<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//TaxonX//DTD Taxonomic Treatment Publishing DTD v0 20100105//EN" "../../nlm/tax-treatment-NS0.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:tp="http://www.plazi.org/taxpub" article-type="rapid-communication" dtd-version="3.0" xml:lang="en">
  <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.1264.153948</article-id>
      <article-id pub-id-type="publisher-id">153948</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Short Communication</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Rutelidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Nomenclature</subject>
          <subject>Systematics</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Brazil</subject>
          <subject>South America</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿Reassessment of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917 reveals <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> Martínez, 1964 as new generic and specific synonym (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Bento</surname>
            <given-names>Matheus</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-9846-9728</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Grossi</surname>
            <given-names>Paschoal</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-6601-5967</uri>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Ruy Vasconcelos da Fonseca</surname>
            <given-names>Claudio</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-1955-288X</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Seidel</surname>
            <given-names>Matthias</given-names>
          </name>
          <email xlink:type="simple">matthias.seidel@nhm-wien.ac.at</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-4913-8778</uri>
          <xref ref-type="aff" rid="A3">3</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Instituto Nacional de Pesquisas da Amazônia (INPA), Coordenação de Biodiversidade, Laboratório de Sistemática e Ecologia de Coleoptera (LASEC), Manaus, Brazil</addr-line>
        <institution>Instituto Nacional de Pesquisas da Amazônia (INPA), Coordenação de Biodiversidade</institution>
        <addr-line content-type="city">Manaus</addr-line>
        <country>Brazil</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Universidade Federal Rural de Pernambuco (UFRPE), PPG em Entomologia, Laboratório de Taxonomia de Insetos (LabTaxIn), Rua Dom Manoel de Medeiros, s/n, 52171-900, Recife, Pernambuco, Brazil</addr-line>
        <institution>Universidade Federal Rural de Pernambuco (UFRPE)</institution>
        <addr-line content-type="city">Recife</addr-line>
        <country>Brazil</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Naturhistorisches Museum Wien, Zweite Zoologische Abteilung, Burgring 7, 1010 Vienna, Austria</addr-line>
        <institution>Naturhistorisches Museum Wien</institution>
        <addr-line content-type="city">Vienna</addr-line>
        <country>Austria</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Matthias Seidel (<email xlink:type="simple">matthias.seidel@nhm.at</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Andrey Frolov</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2025</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>11</day>
        <month>12</month>
        <year>2025</year>
      </pub-date>
      <volume>1264</volume>
      <fpage>61</fpage>
      <lpage>71</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/C89E6850-591A-589F-BD11-7419AB7CB802">C89E6850-591A-589F-BD11-7419AB7CB802</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/17919805">17919805</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/2F02780C-496B-4DC5-B1CD-4556D6131DF5">2F02780C-496B-4DC5-B1CD-4556D6131DF5</uri>
      <history>
        <date date-type="received">
          <day>25</day>
          <month>03</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>22</day>
          <month>09</month>
          <year>2025</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Matheus Bento, Paschoal Grossi, Claudio Ruy Vasconcelos da Fonseca, Matthias Seidel</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">https://zoobank.org/2F02780C-496B-4DC5-B1CD-4556D6131DF5</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p>In this paper, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> Martínez (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>) is synonymized with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus (<bold>syn. nov.</bold>). Hence, the monotypic genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic> Martínez is a new junior synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part></tp:taxon-name></italic> MacLeay (<bold>syn. nov.</bold>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> is redescribed and illustrated based on the type and additional material, including new distributional records in southern Brazil. As a result of our work, the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> now includes 12 genera.</p>
      </abstract>
      <kwd-group>
        <label>Key words:</label>
        <kwd>Bearded scarab beetles</kwd>
        <kwd>monotypic</kwd>
        <kwd>morphology</kwd>
        <kwd>Neotropical region</kwd>
        <kwd>nomenclatural changes</kwd>
        <kwd>taxonomy</kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0EVH">
        <title>Citation</title>
        <p>Bento M, Grossi P, Ruy Vasconcelos da Fonseca C, Seidel M (2025) Reassessment of <italic>Leucothyreus noctivagus</italic> Ohaus, 1917 reveals <italic>Mimogeniates margaridae</italic> Martínez, 1964 as new generic and specific synonym (Scarabaeidae, Rutelinae, Geniatini). ZooKeys 1264: 61–71. <ext-link xlink:href="10.3897/zookeys.1264.153948" ext-link-type="doi" xlink:type="simple">https://doi.org/10.3897/zookeys.1264.153948</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0EGBAC">
      <title>﻿Introduction</title>
