1304]" captionStart="Table 1" captionStartId="2.[114,169,189,211]" captionText="Table 1. Meristic and morphometric data of Betta omega, new species (data from holotype and 4 paratypes)." pageId="1" pageNumber="403">Table 1</tableCitation>
. Body relatively long and stout (body depth at dorsal-fin origin 27.1–28.5% SL); head stout with pointed snout and evenly sloping or with slight convexity at supra-orbital area (head length 28.4–32.1% SL); dorsal fin pointed, situated nearer to caudal fin (predorsal length 62.2–68.9% SL), dorsal-fin base short (dorsal fin base length 12.6–15.4% SL), covering 6½ subdorsal scales; caudal fin rounded with middle rays elongate; anal fin with posterior rays elongate, anal fin base more than half of SL (anal-fin base length 55.3–60.7% SL); pelvic fin falcate with first ray filamentous, relatively long (pelvic fin length 29.8–36.4% SL), reaching up to base of 12th anal fin ray; pectoral fin rounded.
</paragraph>
<paragraph id="8BFA36DFFF92FFA4FCEBF97CFC3EF3E3" blockId="1.[835,1474,1728,1784]" pageId="1" pageNumber="403">Vertebral count: 10–11 + 19–21 (total 29–32, mode 31 or 32, n=5).</paragraph>
<paragraph id="8BFA36DFFF92FFA4FCEBF89CFA10F223" blockId="1.[835,1417,1824,1848]" box="[835,1417,1824,1848]" pageId="1" pageNumber="403">
<emphasis id="B931EACDFF92FFA4FCEBF89CFB91F223" bold="true" box="[835,1032,1824,1848]" pageId="1" pageNumber="403">Live colouration.</emphasis>
See 
<figureCitation id="137E2A5AFF92FFA4FB97F89CFB10F223" box="[1087,1161,1824,1848]" captionStart="Fig" captionStartId="1.[136,171,947,969]" captionTargetBox="[133,1477,189,932]" captionTargetId="figure-218@1.[133,1477,189,932]" captionTargetPageId="1" captionText="Fig. 1. Betta omega, new species – ZRC 59663, 70.0 mm SL holotype, life colouration (photograph by Z. Zakaria)." figureDoi="http://doi.org/10.5281/zenodo.4504616" httpUri="https://zenodo.org/record/4504616/files/figure.png" pageId="1" pageNumber="403">Figs. 1</figureCitation>
, 
<figureCitation id="137E2A5AFF92FFA4FB31F89CFB3FF223" box="[1177,1190,1824,1848]" captionStart="Fig" captionStartId="3.[137,173,1028,1050]" captionTargetBox="[136,775,189,1012]" captionTargetId="figure-736@3.[136,775,189,1012]" captionTargetPageId="3" captionText="Fig. 3. Betta omega, new species – ZRC 59664, 74.0 mm SL paratype, front view of head showing lower jaw and throat pattern (photograph by Z. Zakaria)." figureDoi="http://doi.org/10.5281/zenodo.4504620" httpUri="https://zenodo.org/record/4504620/files/figure.png" pageId="1" pageNumber="403">3</figureCitation>
for live colouration.
</paragraph>
<paragraph id="8BFA36DFFF92FFA6FCEBF8DCFA5DF46E" blockId="1.[834,1475,1888,2104]" lastBlockId="3.[835,1476,189,373]" lastPageId="3" lastPageNumber="405" pageId="1" pageNumber="403">
Head and dorsum of body brown. Eye with unique colouration zones of the 
<taxonomicName id="4C454D5CFF92FFA4FC70F83DFBDBF283" authorityName="Krummenacher" authorityYear="1986" box="[984,1090,1921,1944]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="1" pageNumber="403" phylum="Chordata" rank="species" species="waseri">
<emphasis id="B931EACDFF92FFA4FC70F83DFBDBF283" box="[984,1090,1921,1944]" italics="true" pageId="1" pageNumber="403">B. waseri</emphasis>
</taxonomicName>
group (as defined by Tan, 1998). Mouth with lower and upper lips black, lower jaw black, black marking continuous with two black downward curved bars on throat, ending with bulging rounded tips around near edge of lower jaw (vertical line projection downwards from lower jaw posterior end). Opercle with faint greenish-yellow individuals were caught along every few meters of the ditch, possibly an indication of the territorial behaviour of the fish (Benel Tang, pers. comm.). Syntopic fish species included: 
<taxonomicName id="4C454D5CFF90FFA6FCEBFEA2FBD1F42E" baseAuthorityName="M.C.W.Weber &amp; de Beaufort" baseAuthorityYear="1912" box="[835,1096,285,309]" class="Actinopterygii" family="Cyprinidae" genus="Desmopuntius" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="hexazona">
