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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="038F87AD2B07FFAB2EC55947FB400E30" docLanguage="en" docName="PeerJ.7.e7386.pdf" docOrigin="PeerJ 7" docTitle="Ennominae Duponchel 1845" docType="treatment" docVersion="9" lastPageNumber="29" masterDocId="FFB6FFD52B1FFFB72F375F1EFFC20D03" masterDocTitle="A comprehensive molecular phylogeny of Geometridae (Lepidoptera) with a focus on enigmatic small subfamilies" masterLastPageNumber="39" masterPageNumber="1" pageNumber="25" 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>Friedrich, Egbert</mods:namePart>
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<mods:number>2019-08-27</mods:number>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5767485" ID-GBIF-Taxon="191256158" ID-Zenodo-Dep="5767485" LSID="urn:lsid:plazi:treatment:038F87AD2B07FFAB2EC55947FB400E30" httpUri="http://treatment.plazi.org/id/038F87AD2B07FFAB2EC55947FB400E30" lastPageId="28" lastPageNumber="29" pageId="24" pageNumber="25">
<subSubSection box="[498,855,1625,1651]" pageId="24" pageNumber="25" type="nomenclature">
<paragraph blockId="24.[498,1542,1625,1933]" box="[498,855,1625,1651]" pageId="24" pageNumber="25">
<heading bold="true" box="[498,855,1625,1651]" fontSize="11" level="3" pageId="24" pageNumber="25" reason="6">
<taxonomicName authority="Duponchel, 1845" authorityName="Duponchel" authorityYear="1845" box="[498,855,1625,1651]" class="Insecta" family="Geometridae" genus="Ennominae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="25" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[498,855,1625,1651]" pageId="24" pageNumber="25">Ennominae Duponchel, 1845</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection lastPageId="28" lastPageNumber="29" pageId="24" pageNumber="25" type="discussion">
<paragraph blockId="24.[498,1542,1625,1933]" pageId="24" pageNumber="25">
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[498,633,1667,1693]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
are the most species-rich subfamily of geometrids. The loss of vein M2 on the hindwing is probably the best apomorphy (
<bibRefCitation author="Holloway J." box="[994,1171,1707,1733]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="24" pageNumber="25" 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="[994,1171,1707,1733]" italics="true" pageId="24" pageNumber="25">Holloway, 1994</emphasis>
</bibRefCitation>
), although vein M2 is present as tubular in a few ennomine taxa (
<bibRefCitation author="Staude HS" box="[885,1033,1747,1773]" journalOrPublisher="Metamorphosis" pageId="24" pageNumber="25" pagination="125 - 156" part="12" refId="ref20942" refString="Staude HS. 2001. A revision of the genus Callioratis Felder (Lepidoptera: Geometridae: Diptychinae). Metamorphosis 12: 125 - 156." title="A revision of the genus Callioratis Felder (Lepidoptera: Geometridae: Diptychinae)" type="journal article" year="2001">
<emphasis box="[885,1033,1747,1773]" italics="true" pageId="24" pageNumber="25">Staude, 2001</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[1047,1318,1747,1774]" journalOrPublisher="Stenstrup: Apollo Books" pageId="24" pageNumber="25" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[1047,1318,1747,1774]" italics="true" pageId="24" pageNumber="25">
Skou &amp;
<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="[1142,1250,1747,1773]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
).
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[1342,1477,1747,1773]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
are a morphologically highly diverse subfamily, and attempts to find further synapomorphies shared by all major tribal groups have failed.
</paragraph>
<paragraph blockId="24.[498,1542,1625,1933]" lastBlockId="25.[498,1542,234,1934]" lastPageId="25" lastPageNumber="26" pageId="24" pageNumber="25">
The number of tribes as well as phylogenetic relationships among tribes are still debated (see
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[551,819,1906,1933]" journalOrPublisher="Stenstrup: Apollo Books" pageId="24" pageNumber="25" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[551,819,1906,1933]" italics="true" pageId="24" pageNumber="25">
Skou &amp;
<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="[645,753,1906,1932]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="24" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
for an overview). Moreover, the taxonomic knowledge of this subfamily in tropical regions is still poor.
<bibRefCitation author="Holloway J." box="[992,1184,234,260]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="25" pageNumber="26" 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="[992,1184,234,260]" italics="true" pageId="25" pageNumber="26">Holloway (1994)</emphasis>
</bibRefCitation>
recognized 21 tribes,
<bibRefCitation author="Beljaev EA" journalOrPublisher="Spixiana" pageId="25" pageNumber="26" pagination="215 - 216" part="29" refId="ref18211" refString="Beljaev EA. 2006. A morphological approach to the Ennominae phylogeny (Lepidoptera, Geometridae). Spixiana 29: 215 - 216." title="A morphological approach to the Ennominae phylogeny (Lepidoptera, Geometridae)" type="journal article" year="2006">
<emphasis italics="true" pageId="25" pageNumber="26">Beljaev (2006)</emphasis>
</bibRefCitation>
24 tribes, and
<bibRefCitation accessDate="2018-08-03" author="Forum Herbulot" box="[740,1007,274,300]" pageId="25" pageNumber="26" publicationUrl="http://www.herbulot.de/famgroup.htm" refId="ref18517" refString="Forum Herbulot. 2007. World list of family-group names in Geometridae. Available at http: // www. herbulot. de / famgroup. htm (accessed 3 August 2018)." title="World list of family-group names in Geometridae" type="url" year="2007">
<emphasis box="[740,1007,274,300]" italics="true" pageId="25" pageNumber="26">Forum Herbulot (2007)</emphasis>
</bibRefCitation>
27 tribes. To date, four molecular studies have corroborated the monophyly of
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[878,1013,314,340]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
(
<bibRefCitation author="Yamamoto S &amp; Sota T." box="[1031,1320,314,340]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="25" pageNumber="26" pagination="711 - 723" part="44" refId="ref21575" refString="Yamamoto S, Sota T. 2007. Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 44 (2): 711 - 723 DOI 10.1016 / j. ympev. 2006.12.027." title="Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes" type="journal article" year="2007">
<emphasis box="[1031,1320,314,340]" italics="true" pageId="25" pageNumber="26">Yamamoto &amp; Sota, 2007</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Wahlberg N &amp; Snall N &amp; Viidalepp J &amp; Ruohomaki K &amp; Tammaru T." journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="25" pageNumber="26" pagination="929 - 938" part="55" refId="ref21478" refString="Wahlberg N, Snall N, Viidalepp J, Ruohomaki K, Tammaru T. 2010. The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae). Molecular Phylogenetics and Evolution 55 (3): 929 - 938 DOI 10.1016 / j. ympev. 2010.01.025." title="The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae)" type="journal article" year="2010">
<emphasis italics="true" pageId="25" pageNumber="26">Wahlberg et al., 2010</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Ounap E &amp; Javois J &amp; Viidalepp J &amp; Tammaru T." box="[566,774,354,381]" journalOrPublisher="European Journal of Entomology" pageId="25" pageNumber="26" pagination="267 - 273" part="108" refId="ref19698" refString="Ounap E, Javois J, Viidalepp J, Tammaru T. 2011. Phylogenetic relationships of selected European Ennominae (Lepidoptera: Geometridae). European Journal of Entomology 108 (2): 267 - 273 DOI 10.14411 / eje. 2011.036." title="Phylogenetic relationships of selected European Ennominae (Lepidoptera: Geometridae)" type="journal article" year="2011">
<emphasis box="[566,774,354,381]" italics="true" pageId="25" pageNumber="26">Õunap et al., 2011</emphasis>
</bibRefCitation>
,
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[786,1019,354,380]" journalOrPublisher="PLOS ONE" pageId="25" pageNumber="26" 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="[786,1019,354,380]" italics="true" pageId="25" pageNumber="26">
<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="[786,891,354,380]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al., 2011
</emphasis>
</bibRefCitation>
), with
<bibRefCitation author="Young CJ" box="[1099,1254,354,380]" journalOrPublisher="Zootaxa" pageId="25" pageNumber="26" pagination="1 - 147" part="1264" refId="ref21647" refString="Young CJ. 2006. Molecular relationships of the Australian Ennominae (Lepidoptera: Geometridae) and implications for the phylogeny of the Geometridae from molecular and morphological data. Zootaxa 1264 (1): 1 - 147 DOI 10.11646 / zootaxa. 1264.1.1." title="Molecular relationships of the Australian Ennominae (Lepidoptera: Geometridae) and implications for the phylogeny of the Geometridae from molecular and morphological data" type="journal article" year="2006">
<emphasis box="[1099,1254,354,380]" italics="true" pageId="25" pageNumber="26">Young (2006)</emphasis>
</bibRefCitation>
being the only exception who found
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[635,770,394,420]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
paraphyletic. Moreover, four large-scale taxonomic revisions (without a phylogenetic hypothesis) were published by
<bibRefCitation author="Pitkin L." box="[1151,1303,433,460]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="25" pageNumber="26" pagination="121 - 401" part="135" refId="ref19983" refString="Pitkin L. 2002. Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society 135 (2 - 3): 121 - 401 DOI 10.1046 / j. 1096 - 3642.2002.01200. x." title="Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae)" type="journal article" year="2002">
<emphasis box="[1151,1303,433,460]" italics="true" pageId="25" pageNumber="26">Pitkin (2002)</emphasis>
</bibRefCitation>
for the Neotropical region,
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[588,872,473,500]" journalOrPublisher="Stenstrup: Apollo Books" pageId="25" pageNumber="26" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[588,872,473,500]" italics="true" pageId="25" pageNumber="26">
Skou &amp;
<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="[684,789,473,499]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
(2015)
</emphasis>
</bibRefCitation>
,
<bibRefCitation author="Muller B &amp; Erlacher S &amp; Hausmann A &amp; Rajaei H &amp; Sihvonen P &amp; Skou P." box="[887,1112,473,499]" pageId="25" pageNumber="26" refId="ref19595" refString="Muller B, Erlacher S, Hausmann A, Rajaei H, Sihvonen P, Skou P. 2019. Ennominae II. In: Hausmann A, Sihvonen P, Rajaei H, Skou P, eds. Geometrid Moths of Europe. Vol. 6. Leiden: Brill, 906." title="Ennominae II" type="book" year="2019">
<emphasis box="[887,1112,473,499]" italics="true" pageId="25" pageNumber="26">Müller et al. (2019)</emphasis>
</bibRefCitation>
for the Western Palaearctic region, and
<bibRefCitation author="Holloway J." box="[551,743,513,539]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="25" pageNumber="26" 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="[551,743,513,539]" italics="true" pageId="25" pageNumber="26">Holloway (1994)</emphasis>
</bibRefCitation>
for Borneo. More detailed descriptions of taxonomic changes in
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[498,633,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
will be given by G.