      <p>The bearded scarab tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>) is a diverse group of phytophagous scarabs widely distributed in the Neotropical Region (<xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>). The tribe is currently composed of 338 species in 13 genera (<xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>; <xref ref-type="bibr" rid="B8">Jameson 2008</xref>; <xref ref-type="bibr" rid="B7">Grossi and Vaz-de-Mello 2018</xref>; <xref ref-type="bibr" rid="B4">Ferreira et al. 2019</xref>, <xref ref-type="bibr" rid="B5">2024</xref>). Members of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> are diagnosed by having the labrum vertically positioned in relation to the clypeus, both the labrum and the prementum usually with a medioapical projection, the elytral margins with or without a membranous border, the inner protibial spur present, and the male protarsomeres usually enlarged and densely setose internally (Villatoro and Jameson 2001; <xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>).</p>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic> Martínez, 1964 is a monotypic genus and was described based on <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> Martínez, 1964 from Espírito Santo state (Brazil). The genus was erected for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> based on the labium having a crenulate apical margin and lacking the apical projection, which is characteristic of most geniatine genera (<xref ref-type="bibr" rid="B13">Martínez 1964</xref>). Because of the general form of the maxillary galea, which is “neither enlarged nor keeled at the apex”, <xref ref-type="bibr" rid="B13">Martínez (1964</xref>: 2) thought the genus belonged to a group formed by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Geniates">Geniates</tp:taxon-name-part></tp:taxon-name></italic> Kirby, 1818, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Rhizogeniates">Rhizogeniates</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1909, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Eunanus">Eunanus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1909. However, he classified the genus near to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bolax">Bolax</tp:taxon-name-part></tp:taxon-name></italic> Fischer &amp; Waldheim, 1829 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part></tp:taxon-name></italic> Macleay, 1819 because these genera share laterally carinate abdominal sternites (<xref ref-type="bibr" rid="B13">Martínez 1964</xref>; <xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>). Nevertheless, these character states are broadly distributed across most genera of the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> and uninformative for genus-level taxonomy. Moreover, based on the lack of a medioapical projection on the labrum of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic>, <xref ref-type="bibr" rid="B13">Martínez (1964</xref>: 4) alluded to the superficial similarity of his new genus with <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Anatistini</tp:taxon-name-part></tp:taxon-name>: “This new genus, due to the characters it presents with having the labrum and mentum inerm, fits perfectly in the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spodochlamiini</tp:taxon-name-part></tp:taxon-name> [= <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Anatistini</tp:taxon-name-part></tp:taxon-name>], but we do not think that it should be located in this tribe, since the shape of the maxillae, mandibles, structure of the integument, secondary sexual characters and male genitalia (phallobase and parameres), are all of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> type … . Regarding <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic>, it is not described as <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spodochlamiini</tp:taxon-name-part></tp:taxon-name> [= <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Anatistini</tp:taxon-name-part></tp:taxon-name>] or as <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>, but due to the analysis of the qualitative characters it presents, despite the ‘fundamental’ characters of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spodochlamiini</tp:taxon-name-part></tp:taxon-name> [=<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Anatistini</tp:taxon-name-part></tp:taxon-name>], I consider it more correct to place it in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>”.</p>
      <p>The crenulate apical margin of the labium of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> is a unique feature among geniatine scarabs, and a morphology-based phylogenetic analysis of all <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> genera (<xref ref-type="bibr" rid="B1">Bento 2024</xref>) recovered <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> nested in the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part></tp:taxon-name></italic> clade and forming a well-supported subclade with the type species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kirbyanus">kirbyanus</tp:taxon-name-part></tp:taxon-name></italic> MacLeay, 1819. This analysis provides strong support for the synonymization of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part></tp:taxon-name></italic>. Furthermore, after analysis of the type series of both species, we found evidence that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> is conspecific with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917. Based on these results, we propose the correspondent generic and specific synonyms and redescribe <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic>.</p>
    </sec>
    <sec sec-type="methods" id="SECID0EZKAC">
      <title>﻿Methods</title>
      <p>Type and non-type material used in this study is deposited in the following institutions (acronym and curators in parentheses): <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">Museum für Naturkunde der Humboldt-Universität, Berlin, Germany</named-content> (<bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold>); <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/naturhistorishes-museum">Naturhistorisches Museum Basel, Basel, Switzerland</named-content> (<bold><named-content content-type="dwc:institutional_code" xlink:title="Naturhistorisches Museum Basel, Basel, Switzerland" xlink:href="http://grbio.org/institution/naturhistorishes-museum">NMB</named-content></bold>; Matthias Borer and Isabelle Zürcher); and the Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil (<bold><abbrev content-type="institution" xlink:title="Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil" id="ABBRID0EMLAC">CEIOC</abbrev></bold>; Márcio Felix and Claudia Leal Rodrigues); <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/canadian-museum-nature-3">Canadian Museum of Nature, Ottawa, Canada</named-content> (<bold><named-content content-type="dwc:institutional_code" xlink:title="Canadian Museum of Nature, Ottawa, Canada" xlink:href="http://grbio.org/institution/canadian-museum-nature-3">CMN</named-content></bold>; François Génier); <named-content xlink:type="simple" content-type="institution" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">Coleção Entomológida da Universidade Federal Rural de Pernambuco</named-content> (<bold><named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content></bold>; Paschoal C. Grossi).</p>
