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<document id="A10B52ECDCF74DD3B6B11A2428C9A7C9" ID-CLB-Dataset="2423" ID-DOI="10.11646/zootaxa.5093.1.3" ID-GBIF-Dataset="701b994e-d6dd-4e39-923d-ad55d113d881" ID-ISSN="1175-5326" ID-Zenodo-Dep="5901043" ID-ZooBank="E90AE625-98EA-41D5-9C8C-CEA4B877E687" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1643101911126" checkinUser="plazi" docAuthor="Duarte, Paulo R. M., Grossi, Paschoal C. &amp; Dupuis, Fabien" docDate="2022" docId="03B687B92D7FFFDA76BFFF09FE14A4E1" docLanguage="en" docName="zootaxa.5093.1.3.pdf" docOrigin="Zootaxa 5093 (1)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Bothynus villiersi Endrodi 1968" docType="treatment" docVersion="6" lastPageNumber="52" masterDocId="FF8FFFC12D7DFFD97628FF9EFFA1A041" masterDocTitle="A taxonomic revision of the Bothynus villiersi Endrödi, 1968 species group (Coleoptera: Scarabaeidae: Dynastinae)" masterLastPageNumber="66" masterPageNumber="49" pageNumber="51" updateTime="1699304170453" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="734341E0FC6D9EEF4CEE68D944ABDFE7">A taxonomic revision of the Bothynus villiersi Endrödi, 1968 species group (Coleoptera: Scarabaeidae: Dynastinae)</mods:title>
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<mods:namePart id="58945047B52D7707251E3C0A45AE2A50">Duarte, Paulo R. M.</mods:namePart>
<mods:affiliation id="D51A4F8AE3E808965425D57300D8C8DD">Departamento de Fitossanidade, Universidade Federal Rural de Pernambuco, Rua Manoel de Medeiros, s / n - Dois Irmãos, Recife, Pernambuco, Brazil</mods:affiliation>
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<mods:namePart id="3343D3854D4AE0E0C9D49C9B0CBB48EB">Grossi, Paschoal C.</mods:namePart>
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<mods:affiliation id="670965CE661CBF73731ED81BFA05262F">Departamento de Fitossanidade, Universidade Federal Rural de Pernambuco, Rua Manoel de Medeiros, s / n - Dois Irmãos, Recife, Pernambuco, Brazil &amp; http: // orcid. org / 0000 - 0001 - 6601 - 5967</mods:affiliation>
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<mods:namePart id="D1ECD1E2057C00C4106E4834E50685CA">Dupuis, Fabien</mods:namePart>
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<mods:title id="15BFBCBE92B74D8F72A8ADB2134C7391">Zootaxa</mods:title>
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<mods:date id="907F81EF828ADE3D2C1EEFC7F4535675">2022</mods:date>
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<mods:number id="D83C6D3C7271F5C634E1C4E33C9F78A5">5093</mods:number>
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<treatment id="03B687B92D7FFFDA76BFFF09FE14A4E1" ID-DOI="http://doi.org/10.5281/zenodo.5901047" ID-GBIF-Taxon="192601203" ID-Zenodo-Dep="5901047" LSID="urn:lsid:plazi:treatment:03B687B92D7FFFDA76BFFF09FE14A4E1" httpUri="http://treatment.plazi.org/id/03B687B92D7FFFDA76BFFF09FE14A4E1" lastPageId="3" lastPageNumber="52" pageId="2" pageNumber="51">
<subSubSection id="C30565242D7FFFDB76BFFF09FDBAA0F0" box="[151,539,151,178]" pageId="2" pageNumber="51" type="nomenclature">
<paragraph id="8BA036AF2D7FFFDB76BFFF09FDBAA0F0" blockId="2.[151,895,151,214]" box="[151,539,151,178]" pageId="2" pageNumber="51">
<heading id="D0E881C32D7FFFDB76BFFF09FDBAA0F0" bold="true" box="[151,539,151,178]" fontSize="11" level="1" pageId="2" pageNumber="51" reason="1">
<emphasis id="B96BEABD2D7FFFDB76BFFF09FDBAA0F0" bold="true" box="[151,539,151,178]" pageId="2" pageNumber="51">