<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="[872,951,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="29" phylum="Arthropoda" rank="tribe" status="new tribe" tribe="Brehm">Brehm</taxonomicName>
et al. (2019, unpublished data) and L. MurilloRamos et al. (2019, unpublished data). We here discuss general patterns and give details for taxonomic acts not covered in the other two papers.
</paragraph>
<paragraph blockId="25.[498,1542,234,1934]" pageId="25" pageNumber="26">
Our findings recover
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[784,919,673,699]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
as a monophyletic entity, but results were not highly supported in RAxML (RBS = 67) compared to IQ-TREE (SH-Like =100, UFBoot2 = 99). The lineage comprising
<taxonomicName authorityName="Leach" authorityYear="1815" box="[836,987,752,778]" pageId="25" pageNumber="12" rank="subFamily" subFamily="Geometrinae">Geometrinae</taxonomicName>
and
<taxonomicName authorityName="Guenee. Meyrick" authorityYear="1858" box="[1048,1237,752,778]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="subFamily" subFamily="Oenochrominae">Oenochrominae</taxonomicName>
is recovered as the sister clade of
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[599,737,792,818]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
. In previous studies,
<bibRefCitation author="Wahlberg N &amp; Snall N &amp; Viidalepp J &amp; Ruohomaki K &amp; Tammaru T." box="[986,1247,792,818]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="25" pageNumber="26" pagination="929 - 938" part="55" refId="ref21478" refString="Wahlberg N, Snall N, Viidalepp J, Ruohomaki K, Tammaru T. 2010. The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae). Molecular Phylogenetics and Evolution 55 (3): 929 - 938 DOI 10.1016 / j. ympev. 2010.01.025." title="The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae)" type="journal article" year="2010">
<emphasis box="[986,1247,792,818]" italics="true" pageId="25" pageNumber="26">Wahlberg et al. (2010)</emphasis>
</bibRefCitation>
sampled 49 species of
<taxonomicName authority=", Ounap et al. (2011)" authorityName=", Ounap et al." authorityYear="2011" box="[498,872,832,859]" class="Insecta" family="Geometridae" genus="Ennominae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="genus">
Ennominae,
<bibRefCitation author="Ounap E &amp; Javois J &amp; Viidalepp J &amp; Tammaru T." box="[647,872,832,859]" journalOrPublisher="European Journal of Entomology" pageId="25" pageNumber="26" pagination="267 - 273" part="108" refId="ref19698" refString="Ounap E, Javois J, Viidalepp J, Tammaru T. 2011. Phylogenetic relationships of selected European Ennominae (Lepidoptera: Geometridae). European Journal of Entomology 108 (2): 267 - 273 DOI 10.14411 / eje. 2011.036." title="Phylogenetic relationships of selected European Ennominae (Lepidoptera: Geometridae)" type="journal article" year="2011">
<emphasis box="[647,872,832,859]" italics="true" pageId="25" pageNumber="26">Õunap et al. (2011)</emphasis>
</bibRefCitation>
</taxonomicName>
sampled 33 species, and
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[1167,1417,832,858]" journalOrPublisher="PLOS ONE" pageId="25" pageNumber="26" 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="[1167,1417,832,858]" italics="true" pageId="25" pageNumber="26">
<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="[1167,1272,832,858]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al. (2011)
</emphasis>
</bibRefCitation>
70 species including up to eight markers per species. All these studies supported the division of
<taxonomicName authorityName=", Ounap et al." authorityYear="2011" box="[498,633,912,938]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
into “boarmiine” and “ennomine” moths (
<bibRefCitation author="Holloway J." box="[1128,1306,912,938]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="25" pageNumber="26" 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="[1128,1306,912,938]" italics="true" pageId="25" pageNumber="26">Holloway, 1994</emphasis>
</bibRefCitation>
). This grouping was proposed by
<bibRefCitation author="Forbes WTM" box="[652,809,951,978]" journalOrPublisher="Memoirs of the Cornell University Agricultural Experiment Station" pageId="25" pageNumber="26" pagination="1 - 263" part="274" refId="ref18488" refString="Forbes WTM. 1948. Lepidoptera of New York and neighboring states. II. Memoirs of the Cornell University Agricultural Experiment Station 274: 1 - 263." title="Lepidoptera of New York and neighboring states. II" type="journal article" year="1948">
<emphasis box="[652,809,951,978]" italics="true" pageId="25" pageNumber="26">Forbes (1948)</emphasis>
</bibRefCitation>
and
<bibRefCitation author="Holloway J." box="[871,1063,951,978]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="25" pageNumber="26" 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="[871,1063,951,978]" italics="true" pageId="25" pageNumber="26">Holloway (1994)</emphasis>
</bibRefCitation>
, who suggested close relationships between the tribes
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[717,837,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Boarmiini">Boarmiini</taxonomicName>
, Macariini,
<taxonomicName authorityName="Holloway" authorityYear="1994" box="[975,1096,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Cassymini">Cassymini</taxonomicName>
and
<taxonomicName authorityName="Holloway" authorityYear="1994" box="[1155,1253,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Eutoeini">Eutoeini</taxonomicName>
based on the bifid pupal cremaster and the possession of a fovea in the male forewing. The remaining tribes were defined as “ennomines” based on the loss of a setal comb on male sternum A3 and the presence of a strong furca in male genitalia. Both
<bibRefCitation author="Wahlberg N &amp; Snall N &amp; Viidalepp J &amp; Ruohomaki K &amp; Tammaru T." box="[1065,1319,1111,1137]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="25" pageNumber="26" pagination="929 - 938" part="55" refId="ref21478" refString="Wahlberg N, Snall N, Viidalepp J, Ruohomaki K, Tammaru T. 2010. The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae). Molecular Phylogenetics and Evolution 55 (3): 929 - 938 DOI 10.1016 / j. ympev. 2010.01.025." title="The evolution of female flightlessness among Ennominae of the Holarctic forest zone (Lepidoptera, Geometridae)" type="journal article" year="2010">
<emphasis box="[1065,1319,1111,1137]" italics="true" pageId="25" pageNumber="26">Wahlberg et al. (2010)</emphasis>
</bibRefCitation>
and
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" journalOrPublisher="PLOS ONE" pageId="25" pageNumber="26" 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="25" pageNumber="26">
<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="[1375,1480,1111,1137]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al. (2011)
</emphasis>
</bibRefCitation>
found these two informal groupings to be reciprocally monophyletic.