      <p>The general morphological terms follow <xref ref-type="bibr" rid="B2">Beutel and Lawrence (2005)</xref>, with the adoption of the term <bold>tectum</bold> for the distal portion of the phallobase (non-apodeme <italic>sensu</italic><xref ref-type="bibr" rid="B11">Krell 1996</xref>). Terminology regarding orientation of legs follows <xref ref-type="bibr" rid="B6">Fuhrmann (2021)</xref>, which determines three axes: proximal/distal; inner/outer; and anterior/posterior. Cuticular surface was considered <bold>densely setose</bold> when the surface is completely hidden by setae; <bold>moderately setose</bold> when the surface is covered with setae but not hidden; and <bold>sparsely setose</bold> when the surface has only few setae. Body size was measured from anterior margin of clypeus to apex of pygidium, and the body width was measured at mid-elytra. The surface was considered <bold>densely punctate</bold> when the distance between punctures was less than twice the puncture diameter, <bold>moderately punctate</bold> when the distance was 2–6 times the puncture diameter, and <bold>sparsely punctate</bold> when the distance was more than six puncture diameters.</p>
      <p>A Canon EOS 750D camera provided with an MP-E 65 mm macro lens was used for specimen photographs. All produced images were processed and edited using Helicon Focus (HeliconSoft) and Photoshop (Adobe Inc.). Distributional data were based on collections and published data, including <xref ref-type="bibr" rid="B15">Ohaus (1917)</xref>, <xref ref-type="bibr" rid="B13">Martínez (1964)</xref>, <xref ref-type="bibr" rid="B9">Jameson and Hawkins (2005)</xref>, and <xref ref-type="bibr" rid="B10">Krajcik (2007)</xref>. The geographic coordinates were obtained with Google Maps and the georeferenced points were plotted on the distribution map generated with Simplemappr (<xref ref-type="bibr" rid="B16">Shorthouse 2010</xref>).</p>
    </sec>
    <sec sec-type="﻿Results" id="SECID0ERNAC">
      <title>﻿Results</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">Coleoptera</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family" xlink:type="simple">Scarabaeidae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <label>﻿</label>
          <tp:taxon-name><object-id content-type="arpha">604AF01C-3E12-5B49-B998-87C4CA9F3EDB</object-id>
            <tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part>
            <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part>
          </tp:taxon-name>
          <tp:taxon-authority>Ohaus, 1917</tp:taxon-authority>
          <tp:nomenclature-citation-list>
            <tp:nomenclature-citation>
              <tp:taxon-name>
                <tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name>
              <comment>Ohaus, 1917: 15 [original combination]; <xref ref-type="bibr" rid="B3">Blackwelder 1944</xref>: 249; Machatschke 1974: 409; <xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>: 45; <xref ref-type="bibr" rid="B10">Krajcik 2007</xref>: 77.</comment>
            </tp:nomenclature-citation>
            <tp:nomenclature-citation>
              <tp:taxon-name>
                <tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name>
              <comment>Martínez, 1964: 5 [original combination]; <xref ref-type="bibr" rid="B12">Machatschke 1972</xref>: 359; <xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>: 24, 53, 65, 70; <xref ref-type="bibr" rid="B10">Krajcik 2007</xref>: 89. Syn. nov.</comment>
            </tp:nomenclature-citation>
          </tp:nomenclature-citation-list>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="type material" id="SECID0ENAAE">
          <title>Type material examined.</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic>: <bold>lectotype</bold> • male (here designated) deposited at <bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold> (Fig. <xref ref-type="fig" rid="F1">1A–D</xref>), labeled: “Espírito Santo / Sta. Leopoldina / O. Michaelis” (white, printed) // “berlin” (white, printed in red) // “Cotype” (red, printed) // “Leucothyreus / noctivagus Ohs.” (red, printed) // “SYNTYPUS / Leucothyreus / noctivagus Ohaus, 1817 / labeled by MNHUB 2016” (red, printed). <bold>Paralectotypes</bold>, • same data as lectotype (♂, <bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold>); • same, but “Boa Sorte / F. Sahlb.” (white, printed) // “♀” (♀, <bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold>); • same, but “Rio Itapemirim / E. Esp. S. – Brazil / 5-12-1908 / J. F. Zikán” (white, printed) (♀, <bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold>); • same, but “Petropolis / 15.I.99 / Electr. Licht” (white, printed) (♀, <bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold>); • same but “Minas Gerais / Mar d. Espanha / J. Zikán S.” (white, printed) // “14/11/1909” (white, handwritten) (♀, <bold><named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content></bold>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic>: <bold>Paratype</bold> • female deposited at <bold><named-content content-type="dwc:institutional_code" xlink:title="Canadian Museum of Nature, Ottawa, Canada" xlink:href="http://grbio.org/institution/canadian-museum-nature-3">CMN</named-content></bold>, labeled: “BRASIL. E. Santo / Linhares Sooretama / NOV. 62 A. Martínez” (white, printed) // “PARATIPO” (green, printed) // “Mimogeniates / margaridae / ♀ sp. n. / Martinez-det, 1964” (light green, handwritten) // “H. &amp; A. HOWDEN / COLLECTION / ex. A. Martinez coll.” (white, printed).</p>
          <fig id="F1" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1264.153948.figure1</object-id>
            <object-id content-type="zenodo_dep_id">17919866</object-id>
            <object-id content-type="arpha">0D9D351C-F45A-5CBB-9228-7D90C6367B53</object-id>
            <label>Figure 1.</label>
            <caption>
              <p>Lectotype male (here designated) of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917. <bold>A.</bold> Dorsal view; <bold>B.</bold> Ventral view; <bold>C.</bold> Labels; <bold>D.</bold> Aedeagus in frontal and lateral views. Scale bars: 5 mm (<bold>A, B</bold>); 1 mm (<bold>D</bold>).</p>
            </caption>
            <graphic xlink:href="zookeys-1264-061_article-153948__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1490550.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1490550</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="material" id="SECID0EOFAE">
          <title>Additional material (17 males; 17 females).</title>