<taxonomicName id="4C1F4D2C2D7FFFDB76BFFF09FECDA0F3" authorityName="Endrodi" authorityYear="1968" box="[151,364,151,178]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="villiersi">
<emphasis id="B96BEABD2D7FFFDB76BFFF09FECDA0F3" bold="true" box="[151,364,151,178]" italics="true" pageId="2" pageNumber="51">Bothynus villiersi</emphasis>
</taxonomicName>
species group
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C30565242D7FFFDB76BFFF25FCD2A124" pageId="2" pageNumber="51" type="description">
<paragraph id="8BA036AF2D7FFFDB76BFFF25FCDEA094" blockId="2.[151,895,151,214]" box="[151,895,187,214]" pageId="2" pageNumber="51">
(
<figureCitation id="13242A2A2D7FFFDB76B7FF25FE87A094" box="[159,294,187,213]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="2" pageNumber="51">Figs. 1A–D</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB771AFF25FEDBA094" box="[306,378,187,213]" captionStart="FIGURE 2" captionStartId="4.[151,249,1968,1992]" captionTargetBox="[153,1428,504,1932]" captionTargetId="figure-148@4.[151,1435,489,1945]" captionTargetPageId="4" captionText="FIGURE 2. Holotype dorsal habitus with their respective labels on the right side: A, Bothynus villiersi Endrödi, 1968; B, Bothynus horridus Endrödi, 1968; C, Bothynus alvarengai Martínez, 1983; D, Bothynus spinophallicus (Ratcliffe &amp; Cave, 2021)." figureDoi="http://doi.org/10.5281/zenodo.5901049" httpUri="https://zenodo.org/record/5901049/files/figure.png" pageId="2" pageNumber="51">2A–D</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB77ADFF25FE6BA094" box="[389,458,187,213]" captionStart="FIGURE 3" captionStartId="6.[151,250,1883,1907]" captionTargetBox="[151,1432,277,1856]" captionTargetId="figure-60@6.[151,1435,273,1860]" captionTargetPageId="6" captionText="FIGURE 3. Male dorsal habitus: A, Bothynus villiersi Endrödi, 1968; B, Bothynus horridus Endrödi, 1968; C, Bothynus alvarengai Martínez, 1983; D, Bothynus spinophallicus (Ratcliffe &amp; Cave, 2021); E, Bothynus belemensis new species (holotype)." figureDoi="http://doi.org/10.5281/zenodo.5901053" httpUri="https://zenodo.org/record/5901053/files/figure.png" pageId="2" pageNumber="51">3A–E</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB77FDFF25FDBCA094" box="[469,541,187,213]" captionStart="FIGURE 4" captionStartId="8.[151,250,1044,1068]" captionTargetBox="[158,1428,182,1020]" captionTargetId="figure-27@8.[158,1429,181,1021]" captionTargetPageId="8" captionText="FIGURE 4. Pronotal tubercle: A, Bothynus alvarengai Martínez, 1983; B, Bothynus horridus Endrödi, 1968. Prosternal process: C, Bothynus villiersi Endrödi, 1968; D, Bothynus belemensis new species. Arrows indicate tubercle bilobed in A, tubercle conical in B, prosternal process apex conical in C, and prosternal process apex rounded in D." figureDoi="http://doi.org/10.5281/zenodo.5901055" httpUri="https://zenodo.org/record/5901055/files/figure.png" pageId="2" pageNumber="51">4A–D</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB7401FF25FDD1A097" box="[553,624,187,214]" captionStart="FIGURE 5" captionStartId="10.[151,250,1894,1918]" captionTargetBox="[155,1416,609,1858]" captionTargetId="figure-206@10.[151,1435,597,1870]" captionTargetPageId="10" captionText="FIGURE 5. Spiculum gastrale: A, Bothynus villiersi Endrödi, 1968; B, Bothynus horridus Endrödi, 1968; C, Bothynus alvarengai Martínez, 1983; D, Bothynus belemensis new species. hs = hemisternite." figureDoi="http://doi.org/10.5281/zenodo.5901059" httpUri="https://zenodo.org/record/5901059/files/figure.png" pageId="2" pageNumber="51">5A–D</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB7454FF25FD1BA094" box="[636,698,187,213]" captionStart="FIGURE 6" captionStartId="11.