<emphasis id="B931EACDFF90FFA6FCEBFEA2FBD1F42E" box="[835,1096,285,309]" italics="true" pageId="3" pageNumber="405">Desmopuntius hexazona</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C454D5CFF90FFA6FBFDFEA1FA27F42E" baseAuthorityName="Bleeker" baseAuthorityYear="1851" box="[1109,1470,285,309]" class="Actinopterygii" family="Cyprinidae" genus="Rasbora" kingdom="Animalia" order="Cypriniformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="einthovenii">
<emphasis id="B931EACDFF90FFA6FBFDFEA1FAB4F42E" box="[1109,1325,285,309]" italics="true" pageId="3" pageNumber="405">Rasbora einthovenii</emphasis>
(Cyprinidae)
</taxonomicName>
, 
<taxonomicName id="4C454D5CFF90FFA6FCEBFE82FB66F44E" authorityName="Lacepede" authorityYear="1800" box="[835,1279,317,341]" class="Actinopterygii" family="Synbranchidae" genus="Monopterus" kingdom="Animalia" order="Synbranchiformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="javanensis">
<emphasis id="B931EACDFF90FFA6FCEBFE82FBD8F44E" box="[835,1089,318,341]" italics="true" pageId="3" pageNumber="405">Monopterus javanensis</emphasis>
(Synbranchidae)
</taxonomicName>
, 
<taxonomicName id="4C454D5CFF90FFA6FAA6FE82FA03F44E" authorityName="Sauvage" authorityYear="1884" box="[1294,1434,317,341]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="bellica">
<emphasis id="B931EACDFF90FFA6FAA6FE82FA03F44E" box="[1294,1434,317,341]" italics="true" pageId="3" pageNumber="405">Betta bellica</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C454D5CFF90FFA6FA03FE82FBC2F46E" authorityName="H.H.Tan &amp; S.H.Tan" authorityYear="1996" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="pulchra">
<emphasis id="B931EACDFF90FFA6FA03FE82FC0FF46E" italics="true" pageId="3" pageNumber="405">B. pulchra</emphasis>
(Osphronemidae)
</taxonomicName>
, and 
<taxonomicName id="4C454D5CFF90FFA6FB3DFEE1FA26F46E" baseAuthorityName="Cuvier ex Kuhl &amp; van Hasselt in Cuvier &amp; Valenciennes" baseAuthorityYear="1831" box="[1173,1471,349,373]" class="Actinopterygii" family="Channidae" genus="Channa" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="lucius">
<emphasis id="B931EACDFF90FFA6FB3DFEE1FAA8F46E" box="[1173,1329,349,373]" italics="true" pageId="3" pageNumber="405">Channa lucius</emphasis>
(Channidae)
</taxonomicName>
.
</paragraph>
</subSubSection>
<caption id="DF3A6657FF91FFA7FFDAFF01FB39F5C8" box="[114,1184,189,211]" pageId="2" pageNumber="404">
<paragraph id="8BFA36DFFF91FFA7FFDAFF01FB39F5C8" blockId="2.[114,1184,189,211]" box="[114,1184,189,211]" pageId="2" pageNumber="404">
Table 1. Meristic and morphometric data of 
<taxonomicName id="4C454D5CFF91FFA7FD81FF02FD3BF5C8" authority="Hui &amp; Ahmad, 2018" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[553,674,190,211]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="404" phylum="Chordata" rank="species" species="omega" status="sp. nov.">
<emphasis id="B931EACDFF91FFA7FD81FF02FD3BF5C8" box="[553,674,190,211]" italics="true" pageId="2" pageNumber="404">Betta omega</emphasis>
</taxonomicName>
, 
<taxonomicNameLabel id="A20257B6FF91FFA7FD18FF02FCBDF5C8" box="[688,804,189,211]" pageId="2" pageNumber="404" rank="species">new species</taxonomicNameLabel>
(data from holotype and 4 paratypes).
</paragraph>
</caption>
<paragraph id="8BFA36DFFF91FFA7FE56FEB9FAA3F057" pageId="2" pageNumber="404">
<table id="F945C47FFF91005AFF29FEB9FA1FF057" box="[129,1414,261,1356]" gridcols="5" gridrows="9" pageId="2" pageNumber="404">
<tr id="3575349DFF91005AFF29FEB9FA1FF401" box="[129,1414,261,282]" gridrow="0" pageId="2" pageNumber="404" rowspan-0="1" rowspan-3="1" rowspan-4="1">
<th id="76A45DE1FF91005AFE56FEB9FDC2F401" box="[510,603,261,282]" gridcol="1" gridrow="0" pageId="2" pageNumber="404">
<emphasis id="B931EACDFF91FFA7FE56FEB9FDC2F401" bold="true" box="[510,603,261,282]" pageId="2" pageNumber="404">Holotype</emphasis>
</th>
<th id="76A45DE1FF91005AFD58FEB9FCF1F401" box="[752,872,261,282]" gridcol="2" gridrow="0" pageId="2" pageNumber="404">
<emphasis id="B931EACDFF91FFA7FD51FEB9FCF9F401" bold="true" box="[761,864,261,282]" pageId="2" pageNumber="404">Paratypes</emphasis>
</th>
</tr>
<tr id="3575349DFF91005AFF29FE88FA1FF451" box="[129,1414,308,330]" gridrow="1" pageId="2" pageNumber="404" rowspan-3="1" rowspan-4="1">