          <p>Brazil • Rio de Janeiro, Bom Jardim, Sítio São José, 29.xi.2003, M. Hoffmann col. (♂, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); Brazil • São Paulo, Itú, Fazenda Pau d’Alho, 28–29.X.1965, Martins &amp; Biasi (legs.) (♂, <named-content content-type="dwc:institutional_code" xlink:title="Naturhistorisches Museum Basel, Basel, Switzerland" xlink:href="http://grbio.org/institution/naturhistorishes-museum">NMB</named-content>); Brazil • Espírito Santo, Itapemirim, 5.XII.1908, J. F. Zikán (leg.) (♀, <abbrev content-type="institution" xlink:title="Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil" id="ABBRID0E5FAE">CEIOC</abbrev>); Brazil • Espírito Santo, Linhares, Parque Sooretama, 17–27.X.1962 (♂, <named-content content-type="dwc:institutional_code" xlink:title="Canadian Museum of Nature, Ottawa, Canada" xlink:href="http://grbio.org/institution/canadian-museum-nature-3">CMN</named-content>); Brazil • Espírito Santo, Rio Bonito, XI.1963, A. Maller (leg.), Martínez collection (♂, <named-content content-type="dwc:institutional_code" xlink:title="Canadian Museum of Nature, Ottawa, Canada" xlink:href="http://grbio.org/institution/canadian-museum-nature-3">CMN</named-content>); Brazil, Espírito Santo, Linhares, Flona de Goytacazes, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-40.070167,-19.432306]}" id="NCID0EQGAE">19°25'56.3"S, 40°04'12.6"W</named-content></named-content>, 27.xi.2010, light, DS Martins (♂, ♀, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); • same, but 09.xi.2010 (2♂, 3 ♀, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); • same, but 19.xi.2010 (2 ♀, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); Brazil • Bahia, Encruzilhada, XII.1980, A. Martínez &amp; M. Alvarenga (legs.) (♀, <named-content content-type="dwc:institutional_code" xlink:title="Canadian Museum of Nature, Ottawa, Canada" xlink:href="http://grbio.org/institution/canadian-museum-nature-3">CMN</named-content>); Brazil • Minas Gerais, Pedra Azul, 1974 (♀, <named-content content-type="dwc:institutional_code" xlink:title="Naturhistorisches Museum Basel, Basel, Switzerland" xlink:href="http://grbio.org/institution/naturhistorishes-museum">NMB</named-content>); Brazil • Minas Gerais, Mar de Hespanha, 19.XI.1909, J. F. Zikán (leg.) (♀, <abbrev content-type="institution" xlink:title="Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil" id="ABBRID0EOHAE">CEIOC</abbrev>); Brazil • Minas Gerais, Ipatinga, xi.1989, luz, E. &amp; P. Grossi legs. (5 ♂, 6 ♀, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); Brazil • Minas Gerais, Itumirim, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-44.601600,-23.340000]}" id="NCID0E2HAE">23.3400°S, 44.6016°W</named-content></named-content>, x.2016, FZ Vaz-de-Mello (♂, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); Brazil • Minas Gerais, Lavras, 02.vii.2007, PF Espuri (♂, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); Brazil • Minas Gerais, Berizal, Faz. Veredão, 14.xii.2007, 850 m, Grossi, Rafael &amp; Parizotto legs. (2 ♂, ♀, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>); Brazil • Minas Gerais, Águas Vermelhas, 13.xii.2007, Grossi, Rafael &amp; Parizotto legs. (♂, <named-content content-type="dwc:institutional_code" xlink:title="Coleção Entomológida da Universidade Federal Rural de Pernambuco" xlink:href="http://grbio.org/institution/universidade-federal-rural-de-pernambuco">CERPE</named-content>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="diagnosis" id="SECID0EUIAE">
          <title>Diagnosis.</title>
          <p>Labrum and prementum without medioapical projection (Fig. <xref ref-type="fig" rid="F3">3D1, D4</xref>). Apical margin of prementum crenulated (Fig. <xref ref-type="fig" rid="F3">3D4</xref>). Outer margin of metafemur coarsely crenulated, with each crenulation bearing a short, white seta (Fig. <xref ref-type="fig" rid="F3">3B</xref>). Outer margin of meso- and metatibia with two short, spine-like projections (Fig. <xref ref-type="fig" rid="F3">3A, B</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Differential diagnosis" id="SECID0EKJAE">
          <title>Differential diagnosis.</title>
          <p>This species is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kirbyanus">kirbyanus</tp:taxon-name-part></tp:taxon-name></italic> MacLeay, 1819 (see <xref ref-type="bibr" rid="B9">Jameson and Hawkins 2005</xref>: fig. 7), as both share a similar general appearance of head and clypeus, chaetotaxy, similar form of the male aedeagus, coarsely crenulated outer margin of metafemur (Fig. <xref ref-type="fig" rid="F3">3B</xref>) as well as the outer margin of meso- and metatibia with short, spine-like projections (Fig. <xref ref-type="fig" rid="F3">3A, B</xref>). However, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> is easily distinguished from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kirbyanus">kirbyanus</tp:taxon-name-part></tp:taxon-name></italic> and other species in the genus by the form of the prementum, which is crenulated and lacking a median projection (Fig. <xref ref-type="fig" rid="F3">3A, B</xref>).</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="description" id="SECID0EBLAE">
          <title>Description.</title>
          <p><bold>Male</bold> (Figs <xref ref-type="fig" rid="F1">1A–D</xref>, <xref ref-type="fig" rid="F2">2A–D</xref>). Length 9.5–10.5 mm. Width 4.8–5.3 mm. Body oval, elongate. <bold>Coloration</bold> of head, pronotum, and scutellar shield reddish brown, with weak to strong metallic-green reflections (Figs <xref ref-type="fig" rid="F1">1A</xref>, <xref ref-type="fig" rid="F2">2A, B</xref>); head darker than pronotum. Elytra and legs light brown, with weak greenish reflections. Venter reddish brown, covered with white, scale-like setae. <bold>Pubescence</bold>: lateral surface of pronotum and scutellar shield moderately and irregularly covered with posteriorly decumbent, white, scale-like setae; setae thinner on scutellar shield; thoracic venter and abdomen densely covered with decumbent, white, scale-like setae. <bold>Head</bold> small, shorter than pronotum at middle, with deep, large, densely distributed punctures. Frons as wide as clypeus at base. Frontoclypeal suture complete and slightly sinuous at middle. Clypeus subrectangular, with anterior margin weakly raised. Labrum large, subtriangular, slightly longer than ventral face of clypeus; apex weakly angulated, not projected (Fig. <xref ref-type="fig" rid="F3">3D1</xref>. Labium (Fig. <xref ref-type="fig" rid="F3">3D4</xref>) subrectangular, slightly wider than long; prementum with apical margin coarsely crenulated at middle, not projected; mentum with a transverse, moderate setal brush medially erect and laterally decumbent bearing thick, white setae. Labial palp 3-segmented, with palpomere II distinctly shorter than III. Maxilla densely setose, with thick, decumbent, white setae; palpifer with outer margin strongly curved at base; maxillary palp 4-segmented, with distal palpomere fusiform, longer than palpomere I–III combined, with a small, fusiform dorsal