</paragraph>
<paragraph blockId="25.[498,1542,234,1934]" pageId="25" pageNumber="26">
In our analyses, 653 species with up to 11 markers were sampled, with an emphasis on Neotropical taxa, which so far had been poorly represented in the molecular phylogenetic analyses. Our results recovered the division into two major subclades (
<figureCitation box="[1332,1398,1270,1296]" captionStart="Figure 6" captionStartId="14.[116,181,248,270]" captionTargetId="graphics-17@13.[498,1097,240,1837]" captionTargetPageId="13" captionText="Figure 6 Evolutionary relationships of the subfamily Ennominae. 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 G. Brehm et al., 2019, unpublished data. Full-size DOI: 10.7717/peerj.7386/fig-6" figureDoi="http://doi.org/10.5281/zenodo.5767542" httpUri="https://zenodo.org/record/5767542/files/figure.png" pageId="25" pageNumber="26">Fig. 6</figureCitation>
), a core set of ennomines in a well-supported clade, and a poorly supported larger clade that includes the “boarmiines” among four other lineages usually thought of as &quot;ennomines&quot;. The traditional “ennomines” are thus not found to be monophyletic in our analyses, questioning the utility of such an informal name. Our phylogenetic hypothesis supports the validation of numerous tribes proposed previously, in addition to several unnamed clades. We validate 23 tribes (
<bibRefCitation accessDate="2018-08-03" author="Forum Herbulot" box="[852,1111,1509,1535]" pageId="25" pageNumber="26" publicationUrl="http://www.herbulot.de/famgroup.htm" refId="ref18517" refString="Forum Herbulot. 2007. World list of family-group names in Geometridae. Available at http: // www. herbulot. de / famgroup. htm (accessed 3 August 2018)." title="World list of family-group names in Geometridae" type="url" year="2007">
<emphasis box="[852,1111,1509,1535]" italics="true" pageId="25" pageNumber="26">Forum Herbulot, 2007</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[1126,1398,1509,1536]" journalOrPublisher="Stenstrup: Apollo Books" pageId="25" pageNumber="26" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[1126,1398,1509,1536]" italics="true" pageId="25" pageNumber="26">
Skou &amp;
<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="[1222,1330,1509,1535]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
):
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[498,651,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Gonodontini">Gonodontini</taxonomicName>
,
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[665,780,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
,
<taxonomicName authorityName="Tutt" authorityYear="1896" box="[792,954,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Odontoperini">Odontoperini</taxonomicName>
,
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[967,1117,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
,
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[1131,1250,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
,
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1263,1392,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Campaeini">Campaeini</taxonomicName>
,
<taxonomicName authorityName="Herbulot" authorityYear="1962" box="[1405,1536,1549,1575]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Alsophilini">Alsophilini</taxonomicName>
, Wilemaniini,
<taxonomicName authorityName="Warren" authorityYear="1894" box="[660,842,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Prosopolophini">Prosopolophini</taxonomicName>
,
<taxonomicName authorityName="Janse" authorityYear="1933" box="[855,982,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
,
<taxonomicName authorityName="Herbulot" authorityYear="1963" box="[995,1092,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Theriini">Theriini</taxonomicName>
, Plutodini,
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1229,1340,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Palyadini">Palyadini</taxonomicName>
,
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1353,1515,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Hypochrosini">Hypochrosini</taxonomicName>
,
<taxonomicName authorityName="Stekolnikov &amp; Kuznetzov" authorityYear="1982" box="[498,597,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Apeirini">Apeirini</taxonomicName>
,
<taxonomicName authorityName="Bruand" authorityYear="1846" box="[610,713,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Epionini">Epionini</taxonomicName>
,
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[726,828,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Caberini">Caberini</taxonomicName>
, Macariini,
<taxonomicName authorityName="Holloway" authorityYear="1994" box="[969,1092,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Cassymini">Cassymini</taxonomicName>
,
<taxonomicName authorityName="Warren" authorityYear="1893" box="[1106,1211,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Abraxini">Abraxini</taxonomicName>
,
<taxonomicName authorityName="Holloway" authorityYear="1994" box="[1224,1322,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Eutoeini">Eutoeini</taxonomicName>
and
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[1383,1503,1629,1655]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Boarmiini">Boarmiini</taxonomicName>
.
</paragraph>
<paragraph blockId="25.[498,1542,234,1934]" pageId="25" pageNumber="26">
We hereby propose one new tribe:
<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="[903,1079,1669,1695]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" status="trib. nov." tribe="Drepanogynini">Drepanogynini</taxonomicName>
<emphasis bold="true" box="[1086,1198,1669,1695]" pageId="25" pageNumber="26">
<taxonomicNameLabel box="[1086,1198,1669,1695]" pageId="25" pageNumber="26" rank="tribe">trib. nov.</taxonomicNameLabel>
</emphasis>
(
<tableCitation box="[1215,1301,1669,1695]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="25" pageNumber="26">Table 2</tableCitation>
). Except for the new tribe, most of the groups recovered in this study are in concordance with previous morphological classifications (
<bibRefCitation author="Holloway J." box="[852,1031,1748,1775]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="25" pageNumber="26" 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="[852,1031,1748,1775]" italics="true" pageId="25" pageNumber="26">Holloway, 1994</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Beljaev EA" box="[1045,1196,1748,1774]" journalOrPublisher="Spixiana" pageId="25" pageNumber="26" pagination="215 - 216" part="29" refId="ref18211" refString="Beljaev EA. 2006. A morphological approach to the Ennominae phylogeny (Lepidoptera, Geometridae). Spixiana 29: 215 - 216." title="A morphological approach to the Ennominae phylogeny (Lepidoptera, Geometridae)" type="journal article" year="2006">
<emphasis box="[1045,1196,1748,1774]" italics="true" pageId="25" pageNumber="26">Beljaev, 2006</emphasis>
</bibRefCitation>
,
<bibRefCitation author="Beljaev E." box="[1209,1264,1748,1774]" journalOrPublisher="Vladivostok: Dalnauka" pageId="25" pageNumber="26" part="812" refId="ref18284" refString="Beljaev E. 2016. Annotated catalogue of the insects of Russian Far East. Volume II. Lepidoptera. Vladivostok: Dalnauka, 812." title="Annotated catalogue of the insects of Russian Far East. Volume II. Lepidoptera" type="book" year="2016">
<emphasis box="[1209,1264,1748,1774]" italics="true" pageId="25" pageNumber="26">2016</emphasis>
</bibRefCitation>
;
<bibRefCitation accessDate="2018-08-03" author="Forum Herbulot" box="[1278,1536,1748,1775]" pageId="25" pageNumber="26" publicationUrl="http://www.herbulot.de/famgroup.htm" refId="ref18517" refString="Forum Herbulot. 2007. World list of family-group names in Geometridae. Available at http: // www. herbulot. de / famgroup. htm (accessed 3 August 2018)." title="World list of family-group names in Geometridae" type="url" year="2007">
<emphasis box="[1278,1536,1748,1775]" italics="true" pageId="25" pageNumber="26">Forum Herbulot, 2007</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[498,770,1788,1815]" journalOrPublisher="Stenstrup: Apollo Books" pageId="25" pageNumber="26" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[498,770,1788,1815]" italics="true" pageId="25" pageNumber="26">
Skou &amp;
<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="[594,702,1788,1814]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Muller B &amp; Erlacher S &amp; Hausmann A &amp; Rajaei H &amp; Sihvonen P &amp; Skou P." box="[785,998,1788,1814]" pageId="25" pageNumber="26" refId="ref19595" refString="Muller B, Erlacher S, Hausmann A, Rajaei H, Sihvonen P, Skou P. 2019. Ennominae II. In: Hausmann A, Sihvonen P, Rajaei H, Skou P, eds. Geometrid Moths of Europe. Vol. 6. Leiden: Brill, 906." title="Ennominae II" type="book" year="2019">
<emphasis box="[785,998,1788,1814]" italics="true" pageId="25" pageNumber="26">Müller et al., 2019</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="25.[498,1542,234,1934]" pageId="25" pageNumber="26">
Five known tribes and two further unnamed lineages (E1, E2 in
<figureCitation box="[1291,1357,1828,1854]" captionStart="Figure 6" captionStartId="14.[116,181,248,270]" captionTargetId="graphics-17@13.[498,1097,240,1837]" captionTargetPageId="13" captionText="Figure 6 Evolutionary relationships of the subfamily Ennominae. 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 G. Brehm et al., 2019, unpublished data. Full-size DOI: 10.7717/peerj.7386/fig-6" figureDoi="http://doi.org/10.5281/zenodo.5767542" httpUri="https://zenodo.org/record/5767542/files/figure.png" pageId="25" pageNumber="26">Fig. 6</figureCitation>
) form the core
<taxonomicName authorityName=", Ounap et al." authorityYear="2011" box="[498,636,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
:
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[648,801,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Gonodontini">Gonodontini</taxonomicName>
,
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[814,929,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
,
<taxonomicName authorityName="Tutt" authorityYear="1896" box="[942,1104,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Odontoperini">Odontoperini</taxonomicName>
,
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[1117,1265,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
and
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[1326,1446,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="26" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
.
</paragraph>
<paragraph blockId="25.[498,1542,234,1934]" lastBlockId="26.[498,1542,234,1934]" lastPageId="26" lastPageNumber="27" pageId="25" pageNumber="26">
Several Neotropical clades that conflict with the current tribal classification of
<taxonomicName authorityName=", Ounap et al." authorityYear="2011" box="[1407,1542,1908,1934]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="25" pageNumber="35" phylum="Arthropoda" rank="subFamily" subFamily="Ennominae">Ennominae</taxonomicName>
will be described as new tribes by G.
<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="[933,1012,234,260]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="29" phylum="Arthropoda" rank="tribe" status="new tribe" tribe="Brehm">Brehm</taxonomicName>
et al. (2019, unpublished data).
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1391,1542,234,260]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gonodontini">Gonodontini</taxonomicName>
and
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[550,663,274,300]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
are recovered as sister taxa.
