202 lines
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202 lines
20 KiB
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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.861.34686" ID-GBIF-Dataset="afdd84b2-a5c5-471f-bf87-0747b9f75a39" ID-PMC="PMC6629708" ID-Pensoft-Pub="1313-2970-861-91" ID-PubMed="31333327" ID-ZooBank="802AE175A8104FFCB3FDE289D0CC4191" ModsDocID="1313-2970-861-91" checkinTime="1565091111200" checkinUser="pensoft" docAuthor="Zhang, Jing, Cong, Qian, Shen, Jinhui, Brockmann, Ernst & Grishin, Nick V." docDate="2019" docId="B0EC79CA4EDB591A89B311E5A0A66003" docLanguage="en" docName="ZooKeys 861: 91-105" docOrigin="ZooKeys 861" docSource="http://dx.doi.org/10.3897/zookeys.861.34686" docTitle="Katreinae Grishin, subfam. n." docType="treatment" docUuid="EFD73E63-A0FE-4AB3-B6F2-318977EF7F83" docUuidSource="ZooBank" docVersion="6" lastPageNumber="96" masterDocId="FFFBBB3BFF8CFFA9FFA6FFE1E112FFC6" masterDocTitle="Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae)" masterLastPageNumber="105" masterPageNumber="91" pageNumber="94" updateTime="1668167525244" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Three new subfamilies of skipper butterflies (Lepidoptera, Hesperiidae)</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Zhang, Jing</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Cong, Qian</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Shen, Jinhui</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Brockmann, Ernst</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Grishin, Nick V.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2019</mods:date>
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<mods:detail type="volume">
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<mods:number>861</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>91</mods:start>
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<mods:end>105</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.861.34686</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.861.34686</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-861-91</mods:identifier>
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<mods:identifier type="ZooBank">802AE175A8104FFCB3FDE289D0CC4191</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="159016167" LSID="urn:lsid:zoobank.org:act:EFD73E63-A0FE-4AB3-B6F2-318977EF7F83" httpUri="http://treatment.plazi.org/id/B0EC79CA4EDB591A89B311E5A0A66003" lastPageId="5" lastPageNumber="96" pageId="3" pageNumber="94">
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<subSubSection pageId="3" pageNumber="94" type="nomenclature">
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<paragraph pageId="3" pageNumber="94">
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<taxonomicName LSID="b0ec79ca-4edb-591a-89b3-11e5a0a66003" authority="Grishin" family="Hesperiidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="" order="Lepidoptera" pageId="3" pageNumber="94" rank="subFamily" subFamily="Katreinae">Katreinae Grishin</taxonomicName>
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<taxonomicNameLabel pageId="3" pageNumber="94">subfam. n.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="94" type="type genus">
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<paragraph pageId="3" pageNumber="94">Type genus.</paragraph>
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<paragraph pageId="3" pageNumber="94">
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Katreus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Katreus" order="Lepidoptera" pageId="3" pageNumber="94" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="3" pageNumber="94">Katreus</emphasis>
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</taxonomicName>
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Watson, 1893.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="5" lastPageNumber="96" pageId="4" pageNumber="95" type="diagnosis">
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<paragraph pageId="4" pageNumber="95">
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<pageBreakToken pageId="4" pageNumber="95" start="start">Diagnosis</pageBreakToken>
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.
