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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">2</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:45048D35-BB1D-5CE8-9668-537E44BD4C7E</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:91BD42D4-90F1-4B45-9350-EEF175B1727A</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">ZooKeys</journal-title>
        <abbrev-journal-title xml:lang="en">ZK</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1313-2989</issn>
      <issn pub-type="epub">1313-2970</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zookeys.1198.119765</article-id>
      <article-id pub-id-type="publisher-id">119765</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Review Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Cicadelloidea</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Systematics</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿Revision of the leafhopper genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hemiptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Cicadellidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Typhlocybinae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Webb</surname>
            <given-names>Michael D.</given-names>
          </name>
          <email xlink:type="simple">m.webb@nhm.ac.uk</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-1312-6142</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Xu</surname>
            <given-names>Ye</given-names>
          </name>
          <email xlink:type="simple">yexu@jxau.edu.cn</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-3230-7923</uri>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Department of Life Sciences (Insects), The Natural History Museum, London, SW7 5BD, UK</addr-line>
        <institution>The Natural History Museum</institution>
        <addr-line content-type="city">London</addr-line>
        <country>United Kingdom</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">School of Agricultural Science, Jiangxi Agricultural University, Nanchang 330045, China</addr-line>
        <institution>Jiangxi Agricultural University</institution>
        <addr-line content-type="city">Nanchang</addr-line>
        <country>China</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding authors: Michael D. Webb (<email xlink:type="simple">mdwebb04@gmail.com);</email>Ye Xu (<email xlink:type="simple">yexu@jxau.edu.cn)</email></p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Ilia Gjonov</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2024</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>23</day>
        <month>04</month>
        <year>2024</year>
      </pub-date>
      <volume>1198</volume>
      <fpage>135</fpage>
      <lpage>141</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/74CF9BE9-7386-5BDD-AC73-0C6A8DFEC07F">74CF9BE9-7386-5BDD-AC73-0C6A8DFEC07F</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/4A6656D0-E2FD-4955-A7BE-CC1F4128AE14">4A6656D0-E2FD-4955-A7BE-CC1F4128AE14</uri>
      <history>
        <date date-type="received">
          <day>29</day>
          <month>01</month>
          <year>2024</year>
        </date>
        <date date-type="accepted">
          <day>29</day>
          <month>03</month>
          <year>2024</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Michael D. Webb, Ye Xu</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <self-uri content-type="zoobank" xlink:type="simple">http://zoobank.org/4A6656D0-E2FD-4955-A7BE-CC1F4128AE14</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p>The leafhopper genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Typhlocybinae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Empoascini</tp:taxon-name-part></tp:taxon-name>) is reviewed and a new species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name></italic> Webb &amp; Xu, <bold>sp. nov.</bold> from Brunei and Malaysia, is described based on specimens previously identified as “aberrant specimens” of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska. Images of the types of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> are given for the first time. A checklist and key to known species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> are also provided.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="suborder">Auchenorrhyncha</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>distribution</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="suborder">Homoptera</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>morphology</kwd>
        <kwd>taxonomy</kwd>
      </kwd-group>
      <funding-group>
        <award-group>
          <funding-source>
            <named-content content-type="funder_name">National Natural Science Foundation of China</named-content>
            <named-content content-type="funder_identifier">501100001809</named-content>
            <named-content content-type="funder_doi">http://doi.org/10.13039/501100001809</named-content>