<th id="76A45DE1FF91005AFF29FE88FE07F451" box="[129,414,308,330]" gridcol="0" gridrow="1" pageId="2" pageNumber="404">Standard length (mm)</th>
<td id="76A45DE1FF91005AFE56FE88FDC2F451" box="[510,603,308,330]" gridcol="1" gridrow="1" pageId="2" pageNumber="404">70.0</td>
<td id="76A45DE1FF91005AFD58FE88FCF1F451" box="[752,872,308,330]" gridcol="2" gridrow="1" pageId="2" pageNumber="404">60.4–81.2</td>
</tr>
<tr id="3575349DFF91005AFF29FEDFFA1FF7AD" box="[129,1414,355,694]" gridrow="2" pageId="2" pageNumber="404" rowspan-4="1">
<th id="76A45DE1FF91005AFF29FEDFFE07F7AD" box="[129,414,355,694]" gridcol="0" gridrow="2" pageId="2" pageNumber="404">Meristics Anal-fin rays Dorsal-fin rays Caudal-fin rays Pelvic-fin rays Pectoral-fin rays Subdorsal scales Body depth scales Lateral scales Predorsal scales Postdorsal scales</th>
<td id="76A45DE1FF91005AFE56FEDFFDC2F7AD" box="[510,603,355,694]" gridcol="1" gridrow="2" pageId="2" pageNumber="404">II,29 I,9 ii,6+7,i i,1,4 14 6½ 9½ 32 24 11</td>
<td id="76A45DE1FF91005AFD58FEDFFCF1F7AD" box="[752,872,355,694]" gridcol="2" gridrow="2" pageId="2" pageNumber="404">I–II,28–30 I,8–9 ii,6+7–8,i i,1,4 13–14 6½ 9½ 31–33 23 10–12</td>
<td id="76A45DE1FF91005AFC49FEDFFBEEF7AD" box="[993,1143,355,694]" gridcol="3" gridrow="2" pageId="2" pageNumber="404">Mode 29 or 30 (total) I,9 ii,6+7,i i,1,4 14 6½ 9½ 31 or 32 23 11</td>
</tr>
<tr id="3575349DFF91005AFF29FD73FA1FF179" box="[129,1414,719,1122]" gridrow="3" pageId="2" pageNumber="404">
<th id="76A45DE1FF91005AFF29FD73FE07F179" box="[129,414,719,1122]" gridcol="0" gridrow="3" pageId="2" pageNumber="404">Morphometrics % Standard length Total length Trunk length Predorsal length Postdorsal length Caudal peduncle depth Preanal length Head length Body depth at dorsal-fin origin Pelvic-fin length Anal-fin base length Dorsal-fin base length</th>
<td id="76A45DE1FF91005AFE56FD73FDC2F179" box="[510,603,719,1122]" gridcol="1" gridrow="3" pageId="2" pageNumber="404">143.7 71.3 65.0 22.9 20.0 45.4 30.4 27.9 36.4 56.4 12.6</td>
<td id="76A45DE1FF91005AFD58FD73FCF1F179" box="[752,872,719,1122]" gridcol="2" gridrow="3" pageId="2" pageNumber="404">Range 139.2–147.7 71.2–73.2 62.2–68.9 19.9–24.1 19.2–20.6 42.7–46.0 28.4–32.1 27.1–28.5 29.8–36.4 55.3–60.7 12.6–15.4</td>
<td id="76A45DE1FF91005AFC49FD73FBEEF179" box="[993,1143,719,1122]" gridcol="3" gridrow="3" pageId="2" pageNumber="404">Mean 143.6 72.0 65.6 22.3 20.0 44.1 30.6 27.6 32.3 57.2 13.7</td>
<td id="76A45DE1FF91005AFB78FD73FA1FF179" box="[1232,1414,719,1122]" gridcol="4" gridrow="3" pageId="2" pageNumber="404">Standard deviation 3.1 0.9 2.4 1.7 0.6 1.5 1.4 0.6 2.6 2.0 1.1</td>
</tr>
<tr id="3575349DFF91005AFF29FBC7FA1FF18A" box="[129,1414,1147,1169]" gridrow="4" pageId="2" pageNumber="404">
<th id="76A45DE1FF91005AFF29FBC7FA1FF18A" box="[129,1414,1147,1169]" colspan="5" colspanRight="4" gridcol="0" gridrow="4" pageId="2" pageNumber="404">% Head length</th>
</tr>
<tr id="3575349DFF91005AFF29FB16FA1FF1DB" box="[129,1414,1194,1216]" gridrow="5" pageId="2" pageNumber="404">
<th id="76A45DE1FF91005AFF29FB16FE07F1DB" box="[129,414,1194,1216]" gridcol="0" gridrow="5" pageId="2" pageNumber="404">Orbit diameter</th>
<td id="76A45DE1FF91005AFE56FB16FDC2F1DB" box="[510,603,1194,1216]" gridcol="1" gridrow="5" pageId="2" pageNumber="404">21.1</td>
<td id="76A45DE1FF91005AFD58FB16FCF1F1DB" box="[752,872,1194,1216]" gridcol="2" gridrow="5" pageId="2" pageNumber="404">21.1–25.8</td>
<td id="76A45DE1FF91005AFC49FB16FBEEF1DB" box="[993,1143,1194,1216]" gridcol="3" gridrow="5" pageId="2" pageNumber="404">23.1</td>
<td id="76A45DE1FF91005AFB78FB16FA1FF1DB" box="[1232,1414,1194,1216]" gridcol="4" gridrow="5" pageId="2" pageNumber="404">1.7</td>
</tr>
<tr id="3575349DFF91005AFF29FB65FA1FF1F4" box="[129,1414,1241,1263]" gridrow="6" pageId="2" pageNumber="404">
<th id="76A45DE1FF91005AFF29FB65FE07F1F4" box="[129,414,1241,1263]" gridcol="0" gridrow="6" pageId="2" pageNumber="404">Postorbital length</th>
<td id="76A45DE1FF91005AFE56FB65FDC2F1F4" box="[510,603,1241,1263]" gridcol="1" gridrow="6" pageId="2" pageNumber="404">47.9</td>
