treatments-xml/data/03/8F/87/038F87AD2B0CFFA32EC55D3BFB460C2E.xml
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<document ID-DOI="10.7717/peerj.7386" ID-GBIF-Dataset="41ff7b58-5204-42ec-b85a-75a6904af370" ID-PMC="PMC6716565" ID-PubMed="31523494" ID-Zenodo-Dep="5767530" approvalRequired="1" approvalRequired_for_document="1" checkinTime="1638970172773" checkinUser="carolina" docAuthor="Murillo-Ramos, Leidys, Brehm, Gunnar, Sihvonen, Pasi, Hausmann, Axel, Holm, Sille, Ghanavi, Hamid Reza, Õunap, Erki, Truuverk, Andro, Staude, Hermann, Friedrich, Egbert, Tammaru, Toomas &amp; Wahlberg, Niklas" docDate="2019" docId="038F87AD2B0CFFA32EC55D3BFB460C2E" docLanguage="en" docName="PeerJ.7.e7386.pdf" docOrigin="PeerJ 7" docTitle="Desmobathrinae Meyrick 1886" docType="treatment" docVersion="9" lastPageNumber="20" masterDocId="FFB6FFD52B1FFFB72F375F1EFFC20D03" masterDocTitle="A comprehensive molecular phylogeny of Geometridae (Lepidoptera) with a focus on enigmatic small subfamilies" masterLastPageNumber="39" masterPageNumber="1" pageNumber="20" updateTime="1668138804254" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
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<mods:title>A comprehensive molecular phylogeny of Geometridae (Lepidoptera) with a focus on enigmatic small subfamilies</mods:title>
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<mods:namePart>Murillo-Ramos, Leidys</mods:namePart>
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<mods:namePart>Brehm, Gunnar</mods:namePart>
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<mods:namePart>Sihvonen, Pasi</mods:namePart>
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<mods:namePart>Hausmann, Axel</mods:namePart>
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<mods:namePart>Holm, Sille</mods:namePart>
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<mods:namePart>Ghanavi, Hamid Reza</mods:namePart>
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<mods:namePart>Õunap, Erki</mods:namePart>
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<mods:namePart>Truuverk, Andro</mods:namePart>
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</mods:role>
<mods:namePart>Staude, Hermann</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Friedrich, Egbert</mods:namePart>
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<mods:namePart>Tammaru, Toomas</mods:namePart>
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<mods:date>2019</mods:date>
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<mods:number>2019-08-27</mods:number>
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<mods:identifier type="DOI">10.7717/peerj.7386</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5767471" ID-GBIF-Taxon="191256153" ID-Zenodo-Dep="5767471" LSID="urn:lsid:plazi:treatment:038F87AD2B0CFFA32EC55D3BFB460C2E" httpUri="http://treatment.plazi.org/id/038F87AD2B0CFFA32EC55D3BFB460C2E" lastPageId="20" lastPageNumber="20" pageId="19" pageNumber="20">
<subSubSection box="[498,882,549,575]" pageId="19" pageNumber="20" type="nomenclature">
<paragraph blockId="19.[498,1542,549,1933]" box="[498,882,549,575]" pageId="19" pageNumber="20">
<heading bold="true" box="[498,882,549,575]" fontSize="11" level="3" pageId="19" pageNumber="20" reason="6">
<taxonomicName authority="Meyrick, 1886" authorityName="Meyrick" authorityYear="1886" box="[498,882,549,575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">
<emphasis bold="true" box="[498,882,549,575]" pageId="19" pageNumber="20">Desmobathrinae Meyrick, 1886</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20" type="discussion">
<paragraph blockId="19.[498,1542,549,1933]" pageId="19" pageNumber="20">
Taxa placed in
<taxonomicName authorityName="Meyrick" authorityYear="1886" box="[675,868,591,617]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
were formerly recognized as
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[1209,1398,591,617]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
genera with slender appendages.
<bibRefCitation author="Holloway" box="[742,934,631,657]" firstAuthor="Holloway" journalOrPublisher="Ennominae Malayan Nature Journal" pageId="19" pageNumber="20" pagination="147 - 326" part="49" refId="ref18919" refString="Holloway J. 1996. The moths of Borneo, part 9: Geometridae (incl. Orthostixini), Oenochrominae, Desmobathrinae, Geometrinae. Ennominae Malayan Nature Journal 49: 147 - 326." title="The moths of Borneo, part 9: Geometridae (incl. Orthostixini), Oenochrominae, Desmobathrinae, Geometrinae" type="journal article" year="1996">
<emphasis box="[742,934,631,657]" italics="true" pageId="19" pageNumber="20">Holloway (1996)</emphasis>
</bibRefCitation>
revived
<taxonomicName authorityName="Meyrick" authorityYear="1886" box="[1036,1229,631,657]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
from synonymy with
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[498,687,671,697]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
and divided it into the tribes
<taxonomicName authorityName="Warren" authorityYear="1894" box="[1043,1161,671,697]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Eumeleini">Eumeleini</taxonomicName>
and
<taxonomicName authorityName="Meyrick" authorityYear="1886" box="[1222,1401,671,697]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Desmobathrini">Desmobathrini</taxonomicName>
.