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[995,1146,274,300]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gonodontini">Gonodontini</taxonomicName>
was defined by
<bibRefCitation author="Forbes WTM" box="[1334,1490,274,300]" journalOrPublisher="Memoirs of the Cornell University Agricultural Experiment Station" pageId="26" pageNumber="27" pagination="1 - 263" part="274" refId="ref18488" refString="Forbes WTM. 1948. Lepidoptera of New York and neighboring states. II. Memoirs of the Cornell University Agricultural Experiment Station 274: 1 - 263." title="Lepidoptera of New York and neighboring states. II" type="journal article" year="1948">
<emphasis box="[1334,1490,274,300]" italics="true" pageId="26" pageNumber="27">Forbes (1948)</emphasis>
</bibRefCitation>
and studied by
<bibRefCitation author="Holloway J." box="[626,817,314,340]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="26" pageNumber="27" 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="[626,817,314,340]" italics="true" pageId="26" pageNumber="27">Holloway (1994)</emphasis>
</bibRefCitation>
, who showed synapomorphies shared by
<taxonomicName authority="Hubner, (1823)" authorityName="Hubner" authorityYear="1823" class="Insecta" family="Geometridae" genus="Gonodontis" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[1304,1437,314,340]" italics="true" pageId="26" pageNumber="27">Gonodontis</emphasis>
Hübner, (1823)
</taxonomicName>
,
<taxonomicName authority="Warren, 1898" authorityName="Warren" authorityYear="1898" box="[588,899,354,380]" class="Insecta" family="Geometridae" genus="Xylinophylla" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[588,733,354,380]" italics="true" pageId="26" pageNumber="27">Xylinophylla</emphasis>
Warren, 1898
</taxonomicName>
and
<taxonomicName authority="Warren, 1895" authorityName="Warren" authorityYear="1895" box="[957,1239,354,381]" class="Insecta" family="Geometridae" genus="Xenimpia" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[957,1070,355,380]" italics="true" pageId="26" pageNumber="27">Xenimpia</emphasis>
Warren, 1895
</taxonomicName>
. Our results recovered the genus
<taxonomicName authorityName="Warren" authorityYear="1898" box="[574,719,394,420]" class="Insecta" family="Geometridae" genus="Xylinophylla" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[574,719,394,420]" italics="true" pageId="26" pageNumber="27">Xylinophylla</emphasis>
</taxonomicName>
as sister of
<taxonomicName authorityName="Warren" authorityYear="1895" box="[862,975,394,419]" class="Insecta" family="Geometridae" genus="Xenimpia" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[862,975,394,419]" italics="true" pageId="26" pageNumber="27">Xenimpia</emphasis>
</taxonomicName>
and
<taxonomicName authority="Warren, 1898" authorityName="Warren" authorityYear="1898" box="[1036,1332,394,420]" class="Insecta" family="Geometridae" genus="Psilocladia" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[1036,1159,394,420]" italics="true" pageId="26" pageNumber="27">Psilocladia</emphasis>
Warren, 1898
</taxonomicName>
.
<taxonomicName authorityName="Warren" authorityYear="1898" box="[1344,1467,394,420]" class="Insecta" family="Geometridae" genus="Psilocladia" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[1344,1467,394,420]" italics="true" pageId="26" pageNumber="27">Psilocladia</emphasis>
</taxonomicName>
is an African genus currently unassigned to tribe (see
<bibRefCitation author="Sihvonen P &amp; Staude HS &amp; Mutanen M." box="[1072,1488,433,460]" journalOrPublisher="Systematic Entomology" pageId="26" pageNumber="27" pagination="606 - 627" part="40" refId="ref20789" refString="Sihvonen P, Staude HS, Mutanen M. 2015. Systematic position of the enigmatic African cycad moths: an integrative approach to a nearly century old problem (Lepidoptera: Geometridae, Diptychini). Systematic Entomology 40 (3): 606 - 627 DOI 10.1111 / syen. 12125." title="Systematic position of the enigmatic African cycad moths: an integrative approach to a nearly century old problem (Lepidoptera: Geometridae, Diptychini)" type="journal article" year="2015">
<emphasis box="[1072,1488,433,460]" italics="true" pageId="26" pageNumber="27">
<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="[1072,1180,433,459]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, Staude &amp; Mutanen, 2015
</emphasis>
</bibRefCitation>
for details). Considering the strong support and that the facies and morphology are somewhat similar to other analyzed taxa in
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[889,1042,513,539]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gonodontini">Gonodontini</taxonomicName>
, we formally include
<taxonomicName authorityName="Warren" authorityYear="1898" box="[1297,1420,513,539]" class="Insecta" family="Geometridae" genus="Psilocladia" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[1297,1420,513,539]" italics="true" pageId="26" pageNumber="27">Psilocladia</emphasis>
</taxonomicName>
in
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[498,649,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gonodontini">Gonodontini</taxonomicName>
(
<tableCitation box="[668,755,553,579]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="26" pageNumber="27">Table 2</tableCitation>
).
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[779,892,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
are monophyletic and we formally transfer the African genera
<taxonomicName authority="Warren, 1902" authorityName="Warren" authorityYear="1902" box="[584,880,593,619]" class="Insecta" family="Geometridae" genus="Oedicentra" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[584,710,593,619]" italics="true" pageId="26" pageNumber="27">Oedicentra</emphasis>
Warren, 1902
</taxonomicName>
and
<taxonomicName authority="Janse, 1932" authorityName="Janse" authorityYear="1932" box="[942,1244,593,619]" class="Insecta" family="Geometridae" genus="Hypotephrina" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[942,1102,593,619]" italics="true" pageId="26" pageNumber="27">Hypotephrina</emphasis>
Janse, 1932
</taxonomicName>
, from unassigned to
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[498,611,633,659]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
(
<tableCitation box="[630,717,633,659]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="26" pageNumber="27">Table 2</tableCitation>
). The total number of species, and number of included genera in
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[498,611,673,699]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
are still uncertain (
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[845,1116,673,699]" journalOrPublisher="Stenstrup: Apollo Books" pageId="26" pageNumber="27" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[845,1116,673,699]" italics="true" pageId="26" pageNumber="27">
Skou &amp;
<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="[940,1048,673,699]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Muller B &amp; Erlacher S &amp; Hausmann A &amp; Rajaei H &amp; Sihvonen P &amp; Skou P." box="[1131,1344,673,699]" pageId="26" pageNumber="27" refId="ref19595" refString="Muller B, Erlacher S, Hausmann A, Rajaei H, Sihvonen P, Skou P. 2019. Ennominae II. In: Hausmann A, Sihvonen P, Rajaei H, Skou P, eds. Geometrid Moths of Europe. Vol. 6. Leiden: Brill, 906." title="Ennominae II" type="book" year="2019">
<emphasis box="[1131,1344,673,699]" italics="true" pageId="26" pageNumber="27">Müller et al., 2019</emphasis>
</bibRefCitation>
). Based on morphological examination,
<bibRefCitation author="Beljaev E." box="[833,995,712,738]" journalOrPublisher="Vladivostok: Dalnauka" pageId="26" pageNumber="27" part="812" refId="ref18284" refString="Beljaev E. 2016. Annotated catalogue of the insects of Russian Far East. Volume II. Lepidoptera. Vladivostok: Dalnauka, 812." title="Annotated catalogue of the insects of Russian Far East. Volume II. Lepidoptera" type="book" year="2016">
<emphasis box="[833,995,712,738]" italics="true" pageId="26" pageNumber="27">Beljaev (2016)</emphasis>
</bibRefCitation>
treated
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1089,1222,712,738]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Angeronini">Angeronini</taxonomicName>
as a synonym of
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[1422,1538,712,738]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
. The costal projection on male valva bearing a spine or group of spines was considered as a synapomorphy of the group. Using molecular data,
<bibRefCitation author="Yamamoto S &amp; Sota T." box="[1111,1411,792,819]" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="26" pageNumber="27" pagination="711 - 723" part="44" refId="ref21575" refString="Yamamoto S, Sota T. 2007. Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 44 (2): 711 - 723 DOI 10.1016 / j. ympev. 2006.12.027." title="Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes" type="journal article" year="2007">
<emphasis box="[1111,1411,792,819]" italics="true" pageId="26" pageNumber="27">Yamamoto &amp; Sota (2007)</emphasis>
</bibRefCitation>
showed a close phylogenetic relationship between
<taxonomicName authority="Duponchel, 1829" authorityName="Duponchel" authorityYear="1829" box="[994,1316,832,858]" class="Insecta" family="Geometridae" genus="Angerona" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[994,1107,832,858]" italics="true" pageId="26" pageNumber="27">Angerona</emphasis>
Duponchel, 1829
</taxonomicName>
(
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1332,1470,832,859]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Angeronini">Angeronini</taxonomicName>
) and
<taxonomicName authority="Wehrli, 1953" authorityName="Wehrli" authorityYear="1953" box="[498,823,872,899]" class="Insecta" family="Geometridae" genus="Chariaspilates" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[498,661,872,898]" italics="true" pageId="26" pageNumber="27">Chariaspilates</emphasis>
Wehrli, 1953
</taxonomicName>
(
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[840,959,872,899]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
). Similar results were shown by
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[1341,1513,872,898]" journalOrPublisher="PLOS ONE" pageId="26" pageNumber="27" 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="[1341,1513,872,898]" italics="true" pageId="26" pageNumber="27">
<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="[1341,1446,872,898]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al.
</emphasis>
</bibRefCitation>
<emphasis box="[498,572,912,938]" italics="true" pageId="26" pageNumber="27">(2011)</emphasis>
who recovered
<taxonomicName authorityName="Duponchel" authorityYear="1829" box="[762,875,912,938]" class="Insecta" family="Geometridae" genus="Angerona" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[762,875,912,938]" italics="true" pageId="26" pageNumber="27">Angerona</emphasis>
</taxonomicName>
and
<taxonomicName authority="Curtis, 1826" authorityName="Curtis" authorityYear="1826" box="[935,1187,911,938]" class="Insecta" family="Geometridae" genus="Charissa" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[935,1035,912,938]" italics="true" pageId="26" pageNumber="27">Charissa</emphasis>
Curtis, 1826
</taxonomicName>
as sister taxa, and our results also strongly support treating
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[854,987,951,977]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Angeronini">Angeronini</taxonomicName>
as synonym of
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[1176,1292,951,977]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Gnophini">Gnophini</taxonomicName>
.