sensory area; galea (Fig. <xref ref-type="fig" rid="F3">3D3</xref>) apically large, with two acute teeth. Mandible externally rounded and weakly raised at apex; apical tooth rounded and slightly deflected ventrally; outer face with thick, white setae. Antennae with 10 antennomeres, with club elongate, slightly shorter than antennomeres 2–7 combined. <bold>Pronotum</bold> widest at middle; medially wider than head, with lateral margins evenly curved. Surface moderately covered with deep, setigerous punctures; lateral punctures with decumbent, scale-like, white setae. Anterior bead punctate, barely defined at middle; posterior bead complete and well defined. Anterior angles acute; posterior angles obtuse. <bold>Scutellar shield</bold> small, as wide as long, with posterior margin acute; basal surface with sparse, deep, setigerous punctures bearing thick, decumbent, white setae. <bold>Elytra</bold> shallowly rugopunctate, with three barely defined longitudinal costae. Lateral margins glabrous. <bold>Pygidium</bold> subtriangular and strongly concave posteriorly; whole surface moderately covered with deep, transversely fusiform punctures bearing white, transversely decumbent scale-like setae; posteromedial region with a few thin, erect setae. <bold>Thoracic venter</bold>: prosternum, meso- and metaventrite, and metacoxae densely covered with decumbent, scale-like, white setae; posterior prosternal process triangular, with apex acute. Metaventrite with postcoxal line medially effaced, with a row of thinner, hair-like setae. <bold>Legs</bold>: protibia slender, slightly wider at apex, with three small, acute outer teeth; proximal tooth reduced and largely separated from medial and distal teeth; protarsomeres I–V (Figs <xref ref-type="fig" rid="F1">1A, B</xref>, <xref ref-type="fig" rid="F2">2A, B</xref>, <xref ref-type="fig" rid="F3">3C</xref>) dorsoventrally flattened and densely setose internally; protarsomere I small, slightly surpassing the apex of protibia; II–IV somewhat caliciform and as wide as long; V as long as protarsomeres III and IV combined. Protarsal claws small, shorter than protarsomere V; anterior protarsal claw slender and laterally flattened, with a horizontal, apical cleft; posterior claw simple and shorter than anterior claw. Meso- and metafemur with outer margin coarsely crenulated; each crenulation bearing a short, thin seta. Meso- and metatibia slightly constricted near apex, with outer margin bearing one short, spine-like processes adjacent to each transverse carina (Fig. <xref ref-type="fig" rid="F3">3A, B</xref>); surface moderately covered with thick, hair-like, white setae. Inner meso- and metatibial spur longer than respective tarsomere I. Meso- and metatarsomeres I–IV slightly flattened dorsoventrally and densely setose internally. Meso- and metatarsomere V cylindrical, sparsely setose internally, longer than respective tarsomeres II–IV combined, with a short, acute internobasal tooth. Meso- and metatarsal claws slender, in flexed position as long as respective tarsomere V; anterior claw slightly thicker, with apex narrowly cleft; posterior claw simple. Meso- and metatarsal empodium with two long, apically bifurcate setae. <bold>Abdomen</bold> densely covered with decumbent, scale-like, white setae. Ventrite 1 medially projected, with posterior postcoxal line posteriorly extending to posterior margin of ventrite 1 and bearing dense, hair-like setae. Ventrites 1–4 with wide laterolongitudinal carinae. Ventrite 6 with posterior margin weakly emarginate. <bold>Aedeagus</bold> (Figs <xref ref-type="fig" rid="F1">1D</xref>, <xref ref-type="fig" rid="F2">2C, D</xref>): parameres small, about ½ length of tectum; prolongations subparallel and completely fused together to apex, with a median fusion line barely defined. Apex evenly rounded, as wide as or slightly wider than midpoint, with outer margins slightly converging. Inner surface bearing an internomedial row of setae.</p>
          <fig id="F2" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1264.153948.figure2</object-id>
            <object-id content-type="zenodo_dep_id">17919868</object-id>
            <object-id content-type="arpha">DAB9E2CF-0434-5FC8-960C-0FB143617DE2</object-id>
            <label>Figure 2.</label>
            <caption>
              <p>Male of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917. <bold>A.</bold> Dorsal view; <bold>B.</bold> Head and pronotum showing color variation with weak metallic-green reflections; <bold>C.</bold> Parameres in frontal view; <bold>D.</bold> Aedeagus in lateral view. Scale bars 5 mm (<bold>A</bold>); 1 mm (<bold>B</bold>); 0.2 mm (<bold>C</bold>). Photos by Vitor Nardino.</p>
            </caption>
            <graphic xlink:href="zookeys-1264-061_article-153948__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1490551.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1490551</uri>
            </graphic>
          </fig>
          <fig id="F3" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1264.153948.figure3</object-id>
            <object-id content-type="zenodo_dep_id">17919870</object-id>
            <object-id content-type="arpha">5A710FE2-C62E-5972-981F-7A529EB985BD</object-id>
            <label>Figure 3.</label>
            <caption>
              <p>Paralectotype female of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917. <bold>A.</bold> Dorsal view. <bold>B.</bold> Ventral view; <bold>C.</bold> Pronotum, head, and anterior legs of (left) female and (right) male; <bold>D.</bold> Mouthparts: <bold>D1.</bold> Labrum showing lack of apical projection; <bold>D2.</bold> Left mandible; <bold>D3.</bold> Left maxilla; <bold>D4.</bold> Labium showing crenulate apex. Scale bars: 5 mm (<bold>A, B</bold>); 1 mm (<bold>D</bold>).</p>
            </caption>
            <graphic xlink:href="zookeys-1264-061_article-153948__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1490552.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1490552</uri>
            </graphic>
          </fig>
          <p><bold>Female</bold> (Fig. <xref ref-type="fig" rid="F3">3A–C</xref>). Length 11–11.5 mm. Width 5.8–6 mm. Females very similar to males but with body more robust, pronotum sparsely setose, protibial outer teeth stronger, pro- and mesotarsomeres moderately setose and narrower, and posterior margin of ventrite 6 evenly rounded, not emarginate.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="distribution" id="SECID0ENQAE">
          <title>Distribution (Fig. <xref ref-type="fig" rid="F4">4</xref>).</title>
          <p>Brazil. Bahia: Encruzilhada (new state record). Espírito Santo: Linhares, Rio Bonito, Itapemirim, Santa Leopoldina. Rio de Janeiro: Petrópolis, Boa Sorte, Bom Jardim. Minas Gerais: Mar de Espanha, Águas Vermelhas, Berizal, Itumirim, Ipatinga, Lavras. São Paulo: Itú (new state record).</p>