[151,250,1792,1816]" captionTargetBox="[168,1416,187,1763]" captionTargetId="figure-20@11.[151,1435,181,1768]" captionTargetPageId="11" captionText="FIGURE 6. Aedeagus in lateral view and parameres in dorsal view, respectively: A–B, Bothynus villiersi Endrödi, 1968; C–D, Bothynus horridus Endrödi, 1968; E–F, Bothynus alvarengai Martínez, 1983; G–H, Bothynus belemensis new species. lmp = left medial process, rbp = right basal process, rmp = right medial process. Arrow indicating elongated parameres apex in B." figureDoi="http://doi.org/10.5281/zenodo.5901063" httpUri="https://zenodo.org/record/5901063/files/figure.png" pageId="2" pageNumber="51">6A–J</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB74EEFF25FCACA094" box="[710,781,187,213]" captionStart="FIGURE 7" captionStartId="12.[151,250,1960,1984]" captionTargetBox="[170,1412,193,1925]" captionTargetId="figure-27@12.[157,1430,181,1936]" captionTargetPageId="12" captionText="FIGURE 7. Female dorsal habitus: A, Bothynus villiersi Endrödi, 1968; B, Bothynus horridus Endrödi, 1968; C, Bothynus alvarengai Martínez, 1983; D, Bothynus belemensis new species." figureDoi="http://doi.org/10.5281/zenodo.5901065" httpUri="https://zenodo.org/record/5901065/files/figure.png" pageId="2" pageNumber="51">7A–D</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB7531FF25FCFDA094" box="[793,860,187,213]" captionStart="FIGURE 8" captionStartId="13.[151,250,1808,1832]" captionTargetBox="[157,1430,731,1777]" captionTargetId="figure-202@13.[156,1431,730,1784]" captionTargetPageId="13" captionText="FIGURE 8. Female, detail of the anterior pronotal margin: A, Bothynus villiersi Endrödi, 1968; B, Bothynus belemensis new species. Female, internal view of the tergite 8: C, Bothynus villiersi Endrödi, 1968; D, Bothynus horridus Endrödi, 1968; E, Bothynus alvarengai Martínez, 1983; F, Bothynus belemensis. Arrows indicate arched margin in A, margin sinuous in B, and furrow in C. ip = internal plate." figureDoi="http://doi.org/10.5281/zenodo.5901069" httpUri="https://zenodo.org/record/5901069/files/figure.png" pageId="2" pageNumber="51">8A–F</figureCitation>
;
<figureCitation id="13242A2A2D7FFFDB754FFF25FCD7A094" box="[871,886,187,213]" captionStart="FIGURE 9" captionStartId="14.[151,250,1761,1785]" captionTargetBox="[151,1435,453,1737]" captionTargetId="figure-171@14.[151,1435,453,1737]" captionTargetPageId="14" captionText="FIGURE 9. Geographic distribution of the Bothynus villiersi species group. The star inscribed by a red circle indicates a possible mislabeled record." figureDoi="http://doi.org/10.5281/zenodo.5901071" httpUri="https://zenodo.org/record/5901071/files/figure.png" pageId="2" pageNumber="51">9</figureCitation>
)
</paragraph>
<paragraph id="8BA036AF2D7FFFDB76BFFE9DFCD2A124" blockId="2.[151,1437,259,538]" pageId="2" pageNumber="51">
<emphasis id="B96BEABD2D7FFFDB76BFFE9DFE8EA15C" bold="true" box="[151,303,259,285]" pageId="2" pageNumber="51">Composition</emphasis>
. The
<taxonomicName id="4C1F4D2C2D7FFFDB7744FE9DFD8EA15C" authorityName="Endrodi" authorityYear="1968" box="[364,559,259,285]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="villiersi">
<emphasis id="B96BEABD2D7FFFDB7744FE9DFD8EA15C" box="[364,559,259,285]" italics="true" pageId="2" pageNumber="51">Bothynus villiersi</emphasis>