</paragraph>
<paragraph blockId="19.[498,1542,549,1933]" pageId="19" pageNumber="20">
<taxonomicName authorityName="Meyrick" authorityYear="1886" box="[498,691,711,737]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
species have a pantropical distribution and they apparently (still) lack recognized morphological apomorphies (
<bibRefCitation author="Holloway" box="[983,1162,751,777]" firstAuthor="Holloway" journalOrPublisher="Ennominae Malayan Nature Journal" pageId="19" pageNumber="20" pagination="147 - 326" part="49" refId="ref18919" refString="Holloway J. 1996. The moths of Borneo, part 9: Geometridae (incl. Orthostixini), Oenochrominae, Desmobathrinae, Geometrinae. Ennominae Malayan Nature Journal 49: 147 - 326." title="The moths of Borneo, part 9: Geometridae (incl. Orthostixini), Oenochrominae, Desmobathrinae, Geometrinae" type="journal article" year="1996">
<emphasis box="[983,1162,751,777]" italics="true" pageId="19" pageNumber="20">Holloway, 1996</emphasis>
</bibRefCitation>
). Our phylogenetic analysis has questioned the monophyly of
<taxonomicName authority="sensu Holloway" authorityName="sensu Holloway" box="[854,1238,791,817]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">
Desmobathrinae
<emphasis box="[1055,1117,792,817]" italics="true" pageId="19" pageNumber="20">sensu</emphasis>
Holloway
</taxonomicName>
because some species currently placed in
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[729,918,830,856]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
were embedded within the group (see also
<bibRefCitation author="Sihvonen" etAl="et al." firstAuthor="Sihvonen" journalOrPublisher="PLOS ONE" pageId="19" pageNumber="20" pagination="20356" part="6" refId="ref20700" refString="Sihvonen P, Mutanen M, Kaila L, Brehm G, Hausmann A, Staude HS. 2011. Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae). PLOS ONE 6 (6): e 20356 DOI 10.1371 / journal. pone. 0020356." title="Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae)" type="journal article" year="2011">
<emphasis italics="true" pageId="19" pageNumber="20">
<taxonomicName authority="Murillo-Ramos &amp; Brehm &amp; Sihvonen &amp; Hausmann &amp; Holm &amp; Ghanavi &amp; Õunap &amp; Truuverk &amp; Staude &amp; Friedrich &amp; Tammaru &amp; Wahlberg, 2019" authorityName="Murillo-Ramos &amp; Brehm &amp; Sihvonen &amp; Hausmann &amp; Holm &amp; Ghanavi &amp; Õunap &amp; Truuverk &amp; Staude &amp; Friedrich &amp; Tammaru &amp; Wahlberg" authorityYear="2019" box="[1434,1539,830,856]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al., 2011
</emphasis>
</bibRefCitation>
), and also the phylogenetic position of the tribe
<taxonomicName authorityName="Warren" authorityYear="1894" box="[1203,1321,870,896]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Eumeleini">Eumeleini</taxonomicName>
is unstable (see below).
<taxonomicName authorityName="sensu Holloway" box="[592,785,910,936]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
can be regarded as a monophyletic group after the transfer of
<emphasis box="[498,761,950,976]" italics="true" pageId="19" pageNumber="20">
<taxonomicName authorityName="Herrich-Schaffer" authorityYear="1855" box="[498,650,950,976]" class="Insecta" family="Geometridae" genus="Zanclopteryx" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">Zanclopteryx</taxonomicName>
,
<taxonomicName authorityName="Guest" authorityYear="1887" box="[664,761,950,976]" class="Insecta" family="Geometridae" genus="Nearcha" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">Nearcha</taxonomicName>
</emphasis>
and
<taxonomicName authorityName="Walker" authorityYear="1861" box="[822,912,951,976]" class="Insecta" family="Geometridae" genus="Racasta" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[822,912,951,976]" italics="true" pageId="19" pageNumber="20">Racasta</emphasis>
</taxonomicName>
from
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[987,1176,950,976]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
to
<taxonomicName authorityName="sensu Holloway" box="[1217,1412,950,976]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
, and the removal of
<taxonomicName authorityName="Warren" authorityYear="1894" box="[634,752,990,1016]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Eumeleini">Eumeleini</taxonomicName>
(
<tableCitation box="[770,858,990,1016]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="19" pageNumber="20">Table 2</tableCitation>
).