</paragraph>
<paragraph blockId="26.[498,1542,234,1934]" pageId="26" pageNumber="27">
<bibRefCitation author="Holloway J." box="[530,722,991,1018]" journalOrPublisher="part 11: family Geometridae, subfamily Ennominae" pageId="26" pageNumber="27" 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="[530,722,991,1018]" italics="true" pageId="26" pageNumber="27">Holloway (1994)</emphasis>
</bibRefCitation>
suggested close affinities among
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[1114,1264,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
,
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1277,1383,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Azelinini">Azelinini</taxonomicName>
and
<taxonomicName authorityName="Tutt" authorityYear="1896" box="[498,658,1031,1057]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Odontoperini">Odontoperini</taxonomicName>
on the basis of larval characters. In a morphology-based phylogenetic study,
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[498,782,1071,1098]" journalOrPublisher="Stenstrup: Apollo Books" pageId="26" pageNumber="27" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[498,782,1071,1098]" italics="true" pageId="26" pageNumber="27">
Skou &amp;
<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="[594,699,1071,1097]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
(2015)
</emphasis>
</bibRefCitation>
suggested multiple setae on the proleg on A6 of the larvae as a synapomorphy of the group. Our results also support a close relationship of
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[498,648,1151,1177]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
,
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[662,768,1151,1177]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Azelinini">Azelinini</taxonomicName>
and
<taxonomicName authorityName="Tutt" authorityYear="1896" box="[829,992,1151,1177]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Odontoperini">Odontoperini</taxonomicName>
. These clades will be treated in more detail by G.
<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="[534,675,1191,1217]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="29" phylum="Arthropoda" rank="tribe" status="new tribe" tribe="Brehm">Brehmetal</taxonomicName>
. (2019, unpublisheddata).
</paragraph>
<paragraph blockId="26.[498,1542,234,1934]" pageId="26" pageNumber="27">
Following the ideas of
<bibRefCitation author="Pitkin L." box="[787,933,1230,1257]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="26" pageNumber="27" pagination="121 - 401" part="135" refId="ref19983" refString="Pitkin L. 2002. Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society 135 (2 - 3): 121 - 401 DOI 10.1046 / j. 1096 - 3642.2002.01200. x." title="Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae)" type="journal article" year="2002">
<emphasis box="[787,933,1230,1257]" italics="true" pageId="26" pageNumber="27">Pitkin (2002)</emphasis>
</bibRefCitation>
,
<bibRefCitation author="Beljaev EA" box="[947,1105,1230,1257]" journalOrPublisher="Entomological Review" pageId="26" pageNumber="27" 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="[947,1105,1230,1257]" italics="true" pageId="26" pageNumber="27">Beljaev (2008)</emphasis>
</bibRefCitation>
synonymized the tribes
<taxonomicName authorityName="Bruand" authorityYear="1846" box="[1382,1541,1230,1256]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Ourapterygini">Ourapterygini</taxonomicName>
and
<taxonomicName authorityName="Warren" authorityYear="1894" box="[549,678,1270,1296]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Nephodiini">Nephodiini</taxonomicName>
with
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[746,863,1270,1296]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
. He considered the divided vinculum in male genitalia and the attachment of muscles
<emphasis box="[807,829,1311,1336]" italics="true" pageId="26" pageNumber="27">m</emphasis>
3 as apomorphies of the
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[1114,1230,1310,1336]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
, but did not provide a phylogenetic analysis.
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[752,997,1350,1376]" journalOrPublisher="PLOS ONE" pageId="26" pageNumber="27" 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="[752,997,1350,1376]" italics="true" pageId="26" pageNumber="27">
<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="[752,854,1350,1376]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al. (2011)
</emphasis>
</bibRefCitation>
supported Beljaev s assumptions and recovered
<taxonomicName authority="Treitschke, 1825" authorityName="Treitschke" authorityYear="1825" box="[498,798,1390,1417]" class="Insecta" family="Geometridae" genus="Ennomos" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[498,603,1391,1416]" italics="true" pageId="26" pageNumber="27">Ennomos</emphasis>
Treitschke, 1825
</taxonomicName>
(
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[814,933,1390,1417]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
),
<taxonomicName authority="Leach, 1814" authorityName="Leach" authorityYear="1814" box="[954,1226,1390,1416]" class="Insecta" family="Geometridae" genus="Ourapteryx" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[954,1082,1390,1416]" italics="true" pageId="26" pageNumber="27">Ourapteryx</emphasis>
Leach, 1814
</taxonomicName>
(
<taxonomicName authorityName="Bruand" authorityYear="1846" box="[1242,1405,1390,1417]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Ourapterygini">Ourapterygini</taxonomicName>
) and
<taxonomicName authority="Hubner, 1823" authorityName="Hubner" authorityYear="1823" box="[498,772,1430,1456]" class="Insecta" family="Geometridae" genus="Nephodia" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[498,606,1430,1456]" italics="true" pageId="26" pageNumber="27">Nephodia</emphasis>
Hübner, 1823
</taxonomicName>
(
<taxonomicName authorityName="Warren" authorityYear="1894" box="[788,921,1430,1457]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Nephodiini">Nephodiini</taxonomicName>
) as belonging to the same clade. Our comprehensive analysis confirms those previous findings and we agree with
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[1193,1307,1470,1496]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
as the valid tribal name for this large clade. This clade will be treated in more detail by G.
<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="[1330,1407,1509,1535]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="29" phylum="Arthropoda" rank="tribe" status="new tribe" tribe="Brehm">Brehm</taxonomicName>
et al.
</paragraph>
<paragraph blockId="26.[498,1542,234,1934]" box="[498,789,1549,1576]" pageId="26" pageNumber="27">(2019, unpublisheddata).</paragraph>
<paragraph blockId="26.[498,1542,234,1934]" pageId="26" pageNumber="27">
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[530,659,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Campaeini">Campaeini</taxonomicName>
,
<taxonomicName authorityName="Herbulot" authorityYear="1962" box="[672,803,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Alsophilini">Alsophilini</taxonomicName>
, Wilemaniini and
<taxonomicName authorityName="Warren" authorityYear="1894" box="[1024,1204,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Prosopolophini">Prosopolophini</taxonomicName>
grouped together in a well-supported clade (SH-like = 100, UFBoot2 = 99). Previous molecular analyses have shown an association of
<taxonomicName authorityName="Wehrli" authorityYear="1940" box="[789,899,1669,1695]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Colotoini">Colotoini</taxonomicName>
[=
<taxonomicName authorityName="Warren" authorityYear="1894" box="[944,1128,1668,1695]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Prosopolophini">Prosopolophini</taxonomicName>
] and Wilemaniini (
<bibRefCitation author="Yamamoto S &amp; Sota T." journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="26" pageNumber="27" pagination="711 - 723" part="44" refId="ref21575" refString="Yamamoto S, Sota T. 2007. Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 44 (2): 711 - 723 DOI 10.1016 / j. ympev. 2006.12.027." title="Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes" type="journal article" year="2007">
<emphasis italics="true" pageId="26" pageNumber="27">Yamamoto &amp; Sota, 2007</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[634,875,1709,1735]" journalOrPublisher="PLOS ONE" pageId="26" pageNumber="27" 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="[634,875,1709,1735]" italics="true" pageId="26" pageNumber="27">
<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="[634,739,1709,1735]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al., 2011
</emphasis>
</bibRefCitation>
), although no synapomorphies are known to support synonymization (
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[702,967,1748,1775]" journalOrPublisher="Stenstrup: Apollo Books" pageId="26" pageNumber="27" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[702,967,1748,1775]" italics="true" pageId="26" pageNumber="27">
Skou &amp;
<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="[794,902,1748,1774]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
). The Palaearctic genera
<taxonomicName authority="Blanchard, 1845" authorityName="Blanchard" authorityYear="1845" class="Insecta" family="Geometridae" genus="Compsoptera" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[1258,1409,1749,1775]" italics="true" pageId="26" pageNumber="27">Compsoptera</emphasis>
Blanchard, 1845
</taxonomicName>
,
<taxonomicName authority="Agassiz, 1847" authorityName="Agassiz" authorityYear="1847" box="[570,856,1788,1814]" class="Insecta" family="Geometridae" genus="Apochima" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[570,688,1788,1814]" italics="true" pageId="26" pageNumber="27">Apochima</emphasis>
Agassiz, 1847
</taxonomicName>
,
<taxonomicName authority="Prout, 1915" authorityName="Prout" authorityYear="1915" box="[869,1133,1788,1815]" class="Insecta" family="Geometridae" genus="Dasycorsa" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[869,985,1789,1814]" italics="true" pageId="26" pageNumber="27">Dasycorsa</emphasis>
Prout, 1915
</taxonomicName>
,
<taxonomicName authority="Anker, 1854" authorityName="Anker" authorityYear="1854" box="[1147,1461,1788,1815]" class="Insecta" family="Geometridae" genus="Chondrosoma" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[1147,1309,1788,1814]" italics="true" pageId="26" pageNumber="27">Chondrosoma</emphasis>
Anker, 1854
</taxonomicName>
and
<taxonomicName authority="Nupponen &amp; Sihvonen, 2013" authorityName="Nupponen &amp; Sihvonen" authorityYear="2013" box="[498,967,1828,1855]" class="Insecta" family="Geometridae" genus="Dorsispina" higherTaxonomySource="GBIF" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="genus">
<emphasis box="[498,622,1829,1854]" italics="true" pageId="26" pageNumber="27">Dorsispina</emphasis>
Nupponen &amp; Sihvonen, 2013
</taxonomicName>
, are potentially part of the same complex (
<bibRefCitation author="Skou P &amp; Sihvonen P." journalOrPublisher="Stenstrup: Apollo Books" pageId="26" pageNumber="27" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis italics="true" pageId="26" pageNumber="27">
Skou &amp;
<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="[498,606,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
,
<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="[689,798,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
pers. obs.), but they were not included in the current study.