</taxonomicName>
species group includes five species:
<taxonomicName id="4C1F4D2C2D7FFFDB75E9FE9DFA8BA15C" authority="Endrodi, 1968" authorityName="Endrodi" authorityYear="1968" box="[961,1322,259,285]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="villiersi">
<emphasis id="B96BEABD2D7FFFDB75E9FE9DFB25A15C" box="[961,1156,259,285]" italics="true" pageId="2" pageNumber="51">Bothynus villiersi</emphasis>
<bibRefCitation id="EF8E4B5E2D7FFFDB72A1FE9DFA8BA15C" author="Endrodi, S." box="[1161,1322,259,285]" pageId="2" pageNumber="51" pagination="161 - 177" refId="ref10112" refString="Endrodi, S. (1968). Neue Arten der Pentodontini. Folia Entomologica Hungarica, Series Nova, 21 (12), 161 - 177." type="journal article" year="1968">Endrödi, 1968</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C1F4D2C2D7FFFDB731CFE9DFE05A100" authority="Endrodi, 1968" authorityName="Endrodi" authorityYear="1968" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="horridus">
<emphasis id="B96BEABD2D7FFFDB731CFE9DFF56A100" italics="true" pageId="2" pageNumber="51">Bothynus horridus</emphasis>
<bibRefCitation id="EF8E4B5E2D7FFFDB7728FEB9FE05A100" author="Endrodi, S." box="[256,420,295,321]" pageId="2" pageNumber="51" pagination="161 - 177" refId="ref10112" refString="Endrodi, S. (1968). Neue Arten der Pentodontini. Folia Entomologica Hungarica, Series Nova, 21 (12), 161 - 177." type="journal article" year="1968">Endrödi, 1968</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C1F4D2C2D7FFFDB7799FEB9FCF0A100" authority="Martinez, 1983" authorityName="Martinez" authorityYear="1983" box="[433,849,295,321]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="alvarengai">
<emphasis id="B96BEABD2D7FFFDB7799FEB9FD38A100" box="[433,665,295,321]" italics="true" pageId="2" pageNumber="51">Bothynus alvarengai</emphasis>
<bibRefCitation id="EF8E4B5E2D7FFFDB748AFEB9FCF0A100" author="Martinez, A." box="[674,849,295,321]" pageId="2" pageNumber="51" pagination="297 - 300" refId="ref10673" refString="Martinez, A. (1983) Nueva especie de Bothynus de Brasil (Coleoptera, Scarabaeidae, Dynastinae, Pentodontini). Revista Brasileira de Entomologia, 27, 297 - 300." type="journal article" year="1983">Martínez, 1983</bibRefCitation>
</taxonomicName>
;
<taxonomicName id="4C1F4D2C2D7FFFDB7576FEB9FA36A100" authority="(Ratcliffe &amp; Cave, 2021)" authorityName="Duarte &amp; Grossi &amp; Dupuis" authorityYear="2022" baseAuthorityName="Ratcliffe &amp; Cave" baseAuthorityYear="2021" box="[862,1431,295,321]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="spinophallicus">
<emphasis id="B96BEABD2D7FFFDB7576FEB9FBD0A100" box="[862,1137,295,321]" italics="true" pageId="2" pageNumber="51">Bothynus spinophallicus</emphasis>
(Ratcliffe &amp; Cave, 2021)
</taxonomicName>
; and
<taxonomicName id="4C1F4D2C2D7FFFDB76EFFED5FD7BA124" authority="Duarte, Grossi, &amp; Dupuis" authorityName="Duarte, Grossi, &amp; Dupuis" authorityYear="2022" box="[199,730,331,358]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="belemensis" status="sp. nov.">
<emphasis id="B96BEABD2D7FFFDB76EFFED5FE10A124" box="[199,433,331,357]" italics="true" pageId="2" pageNumber="51">Bothynus belemensis</emphasis>
Duarte, Grossi, &amp; Dupuis
</taxonomicName>
,
<taxonomicNameLabel id="A25857C62D7FFFDB74CDFED2FCCFA124" box="[741,878,331,358]" pageId="2" pageNumber="51" rank="species">new species</taxonomicNameLabel>
.