<taxonomicName authorityName="sensu Holloway" box="[883,1076,990,1016]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
as circumscribed here are an independent lineage that is sister to all
<taxonomicName authorityName="Leach" authorityYear="1815" box="[962,1113,1030,1056]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="family">Geometridae</taxonomicName>
except
<taxonomicName authorityName="Meyrick" authorityYear="1892" box="[1204,1326,1030,1056]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="9" phylum="Arthropoda" rank="subFamily" subFamily="Sterrhinae">Sterrhinae</taxonomicName>
,
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[1339,1471,1030,1056]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Larentiinae">Larentiinae</taxonomicName>
and
<taxonomicName authorityName="Fletcher" authorityYear="1953" box="[498,651,1070,1096]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Archiearinae">Archiearinae</taxonomicName>
.
</paragraph>
<paragraph blockId="19.[498,1542,549,1933]" lastBlockId="20.[498,1430,234,301]" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20">
The monobasic
<taxonomicName authorityName="Warren" authorityYear="1894" box="[720,838,1110,1136]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Eumeleini">Eumeleini</taxonomicName>
has had a dynamic taxonomic history:
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[1306,1405,1110,1136]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[1306,1405,1110,1136]" italics="true" pageId="19" pageNumber="20">Eumelea</emphasis>
</taxonomicName>
was transferred from
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[701,890,1149,1175]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
<emphasis box="[899,923,1149,1175]" italics="true" pageId="19" pageNumber="20">s.l</emphasis>
. to
<taxonomicName authorityName="sensu Holloway" box="[970,1163,1149,1175]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Desmobathrinae">Desmobathrinae</taxonomicName>
based on the pupal cremaster (
<bibRefCitation author="Holloway" box="[508,687,1189,1215]" firstAuthor="Holloway" journalOrPublisher="Ennominae Malayan Nature Journal" pageId="19" pageNumber="20" pagination="147 - 326" part="49" refId="ref18919" refString="Holloway J. 1996. The moths of Borneo, part 9: Geometridae (incl. Orthostixini), Oenochrominae, Desmobathrinae, Geometrinae. Ennominae Malayan Nature Journal 49: 147 - 326." title="The moths of Borneo, part 9: Geometridae (incl. Orthostixini), Oenochrominae, Desmobathrinae, Geometrinae" type="journal article" year="1996">
<emphasis box="[508,687,1189,1215]" italics="true" pageId="19" pageNumber="20">Holloway, 1996</emphasis>
</bibRefCitation>
), whereas
<bibRefCitation author="Beljaev" box="[812,976,1189,1215]" firstAuthor="Beljaev" journalOrPublisher="Entomological Review" pageId="19" pageNumber="20" pagination="50 - 60" part="88" refId="ref18236" refString="Beljaev EA. 2008. A new concept of the generic composition of the geometrid moth tribe Ennomini (Lepidoptera, Geometridae) based on functional morphology of the male genitalia. Entomological Review 88 (1): 50 - 60 DOI 10.1134 / S 0013873808010089." title="A new concept of the generic composition of the geometrid moth tribe Ennomini (Lepidoptera, Geometridae) based on functional morphology of the male genitalia" type="journal article" year="2008">
<emphasis box="[812,976,1189,1215]" italics="true" pageId="19" pageNumber="20">Beljaev (2008)</emphasis>
</bibRefCitation>
pointed out that
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[1180,1279,1189,1215]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[1180,1279,1189,1215]" italics="true" pageId="19" pageNumber="20">Eumelea</emphasis>
</taxonomicName>
could be a member of
<taxonomicName authorityName="Leach" authorityYear="1815" box="[498,649,1229,1255]" pageId="19" pageNumber="12" rank="subFamily" subFamily="Geometrinae">Geometrinae</taxonomicName>
based on the skeleto-muscular structure of the male genitalia. Molecular studies (
<bibRefCitation author="Sihvonen" box="[597,839,1269,1295]" etAl="et al." firstAuthor="Sihvonen" journalOrPublisher="PLOS ONE" pageId="19" pageNumber="20" pagination="20356" part="6" refId="ref20700" refString="Sihvonen P, Mutanen M, Kaila L, Brehm G, Hausmann A, Staude HS. 2011. Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae). PLOS ONE 6 (6): e 20356 DOI 10.1371 / journal. pone. 0020356." title="Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae)" type="journal article" year="2011">