          <fig id="F4" position="float" orientation="portrait">
            <object-id content-type="doi">10.3897/zookeys.1264.153948.figure4</object-id>
            <object-id content-type="zenodo_dep_id">17919872</object-id>
            <object-id content-type="arpha">911C2922-96F0-524E-A269-F5A7555181A4</object-id>
            <label>Figure 4.</label>
            <caption>
              <p>Distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917.</p>
            </caption>
            <graphic xlink:href="zookeys-1264-061_article-153948__-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1490553.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1490553</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="﻿Discussion" id="SECID0EPRAE">
      <title>﻿Discussion</title>
      <p><xref ref-type="bibr" rid="B13">Martínez (1964)</xref> erected the monotypic genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic> for his new species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> based on the crenulated, unprojected prementum, as well as the unprojected apical margin of labrum. Herein, based on the analysis of the type series and additional material, we synonymize this species with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917 (new synonym).</p>
      <p>In the original description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic>, <xref ref-type="bibr" rid="B15">Ohaus (1917)</xref> compared this species to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niveicollis">niveicollis</tp:taxon-name-part></tp:taxon-name></italic> Laporte, 1840. According to <xref ref-type="bibr" rid="B9">Jameson and Hawkins (2005)</xref>, Ohaus was confused about the identity of these species and even mixed both species within an identified series of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niveicollis">niveicollis</tp:taxon-name-part></tp:taxon-name></italic>, which is reflective of their similarity. However, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> is easily distinguished from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niveicollis">niveicollis</tp:taxon-name-part></tp:taxon-name></italic> by the lack of scale-like setae on the elytral surface (covered with scale-like setae in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niveicollis">niveicollis</tp:taxon-name-part></tp:taxon-name></italic>) and the lack of a medioapical projection on the labium (present in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="niveicollis">niveicollis</tp:taxon-name-part></tp:taxon-name></italic>).</p>
      <p>As revealed by a phylogenetic analysis (<xref ref-type="bibr" rid="B1">Bento 2024</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> is nested within the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part></tp:taxon-name></italic> clade, and forms a subclade with the type species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kirbyanus">kirbyanus</tp:taxon-name-part></tp:taxon-name></italic>. This clade was strongly supported by 10 synapomorphies (Bremer support value = 83), including the coarsely crenulated outer margin of metafemur and the outer margin of meso- and metatibia with short, spine-like projections appeared as exclusive synapomorphies. Additional synapomorphies are: maxillary tooth III pointed, pedicel shorter than antennomere 3, scale-like setae on pronotal surface, posterior margin of pronotum beaded, and prosternal process projected between procoxae.</p>
      <p>The close morphological similarity and phylogenetic relationship between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kirbyanus">kirbyanus</tp:taxon-name-part></tp:taxon-name></italic> suggest that the median projection on the apical margins of the labrum and prementum—a diagnostic trait of the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>—has been lost independently multiple times during the tribe’s evolutionary history. Although uncommonly found in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>, the absence of medioapical projections on the labrum and prementum, a character state based on which <xref ref-type="bibr" rid="B13">Martínez (1964)</xref> erected <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic>, should not be used alone to define higher taxa. The aforementioned phylogenetic analysis provides support that this character state may have been independently derived in many unrelated lineages of geniatine scarabs (<xref ref-type="bibr" rid="B1">Bento 2024</xref>).</p>
      <p>Therefore, based on the synonymy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="margaridae">margaridae</tp:taxon-name-part></tp:taxon-name></italic> with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">L.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="noctivagus">noctivagus</tp:taxon-name-part></tp:taxon-name></italic>, the monotypic genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mimogeniates">Mimogeniates</tp:taxon-name-part></tp:taxon-name></italic> Martínez, 1964 is here synonymized within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Leucothyreus">Leucothyreus</tp:taxon-name-part></tp:taxon-name></italic> Macleay, 1819 (new synonym). As a result of our work, the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> includes 12 genera and 337 species.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgements</title>
      <p>We are grateful to the curators of the collections for providing us with material for this study. We thank Vitor Nardino for providing all images of Figure <xref ref-type="fig" rid="F2">2</xref>. The editor and anonymous reviewers are acknowledged for their valuable contributions to the manuscript.</p>
    </ack>
    <sec sec-type="﻿Additional information" id="SECID0EVZAE">
      <title>﻿Additional information</title>
      <sec sec-type="Conflict of interest" id="SECID0EZZAE">
        <title>Conflict of interest</title>
        <p>The authors have declared that no competing interests exist.</p>
      </sec>
      <sec sec-type="Ethical statement" id="SECID0E5ZAE">
        <title>Ethical statement</title>
        <p>No ethical statement was reported.</p>
      </sec>
      <sec sec-type="Use of AI" id="SECID0ED1AE">
        <title>Use of AI</title>
        <p>No use of AI was reported.</p>
      </sec>
      <sec sec-type="Funding" id="SECID0EI1AE">
        <title>Funding</title>