</paragraph>
<paragraph blockId="26.[498,1542,234,1934]" lastBlockId="27.[498,1542,234,1935]" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[498,625,1908,1934]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="26" pageNumber="27" phylum="Arthropoda" rank="tribe" tribe="Campaeini">Campaeini</taxonomicName>
is a small group including four genera with Oriental, Palaearctic and Nearctic distribution, apparently closely related to
<taxonomicName authorityName="Herbulot" authorityYear="1962" box="[990,1119,234,260]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Alsophilini">Alsophilini</taxonomicName>
and
<taxonomicName authorityName="Warren" authorityYear="1894" box="[1180,1362,234,260]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Prosopolophini">Prosopolophini</taxonomicName>
, but currently accepted as a tribe (
<bibRefCitation accessDate="2018-08-03" author="Forum Herbulot" box="[723,977,274,300]" pageId="27" pageNumber="28" publicationUrl="http://www.herbulot.de/famgroup.htm" refId="ref18517" refString="Forum Herbulot. 2007. World list of family-group names in Geometridae. Available at http: // www. herbulot. de / famgroup. htm (accessed 3 August 2018)." title="World list of family-group names in Geometridae" type="url" year="2007">
<emphasis box="[723,977,274,300]" italics="true" pageId="27" pageNumber="28">Forum Herbulot, 2007</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[989,1252,274,301]" journalOrPublisher="Stenstrup: Apollo Books" pageId="27" pageNumber="28" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[989,1252,274,301]" italics="true" pageId="27" pageNumber="28">
Skou &amp;
<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="[1079,1187,274,300]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
). Our results support the close phylogenetic affinities among these tribes, but due to the limited number of sampled taxa, we do not propose any formal changes.
</paragraph>
<paragraph blockId="27.[498,1542,234,1935]" box="[530,1526,393,420]" pageId="27" pageNumber="28">
The genus
<taxonomicName authority="Walker, 1858" authorityName="Walker" authorityYear="1858" box="[658,912,393,420]" class="Insecta" family="Geometridae" genus="Declana" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[658,750,394,420]" italics="true" pageId="27" pageNumber="28">Declana</emphasis>
Walker, 1858
</taxonomicName>
is recovered as an isolated clade sister to
<taxonomicName authorityName="Janse" authorityYear="1933" box="[1396,1521,394,420]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
.
</paragraph>
<paragraph blockId="27.[498,1542,234,1935]" pageId="27" pageNumber="28">
This genus is endemic to New Zealand, but to date has not been assigned to tribe. According to our results,
<taxonomicName authorityName="Walker" authorityYear="1858" box="[667,759,473,499]" class="Insecta" family="Geometridae" genus="Declana" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[667,759,473,499]" italics="true" pageId="27" pageNumber="28">Declana</emphasis>
</taxonomicName>
could well be defined as its own tribe. However, the delimitation of this tribe is beyond the scope of our paper and more genera from Australia and New Zealand should first be examined. Aclose relationship between
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[1236,1381,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
and
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1442,1542,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Lithinini">Lithinini</taxonomicName>
was suggested by
<bibRefCitation author="Pitkin L." box="[703,852,593,619]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="27" pageNumber="28" pagination="121 - 401" part="135" refId="ref19983" refString="Pitkin L. 2002. Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society 135 (2 - 3): 121 - 401 DOI 10.1046 / j. 1096 - 3642.2002.01200. x." title="Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae)" type="journal article" year="2002">
<emphasis box="[703,852,593,619]" italics="true" pageId="27" pageNumber="28">Pitkin (2002)</emphasis>
</bibRefCitation>
, based on the similar pair of processes of the anellus in the male genitalia. Pitkin also noted a morphological similarity in the male genitalia (processes of the juxta) shared by
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[879,1022,673,699]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
and
<taxonomicName authorityName="Janse" authorityYear="1933" box="[1078,1201,673,699]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
. In a study of the Australasian fauna,
<bibRefCitation author="Young CJ" box="[575,728,712,738]" journalOrPublisher="Zootaxa" pageId="27" pageNumber="28" pagination="1 - 141" part="1736" refId="ref21693" refString="Young CJ. 2008. Characterization of the Australian Nacophorini using adult morphology, and phylogeny of the Geometridae based on morphological characters. Zootaxa 1736 (1): 1 - 141 DOI 10.11646 / zootaxa. 1736.1.1." title="Characterization of the Australian Nacophorini using adult morphology, and phylogeny of the Geometridae based on morphological characters" type="journal article" year="2008">
<emphasis box="[575,728,712,738]" italics="true" pageId="27" pageNumber="28">Young (2008)</emphasis>
</bibRefCitation>
suggested the synonymization of
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[1114,1257,712,738]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
and
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1316,1416,712,738]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Lithinini">Lithinini</taxonomicName>
. This was further corroborated by
<bibRefCitation author="Sihvonen P &amp; Staude HS &amp; Mutanen M." box="[769,1176,752,779]" journalOrPublisher="Systematic Entomology" pageId="27" pageNumber="28" pagination="606 - 627" part="40" refId="ref20789" refString="Sihvonen P, Staude HS, Mutanen M. 2015. Systematic position of the enigmatic African cycad moths: an integrative approach to a nearly century old problem (Lepidoptera: Geometridae, Diptychini). Systematic Entomology 40 (3): 606 - 627 DOI 10.1111 / syen. 12125." title="Systematic position of the enigmatic African cycad moths: an integrative approach to a nearly century old problem (Lepidoptera: Geometridae, Diptychini)" type="journal article" year="2015">
<emphasis box="[769,1176,752,779]" italics="true" pageId="27" pageNumber="28">
<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="[769,873,752,778]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, Staude &amp; Mutanen (2015)
</emphasis>
</bibRefCitation>
who found that
<taxonomicName authorityName="Janse" authorityYear="1933" box="[1362,1482,752,778]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
were nested within some
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[727,870,792,818]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
and
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[929,1029,792,818]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Lithinini">Lithinini</taxonomicName>
. However, none of the studies proposed formal taxonomic changes because of limited taxon sampling. In contrast, samples in our analyses cover all biogeographic regions and the results suggest that true
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[1329,1472,872,898]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
is a clade which comprises almost exclusively New World species. This clade is clearly separate from Old World “nacophorines” (cf.
<bibRefCitation author="Young CJ" box="[922,1061,952,978]" journalOrPublisher="Spixiana" pageId="27" pageNumber="28" pagination="199 - 200" part="26" refId="ref21625" refString="Young CJ. 2003. The place of the Australian Nacophorini in the Geometridae. Spixiana 26: 199 - 200." title="The place of the Australian Nacophorini in the Geometridae" type="journal article" year="2003">
<emphasis box="[922,1061,952,978]" italics="true" pageId="27" pageNumber="28">Young, 2003</emphasis>
</bibRefCitation>
) that are intermixed with
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[1364,1462,951,977]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Lithinini">Lithinini</taxonomicName>
and
<taxonomicName authorityName="Janse" authorityYear="1933" box="[498,621,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
. We here formally transfer Old World nacophorines to
<taxonomicName authorityName="Janse" authorityYear="1933" box="[1233,1353,991,1017]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
and synonymize
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[498,596,1031,1057]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" status="syn. nov." tribe="Lithinini">Lithinini</taxonomicName>
<emphasis bold="true" box="[604,710,1031,1057]" pageId="27" pageNumber="28">
<taxonomicNameLabel box="[604,710,1031,1057]" pageId="27" pageNumber="28">syn. nov.</taxonomicNameLabel>
</emphasis>
with
<taxonomicName authorityName="Janse" authorityYear="1933" box="[776,896,1031,1057]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Diptychini">Diptychini</taxonomicName>
(
<tableCitation box="[914,998,1031,1057]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="27" pageNumber="28">Table 2</tableCitation>
). Further formal taxonomic changes in the
<taxonomicName baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[498,641,1071,1097]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">Nacophorini</taxonomicName>
complex are provided by G.
<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="[975,1052,1071,1097]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="29" phylum="Arthropoda" rank="tribe" status="new tribe" tribe="Brehm">Brehm</taxonomicName>
et al. (2019, unpublished data).