<emphasis box="[597,839,1269,1295]" italics="true" pageId="19" pageNumber="20">
<taxonomicName authority="Murillo-Ramos &amp; Brehm &amp; Sihvonen &amp; Hausmann &amp; Holm &amp; Ghanavi &amp; Õunap &amp; Truuverk &amp; Staude &amp; Friedrich &amp; Tammaru &amp; Wahlberg, 2019" authorityName="Murillo-Ramos &amp; Brehm &amp; Sihvonen &amp; Hausmann &amp; Holm &amp; Ghanavi &amp; Õunap &amp; Truuverk &amp; Staude &amp; Friedrich &amp; Tammaru &amp; Wahlberg" authorityYear="2019" box="[597,702,1269,1295]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al., 2011
</emphasis>
</bibRefCitation>
,
<bibRefCitation author="Ban X &amp; Jiang N &amp; Cheng R &amp; Xue D &amp; Han H." box="[854,1037,1269,1295]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="19" pageNumber="20" pagination="653 - 672" part="184" refId="ref18087" refString="Ban X, Jiang N, Cheng R, Xue D, Han H. 2018. Tribal classification and phylogeny of Geometrinae (Lepidoptera: Geometridae) inferred from seven gene regions. Zoological Journal of the Linnean Society 184 (3): 653 - 672 DOI 10.1093 / zoolinnean / zly 013." title="Tribal classification and phylogeny of Geometrinae (Lepidoptera: Geometridae) inferred from seven gene regions" type="journal article" year="2018">
<emphasis box="[854,1037,1269,1295]" italics="true" pageId="19" pageNumber="20">Ban et al., 2018</emphasis>
</bibRefCitation>
) suggested that
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[1230,1329,1269,1295]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[1230,1329,1269,1295]" italics="true" pageId="19" pageNumber="20">Eumelea</emphasis>
</taxonomicName>
was part of
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[498,687,1309,1335]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
<emphasis box="[696,742,1310,1335]" italics="true" pageId="19" pageNumber="20">s.str</emphasis>
., but these findings were not well-supported and no formal taxonomic changes were proposed. Our analyses with IQTREE and RAxML recovered
<taxonomicName authorityName="Warren" authorityYear="1894" box="[498,616,1388,1414]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="tribe" tribe="Eumeleini">Eumeleini</taxonomicName>
in two very different positions, either as sister to
<taxonomicName authorityName="Leach" authorityYear="1815" box="[1198,1349,1388,1414]" pageId="19" pageNumber="12" rank="subFamily" subFamily="Geometrinae">Geometrinae</taxonomicName>
(SH-like = 93.6, UFBoot2 = 71) (
<figureCitation box="[689,763,1428,1454]" captionStart="Figure 4" captionStartId="10.[116,181,1038,1060]" captionTargetBox="[90,1342,236,978]" captionTargetId="graphics-225@10.[90,1168,244,977]" captionText="Figure 4 Phylogenetic relationships of the subfamilies Archierinae, Desmobathrinae, Epidesmiinae subfam. nov., Oenochrominae. Numbers above branches are SH-aLRT support (%)/ultrafast bootstrap support, UFBoot2(%), for nodes to the right of the numbers. Values of SH ≥ 80 and UFBoot2 ≥ 95 indicate well-supported clades (Trifinopoulos &amp; Minh, 2018). Taxonomic changes are indicated by a symobolized arrow&gt;. *Formal taxonomic treatment will be dealt with in P. Sihvonen et al., 2019, unpublished data. Full-size DOI: 10.7717/peerj.7386/fig-4" figureDoi="http://doi.org/10.5281/zenodo.5767538" httpUri="https://zenodo.org/record/5767538/files/figure.png" pageId="19" pageNumber="20">Figs. 4</figureCitation>
and
<figureCitation box="[820,834,1428,1455]" captionStart="Figure 5" captionStartId="11.[116,181,1827,1849]" captionTargetBox="[90,1274,236,1771]" captionTargetId="graphics-20@11.[90,880,244,1768]" captionText="Figure 5 Evolutionary relationships of the subfamily Geometrinae. Numbers above branches are SH-aLRT support (%)/ultrafast bootstrap support, UFBoot2(%), for nodes to the right of the numbers. Values of SH ≥ 80 and UFBoot2 ≥ 95 indicate well-supported clades (Trifinopoulos &amp; Minh, 2018). Taxonomic changes are indicated by a symobolized arrow&gt;. § New subfamily. Full-size DOI: 10.7717/peerj.7386/fig-5" figureDoi="http://doi.org/10.5281/zenodo.5767540" httpUri="https://zenodo.org/record/5767540/files/figure.png" pageId="19" pageNumber="20">5</figureCitation>
), or as sister of