<td id="76A45DE1FF91005AFD58FB65FCF1F1F4" box="[752,872,1241,1263]" gridcol="2" gridrow="6" pageId="2" pageNumber="404">47.9–51.9</td>
<td id="76A45DE1FF91005AFC49FB65FBEEF1F4" box="[993,1143,1241,1263]" gridcol="3" gridrow="6" pageId="2" pageNumber="404">50.0</td>
<td id="76A45DE1FF91005AFB78FB65FA1FF1F4" box="[1232,1414,1241,1263]" gridcol="4" gridrow="6" pageId="2" pageNumber="404">2.0</td>
</tr>
<tr id="3575349DFF91005AFF29FABBFA1FF005" box="[129,1414,1287,1310]" gridrow="7" pageId="2" pageNumber="404">
<th id="76A45DE1FF91005AFF29FABBFE07F005" box="[129,414,1287,1310]" gridcol="0" gridrow="7" pageId="2" pageNumber="404">Interorbital width</th>
<td id="76A45DE1FF91005AFE56FABBFDC2F005" box="[510,603,1287,1310]" gridcol="1" gridrow="7" pageId="2" pageNumber="404">43.7</td>
<td id="76A45DE1FF91005AFD58FABBFCF1F005" box="[752,872,1287,1310]" gridcol="2" gridrow="7" pageId="2" pageNumber="404">39.2–45.7</td>
<td id="76A45DE1FF91005AFC49FABBFBEEF005" box="[993,1143,1287,1310]" gridcol="3" gridrow="7" pageId="2" pageNumber="404">42.6</td>
<td id="76A45DE1FF91005AFB78FABBFA1FF005" box="[1232,1414,1287,1310]" gridcol="4" gridrow="7" pageId="2" pageNumber="404">2.4</td>
</tr>
<tr id="3575349DFF91005AFF29FA8AFA1FF057" box="[129,1414,1334,1356]" gridrow="8" pageId="2" pageNumber="404">
<th id="76A45DE1FF91005AFF29FA8AFE07F057" box="[129,414,1334,1356]" gridcol="0" gridrow="8" pageId="2" pageNumber="404">Snout length</th>
<td id="76A45DE1FF91005AFE56FA8AFDC2F057" box="[510,603,1334,1356]" gridcol="1" gridrow="8" pageId="2" pageNumber="404">27.7</td>
<td id="76A45DE1FF91005AFD58FA8AFCF1F057" box="[752,872,1334,1356]" gridcol="2" gridrow="8" pageId="2" pageNumber="404">20.9–27.7</td>
<td id="76A45DE1FF91005AFC49FA8AFBEEF057" box="[993,1143,1334,1356]" gridcol="3" gridrow="8" pageId="2" pageNumber="404">24.2</td>
<td id="76A45DE1FF91005AFB78FA8AFA1FF057" box="[1232,1414,1334,1356]" gridcol="4" gridrow="8" pageId="2" pageNumber="404">2.5</td>
</tr>
</table>
</paragraph>
<caption id="DF3A6657FF91FFA7FFDAF7B6FC67FD3B" ID-DOI="http://doi.org/10.5281/zenodo.4504618" ID-Zenodo-Dep="4504618" box="[114,1022,2058,2080]" httpUri="https://zenodo.org/record/4504618/files/figure.png" pageId="2" pageNumber="404" startId="2.[114,149,2058,2080]" targetBox="[114,1452,1458,2052]" targetPageId="2">
<paragraph id="8BFA36DFFF91FFA7FFDAF7B6FC67FD3B" blockId="2.[114,1022,2058,2080]" box="[114,1022,2058,2080]" pageId="2" pageNumber="404">
Fig. 2. 
<taxonomicName id="4C454D5CFF91FFA7FF13F7B6FEAAFD04" authority="Hui &amp; Ahmad, 2018" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[187,307,2058,2079]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="2" pageNumber="404" phylum="Chordata" rank="species" species="omega" status="sp. nov.">
<emphasis id="B931EACDFF91FFA7FF13F7B6FEAAFD04" box="[187,307,2058,2079]" italics="true" pageId="2" pageNumber="404">Betta omega</emphasis>
</taxonomicName>
, 
<taxonomicNameLabel id="A20257B6FF91FFA7FEE9F7B6FE2FFD3B" box="[321,438,2058,2080]" pageId="2" pageNumber="404" rank="species">new species</taxonomicNameLabel>
– ZRC 59663, 70.0 mm SL holotype, preserved specimen.
</paragraph>
</caption>
<subSubSection id="C35F6554FF90FFA6FCEBFE21FA58F72E" pageId="3" pageNumber="405" type="etymology">
<paragraph id="8BFA36DFFF90FFA6FCEBFE21FA58F72E" blockId="3.[834,1475,413,565]" pageId="3" pageNumber="405">
<emphasis id="B931EACDFF90FFA6FCEBFE21FC5FF4AE" bold="true" box="[835,966,413,437]" pageId="3" pageNumber="405">Etymology.</emphasis>
From the Greek 
<emphasis id="B931EACDFF90FFA6FB23FE22FB41F4AE" box="[1163,1240,414,437]" italics="true" pageId="3" pageNumber="405">Omega</emphasis>
/ Ω (upper case), the last letter of the Greek alphabet; in allusion to the unique throat pattern and in reference to the last members of this species in the quickly disappearing black water habitat 
<typeStatus id="54FE887DFF90FFA6FAD4FE42FA30F70D" box="[1404,1449,510,534]" pageId="3" pageNumber="405">type</typeStatus>
in 
<collectingCountry id="F352764FFF90FFA6FCEAFDA1FC3DF72E" box="[834,932,541,565]" name="Malaysia" pageId="3" pageNumber="405">Malaysia</collectingCountry>
and Southeast Asia. Used as a noun in apposition.