</paragraph>
</subSubSection>
<subSubSection id="C30565242D7FFFDB76EFFEF1FD48A25B" pageId="2" pageNumber="51" type="diagnosis">
<paragraph id="8BA036AF2D7FFFDB76EFFEF1FD48A25B" blockId="2.[151,1437,259,538]" pageId="2" pageNumber="51">
<emphasis id="B96BEABD2D7FFFDB76EFFEF1FE9BA1C8" bold="true" box="[199,314,367,393]" pageId="2" pageNumber="51">Diagnosis</emphasis>
. The
<taxonomicName id="4C1F4D2C2D7FFFDB7756FEF1FDE4A1C8" authorityName="Endrodi" authorityYear="1968" box="[382,581,367,393]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="villiersi">
<emphasis id="B96BEABD2D7FFFDB7756FEF1FDE4A1C8" box="[382,581,367,393]" italics="true" pageId="2" pageNumber="51">Bothynus villiersi</emphasis>
</taxonomicName>
species group has an exclusive set of characters within the genus: clypeus obliquely inclined downward in lateral view (
<figureCitation id="13242A2A2D7FFFDB748CFE0DFCA1A1EC" box="[676,768,403,429]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="2" pageNumber="51">Fig. 1A</figureCitation>
); frontoclypeal suture complete bearing a transverse ridge reaching the lateral margins of head (
<figureCitation id="13242A2A2D7FFFDB7412FE29FD30A190" box="[570,657,439,465]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="2" pageNumber="51">Fig. 1B</figureCitation>
); maxillary palpomere 1 short (three times shorter than palpomere 2) (
<figureCitation id="13242A2A2D7FFFDB7688FE45FF54A1B4" box="[160,245,475,501]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="2" pageNumber="51">Fig. 1C</figureCitation>
); labium with a conical process on disc in lateral view (
<figureCitation id="13242A2A2D7FFFDB7548FE45FC16A1B4" box="[864,951,475,501]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="2" pageNumber="51">Fig. 1D</figureCitation>
); parameres asymmetrical, provided of 1–2 process on inner lateral margins (
<figureCitation id="13242A2A2D7FFFDB7424FE61FD7AA25B" box="[524,731,511,538]" captionStart="FIGURE 6" captionStartId="11.[151,250,1792,1816]" captionTargetBox="[168,1416,187,1763]" captionTargetId="figure-20@11.[151,1435,181,1768]" captionTargetPageId="11" captionText="FIGURE 6. Aedeagus in lateral view and parameres in dorsal view, respectively: A–B, Bothynus villiersi Endrödi, 1968; C–D, Bothynus horridus Endrödi, 1968; E–F, Bothynus alvarengai Martínez, 1983; G–H, Bothynus belemensis new species. lmp = left medial process, rbp = right basal process, rmp = right medial process. Arrow indicating elongated parameres apex in B." figureDoi="http://doi.org/10.5281/zenodo.5901063" httpUri="https://zenodo.org/record/5901063/files/figure.png" pageId="2" pageNumber="51">Fig. 6B, D, F, H, J</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="DF6066272D7FFFDB76BFF886FF11A7DD" ID-DOI="http://doi.org/10.5281/zenodo.5901045" ID-Zenodo-Dep="5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="2" pageNumber="51" startId="2.[151,250,1816,1840]" targetBox="[165,1429,573,1780]" targetPageId="2">
<paragraph id="8BA036AF2D7FFFDB76BFF886FF11A7DD" blockId="2.[151,1437,1816,1948]" pageId="2" pageNumber="51">