          </funding-source>
        </award-group>
      </funding-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0E5G">
        <title>Citation</title>
        <p>Webb MD, Xu Y (2024) Revision of the leafhopper genus <italic>Smyga</italic> (Hemiptera, Cicadellidae, Typhlocybinae). ZooKeys 1198: 135–141. <ext-link xlink:href="10.3897/zookeys.1198.119765" ext-link-type="doi" xlink:type="simple">https://doi.org/10.3897/zookeys.1198.119765</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0EEAAC">
      <title>﻿Introduction</title>
      <p>The typhlocybine leafhopper genus, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995, was described for five species from Brunei and Malaysia (Sarawak and Sabah) (<xref ref-type="bibr" rid="B1">Dworakowska 1995</xref>; <xref ref-type="bibr" rid="B2">Knight 2010</xref>). The genus is recognized by a brown spot on the fore margin of the head and sinuate transverse brown band on the pronotum (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F1">9</xref>), a well-developed coronal suture extended onto the face (Fig. <xref ref-type="fig" rid="F1">2</xref>), and a hind wing with veins MP and CuA free distally. Although species are very similar externally, there are considerable differences in their male genitalia, particularly with respect to the aedeagus, so much so that <xref ref-type="bibr" rid="B1">Dworakowska (1995)</xref> tentatively regarded one species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic>, as having an aberrant form with a remarkably short aedeagal shaft. However, the discovery of two new species from Papua New Guinea in <xref ref-type="bibr" rid="B4">Xu’s (2019)</xref> PhD thesis, with similar aedeagus, suggests that the specimens described and illustrated by Dworakowska as an aberrant form represent a separate species. This new species is described and illustrated below and a checklist and key to the known species of the genus are presented.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EYBAC">
      <title>﻿Materials and methods</title>
      <p>Morphological terminology used in this work follows <xref ref-type="bibr" rid="B5">Xu et al. (2021)</xref>. Studied specimens are deposited in the insect collection of the <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/natural-history-museum-london">Natural History Museum, London, UK</named-content> (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>).</p>
    </sec>
    <sec sec-type="﻿Taxonomy" id="SECID0EGCAC">
      <title>﻿Taxonomy</title>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Hemiptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Cicadellidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">17410AFD-B79D-50AE-8974-05195CD017A8</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>Dworakowska, 1995</tp:taxon-authority>
          <tp:nomenclature-citation-list>
            <tp:nomenclature-citation>
              <tp:taxon-name>
                <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part>
              </tp:taxon-name>
              <comment>Dworakowska, 1995: 151.</comment>
            </tp:nomenclature-citation>
          </tp:nomenclature-citation-list>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="type species" id="SECID0EUDAC">
          <title>Type species.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995 by original designation.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="description" id="SECID0EFEAC">
          <title>Description.</title>
          <p>Body relatively robust. Pale yellow; head with a brown patch anteriorly (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F1">9</xref>); face with anteclypeus brown distally (Fig. <xref ref-type="fig" rid="F1">2</xref>), sometimes with more extensive brown marking (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="divergens">divergens</tp:taxon-name-part></tp:taxon-name></italic>); pronotum with a brown transverse narrow band at midlength, pale yellow anterior to band and silvery posterior to band (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F1">9</xref>); mesonotum with yellow to brown basal triangles (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F1">9</xref>).</p>
          <p>Head including eyes broader than pronotum in dorsal view, crown short and broad, round anteriorly, length along midline shorter than one-half width between eyes; coronal suture well developed, extended onto face, and terminating at level of antennal bases (Fig. <xref ref-type="fig" rid="F1">2</xref>). Ocelli distinct, well separated from eyes (Fig. <xref ref-type="fig" rid="F1">2</xref>). Face moderately broad; lateral frontal suture well developed, curved mesad above antennal pit and meeting coronal suture at midline ventromesad of ocelli; anteclypeus slightly convex, not expanded (Fig. <xref ref-type="fig" rid="F1">2</xref>). Pronotum large with sinuate transverse depression (Figs <xref ref-type="fig" rid="F1">1</xref>, <xref ref-type="fig" rid="F1">9</xref>). Forewing narrow, rounded apically; apical cells occupying almost one-third of total length; vein R2 and RM dissociated at bases, joined by cross-vein, both arising from r cell; vein ScP+RA is not detectable; vein MCu almost parallel with vein RM apically. Hindwing with MP+CuA confluent. Front femur seta AM1 stout, situated near ventral margin; intercalary row with one large basal seta and eight or nine smaller setae near tip of femur. Hind femur with macrosetal formula 2 + 1 + 1; tibia row AV with six or seven preapical macrosetae.</p>