</paragraph>
</subSubSection>
<subSubSection id="C35F6554FF90FFA6FCEBFDE1FE96F3D9" pageId="3" pageNumber="405" type="discussion">
<paragraph id="8BFA36DFFF90FFA6FCEBFDE1FA34F68E" blockId="3.[835,1476,605,917]" pageId="3" pageNumber="405">
<emphasis id="B931EACDFF90FFA6FCEBFDE1FC2BF76E" bold="true" box="[835,946,605,629]" pageId="3" pageNumber="405">Remarks.</emphasis>
<taxonomicName id="4C454D5CFF90FFA6FC10FDE2FBA4F76E" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[952,1085,606,629]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FC10FDE2FBA4F76E" box="[952,1085,606,629]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
can be distinguished further from 
<taxonomicName id="4C454D5CFF90FFA6FA03FDE2FC13F78E" authorityName="Krummenacher" authorityYear="1986" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="waseri">
<emphasis id="B931EACDFF90FFA6FA03FDE2FC13F78E" italics="true" pageId="3" pageNumber="405">B. waseri</emphasis>
</taxonomicName>
in the following characters: “Ω /omega” shape throat pattern (vs. two tear-drop shaped black marks below but not connected to black lower lip; see 
<figureCitation id="137E2A5AFF90FFA6FB1BFD01FB6FF7CE" box="[1203,1270,701,725]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="3" pageNumber="405">Fig. 4</figureCitation>
); absence of black lower margin of operculum (vs. presence); opercle without any distinct brown markings and with light greenish-yellow iridescence (vs. several short black streaks, and without iridescence); deeper body (27.1–28.5% SL, vs. 23.1–27.6); dorsal fin begins at lateral scale 15 (vs. 17–18); anterior anal-fin base at vertical through lateral scale 6 (vs. 8–9).
</paragraph>
<caption id="DF3A6657FF90FFA6FF21FBB8FE03F148" ID-DOI="http://doi.org/10.5281/zenodo.4504620" ID-Zenodo-Dep="4504620" httpUri="https://zenodo.org/record/4504620/files/figure.png" pageId="3" pageNumber="405" startId="3.[137,173,1028,1050]" targetBox="[136,775,189,1012]" targetPageId="3">
<paragraph id="8BFA36DFFF90FFA6FF21FBB8FE03F148" blockId="3.[136,775,1027,1107]" pageId="3" pageNumber="405">
Fig. 3. 
<taxonomicName id="4C454D5CFF90FFA6FF7DFBB8FECCF102" authority="Hui &amp; Ahmad, 2018" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[213,341,1028,1049]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega" status="sp. nov.">
<emphasis id="B931EACDFF90FFA6FF7DFBB8FECCF102" box="[213,341,1028,1049]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
, 
<taxonomicNameLabel id="A20257B6FF90FFA6FECDFBB8FE46F101" box="[357,479,1028,1050]" pageId="3" pageNumber="405" rank="species">new species</taxonomicNameLabel>
– ZRC 59664, 74.0 mm SL paratype, front view of head showing lower jaw and throat pattern (photograph by Z. Zakaria).
</paragraph>
</caption>
<paragraph id="8BFA36DFFF90FFA6FF21FB36FDEFF0D9" blockId="3.[137,777,1162,1474]" pageId="3" pageNumber="405">iridescence, iridescence extending to side of body (up to 10 scale rows) above belly area. Body yellowish-brown, with faint dark patches just above anal fin sheath scales. Dorsal fin brownish with up to 10 transverse bars on interradial membranes. Caudal fin brownish with up to 24 transverse bars on interradial membranes. Anal fin brownish, without distal dark border but with narrow white margin. Pelvic fin yellowish-brown with whitish interradial membranes, distal part of filamentous pelvic-fin ray iridescent whitish-green. Pectoral fin hyaline with black sub-basal bar.</paragraph>
<paragraph id="8BFA36DFFF90FFA6FF21FA56FE96F3D9" blockId="3.[136,776,1514,1730]" pageId="3" pageNumber="405">
<emphasis id="B931EACDFF90FFA6FF21FA56FE12F319" bold="true" box="[137,395,1514,1538]" pageId="3" pageNumber="405">Preserved colouration.</emphasis>
Preserved colouration illustrated in 
<figureCitation id="137E2A5AFF90FFA6FF21F9B6FF57F339" box="[137,206,1546,1570]" captionStart="Fig" captionStartId="2.[114,149,2058,2080]" captionTargetBox="[114,1452,1458,2052]" captionTargetId="figure-12@2.[114,1452,1458,2052]" captionTargetPageId="2" captionText="Fig. 2. Betta omega, new species – ZRC 59663, 70.0 mm SL holotype, preserved specimen." figureDoi="http://doi.org/10.5281/zenodo.4504618" httpUri="https://zenodo.org/record/4504618/files/figure.png" pageId="3" pageNumber="405">Fig. 2</figureCitation>
. Head and body dorsum dark brown. Opercle area with brown mottling. Throat pattern indistinct. Lateral of body with indistinct brown central stripe. Faint brown stripe above anal-fin base. Rest of body yellowish-brown on lateral and cream-yellow on ventrum. Fin rays light brownish and rest hyaline.