<emphasis id="B96BEABD2D7FFFDB76BFF886FEADA771" bold="true" box="[151,268,1816,1840]" pageId="2" pageNumber="51">FIGURE 1</emphasis>
. Morphological characters of the
<taxonomicName id="4C1F4D2C2D7FFFDB7448F886FCB0A771" authorityName="Endrodi" authorityYear="1968" box="[608,785,1816,1840]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="51" phylum="Arthropoda" rank="species" species="villiersi">
<emphasis id="B96BEABD2D7FFFDB7448F886FCB0A771" box="[608,785,1816,1840]" italics="true" pageId="2" pageNumber="51">Bothynus villiersi</emphasis>
</taxonomicName>
species group:
<emphasis id="B96BEABD2D7FFFDB7586F886FC61A771" bold="true" box="[942,960,1816,1840]" pageId="2" pageNumber="51">A</emphasis>
, head in lateral view (mouthparts removed);
<emphasis id="B96BEABD2D7FFFDB73ADF886FA37A771" bold="true" box="[1413,1430,1816,1840]" pageId="2" pageNumber="51">B</emphasis>
, head in frontal view;
<emphasis id="B96BEABD2D7FFFDB775AF8A2FE25A715" bold="true" box="[370,388,1852,1876]" pageId="2" pageNumber="51">C</emphasis>
, right maxilla in ventral view;
<emphasis id="B96BEABD2D7FFFDB74EAF8A2FD75A715" bold="true" box="[706,724,1852,1876]" pageId="2" pageNumber="51">D</emphasis>
, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D.
</paragraph>
</caption>
<subSubSection id="C30565242D7EFFDA76EFFF08FE14A4E1" pageId="3" pageNumber="52" type="description">
<paragraph id="8BA036AF2D7EFFDA76EFFF08FB93A455" blockId="3.[151,1437,150,1184]" pageId="3" pageNumber="52">
<emphasis id="B96BEABD2D7EFFDA76EFFF08FEF0A0F1" bold="true" box="[199,337,150,176]" pageId="3" pageNumber="52">Description</emphasis>
. The
<taxonomicName id="4C1F4D2C2D7EFFDA77B1FF09FDB3A0F0" authorityName="Endrodi" authorityYear="1968" box="[409,530,151,177]" class="Insecta" family="Dynastidae" genus="Bothynus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="52" phylum="Arthropoda" rank="species" species="villiersi">
<emphasis id="B96BEABD2D7EFFDA77B1FF09FDB3A0F0" box="[409,530,151,177]" italics="true" pageId="3" pageNumber="52">B. villiersi</emphasis>
</taxonomicName>
species group is characterized by oval-shaped species in dorsal view, convex dorsally in lateral view, ranging from reddish brown to dark reddish coloration.
<emphasis id="B96BEABD2D7EFFDA7242FF27FB0BA092" bold="true" box="[1130,1194,185,211]" pageId="3" pageNumber="52">Head</emphasis>
: Clypeus triangular, obliquely inclined downward in lateral view (
<figureCitation id="13242A2A2D7EFFDA7484FF42FCA9A0B7" box="[684,776,220,246]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="3" pageNumber="52">Fig. 1A</figureCitation>
), with 2 raised tapered teeth on anterior margin; surface mostly transversely rugose, glabrous. Frontoclypeal suture with a raised, transverse, complete ridge that reaches the lateral margins of head (
<figureCitation id="13242A2A2D7EFFDA778DFEBFFE5DA17A" box="[421,508,289,315]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="3" pageNumber="52">Fig. 1B</figureCitation>
). Frons glabrous, transversely rugose or rugopunctate except by a posterior smooth area. Ocular canthus subtriangular, transverse, with scarce setae scattered along of anterior margin.