          <p>Male basal abdominal sternal apodemes (2S apodemes) well developed (Fig. <xref ref-type="fig" rid="F1">3</xref>). Male pygofer elongated, strongly narrowing caudad, posterior margin acute with few rigid microsetae distally, dorsal margin with macrosetae, long fine ventrolateral setae present, ventral appendage absent (Figs <xref ref-type="fig" rid="F1">4</xref>, <xref ref-type="fig" rid="F1">10</xref>). Anal tube process elongate, extended to ventral margin of genital capsule (Figs <xref ref-type="fig" rid="F1">4</xref>, <xref ref-type="fig" rid="F1">10</xref>). Subgenital plate broadest near base (Fig. <xref ref-type="fig" rid="F1">10</xref>) or subapically (Fig. <xref ref-type="fig" rid="F1">4</xref>), all categories of setae well differentiated; basal setae encompassing mid-length of plate; marginal setae well defined; macrosetae uniserate; feeble microsetae arranged in 2–4 irregular rows apically (Fig. <xref ref-type="fig" rid="F1">4</xref>). Connective with media sclerotization, anterior margin and posterior margin deeply emarginated (Fig. <xref ref-type="fig" rid="F1">7</xref>). Style short, sinuate, with tiny teeth and microsetae preapically (Fig. <xref ref-type="fig" rid="F1">11</xref>). Aedeagus shaft short, tubular, with basal apodeme long (Figs <xref ref-type="fig" rid="F1">5</xref>, <xref ref-type="fig" rid="F1">13</xref>) or laterally compressed with basal apodeme short (Figs <xref ref-type="fig" rid="F1">6</xref>, <xref ref-type="fig" rid="F1">12</xref>); gonopore apical on ventral surface (Fig. <xref ref-type="fig" rid="F1">12</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="notes" id="SECID0EGIAC">
          <title>Notes.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> superficially resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dapitana">Dapitana</tp:taxon-name-part></tp:taxon-name></italic> Mahmood, 1967 in the features of the head and wings (coronal suture extended onto face and terminating at level of antennal bases (Fig. <xref ref-type="fig" rid="F1">2</xref>), forewing with vein R2 and RM dissociated at bases, joined by cross-vein, both arising from r cell and vein ScP+RA is not detectable). It differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dapitana">Dapitana</tp:taxon-name-part></tp:taxon-name></italic> in color pattern (see generic description) and in having the male pygofer with one or two dorsal macrosetae and long fine ventrolateral setae (Figs <xref ref-type="fig" rid="F1">4</xref>, <xref ref-type="fig" rid="F1">10</xref>) (pygofer without macrosetae and long fine setae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dapitana">Dapitana</tp:taxon-name-part></tp:taxon-name></italic>). Both genera occur on both sides of Wallace’s Line, separating the Oriental from Australian regions.</p>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> includes five previously known species, all from Borneo, described by <xref ref-type="bibr" rid="B1">Dworakowska (1995)</xref>. In her treatment of the type species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic>, Dworakowska also described and illustrated some “aberrant specimens” that she excluded from the type series because she considered the small, ventrally positioned aedeagal shaft to be “not functional.” Here we recognize these specimens as a valid species: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., as similar specimens of two new species have been seen in <xref ref-type="bibr" rid="B4">Xu’s (2019)</xref> PhD thesis.</p>
          <p>As most <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> species are from similar localities in Borneo (see Checklist) and as males are needed for identification, the female paratypes of two species (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part></tp:taxon-name></italic>) must be regarded as of uncertain identity (see also comments under <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part></tp:taxon-name></italic>). It is also of some interest that, compared to males, the number of known females is very low.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="distribution" id="SECID0EQMAC">
          <title>Distribution.</title>
          <p>Oriental Region (Brunei, Malaysia).</p>
          <sec sec-type="﻿Checklist to species of Smyga" id="SECID0EVMAC">