        <p>MS studied the <named-content content-type="dwc:institutional_code" xlink:title="Museum für Naturkunde der Humboldt-Universität, Berlin, Germany" xlink:href="http://grbio.org/institution/museum-f%C3%BCr-naturkunde-2">MFNB</named-content> collection with financial support of the SYNTHESYS project DE-TAF-6712. This research is part of MB’s doctoral thesis, which was financed in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES, Brazil) through doctoral (Finance Code 001) and sandwich doctorate abroad (PDSE; code 88881.690179/2022-01) scholarships, as well as by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq, Brazil) and the Fundação de Amparo à Pesquisa do Estado do Amazonas (FAPEAM-POSGRAD, Brazil). PCG was funded by CNPq, Process numbers: 309786/2019-3 and 312917/2022-8 PQ2.</p>
      </sec>
      <sec sec-type="Author contributions" id="SECID0ET1AE">
        <title>Author contributions</title>
        <p>MB: Conceptualization, Methodology, Investigation, Resources, Data Curation, Writing of original draft, Writing, Review and Editing, Project administration, Funding Acquisition. MS: Conceptualization, Investigation, Resources, Supervision, Writing, Review and Editing, Funding Acquisition. PG: Data Curation, Review. CRVF: Supervision, Review.</p>
      </sec>
      <sec sec-type="Author ORCIDs" id="SECID0EY1AE">
        <title>Author ORCIDs</title>
        <p>Matheus Bento <ext-link xlink:href="https://orcid.org/0000-0001-9846-9728" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0001-9846-9728</ext-link></p>
        <p>Paschoal Grossi <ext-link xlink:href="https://orcid.org/0000-0001-6601-5967" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0001-6601-5967</ext-link></p>
        <p>Claudio Ruy Vasconcelos da Fonseca <ext-link xlink:href="https://orcid.org/0000-0002-1955-288X" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-1955-288X</ext-link></p>
        <p>Matthias Seidel <ext-link xlink:href="https://orcid.org/0000-0002-4913-8778" ext-link-type="uri" xlink:type="simple">https://orcid.org/0000-0002-4913-8778</ext-link></p>
      </sec>
      <sec sec-type="Data availability" id="SECID0EU2AE">
        <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>
    <ref-list>
      <title>﻿References</title>
      <ref id="B1">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Bento</surname><given-names>M</given-names></name></person-group> (<year>2024</year>) <source>Systematics of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> Burmeister, 1844 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>). Ph.D.</source><publisher-name>thesis</publisher-name>, <publisher-loc>National Institute for Amazonian Research, Manaus, Brazil</publisher-loc>, <size units="page">514 pp</size>.</mixed-citation>
      </ref>
      <ref id="B2">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Beutel</surname><given-names>RG</given-names></name><name name-style="western"><surname>Lawrence</surname><given-names>JF</given-names></name></person-group> (<year>2005</year>) <article-title><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>, morphology.</article-title> In: <person-group><name name-style="western"><surname>Beutel</surname><given-names>RG</given-names></name><name name-style="western"><surname>Leschen</surname><given-names>RAB</given-names></name></person-group> (<role>Eds</role>) <issue-title>Coleoptera, Beetles.</issue-title><source>Volume 1: Morphology and Systematics (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Archostemata</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Adephaga</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Myxophaga</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Polyphaga</tp:taxon-name-part></tp:taxon-name></italic> partim), 2<sup>nd</sup> edition. Walter de Gruyter, Berlin–New York</source>, <fpage>35</fpage>–<lpage>40</lpage>. <ext-link xlink:href="10.1515/9783110904550" ext-link-type="doi" xlink:type="simple">https://doi.org/10.1515/9783110904550</ext-link></mixed-citation>
      </ref>
      <ref id="B3">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Blackwelder</surname><given-names>RE</given-names></name></person-group> (<year>1944</year>) <article-title>Checklist of the coleopterous insects of Mexico, Central America the West Indies, and South America. Parts 1–6.</article-title><source>Bulletin – United States National Museum</source><volume>185</volume>: <fpage>1</fpage>–<lpage>1492</lpage>. <ext-link xlink:href="10.5479/si.03629236.185.i" ext-link-type="doi" xlink:type="simple">https://doi.org/10.5479/si.03629236.185.i</ext-link></mixed-citation>
      </ref>
      <ref id="B4">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Ferreira</surname><given-names>AS</given-names></name><name name-style="western"><surname>Bravo</surname><given-names>F</given-names></name><name name-style="western"><surname>Grossi</surname><given-names>PC</given-names></name><name name-style="western"><surname>Seidel</surname><given-names>M</given-names></name></person-group> (<year>2019</year>) <article-title>Seven new species and new distributional records of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Trizogeniates</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1917 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>) with a key and illustrated checklist of Brazilian species.</article-title><source>Zootaxa</source><volume>4667</volume>(<issue>1</issue>): <fpage>001</fpage>–<lpage>069</lpage>. <ext-link xlink:href="10.11646/zootaxa.4667.1.1" ext-link-type="doi" xlink:type="simple">https://doi.org/10.11646/zootaxa.4667.1.1</ext-link></mixed-citation>
      </ref>
      <ref id="B5">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Ferreira</surname><given-names>AS</given-names></name><name name-style="western"><surname>Grossi</surname><given-names>P</given-names></name><name name-style="western"><surname>Seidel</surname><given-names>M</given-names></name></person-group> (<year>2024</year>) <article-title>Taxonomic revision of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Eunanus</tp:taxon-name-part></tp:taxon-name></italic> Ohaus, 1909 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>), with first records and two new species from Brazil.</article-title><source>Austral Entomology</source><volume>63</volume>(<issue>1</issue>): <fpage>49</fpage>–<lpage>71</lpage>. <ext-link xlink:href="10.1111/aen.12681" ext-link-type="doi" xlink:type="simple">https://doi.org/10.1111/aen.12681</ext-link></mixed-citation>
      </ref>
      <ref id="B6">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Fuhrmann</surname><given-names>J</given-names></name></person-group> (<year>2021</year>) <article-title><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Zagurya</tp:taxon-name-part></tp:taxon-name></italic>, a new genus from Brazil with notes on claw movement in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Macrodactylini</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Melolonthinae</tp:taxon-name-part></tp:taxon-name>).</article-title><source>Zootaxa</source><volume>5016</volume>(<issue>3</issue>): <fpage>395</fpage>–<lpage>406</lpage>. <ext-link xlink:href="10.11646/zootaxa.5016.3.5" ext-link-type="doi" xlink:type="simple">https://doi.org/10.11646/zootaxa.5016.3.5</ext-link></mixed-citation>