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1016,1113,1428,1454]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[1016,1113,1428,1454]" italics="true" pageId="19" pageNumber="20">Plutodes</emphasis>
</taxonomicName>
in
<taxonomicName baseAuthorityName="Abraham et al." baseAuthorityYear="2001" box="[1150,1285,1428,1454]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
(RBS = 60) (Data S3). The examination of morphological details suggests that the position as sister to
<taxonomicName authorityName="Leach" authorityYear="1815" box="[498,649,1508,1534]" pageId="19" pageNumber="12" rank="subFamily" subFamily="Geometrinae">Geometrinae</taxonomicName>
is more plausible: hindwing vein M2 is present and tubular; anal margin of the hindwing is elongated; and large coremata originate from the saccus (
<bibRefCitation author="Holloway J." box="[1357,1536,1548,1574]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="19" pageNumber="20" refId="ref18891" refString="Holloway J. 1994. The moths of Borneo, part 11: family Geometridae, subfamily Ennominae. Malayan Nature Journal 47: 1 - 309." title="The moths of Borneo" type="book" year="1994">
<emphasis box="[1357,1536,1548,1574]" italics="true" pageId="19" pageNumber="20">Holloway, 1994</emphasis>
</bibRefCitation>
, our observations). The morphology of
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[957,1056,1588,1614]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[957,1056,1588,1614]" italics="true" pageId="19" pageNumber="20">Eumelea</emphasis>
</taxonomicName>
is partly unusual, and for that reason we illustrate selected structures (Data S4), which include for instance the following: antennae and legs of both sexes are very long; forewing vein Sc (homology unclear) reaches wing margin; in male genitalia coremata are extremely large and branched; uncus is crossshaped (cruciform); tegumen is narrow and it extends ventrally beyond the point of articulation with vinculum; saccus arms are extremely long, looped; and vesica is with lateral rows of cornuti. However, the green geoverdin pigment concentration of
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[1418,1517,1827,1853]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[1418,1517,1827,1853]" italics="true" pageId="19" pageNumber="20">Eumelea</emphasis>
</taxonomicName>
is low in comparison to
<taxonomicName authorityName="Leach" authorityYear="1815" box="[760,911,1867,1893]" pageId="19" pageNumber="12" rank="subFamily" subFamily="Geometrinae">Geometrinae</taxonomicName>
(
<bibRefCitation author="Cook MA &amp; Harwood LM &amp; Scoble MJ &amp; McGavin GC" box="[929,1125,1867,1893]" journalOrPublisher="Lepidoptera: Geometridae" pageId="19" pageNumber="20" pagination="43 - 51" part="22" refId="ref18378" refString="Cook MA, Harwood LM, Scoble MJ, McGavin GC. 1994. The chemistry and systematic importance of the green wing pigment in emerald moths (Lepidoptera: Geometridae, Geometrinae). Biochemical systematics and ecology 22 (1): 43 - 51 DOI 10.1016 / 0305 - 1978 (94) 90113 - 9." title="The chemistry and systematic importance of the green wing pigment in emerald moths" type="book" year="1994">
<emphasis box="[929,1125,1867,1893]" italics="true" pageId="19" pageNumber="20">Cook et al., 1994</emphasis>
</bibRefCitation>
). We tentatively conclude that
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[498,597,1906,1932]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis box="[498,597,1906,1932]" italics="true" pageId="19" pageNumber="20">Eumelea</emphasis>
</taxonomicName>
is probably indeed associated with
<taxonomicName authorityName="Leach" authorityYear="1815" box="[1018,1172,1906,1932]" pageId="19" pageNumber="12" rank="subFamily" subFamily="Geometrinae">Geometrinae</taxonomicName>
. However, since eleven genetic markers were not sufficient to clarify the phylogenetic affinities of
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[1284,1383,234,260]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis box="[1284,1383,234,260]" italics="true" pageId="20" pageNumber="21">Eumelea</emphasis>
</taxonomicName>
, we provisionally place the genus as
<emphasis box="[880,1034,274,300]" italics="true" pageId="20" pageNumber="21">incertae sedis</emphasis>
(
<tableCitation box="[1052,1139,274,300]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="20" pageNumber="21">Table 2</tableCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>