            <title>﻿Checklist to species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic></title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name></italic> Webb &amp; Xu, sp. nov. (Brunei, Ulu Temburong; Sarawak, Gunong Mulu National Park)</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995: 153–155, figs 168–177 (Brunei, Ulu Temburong)</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="exhibita">exhibita</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995: 155, figs 198–205 (Brunei, Ulu Temburong)</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niema">niema</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995: 155–156, figs 206–212 (Sabah)</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ziewa">ziewa</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995: 155, figs 188–192 (Brunei, Bukit Sulang)</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska, 1995: 155, figs 193–197 (Sarawak, Gunung Mulu National Park)</p>
          </sec>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <sec sec-type="﻿Key to species Smyga Dworakowska (males)" id="SECID0EOPAC">
        <title>﻿Key to species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> Dworakowska (males)</title>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table id="TID0ESFAC" rules="all">
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1">Aedeagus with basal apodeme (dorsal apodeme) very long (Fig. <xref ref-type="fig" rid="F1">5</xref>)</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Aedeagus with basal apodeme very short (Fig. <xref ref-type="fig" rid="F1">13</xref>)</td>
                <td rowspan="1" colspan="1">
                  <bold>2</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">Aedeagus without processes (Figs <xref ref-type="fig" rid="F1">12</xref>, <xref ref-type="fig" rid="F1">13</xref>)</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Aedeagus with processes</td>
                <td rowspan="1" colspan="1">
                  <bold>3</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">3</td>
                <td rowspan="1" colspan="1">Aedeagus with processes near base of shaft</td>
                <td rowspan="1" colspan="1">
                  <bold>4</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Aedeagus with processes at apex of shaft</td>
                <td rowspan="1" colspan="1">
                  <bold>5</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">4</td>
                <td rowspan="1" colspan="1">Aedeagus with processes near base of shaft long</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Aedeagus with processes near base of shaft short</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ziewa">ziewa</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">5</td>
                <td rowspan="1" colspan="1">Aedeagus with short apical processes</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="exhibita">exhibita</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Aedeagus with long apical processes</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niema">niema</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Hemiptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Cicadellidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">ECCEBE61-6F2E-5A95-9651-E20955DFF1C3</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part>
            <object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/23A87192-5521-448F-B080-84EC4CCEC98D</object-id>
          </tp:taxon-name>
          <tp:taxon-authority>Webb &amp; Xu</tp:taxon-authority>
          <tp:taxon-status>sp. nov.</tp:taxon-status>
          <xref ref-type="fig" rid="F1">Figs 1–8</xref>
          <tp:nomenclature-citation-list>
            <tp:nomenclature-citation>
              <tp:taxon-name>
                <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name>
              <comment>Dworakowska, 1995: 153, figs 178–182, in part.</comment>
            </tp:nomenclature-citation>
          </tp:nomenclature-citation-list>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="material" id="SECID0E2IAE">
          <title>Material examined.</title>
          <p><bold><italic>Holotype</italic>.</bold> ♂, Brunei, Ulu Temburong; 300 m elev.; Feb–Mar. 1982; M.C. Day leg.; B.M. 1983-75 (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>).</p>
          <p><bold><italic>Paratypes</italic>.</bold> Brunei, 8 ♂♂, same data as holotype; 10 ♂♂, same data as holotype except Ulu Temburong ridge; (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>). Malaysia, Sarawak, Gunung Mulu National Park: 5 ♂♂, Gunung Api, 900 m elev., site 25, April; montane forest, mv and act light trap; J.D. Holloway leg.; 2 ♂♂, nr Long Melinau, 50 m elev., site 17, May; low secondary forest, mv light on river-bank; J.D. Holloway; 7 ♂♂, W. Melinau Gorge leg., 250 m elev., site 23, April; limestone forest, canopy/understory, mv light; J.D. Holloway leg.; 1 ♂, W. Melinau Gorge leg., Kerankas, 150 m elev., Mar.–Apr. 1978; J.D. Holloway leg.; 2 ♂♂, camp 2.5, 30 Apr. 1978, V.F. Eastop. All paratypes from Sarawak were collected on the RGS Mulu Expedition, B.M. 1978-206 (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="description" id="SECID0EZJAE">