</paragraph>
</subSubSection>
<subSubSection id="C35F6554FF90FFA1FF23F956FB20F241" lastPageId="4" lastPageNumber="406" pageId="3" pageNumber="405" type="distribution">
<paragraph id="8BFA36DFFF90FFA6FF23F956FE5BF279" blockId="3.[135,777,1770,1890]" pageId="3" pageNumber="405">
<emphasis id="B931EACDFF90FFA6FF23F956FE82F219" bold="true" box="[139,283,1770,1794]" pageId="3" pageNumber="405">Distribution.</emphasis>
<taxonomicName id="4C454D5CFF90FFA6FEB6F957FE3BF219" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[286,418,1771,1794]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FEB6F957FE3BF219" box="[286,418,1771,1794]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
is recorded only from the remnant black water habitats near Pekan Nanas in 
<collectingRegion id="4981F83DFF90FFA6FDE4F8B6FD13F239" box="[588,650,1802,1826]" country="Malaysia" name="Johor" pageId="3" pageNumber="405">Johor</collectingRegion>
, Peninsular 
<collectingCountry id="F352764FFF90FFA6FF22F896FF6AF259" box="[138,243,1834,1858]" name="Malaysia" pageId="3" pageNumber="405">Malaysia</collectingCountry>
. There is a likelihood that this species is near extinction or already extinct.
</paragraph>
<paragraph id="8BFA36DFFF90FFA6FF21F836FAD0F00E" blockId="3.[137,776,1930,2114]" lastBlockId="3.[834,1476,957,1301]" pageId="3" pageNumber="405">
<emphasis id="B931EACDFF90FFA6FF21F836FE8AF2B9" bold="true" box="[137,275,1930,1954]" pageId="3" pageNumber="405">Field notes.</emphasis>
The habitat of 
<taxonomicName id="4C454D5CFF90FFA6FE62F837FDC1F2B9" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[458,600,1931,1954]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FE62F837FDC1F2B9" box="[458,600,1931,1954]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
was located in remnant black water swamps, present as roadside ditches along the Pontian Kecil-Pekan 
<collectingRegion id="4981F83DFF90FFA6FE4FF877FDB3F2F9" box="[487,554,1995,2018]" country="Thailand" name="Nan" pageId="3" pageNumber="405">Nanas</collectingRegion>
road; also as small streams flowing through oil palm estates. These dug ditches are usually overgrown with vegetation. Fishes were caught using a simple earthworm-baited hand-line. Only a few 
<taxonomicName id="4C454D5CFF90FFA6FCEBFC02FC51F6CE" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[835,968,958,981]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FCEBFC02FC51F6CE" box="[835,968,958,981]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
can be further distinguished from 
<taxonomicName id="4C454D5CFF90FFA6FA9CFC02FA19F6CE" authorityName="P.K.L.Ng &amp; Kottelat" authorityYear="1994" box="[1332,1408,958,981]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="tomi">
<emphasis id="B931EACDFF90FFA6FA9CFC02FA19F6CE" box="[1332,1408,958,981]" italics="true" pageId="3" pageNumber="405">B. tomi</emphasis>
</taxonomicName>
in the following characters: “Ω /omega” shape throat pattern (vs. throat with two black oval spots which do not merge with black lower lip; see 
<figureCitation id="137E2A5AFF90FFA6FBB7FBA1FBF8F12E" box="[1055,1121,1053,1077]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="3" pageNumber="405">Fig. 4</figureCitation>
); absence of black lower margin of operculum (vs. presence); opercle without any distinct brown markings and with light greenish-yellow iridescence (vs. several short black streaks, and without iridescence); distal margin of anal fin without black or coloured band (vs. presence); anterior anal-fin base at vertical through lateral scale 6 (vs. 7–8); subdorsal scales 6½ (vs. 5–5½); postdorsal scales 10–12, mode 11 (vs. 10–10½, mode 10).
</paragraph>
<paragraph id="8BFA36DFFF90FFA6FCEBFA82FA2FF36E" blockId="3.[835,1475,1341,1653]" pageId="3" pageNumber="405">
<taxonomicName id="4C454D5CFF90FFA6FCEBFA82FC5EF04E" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[835,967,1342,1365]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FCEBFA82FC5EF04E" box="[835,967,1342,1365]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
can be further distinguished from 
<taxonomicName id="4C454D5CFF90FFA6FA82FA82FA59F04E" authorityName="P.K.L.Ng &amp; Kottelat" authorityYear="1994" box="[1322,1472,1341,1365]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="spilotogena">
<emphasis id="B931EACDFF90FFA6FA82FA82FA59F04E" box="[1322,1472,1341,1365]" italics="true" pageId="3" pageNumber="405">B. spilotogena</emphasis>
</taxonomicName>
in the following characters: “Ω /omega” shape throat pattern (vs. throat with two very broad black oval spots which may appear joined medially, spots do not merge with black lower lip; see 
<figureCitation id="137E2A5AFF90FFA6FC3CFA01FC4CF0CE" box="[916,981,1469,1493]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="3" pageNumber="405">Fig. 4</figureCitation>
); absence of black lower margin of operculum (vs. presence); opercle without any distinct brown markings and with light greenish-yellow iridescence (vs. distinctly covered with several large black spots, without iridescence); anterior anal-fin base at vertical through lateral scale 6 (vs. 7–8, mode 7); subdorsal scales 6½ (vs. 5½–6, mode 5½).