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</paragraph>
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<paragraph lastPageId="5" lastPageNumber="96" pageId="4" pageNumber="95">
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In appearance, most similar to
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Celaenorrhinus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Celaenorrhinus" order="Lepidoptera" pageId="4" pageNumber="95" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="4" pageNumber="95">Celaenorrhinus</emphasis>
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</taxonomicName>
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<normalizedToken originalValue="Hübner">Huebner</normalizedToken>
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, [1819] and its relatives (
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<bibRefCitation author="Evans, WH" journalOrPublisher="British Museum (Natural History), London" pageId="11" pageNumber="102" refId="B8" refString="Evans, WH, 1937. A Catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum. . British Museum (Natural History), London" title="A Catalogue of the African Hesperiidae indicating the classification and nomenclature adopted in the British Museum." year="1937">Evans 1937</bibRefCitation>
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), and was placed in
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<taxonomicName lsidName="" pageId="4" pageNumber="95" rank="tribe" tribe="Celaenorrhinini">Celaenorrhinini</taxonomicName>
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Swinhoe, 1912 by
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<bibRefCitation DOI="https://doi.org/10.1111/j.1096-0031.2008.00218.x" author="Warren, AD" journalOrPublisher="Cladistics" pageId="12" pageNumber="103" pagination="642 - 676" refId="B26" refString="Warren, AD, Ogawa, JR, Brower, AVZ, 2008. Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea). . Cladistics 24: 642 - 676" title="Phylogenetic relationships of subfamilies and circumscription of tribes in the family Hesperiidae (Lepidoptera: Hesperioidea)." url="https://doi.org/10.1111/j.1096-0031.2008.00218.x" volume="24" year="2008">Warren et al. (2008</bibRefCitation>
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,
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<bibRefCitation DOI="https://doi.org/10.1111/j.1365-3113.2008.00463.x" author="Warren, AD" journalOrPublisher="Systematic Entomology" pageId="12" pageNumber="103" pagination="467 - 523" refId="B27" refString="Warren, AD, Ogawa, JR, Brower, AVZ, 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. . Systematic Entomology 34: 467 - 523" title="Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data." url="https://doi.org/10.1111/j.1365-3113.2008.00463.x" volume="34" year="2009">2009</bibRefCitation>
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) but differs by longer apiculus of antennae and hindwing produced at vein 1A+2A. Morphologically, distinguished from all
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<taxonomicName family="Hesperiidae" lsidName="" pageId="4" pageNumber="95" rank="family">Hesperiidae</taxonomicName>
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by the
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<pageBreakToken pageId="5" pageNumber="96" start="start">combination</pageBreakToken>
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of the following characters. Abdomen short, shorter than inner margin of hindwing. Antennal club arcuate, bent in the middle, apiculus long, pointed. Second segment of palpi protrudes partly forward and partly upward (at an angle between the axis of the body and the axis perpendicular to it, =sub-erect). Males with hair pencil on hind tibiae, without stigmas or brands on wings. Forewing discal cell long, about 2/3 of the costa; vein M2 originates about midway between or closer to M1 than to M3 and vein CuA2 originates closer to the base of wing than to the end of discal cell. Hindwing produced at vein 1A+2A, vein 3A much shorter than vein CuA2. Male genitalia with a well-developed gnathos, which is not smaller than uncus, uncus bulging dorsad in lateral view, with small or tiny arms distant from each other, tegumen robust, extends caudad for the length of uncus, harpe longer than sacculus. See
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<bibRefCitation pageId="5" pageNumber="96" refId="B17">Larsen (2005</bibRefCitation>
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: 469-471) for illustrations of all representative species in this subfamily. In DNA, a combination of the following base pairs in the nuclear genome is diagnostic: aly528.10.2:G940C, aly925.27.5:A3610T, aly84.77.5:T1651G, aly595.14.2:G184C, aly2284.22.2:G967C, and in COI barcode region: C235T, A335T, C347T, and T349A.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="5" pageNumber="96" type="genera included">
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<paragraph pageId="5" pageNumber="96">Genera included.</paragraph>
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<paragraph pageId="5" pageNumber="96">
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Katreus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Katreus" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Katreus</emphasis>
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</taxonomicName>
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with its invalid synonym
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<taxonomicName class="Insecta" family="Tortricidae" genus="Choristoneura" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Choristoneura" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Choristoneura</emphasis>
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</taxonomicName>
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Mabille, 1889 (junior homonym of
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<taxonomicName class="Insecta" family="Tortricidae" genus="Choristoneura" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Choristoneura" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Choristoneura</emphasis>
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</taxonomicName>
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Lederer 1859 in