          <title>Description.</title>
          <p>Body length: 3.5–4.4 mm.</p>
          <p>Color as in generic description (Fig. <xref ref-type="fig" rid="F1">1</xref>).</p>
          <p>Male 2S apodemes not extending to end of segment V (Fig. <xref ref-type="fig" rid="F1">3</xref>). Anal-tube appendage hook-shaped at apex (Fig. <xref ref-type="fig" rid="F1">4</xref>). Male pygofer strongly emarginate dorsally in profile, and posteriorly extended into sharp point with spine-like setae; ventral appendage absent; bearing one long macroseta near dorsal margin (Fig. <xref ref-type="fig" rid="F1">4</xref>); dorsal bridge with a posterior lobe-like projection each side of midline, in dorsal view. Subgenital plate surpassing pygofer lobe, broadened medially in later view, all categories of setae represented (Fig. <xref ref-type="fig" rid="F1">4</xref>). Connective anterior margin with undeveloped median lobe, posterior margin notched medially (Fig. <xref ref-type="fig" rid="F1">7</xref>). Style sinuate (Figs <xref ref-type="fig" rid="F1">4</xref>, <xref ref-type="fig" rid="F1">8</xref>). Aedeagus without preatrium, basal apodeme well developed, plate-like; shaft with pair of lamellate lateral processes; gonopore subapical, ventrad (Figs <xref ref-type="fig" rid="F1">5</xref>, <xref ref-type="fig" rid="F1">6</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="etymology" id="SECID0EKLAE">
          <title>Etymology.</title>
          <p>The name is derived from the Latin words <italic>brevis</italic> (short) and <italic>penis</italic>, which refers to the very short aedeagal shaft.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="distribution" id="SECID0EULAE">
          <title>Distribution.</title>
          <p>Brunei, Malaysia.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="notes" id="SECID0EZLAE">
          <title>Notes.</title>
          <p>Specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name></italic> were considered aberrant specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> by <xref ref-type="bibr" rid="B1">Dworakowska (1995)</xref> and of uncertain status. However, based on the high degree of difference between these two species and the discovery of two similar new species from Papua New Guinea noted in <xref ref-type="bibr" rid="B4">Xu’s (2019)</xref> PhD thesis, we describe the above specimens as a new species. All type specimens listed here (with some amendments) were identified by I. Dworakowska as the aberrant form of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic>. All are male and most have a Dworakowska genitalia dissection affixed to a card placed beneath the specimen. Three females (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>) with identical data to the males could be the same species but, as three other species are also from these localities (see Checklist), their identity is uncertain. See also the concluding comments under the genus description. In <xref ref-type="bibr" rid="B1">Dworakowska’s (1995)</xref> figure of the head and thorax there are some fine lines in the basal triangles of the mesonotum. These are seen in some specimens, but it is not clear if they are markings or perhaps caused by shrinkage of internal tissue during drying.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Hemiptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Cicadellidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">C2549A40-24FB-5273-B01C-E13C8D55D3A2</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>Dworakowska</tp:taxon-authority>
          <xref ref-type="fig" rid="F1">Figs 9–13</xref>
          <tp:nomenclature-citation-list>
            <tp:nomenclature-citation>
              <tp:taxon-name>
                <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name>
              <comment>Dworakowska, 1995: 153–155, figs 168–177.</comment>
            </tp:nomenclature-citation>
          </tp:nomenclature-citation-list>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="notes" id="SECID0EGPAE">
          <title>Notes.</title>
          <p>The type series was originally stated as holotype male from Brunei, Ulu Temburong and one paratype male and two females from three different localities in Sarawak, Gunung Mulu National Park; however, the paratype male is in fact female. The identity of these three females must remain uncertain until females in the genus can be identified, particularly as other species are sympatric with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic> (see Checklist).</p>
          <fig id="F1" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1198.119765.figures1-13</object-id>