<emphasis id="B96BEABD2D7EFFDA732DFEDAFA35A11F" bold="true" box="[1285,1428,324,350]" pageId="3" pageNumber="52">Mouthparts</emphasis>
: Mandibles with 3 marginal teeth exposed laterally to clypeus; apical and medial teeth mostly triangular, basal tooth lobed. Maxillary galea shorter than lacinia, usually devoid of teeth, rarely provided with a small tooth on apex. Maxillary palpomere 1 short, about 2 times shorter than subsequent palpomere, hidden under lateral stipe (
<figureCitation id="13242A2A2D7EFFDA7688FE51FF59A1A8" box="[160,248,463,489]" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="3" pageNumber="52">Fig. 1C</figureCitation>
); palpomere 4 clavate. Labium subtriangular, with a conical process on disc visible in lateral view (
<figureCitation id="13242A2A2D7EFFDA7347FE51FF1AA24A" captionStart="FIGURE 1" captionStartId="2.[151,250,1816,1840]" captionTargetBox="[165,1429,573,1780]" captionTargetId="figure-158@2.[157,1430,561,1792]" captionTargetPageId="2" captionText="FIGURE 1. Morphological characters of the Bothynus villiersi species group: A, head in lateral view (mouthparts removed); B, head in frontal view; C, right maxilla in ventral view; D, head, prothoracic legs and pronotum in lateral view. Arrows indicate oblique downward clypeus in A, complete frontoclypeal ridge in B, palpomere 1 short in C, and conical process of mentum in D." figureDoi="http://doi.org/10.5281/zenodo.5901045" httpUri="https://zenodo.org/record/5901045/files/figure.png" pageId="3" pageNumber="52">Fig. 1D</figureCitation>
), surrounded by scarce setae.
<emphasis id="B96BEABD2D7EFFDA740AFE6FFD34A24A" bold="true" box="[546,661,497,523]" pageId="3" pageNumber="52">Antennae</emphasis>
: Comprised of 10 antennomeres; club oval, subequal in length to antennomeres 2–7 combined; lamellae 8–10 decreasing in size.
<emphasis id="B96BEABD2D7EFFDA7540FD8AFC4BA26F" bold="true" box="[872,1002,532,558]" pageId="3" pageNumber="52">Pronotum:</emphasis>
Bearing a tubercle raised at middle of anterior margin; anterior middle area concave; posterior area strongly convex in lateral view.
<emphasis id="B96BEABD2D7EFFDA728FFDA9FB5CA210" bold="true" box="[1191,1277,567,593]" pageId="3" pageNumber="52">Elytra:</emphasis>
Striae usually well marked; punctures ocellated, moderate to dense, irregularly scattered on interstriae; striae bearing punctures separated about 1–2 puncture diameters.
<emphasis id="B96BEABD2D7EFFDA7472FDE2FCAEA2D7" bold="true" box="[602,783,636,662]" pageId="3" pageNumber="52">Scutellar plate:</emphasis>
Triangular, smooth.
<emphasis id="B96BEABD2D7EFFDA75DBFDE2FB8AA2D7" bold="true" box="[1011,1067,636,662]" pageId="3" pageNumber="52">Legs</emphasis>
: Protarsal claws strongly curved; protarsomere 5 elongate, subequal in length to tarsomeres 3–4 combined; protarsomeres 1–4 subequal in length. Protibia with 3 teeth on outer margin; margin slightly sinuous just behind the basal tooth. Meso- and metatarsomeres 1–4 decreasing gradually in size. Mesotibia usually with 2 weakly crenulate carinae on outer surface; basal carina short, transversely projected; medial carina long, obliquely projected; apex truncate; carinae and apex covered with stout spinule-like setae. Mesofemur subrectangular. Metatibia triangular, wide apically. Metafemur oval-shaped, wider compared to mesofemur.