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<taxonomicName lsidName="" order="Lepidoptera" pageId="5" pageNumber="96" rank="order">Lepidoptera</taxonomicName>
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:
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<taxonomicName family="Tortricidae" lsidName="" pageId="5" pageNumber="96" rank="family">Tortricidae</taxonomicName>
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) and subjective synonyms
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Loxolexis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Loxolexis" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Loxolexis</emphasis>
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</taxonomicName>
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Karsch, 1895 and
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Daratus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Daratus" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Daratus</emphasis>
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</taxonomicName>
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Lindsey, 1925 (replacement name for
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<taxonomicName class="Insecta" family="Tortricidae" genus="Choristoneura" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Choristoneura" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Choristoneura</emphasis>
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</taxonomicName>
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) (
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Sequenced specimens from the new Hesperiidae subfamilies. DNA sample numbers are given for each specimen, additional data are in the Suppl. material 1: Table S 1 a Ortholexis holocausta syntype, NVG- 18053 C 02 b Ortholexis hollandi, NVG- 18082 A 08 c Ortholexis melichroptera, holotype of Acallopistes dimidia Holland, 1896; NVG- 18053 C 05 d Ortholexis melichroptera, holotype, NVG- 18053 A 06 e Katreus johnstonii, NVG- 18053 B 05 f Chamunda chamunda, NVG- 18086 E 02 g Barca bicolor, NVG- 17069 C 10 h Apostictopterus fuliginosus, NVG- 17069 C 12." figureDoi="10.3897/zookeys.861.34686.figure1" httpUri="https://binary.pensoft.net/fig/315261" pageId="5" pageNumber="96">Fig. 1e</figureCitation>
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); and
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Ortholexis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ortholexis" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Ortholexis</emphasis>
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</taxonomicName>
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Karsch, 1895 with its subjective synonym
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Acallopistes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Acallopistes" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Acallopistes</emphasis>
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</taxonomicName>
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Holland, 1896 (
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<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Sequenced specimens from the new Hesperiidae subfamilies. DNA sample numbers are given for each specimen, additional data are in the Suppl. material 1: Table S 1 a Ortholexis holocausta syntype, NVG- 18053 C 02 b Ortholexis hollandi, NVG- 18082 A 08 c Ortholexis melichroptera, holotype of Acallopistes dimidia Holland, 1896; NVG- 18053 C 05 d Ortholexis melichroptera, holotype, NVG- 18053 A 06 e Katreus johnstonii, NVG- 18053 B 05 f Chamunda chamunda, NVG- 18086 E 02 g Barca bicolor, NVG- 17069 C 10 h Apostictopterus fuliginosus, NVG- 17069 C 12." figureDoi="10.3897/zookeys.861.34686.figure1" httpUri="https://binary.pensoft.net/fig/315261" pageId="5" pageNumber="96">Fig. 1a-d</figureCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection pageId="5" pageNumber="96" type="comments">
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<paragraph pageId="5" pageNumber="96">Comments.</paragraph>
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<paragraph pageId="5" pageNumber="96">
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Taxonomy of these skippers has been confusing until it was resolved by
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<bibRefCitation DOI="https://doi.org/10.11646/zootaxa.3033.1.1" author="Cock, MJ" journalOrPublisher="Zootaxa" pageId="10" pageNumber="101" pagination="1 - 67" refId="B3" refString="Cock, MJ, Congdon, TC, 2011. Observations on the biology of Afrotropical Hesperiidae (Lepidoptera) principally from Kenya. Part 3. Pyrginae: Celaenorrhinini. . Zootaxa 3033: 1 - 67" title="Observations on the biology of Afrotropical Hesperiidae (Lepidoptera) principally from Kenya. Part 3. Pyrginae: Celaenorrhinini." url="https://doi.org/10.11646/zootaxa.3033.1.1" volume="3033" year="2011">Cock and Congdon (2011)</bibRefCitation>
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. For the most part, they were all placed in the genus
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Katreus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Katreus" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Katreus</emphasis>
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</taxonomicName>
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, until Larsen emphasized the differences in genitalia of those species placed in
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Ortholexis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ortholexis" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Ortholexis</emphasis>
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</taxonomicName>
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from true
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Katreus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Katreus" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Katreus</emphasis>
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</taxonomicName>
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(
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<bibRefCitation pageId="5" pageNumber="96" refId="B17">Larsen 2005</bibRefCitation>