            <object-id content-type="arpha">CDF5DB21-FF05-5D66-8B44-75F5072DB3AE</object-id>
            <label>Figures 1–13.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part></tp:taxon-name></italic> species <bold>1–8</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevipenis">brevipenis</tp:taxon-name-part></tp:taxon-name></italic><bold>1</bold> head and thorax, dorsal view (holotype) <bold>2</bold> face (holotype) <bold>3–8</bold> male genitalia after <xref ref-type="bibr" rid="B1">Dworakowska’s (1995)</xref> figures of aberrant form of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic><bold>3</bold> basal sternal apodemes <bold>4</bold> male genital capsule, left lateral view <bold>5</bold> aedeagus left lateral view <bold>6</bold> aedeagus, ventral view <bold>7</bold> connective <bold>8</bold> connective, style and subgenital plate <bold>9–13</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="distincta">distincta</tp:taxon-name-part></tp:taxon-name></italic><bold>9</bold> habitus (holotype) <bold>10–13</bold> male genitalia, after <xref ref-type="bibr" rid="B1">Dworakowska (1995)</xref><bold>10</bold> genital capsule <bold>11</bold> style <bold>12</bold> aedeagus ventral view <bold>13</bold> aedeagus, left lateral view.</p>
            </caption>
            <graphic xlink:href="zookeys-1198-135_article-119765__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1031824.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1031824</uri>
            </graphic>
          </fig>
          <p>Aberrant specimens regarded as possibly this species in its original description are described above as a new species. Therefore, this species is known with certainty only from a single male, the holotype (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>).</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order" xlink:type="simple">Hemiptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Cicadellidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">1FA8216A-07AB-5460-9B4F-43E1E241493D</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>Dworakowska</tp:taxon-authority>
          <tp:nomenclature-citation-list>
            <tp:nomenclature-citation>
              <tp:taxon-name>
                <tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">Smyga</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part></tp:taxon-name>
              <comment>Dworakowska, 1995: 155, figs 193–197.</comment>
            </tp:nomenclature-citation>
          </tp:nomenclature-citation-list>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="notes">
          <title>Notes.</title>
          <p>
           The type series was originally stated to include a holotype male from Sarawak, Gunung Mulu National Park and one male and two female paratypes from Brunei, Ulu Temburong. The identity of these two females must remain uncertain until females in the genus can be identified, particularly as other species are sympatric with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Smyga">S.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="zonata">zonata</tp:taxon-name-part></tp:taxon-name></italic> (see Checklist). Therefore, this species is known with certainty only from two male specimens (<named-content content-type="dwc:institutional_code" xlink:title="Natural History Museum, London, UK" xlink:href="http://grbio.org/institution/natural-history-museum-london">NHM</named-content>).</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgements</title>
      <p>For their helpful comments we give our sincere thanks to Dr I. Dworakowska and Dr D. Dmitriev.</p>
    </ack>
    <sec sec-type="﻿Additional information" id="SECID0EEVAE">
      <title>﻿Additional information</title>
      <sec sec-type="Conflict of interest" id="SECID0EIVAE">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="SECID0ENVAE">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Funding" id="SECID0ESVAE">
        <title>Funding</title>
        <p>This work was supported by the National Natural Science Foundation of China (32160121).</p>
      </sec>
      <sec sec-type="Author contributions" id="SECID0EXVAE">
        <title>Author contributions</title>
        <p>Writing - original draft: MW. Writing - review and editing: YX.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="SECID0E3VAE">
        <title>Author ORCIDs</title>
        <p>Michael D. Webb <ext-link xlink:href="https://orcid.org/0000-0002-1312-6142" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-1312-6142</ext-link></p>
        <p>Ye Xu <ext-link xlink:href="https://orcid.org/0000-0003-3230-7923" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0003-3230-7923</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="SECID0EMWAE">
        <title>Data availability</title>
        <p>All of the data that support the findings of this study are available in the main text.</p>
      </sec>
    </sec>
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</article>