</paragraph>
<paragraph id="8BFA36DFFF90FFA6FCEBF922FA23F24E" blockId="3.[834,1475,1693,1877]" pageId="3" pageNumber="405">
<taxonomicName id="4C454D5CFF90FFA6FCEBF922FC79F3AE" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[835,992,1694,1717]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FCEBF922FC79F3AE" box="[835,992,1694,1717]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
can be further distinguished from 
<taxonomicName id="4C454D5CFF90FFA6FA01F922FC46F3CE" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="chloropharynx">
<emphasis id="B931EACDFF90FFA6FA01F922FC46F3CE" italics="true" pageId="3" pageNumber="405">B. chloropharynx</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C454D5CFF90FFA6FBA7F902FB07F3CE" authorityName="Tan" authorityYear="2009" box="[1039,1182,1725,1749]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="pardalotos">
<emphasis id="B931EACDFF90FFA6FBA7F902FB07F3CE" box="[1039,1182,1725,1749]" italics="true" pageId="3" pageNumber="405">B. pardalotos</emphasis>
</taxonomicName>
in the following characters: “Ω /omega” shape throat pattern (vs. throat with two cream blotches on a black throat; see 
<figureCitation id="137E2A5AFF90FFA6FB3DF941FB4EF20E" box="[1173,1239,1789,1813]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="3" pageNumber="405">Fig. 4</figureCitation>
); opercle without any distinct brown markings and with light greenish-yellow iridescence (vs. heavily spotted opercle of 
<taxonomicName id="4C454D5CFF90FFA6FAB1F882FA33F24E" authorityName="Tan" authorityYear="2009" box="[1305,1450,1853,1877]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="pardalotos">
<emphasis id="B931EACDFF90FFA6FAB1F882FA33F24E" box="[1305,1450,1853,1877]" italics="true" pageId="3" pageNumber="405">B. pardalotos</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8BFA36DFFF90FFA6FCEBF8C2FA5AFD2E" blockId="3.[835,1476,1917,2101]" pageId="3" pageNumber="405">
<taxonomicName id="4C454D5CFF90FFA6FCEBF8C2FC52F28E" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[835,971,1918,1941]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="3" pageNumber="405" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF90FFA6FCEBF8C2FC52F28E" box="[835,971,1918,1941]" italics="true" pageId="3" pageNumber="405">Betta omega</emphasis>
</taxonomicName>
can be further distinguished from 
<emphasis id="B931EACDFF90FFA6FAE1F8C2FAE7F28E" box="[1353,1406,1918,1941]" italics="true" pageId="3" pageNumber="405">B. pi</emphasis>
in the following characters: “Ω /omega” shape throat pattern (vs. throat with π-shaped black pattern; see 
<figureCitation id="137E2A5AFF90FFA6FABAF801FAC0F2CE" box="[1298,1369,1981,2005]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="3" pageNumber="405">Fig. 4</figureCitation>
); opercle without any distinct brown markings and with light greenishyellow iridescence (vs. opercle with several black blotches of variable size, without iridescence); distal margin of anal
</paragraph>
<caption id="DF3A6657FF97FFA1FFD9F924FCA5F3D1" ID-DOI="http://doi.org/10.5281/zenodo.4504622" ID-Zenodo-Dep="4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="4" pageNumber="406" startId="4.[113,148,1688,1710]" targetBox="[112,1451,190,1673]" targetPageId="4">
<paragraph id="8BFA36DFFF97FFA1FFD9F924FCA5F3D1" blockId="4.[113,1451,1688,1738]" pageId="4" pageNumber="406">
Fig. 4. Schematic diagrams of throat patterns of the 
<taxonomicName id="4C454D5CFF97FFA1FDF4F924FD4DF3B6" authorityName="Krummenacher" authorityYear="1986" box="[604,724,1688,1709]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="waseri">
<emphasis id="B931EACDFF97FFA1FDF4F924FD4DF3B6" box="[604,724,1688,1709]" italics="true" pageId="4" pageNumber="406">Betta waseri</emphasis>
</taxonomicName>
species group. a, 
<taxonomicName id="4C454D5CFF97FFA1FCD5F924FC4EF3B6" authorityName="Krummenacher" authorityYear="1986" box="[893,983,1688,1709]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="waseri">
<emphasis id="B931EACDFF97FFA1FCD5F924FC4EF3B6" box="[893,983,1688,1709]" italics="true" pageId="4" pageNumber="406">B. waseri</emphasis>
</taxonomicName>
; b, 
<taxonomicName id="4C454D5CFF97FFA1FC52F924FB09F3B6" authorityName="P.K.L.Ng &amp; Kottelat" authorityYear="1994" box="[1018,1168,1688,1709]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="hipposideros">
<emphasis id="B931EACDFF97FFA1FC52F924FB09F3B6" box="[1018,1168,1688,1709]" italics="true" pageId="4" pageNumber="406">B. hipposideros</emphasis>
</taxonomicName>
; c, 