      </ref>
      <ref id="B7">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Grossi</surname><given-names>PC</given-names></name><name name-style="western"><surname>Vaz-de-Mello</surname><given-names>FZ</given-names></name></person-group> (<year>2018</year>) <article-title>Revision of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Evanos</tp:taxon-name-part></tp:taxon-name></italic> Laporte de Castelnau, 1840, with the description of a second species (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Scarabaeoidea</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rutelidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>), Annales de la Société Entomologique de France (N.S.</article-title><source>)</source><volume>54</volume>(<issue>6</issue>): <fpage>489</fpage>–<lpage>496</lpage>. <ext-link xlink:href="10.1080/00379271.2018.1514984" ext-link-type="doi" xlink:type="simple">https://doi.org/10.1080/00379271.2018.1514984</ext-link></mixed-citation>
      </ref>
      <ref id="B8">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Jameson</surname><given-names>ML</given-names></name></person-group> (<year>2008</year>) <article-title>Review of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Microchilus</tp:taxon-name-part></tp:taxon-name></italic> Blanchard (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>).</article-title><source><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Insecta</tp:taxon-name-part></tp:taxon-name> Mundi</source><volume>0025</volume>: <fpage>1</fpage>–<lpage>14</lpage>.</mixed-citation>
      </ref>
      <ref id="B9">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Jameson</surname><given-names>ML</given-names></name><name name-style="western"><surname>Hawkins</surname><given-names>SJ</given-names></name></person-group> (<year>2005</year>) <article-title>Synopsis of the genera of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>) with an annotated catalog of species.</article-title><source>Zootaxa</source><volume>874</volume>(<issue>1</issue>): <fpage>1</fpage>–<lpage>58</lpage>. <ext-link xlink:href="10.11646/zootaxa.874.1.1" ext-link-type="doi" xlink:type="simple">https://doi.org/10.11646/zootaxa.874.1.1</ext-link></mixed-citation>
      </ref>
      <ref id="B10">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Krajcik</surname><given-names>M</given-names></name></person-group> (<year>2007</year>) <article-title>Checklist of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Scarabaeoidea</tp:taxon-name-part></tp:taxon-name> of the World. 2. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>. Animma.</article-title><source>x</source><volume>4</volume>: <fpage>1</fpage>–<lpage>273</lpage>.</mixed-citation>
      </ref>
      <ref id="B11">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Krell</surname><given-names>FT</given-names></name></person-group> (<year>1996</year>) <article-title>Die Kopulationsorgane des Maikäfers <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Melolontha</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">melolontha</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Insecta</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>). Ein Beitrag zur vergleichenden und funktionellen Anatomie der ektodermalen Genitalien der <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>. Stuttgarter Beiträge zur Naturkunde.</article-title><source>Serie A, Biologie</source><volume>537</volume>: <fpage>1</fpage>–<lpage>101</lpage>.</mixed-citation>
      </ref>
      <ref id="B12">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Machatschke</surname><given-names>JW</given-names></name></person-group> (<year>1972</year>) <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Scarabaeoidea</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Melolonthidae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>. Coleopterorum Catalogus (Supplementa 66): 1–361.</mixed-citation>
      </ref>
      <ref id="B13">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Martínez</surname><given-names>M</given-names></name></person-group> (<year>1964</year>) <article-title><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name> nuevos o poco conocidos (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>, Scarabaiedae, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>).</article-title><source>Comunicaciones del Museo Artgentino de Ciencias Naturales Bernardino Rivadavia</source><volume>1</volume>: <fpage>1</fpage>–<lpage>18</lpage>.</mixed-citation>
      </ref>
      <ref id="B14">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Ohaus</surname><given-names>F</given-names></name></person-group> (<year>1909</year>) <article-title>Neue <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name> lamellicornia aus Argentinien.</article-title><source>Deutsche Entomologische Zeitschrift</source><volume>1909</volume>: <fpage>425</fpage>–<lpage>447</lpage>.</mixed-citation>
      </ref>
      <ref id="B15">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Ohaus</surname><given-names>F</given-names></name></person-group> (<year>1917</year>) <article-title>Neue Geniatinen (Col. lamell. Rutelin.).</article-title><source>Stettiner Entomologische Zeitung</source><volume>78</volume>: <fpage>3</fpage>–<lpage>53</lpage>.</mixed-citation>
      </ref>
      <ref id="B16">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Shorthouse</surname><given-names>DP</given-names></name></person-group> (<year>2010</year>) SimpleMappr, an online tool to produce publication-quality point maps. <ext-link xlink:href="http://www.simplemappr.net" ext-link-type="uri" xlink:type="simple">http://www.simplemappr.net</ext-link> [Accessed on 10 Jul 2022]</mixed-citation>
      </ref>
      <ref id="B17">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Villatoro</surname><given-names>K</given-names></name></person-group> (<year>2002</year>) <article-title>Revision of the Neotropical genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Trizogeniates</tp:taxon-name-part></tp:taxon-name></italic> Ohaus (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Coleoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scarabaeidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rutelinae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Geniatini</tp:taxon-name-part></tp:taxon-name>).</article-title><source>Entomotrópica</source><volume>17</volume>(<issue>3</issue>): <fpage>225</fpage>–<lpage>294</lpage>.</mixed-citation>
      </ref>
    </ref-list>
  </back>
</article>