</paragraph>
<paragraph blockId="27.[498,1542,234,1935]" pageId="27" pageNumber="28">
<taxonomicName authority="Hubner 1825" authorityName="Hubner" authorityYear="1825" box="[530,768,1111,1138]" class="Insecta" family="Geometridae" genus="Theria" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[530,607,1111,1137]" italics="true" pageId="27" pageNumber="28">Theria</emphasis>
Hübner 1825
</taxonomicName>
, the only representative of
<taxonomicName authorityName="Herbulot" authorityYear="1963" box="[1079,1174,1111,1137]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Theriini">Theriini</taxonomicName>
in this study, clustered together with
<taxonomicName authority="Hubner, 1825" authorityName="Hubner" authorityYear="1825" box="[559,874,1151,1178]" class="Insecta" family="Geometridae" genus="Lomographa" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[559,705,1151,1177]" italics="true" pageId="27" pageNumber="28">Lomographa</emphasis>
Hübner, 1825
</taxonomicName>
(
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[891,978,1151,1177]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Baptini">Baptini</taxonomicName>
in
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[1018,1289,1151,1177]" journalOrPublisher="Stenstrup: Apollo Books" pageId="27" pageNumber="28" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[1018,1289,1151,1177]" italics="true" pageId="27" pageNumber="28">
Skou &amp;
<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="[1114,1222,1151,1177]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
), in a well-supported clade, agreeing with the molecular results of
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[1026,1279,1191,1217]" journalOrPublisher="PLOS ONE" pageId="27" pageNumber="28" 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="[1026,1279,1191,1217]" italics="true" pageId="27" pageNumber="28">
<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="[1026,1131,1191,1217]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al. (2011)
</emphasis>
</bibRefCitation>
. The placement of
<taxonomicName authorityName="Hubner" authorityYear="1825" box="[498,644,1230,1256]" class="Insecta" family="Geometridae" genus="Lomographa" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[498,644,1230,1256]" italics="true" pageId="27" pageNumber="28">Lomographa</emphasis>
</taxonomicName>
in
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[685,785,1230,1256]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Caberini">Caberini</taxonomicName>
(
<bibRefCitation author="Rindge FH" box="[804,954,1230,1257]" journalOrPublisher="American Museum Novitates" pageId="27" pageNumber="28" pagination="1 - 18" part="2673" refId="ref20457" refString="Rindge FH. 1979. A revision of the North American moths of the genus Lomographa (Lepidoptera, Geometridae). American Museum Novitates 2673: 1 - 18." title="A revision of the North American moths of the genus Lomographa (Lepidoptera, Geometridae)" type="journal article" year="1979">
<emphasis box="[804,954,1230,1257]" italics="true" pageId="27" pageNumber="28">Rindge, 1979</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Pitkin L." box="[969,1109,1230,1257]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="27" pageNumber="28" pagination="121 - 401" part="135" refId="ref19983" refString="Pitkin L. 2002. Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society 135 (2 - 3): 121 - 401 DOI 10.1046 / j. 1096 - 3642.2002.01200. x." title="Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae)" type="journal article" year="2002">
<emphasis box="[969,1109,1230,1257]" italics="true" pageId="27" pageNumber="28">Pitkin, 2002</emphasis>
</bibRefCitation>
) is not supported by our study nor by that of
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[622,875,1270,1296]" journalOrPublisher="PLOS ONE" pageId="27" pageNumber="28" 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="[622,875,1270,1296]" italics="true" pageId="27" pageNumber="28">
<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="[622,727,1270,1296]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al. (2011)
</emphasis>
</bibRefCitation>
. The monophyly of
<taxonomicName authorityName="Hubner" authorityYear="1825" box="[1119,1265,1270,1296]" class="Insecta" family="Geometridae" genus="Lomographa" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1119,1265,1270,1296]" italics="true" pageId="27" pageNumber="28">Lomographa</emphasis>
</taxonomicName>
has not been tested before, but we show that one Neotropical and one Palaearctic
<taxonomicName box="[1222,1456,1310,1336]" class="Insecta" family="Geometridae" genus="Lomographa" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis box="[1222,1368,1310,1336]" italics="true" pageId="27" pageNumber="28">Lomographa</emphasis>
species
</taxonomicName>
indeed group together. Our results show that
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[952,1052,1350,1376]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Caberini">Caberini</taxonomicName>
are not closely related to the
<taxonomicName authorityName="Herbulot" authorityYear="1963" box="[1405,1500,1350,1376]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Theriini">Theriini</taxonomicName>
+
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[498,583,1390,1416]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Baptini">Baptini</taxonomicName>
clade, unlike in earlier morphology-based hypotheses (
<bibRefCitation author="Rindge FH" box="[1226,1373,1390,1416]" journalOrPublisher="American Museum Novitates" pageId="27" pageNumber="28" pagination="1 - 18" part="2673" refId="ref20457" refString="Rindge FH. 1979. A revision of the North American moths of the genus Lomographa (Lepidoptera, Geometridae). American Museum Novitates 2673: 1 - 18." title="A revision of the North American moths of the genus Lomographa (Lepidoptera, Geometridae)" type="journal article" year="1979">
<emphasis box="[1226,1373,1390,1416]" italics="true" pageId="27" pageNumber="28">Rindge, 1979</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Pitkin L." box="[1387,1526,1390,1416]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="27" pageNumber="28" pagination="121 - 401" part="135" refId="ref19983" refString="Pitkin L. 2002. Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society 135 (2 - 3): 121 - 401 DOI 10.1046 / j. 1096 - 3642.2002.01200. x." title="Neotropical Ennomine moths: a review of the genera (Lepidoptera: Geometridae)" type="journal article" year="2002">
<emphasis box="[1387,1526,1390,1416]" italics="true" pageId="27" pageNumber="28">Pitkin, 2002</emphasis>
</bibRefCitation>
). Morphologically,
<taxonomicName authorityName="Herbulot" authorityYear="1963" box="[704,799,1430,1456]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Theriini">Theriini</taxonomicName>
and
<taxonomicName authorityName="Forbes" authorityYear="1948" box="[857,942,1430,1456]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Baptini">Baptini</taxonomicName>
are dissimilar, therefore we recognize them as valid tribes (see description and illustrations in
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[996,1267,1470,1496]" journalOrPublisher="Stenstrup: Apollo Books" pageId="27" pageNumber="28" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[996,1267,1470,1496]" italics="true" pageId="27" pageNumber="28">
Skou &amp;
<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="[1092,1200,1470,1496]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
, 2015
</emphasis>
</bibRefCitation>
).
</paragraph>
<paragraph blockId="27.[498,1542,234,1935]" pageId="27" pageNumber="28">
According to our results, 11 molecular markers were not enough to infer phylogenetic affinities of Plutodini (represented by one species of
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1119,1216,1549,1575]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1119,1216,1549,1575]" italics="true" pageId="27" pageNumber="28">Plutodes</emphasis>
</taxonomicName>
). Similar results were found by
<bibRefCitation author="Sihvonen P &amp; Mutanen M &amp; Kaila L &amp; Brehm G &amp; Hausmann A &amp; Staude HS" box="[613,867,1589,1615]" journalOrPublisher="PLOS ONE" pageId="27" pageNumber="28" 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="[613,867,1589,1615]" italics="true" pageId="27" pageNumber="28">
<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="[613,718,1589,1615]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
et al. (2011)
</emphasis>
</bibRefCitation>
, who in some analyses recovered
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1269,1366,1589,1615]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1269,1366,1589,1615]" italics="true" pageId="27" pageNumber="28">Plutodes</emphasis>
</taxonomicName>
as sister of
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[498,597,1629,1655]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[498,597,1629,1655]" italics="true" pageId="27" pageNumber="28">Eumelea</emphasis>
</taxonomicName>
. Our analyses are congruent with those findings. IQ-TREE results suggest that
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[553,650,1669,1695]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[553,650,1669,1695]" italics="true" pageId="27" pageNumber="28">Plutodes</emphasis>
</taxonomicName>
is sister to
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[787,898,1669,1695]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Palyadini">Palyadini</taxonomicName>
, but RAxML analyses recovered
<taxonomicName authorityName="Duncan &amp; Westwood" authorityYear="1841" box="[1287,1386,1669,1695]" class="Insecta" family="Geometridae" genus="Eumelea" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1287,1386,1669,1695]" italics="true" pageId="27" pageNumber="28">Eumelea</emphasis>
</taxonomicName>
as the most probable sister of
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[777,874,1709,1735]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[777,874,1709,1735]" italics="true" pageId="27" pageNumber="28">Plutodes</emphasis>
</taxonomicName>
. Given that our analyses are not in agreement on the sister-group affinities of
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[782,879,1748,1774]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[782,879,1748,1774]" italics="true" pageId="27" pageNumber="28">Plutodes</emphasis>
</taxonomicName>
, we do not make any assumptions about its phylogenetic position. Instead, we emphasize that further work needs to be done to clarify the phylogenetic positions of
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[801,898,1828,1854]" class="Insecta" family="Geometridae" genus="Plutodes" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[801,898,1828,1854]" italics="true" pageId="27" pageNumber="28">Plutodes</emphasis>
</taxonomicName>
and related groups.