<emphasis id="B96BEABD2D7EFFDA7424FCD2FDFDA327" bold="true" box="[524,604,844,870]" pageId="3" pageNumber="52">Venter</emphasis>
: Prosternal process columnar, subcylindrical, raised between procoxae. Mesoventrite nearly completely rugopunctate, densely setose. Metaventrite coarsely rugopunctate, densely setose on sides, finely punctate and glabrous on disc.
<emphasis id="B96BEABD2D7EFFDA74B1FC0FFCAFA3EA" bold="true" box="[665,782,913,939]" pageId="3" pageNumber="52">Abdomen</emphasis>
: Tergite 7 with two paired rows of striae confined on discal area; striae well marked anteriorly, becoming finely marked posteriorly. Tergite 8 strongly convex lateral view; surface glabrous, rugose on sides and on a transverse area near to anterior margin, disc finely punctate.
<emphasis id="B96BEABD2D7EFFDA7309FC48FA35A3B1" bold="true" box="[1313,1428,982,1008]" pageId="3" pageNumber="52">Aedeagus</emphasis>
: Parameres asymmetrical, with 1–2 process on inner margins (
<figureCitation id="13242A2A2D7EFFDA7561FC67FB85A455" box="[841,1060,1017,1044]" captionStart="FIGURE 6" captionStartId="11.[151,250,1792,1816]" captionTargetBox="[168,1416,187,1763]" captionTargetId="figure-20@11.[151,1435,181,1768]" captionTargetPageId="11" captionText="FIGURE 6. Aedeagus in lateral view and parameres in dorsal view, respectively: A–B, Bothynus villiersi Endrödi, 1968; C–D, Bothynus horridus Endrödi, 1968; E–F, Bothynus alvarengai Martínez, 1983; G–H, Bothynus belemensis new species. lmp = left medial process, rbp = right basal process, rmp = right medial process. Arrow indicating elongated parameres apex in B." figureDoi="http://doi.org/10.5281/zenodo.5901063" httpUri="https://zenodo.org/record/5901063/files/figure.png" pageId="3" pageNumber="52">Figs. 6B, D, F, H, J</figureCitation>
).
</paragraph>
<paragraph id="8BA036AF2D7EFFDA76EFFB82FE14A4E1" blockId="3.[151,1437,150,1184]" pageId="3" pageNumber="52">
<emphasis id="B96BEABD2D7EFFDA76EFFB82FE13A477" bold="true" box="[199,434,1052,1078]" pageId="3" pageNumber="52">Sexual dimorphism</emphasis>
. Females are similar to males, except by the pronotum weakly convex in lateral view, devoid of tubercle and concavity absent or shallow; tergite 8 flattened or weakly convex in lateral view, with an internalized plate on posterior margin (
<figureCitation id="13242A2A2D7EFFDA7467FBFFFD67A43A" box="[591,710,1121,1147]" captionStart="FIGURE 8" captionStartId="13.[151,250,1808,1832]" captionTargetBox="[157,1430,731,1777]" captionTargetId="figure-202@13.[156,1431,730,1784]" captionTargetPageId="13" captionText="FIGURE 8. Female, detail of the anterior pronotal margin: A, Bothynus villiersi Endrödi, 1968; B, Bothynus belemensis new species. Female, internal view of the tergite 8: C, Bothynus villiersi Endrödi, 1968; D, Bothynus horridus Endrödi, 1968; E, Bothynus alvarengai Martínez, 1983; F, Bothynus belemensis. Arrows indicate arched margin in A, margin sinuous in B, and furrow in C. ip = internal plate." figureDoi="http://doi.org/10.5281/zenodo.5901069" httpUri="https://zenodo.org/record/5901069/files/figure.png" pageId="3" pageNumber="52">Fig. 8C–F</figureCitation>
); ventrite 6 triangular, complete on posterior margin and surface completely rugopunctate.
</paragraph>
</subSubSection>
</treatment>
</document>
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