<taxonomicName id="4C454D5CFF97FFA1FB1AF924FB6EF3B6" authorityName="P.K.L.Ng &amp; Kottelat" authorityYear="1994" box="[1202,1271,1688,1709]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="tomi">
<emphasis id="B931EACDFF97FFA1FB1AF924FB6EF3B6" box="[1202,1271,1688,1709]" italics="true" pageId="4" pageNumber="406">B. tomi</emphasis>
</taxonomicName>
; d, 
<taxonomicName id="4C454D5CFF97FFA1FAB2F924FA3DF3B6" authorityName="P.K.L.Ng &amp; Kottelat" authorityYear="1994" box="[1306,1444,1688,1709]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="spilotogena">
<emphasis id="B931EACDFF97FFA1FAB2F924FA3DF3B6" box="[1306,1444,1688,1709]" italics="true" pageId="4" pageNumber="406">B. spilotogena</emphasis>
</taxonomicName>
; e, 
<taxonomicName id="4C454D5CFF97FFA1FF21F909FEABF3D2" box="[137,306,1716,1738]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="chloropharynx">
<emphasis id="B931EACDFF97FFA1FF21F909FEABF3D2" box="[137,306,1716,1738]" italics="true" pageId="4" pageNumber="406">B. chloropharynx</emphasis>
</taxonomicName>
; f, 
<taxonomicName id="4C454D5CFF97FFA1FEF1F909FE2BF3D1" authorityName="Tan" authorityYear="1998" box="[345,434,1717,1738]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="renata">
<emphasis id="B931EACDFF97FFA1FEF1F909FE2BF3D1" box="[345,434,1717,1738]" italics="true" pageId="4" pageNumber="406">B. renata</emphasis>
</taxonomicName>
; g, 
<emphasis id="B931EACDFF97FFA1FE74F909FD93F3D1" box="[476,522,1717,1738]" italics="true" pageId="4" pageNumber="406">B. pi</emphasis>
; h, 
<taxonomicName id="4C454D5CFF97FFA1FD9CF909FD2FF3D2" box="[564,694,1716,1738]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="pardalatos">
<emphasis id="B931EACDFF97FFA1FD9CF909FD2FF3D2" box="[564,694,1716,1738]" italics="true" pageId="4" pageNumber="406">B. pardalatos</emphasis>
</taxonomicName>
; i, 
<taxonomicName id="4C454D5CFF97FFA1FD73F909FCACF3D1" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[731,821,1717,1738]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF97FFA1FD73F909FCACF3D1" box="[731,821,1717,1738]" italics="true" pageId="4" pageNumber="406">B. omega</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BFA36DFFF97FFA1FFD9F8BEFE51F261" blockId="4.[113,753,1794,1914]" pageId="4" pageNumber="406">fin without black or coloured band (vs. presence); shorter head length (28.4–32.1% SL, vs. 32.3–34.4); deeper caudal peduncle depth (19.2–20.6% SL, vs. 16.3–18.4); subdorsal scales 6 ½ (vs. 5½–6, mode 6).</paragraph>
<paragraph id="8BFA36DFFF97FFA1FFD9F81EFD68FD01" blockId="4.[113,753,1954,2074]" pageId="4" pageNumber="406">
<taxonomicName id="4C454D5CFF97FFA1FFD9F81EFF61F2A2" authorityName="Hui &amp; Ahmad" authorityYear="2018" box="[113,248,1954,1977]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="omega">
<emphasis id="B931EACDFF97FFA1FFD9F81EFF61F2A2" box="[113,248,1954,1977]" italics="true" pageId="4" pageNumber="406">Betta omega</emphasis>
</taxonomicName>
can be further distinguished from 
<taxonomicName id="4C454D5CFF97FFA1FDD8F81EFD4CF2A2" authorityName="Tan" authorityYear="1998" box="[624,725,1954,1977]" class="Actinopterygii" family="Osphronemidae" genus="Betta" kingdom="Animalia" order="Perciformes" pageId="4" pageNumber="406" phylum="Chordata" rank="species" species="renata">
<emphasis id="B931EACDFF97FFA1FDD8F81EFD4CF2A2" box="[624,725,1954,1977]" italics="true" pageId="4" pageNumber="406">B. renata</emphasis>
</taxonomicName>
in the following characters: “Ω /omega” shape throat pattern (vs. a kidney-shaped black mark on throat, not connected with black lower lip; see 
<figureCitation id="137E2A5AFF97FFA1FED0F7BEFE21FD01" box="[376,440,2050,2074]" captionStart="Fig" captionStartId="4.[113,148,1688,1710]" captionTargetBox="[112,1451,190,1673]" captionTargetId="figure-13@4.[97,1471,172,1679]" captionTargetPageId="4" captionText="Fig. 4. Schematic diagrams of throat patterns of the Betta waseri species group. a, B. waseri; b, B. hipposideros; c, B. tomi; d, B. spilotogena; e, B. chloropharynx; f, B. renata; g, B. pi; h, B. pardalatos; i, B. omega." figureDoi="http://doi.org/10.5281/zenodo.4504622" httpUri="https://zenodo.org/record/4504622/files/figure.png" pageId="4" pageNumber="406">Fig. 4</figureCitation>
); opercle without any distinct
</paragraph>
<paragraph id="8BFA36DFFF97FFA1FC83F8BEFB20F241" blockId="4.[811,1453,1794,1882]" pageId="4" pageNumber="406">brown markings and with light greenish-yellow iridescence (vs. several black spots on opercle); absence of black lower margin of operculum (vs. presence).</paragraph>
</subSubSection>
</treatment>
</document>