</paragraph>
<paragraph blockId="27.[498,1542,234,1935]" lastBlockId="28.[498,1542,234,819]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[530,690,1868,1894]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Hypochrosini">Hypochrosini</taxonomicName>
is only recovered in a well-defined lineage if the genera
<taxonomicName authority="Gistl, 1848" authorityName="Gistl" authorityYear="1848" box="[1334,1542,1868,1895]" class="Insecta" family="Geometridae" genus="Apeira" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1334,1412,1869,1895]" italics="true" pageId="27" pageNumber="28">Apeira</emphasis>
Gistl, 1848
</taxonomicName>
(
<taxonomicName authorityName="Stekolnikov &amp; Kuznetzov" authorityYear="1982" box="[506,609,1908,1935]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Apeirini">Apeirini</taxonomicName>
),
<taxonomicName authority="Duponchel, 1829" authorityName="Duponchel" authorityYear="1829" box="[630,918,1908,1934]" class="Insecta" family="Geometridae" genus="Epione" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[630,709,1909,1934]" italics="true" pageId="27" pageNumber="28">Epione</emphasis>
Duponchel, 1829
</taxonomicName>
(
<taxonomicName authorityName="Bruand" authorityYear="1846" box="[935,1041,1908,1935]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Epionini">Epionini</taxonomicName>
),
<taxonomicName authorityName="Warren" authorityYear="1895" box="[1063,1189,1908,1934]" class="Insecta" family="Geometridae" genus="Sericosema" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1063,1189,1908,1934]" italics="true" pageId="27" pageNumber="28">Sericosema</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Duponchel" authorityYear="1845" box="[1206,1312,1908,1935]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Caberini">Caberini</taxonomicName>
),
<taxonomicName authorityName="Hubner" authorityYear="1825" box="[1333,1411,1908,1934]" class="Insecta" family="Geometridae" genus="Ithysia" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="genus">
<emphasis box="[1333,1411,1908,1934]" italics="true" pageId="27" pageNumber="28">Ithysia</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Herbulot" authorityYear="1963" box="[1428,1529,1908,1935]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="27" pageNumber="28" phylum="Arthropoda" rank="tribe" tribe="Theriini">Theriini</taxonomicName>
),
<taxonomicName authority="Walker, 1866" authorityName="Walker" authorityYear="1866" box="[498,749,234,261]" class="Insecta" family="Geometridae" genus="Capasa" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[498,583,235,261]" italics="true" pageId="28" pageNumber="29">Capasa</emphasis>
Walker, 1866
</taxonomicName>
(unassigned) and
<taxonomicName authority="Warren, 1894" authorityName="Warren" authorityYear="1894" box="[968,1251,234,261]" class="Insecta" family="Geometridae" genus="Omizodes" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[968,1081,234,260]" italics="true" pageId="28" pageNumber="29">Omizodes</emphasis>
Warren, 1894
</taxonomicName>
(unassigned) were transferred to
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[663,826,274,300]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Hypochrosini">Hypochrosini</taxonomicName>
.
<bibRefCitation author="Skou P &amp; Sihvonen P." box="[836,1114,274,301]" journalOrPublisher="Stenstrup: Apollo Books" pageId="28" pageNumber="29" refId="ref20843" refString="Skou P, Sihvonen P. 2015. The Geometrid moths of Europe. Vol. 5: Ennominae I. Stenstrup: Apollo Books." title="The Geometrid moths of Europe. Vol. 5: Ennominae I" type="book" year="2015">
<emphasis box="[836,1114,274,301]" italics="true" pageId="28" pageNumber="29">
Skou &amp;
<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="[928,1033,274,300]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="22" phylum="Arthropoda" rank="subFamily" status="new subfamily" subFamily="Sihvonen">Sihvonen</taxonomicName>
(2015)
</emphasis>
</bibRefCitation>
already suggested aclose association of
<taxonomicName authorityName="Bruand" authorityYear="1846" box="[530,633,314,340]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Epionini">Epionini</taxonomicName>
,
<taxonomicName authorityName="Stekolnikov &amp; Kuznetzov" authorityYear="1982" box="[646,743,314,340]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Apeirini">Apeirini</taxonomicName>
and
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[804,967,314,340]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Hypochrosini">Hypochrosini</taxonomicName>
. We think that synonymizing these tribes is desirable. However, due to the limited number of sampled taxa we do not propose any formal changes until more data becomes available. We do suggest, however, formal taxonomic changes for the genera
<taxonomicName authorityName="Walker" authorityYear="1866" box="[893,978,434,460]" class="Insecta" family="Geometridae" genus="Capasa" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[893,978,434,460]" italics="true" pageId="28" pageNumber="29">Capasa</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Warren" authorityYear="1894" box="[1034,1147,433,459]" class="Insecta" family="Geometridae" genus="Omizodes" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[1034,1147,433,459]" italics="true" pageId="28" pageNumber="29">Omizodes</emphasis>
</taxonomicName>
from unassigned to
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1382,1542,433,459]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Hypochrosini">Hypochrosini</taxonomicName>
(
<tableCitation box="[508,596,473,499]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionText="Table 2 Summary of formally proposed taxonomic changes." pageId="28" pageNumber="29">Table 2</tableCitation>
).
</paragraph>
<paragraph blockId="28.[498,1542,234,819]" pageId="28" pageNumber="29">
The southern African genus
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[861,1019,514,539]" class="Insecta" family="Geometridae" genus="Drepanogynis" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[861,1019,514,539]" italics="true" pageId="28" pageNumber="29">Drepanogynis</emphasis>
</taxonomicName>
is paraphyletic and has earlier been classified as belonging in
<taxonomicName baseAuthorityName="sensu Beljaev" baseAuthorityYear="2008" box="[680,799,553,579]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Ennomini">Ennomini</taxonomicName>
, and later in
<taxonomicName authority="(Kruger, 2002)" baseAuthorityName="Kruger" baseAuthorityYear="2002" box="[951,1274,553,580]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Nacophorini">
Nacophorini (
<bibRefCitation author="Kruger M." bookContentInfo="442 figs." box="[1116,1263,553,579]" journalOrPublisher="Transvaal Museum Monograph" pageId="28" pageNumber="29" pagination="1 - 220" part="13" refId="ref19244" refString="Kruger M. 2002. Revision of Afrotropical Ennominae of the Drepanogynis group IV: the genus Drepanogynis Guenee (Lepidoptera: Geometridae). Transvaal Museum Monograph 13: 1 - 220 incl. 442 figs." title="Revision of Afrotropical Ennominae of the Drepanogynis group IV: the genus Drepanogynis Guenee (Lepidoptera: Geometridae)" type="book chapter" year="2002">
<emphasis box="[1116,1263,553,579]" italics="true" pageId="28" pageNumber="29">Krüger, 2002</emphasis>
</bibRefCitation>
)
</taxonomicName>
. In our phylogeny, it is intermixed with the genera
<taxonomicName authority="Warren, 1899" authorityName="Warren" authorityYear="1899" box="[826,1157,593,619]" class="Insecta" family="Geometridae" genus="Sphingomima" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[826,985,593,619]" italics="true" pageId="28" pageNumber="29">Sphingomima</emphasis>
Warren, 1899
</taxonomicName>
, and
<taxonomicName authority="Walker, 1855" authorityName=", Walker" authorityYear="1855" box="[1223,1493,593,620]" class="Insecta" family="Geometridae" genus="Thenopa" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[1223,1324,593,619]" italics="true" pageId="28" pageNumber="29">Thenopa</emphasis>
Walker, 1855
</taxonomicName>
.
</paragraph>
<subSection pageId="28" pageNumber="29" type="multiple">
<paragraph blockId="28.[498,1542,234,819]" pageId="28" pageNumber="29">
<taxonomicName authority="Prout, 1917" authorityName="Prout" authorityYear="1917" box="[498,913,633,659]" class="Insecta" family="Geometridae" genus="Hebdomophruda" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="errans">
<emphasis box="[498,772,633,659]" italics="true" pageId="28" pageNumber="29">Hebdomophruda errans</emphasis>
Prout, 1917
</taxonomicName>
also clusters together with these taxa, apart from other
<taxonomicName box="[498,956,673,699]" class="Insecta" family="Geometridae" genus="Hebdomophruda" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis box="[498,694,673,699]" italics="true" pageId="28" pageNumber="29">Hebdomophruda</emphasis>
Warren, 1897 species
</taxonomicName>
, which suggests that this genus is polyphyletic.
</paragraph>
</subSection>
<paragraph blockId="28.[498,1542,234,819]" pageId="28" pageNumber="29">
These genera form a clade sister to the lineage that comprises several
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[1289,1449,712,738]" class="Insecta" family="Geometridae" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="tribe" tribe="Hypochrosini">Hypochrosini</taxonomicName>
species. Considering that our analysis strongly supports this clade, we place
<taxonomicName authorityName=", Walker" authorityYear="1855" box="[1301,1402,752,778]" class="Insecta" family="Geometridae" genus="Thenopa" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[1301,1402,752,778]" italics="true" pageId="28" pageNumber="29">Thenopa</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Warren" authorityYear="1899" box="[498,657,792,818]" class="Insecta" family="Geometridae" genus="Sphingomima" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[498,657,792,818]" italics="true" pageId="28" pageNumber="29">Sphingomima</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Guenee" authorityYear="1858" box="[719,877,793,818]" class="Insecta" family="Geometridae" genus="Drepanogynis" kingdom="Animalia" order="Lepidoptera" pageId="28" pageNumber="29" phylum="Arthropoda" rank="genus">
<emphasis box="[719,877,793,818]" italics="true" pageId="28" pageNumber="29">Drepanogynis</emphasis>
</taxonomicName>
in a tribe of their own.
</paragraph>
</subSubSection>
</treatment>
</document>