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). Indeed, the two genera are quite distinct in our genomic analysis. A recent study based on several genes placed this group (only
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Ortholexis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ortholexis holocausta" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="species" species="holocausta">
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<emphasis italics="true" pageId="5" pageNumber="96">Ortholexis holocausta</emphasis>
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</taxonomicName>
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(Mabille, 1891) was included in that study) as a sister of
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<taxonomicName lsidName="" pageId="5" pageNumber="96" rank="tribe" tribe="Pyrrhopygini">Pyrrhopygini</taxonomicName>
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Mabille, 1877 (
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<bibRefCitation DOI="https://doi.org/10.1186/s12862-017-1016-x" author="Sahoo, RK" journalOrPublisher="Annales de la Societe entomologique de Belgique" pageId="12" pageNumber="103" refId="B21" refString="Sahoo, RK, Warren, AD, Collins, SC, Kodandaramaiah, U, 2017. . https://doi.org/10.1186/s12862-017-1016-x" url="https://doi.org/10.1186/s12862-017-1016-x" year="2017">Sahoo et al. 2017</bibRefCitation>
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), probably due to an insufficient number of genes included. In their study,
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<taxonomicName class="Insecta" family="Hesperiidae" genus="Euschemon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Euschemon" order="Lepidoptera" pageId="5" pageNumber="96" phylum="Arthropoda" rank="genus">
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<emphasis italics="true" pageId="5" pageNumber="96">Euschemon</emphasis>
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</taxonomicName>
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Doubleday, 1846 grouped with
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<taxonomicName lsidName="" pageId="5" pageNumber="96" rank="subFamily" subFamily="Eudaminae">Eudaminae</taxonomicName>
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instead of being sister to all other
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<taxonomicName family="Hesperiidae" lsidName="" pageId="5" pageNumber="96" rank="family">Hesperiidae</taxonomicName>
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with exclusion of
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<taxonomicName lsidName="" pageId="5" pageNumber="96" rank="subFamily" subFamily="Coeliadinae">Coeliadinae</taxonomicName>
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Evans, 1937 (
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<bibRefCitation DOI="https://doi.org/10.1111/j.1365-3113.2008.00463.x" author="Warren, AD" journalOrPublisher="Systematic Entomology" pageId="12" pageNumber="103" pagination="467 - 523" refId="B27" refString="Warren, AD, Ogawa, JR, Brower, AVZ, 2009. Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data. . Systematic Entomology 34: 467 - 523" title="Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data." url="https://doi.org/10.1111/j.1365-3113.2008.00463.x" volume="34" year="2009">Warren et al. 2009</bibRefCitation>
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;
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<bibRefCitation DOI="https://doi.org/10.1080/23802359.2017.1292478" author="Zhang, J" journalOrPublisher="Mitochondrial DNA Part B: Resources" pageId="12" pageNumber="103" pagination="136 - 138" refId="B29" refString="Zhang, J, Cong, Q, Shen, J, Fan, XL, Wang, M, Grishin, NV, 2017. The complete mitogenome of Euschemon rafflesia (Lepidoptera: Hesperiidae). . Mitochondrial DNA Part B: Resources 2: 136 - 138" title="The complete mitogenome of Euschemon rafflesia (Lepidoptera: Hesperiidae)." url="https://doi.org/10.1080/23802359.2017.1292478" volume="2" year="2017">Zhang et al. 2017</bibRefCitation>
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;
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<bibRefCitation DOI="https://doi.org/10.1186/s12862-018-1216-z" author="Toussaint, EFA" journalOrPublisher="Current Genomics" pageId="12" pageNumber="103" refId="B25" refString="Toussaint, EFA, Breinholt, JW, Earl, C, Warren, AD, Brower, AVZ, Yago, M, Dexter, KM, Espeland, M, Pierce, NE, Lohman, DJ, Kawahara, AY, 2018. . https://doi.org/10.1186/s12862-018-1216-z" url="https://doi.org/10.1186/s12862-018-1216-z" year="2018">Toussaint et al. 2018</bibRefCitation>
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); such problems are expected from smaller datasets. We find (
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<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Phylogenetic trees. The trees are constructed from protein-coding regions of a nuclear genome b Z-chromosome, and c mitochondrial genome. The trees are rooted with Pterourus glaucus (NVG- 1670). Specimen names are not shown in the Z-chromosome tree and can be deduced from the nuclear tree by corresponding dotted lines. Details about specimens are in Suppl. material 1: Table S 1. Sections of the tree corresponding to different subfamilies are highlighted in different colors. Names of new subfamilies and specimens in them are highlighted yellow. Names of other subfamilies are shown by their clades in the nuclear tree." figureDoi="10.3897/zookeys.861.34686.figure2" httpUri="https://binary.pensoft.net/fig/315262" pageId="5" pageNumber="96">Fig. 2</figureCitation>
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) that the
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<taxonomicName lsidName="" pageId="5" pageNumber="96" rank="subFamily" subFamily="Katreinae">Katreinae</taxonomicName>
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subfam. n. is an ancient and unique Afrotropical lineage that diverged from other
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<taxonomicName family="Hesperiidae" lsidName="" pageId="5" pageNumber="96" rank="family">Hesperiidae</taxonomicName>
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at the time when the family was diversifying into subfamilies.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |