Add updates up until 2025-04-15 19:55:58
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<document id="BFFF4844DC1E0C2CC5613F863B2D36E0" ID-DOI="10.11646/zootaxa.5604.4.1" ID-ISSN="1175-5326" ID-Zenodo-Dep="15213967" ID-ZooBank="6A95109D-6F33-4DE7-9D47-6A722DD26918" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="carolina" IM.materialsCitations_approvedBy="carolina" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch,operationResults" IM.taxonomicNames_approvedBy="carolina" IM.treatments_approvedBy="carolina" checkinTime="1744660517245" checkinUser="plazi" docAuthor="Kohlmann, Bert & Solís, Ángel" docDate="2025" docId="03AA23621478FF8DFF28AADCFC71A917" docLanguage="en" docName="zootaxa.5604.4.1.pdf" docOrigin="Zootaxa 5604 (4)" docSource="https://doi.org/10.11646/zootaxa.5604.4.1" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Onthophagus marginatus" docType="treatment" docVersion="1" lastPageNumber="428" masterDocId="FF935B1A1463FF96FFEFAE2EFF91AE3E" masterDocTitle="A review of the species groups of the Western Hemisphere Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) using COI barcoding and gene trees" masterLastPageNumber="447" masterPageNumber="401" pageNumber="428" updateTime="1744746916744" updateUser="carolina" zenodo-license-document="CLOSED">
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<mods:titleInfo id="8F57A2AD9E9792B513CC55A06F059D14">
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<mods:title id="DDC15E8AB80FCF86F432C0A7A4A0A11C">A review of the species groups of the Western Hemisphere Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) using COI barcoding and gene trees</mods:title>
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<mods:namePart id="1E5245B8415B2B853F626CAC81A0454D">Kohlmann, Bert</mods:namePart>
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<mods:affiliation id="BC06046392EF9CF9FBE7BB1F7C86B13A">BioAlfa Barcoding Project Santo Domingo de Heredia, Costa Rica.</mods:affiliation>
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<mods:nameIdentifier id="E91BA51FA0D3BF1FA505B7551CC78168" type="email">bkohlmann64@gmail.com</mods:nameIdentifier>
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<mods:namePart id="7E6C1D433E6B05C47526ED0E5E2D5DBA">Solís, Ángel</mods:namePart>
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<mods:affiliation id="777FB8648ED5AC5AB102ACD170FCAB5D">BioAlfa Barcoding Project Santo Domingo de Heredia, Costa Rica.</mods:affiliation>
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<mods:nameIdentifier id="55E2F96FDDE31E13EC57805507B8A601" type="email">bkohlmann64@gmail.com</mods:nameIdentifier>
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<mods:title id="E5A05F440EA670516E602046BFA72FA8">Zootaxa</mods:title>
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<mods:date id="F741AB3632A313413FBA903FC86C9EA0">2025</mods:date>
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<mods:number id="F1CFA104256C309A412867A7DD7B7CC3">2025-03-17</mods:number>
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<treatment id="03AA23621478FF8DFF28AADCFC71A917" LSID="urn:lsid:plazi:treatment:03AA23621478FF8DFF28AADCFC71A917" httpUri="http://treatment.plazi.org/id/03AA23621478FF8DFF28AADCFC71A917" lastPageNumber="428" pageId="27" pageNumber="428">
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<subSubSection id="C319C1FF1478FF8DFF28AADCFD77AB33" box="[199,742,1266,1293]" pageId="27" pageNumber="428" type="nomenclature">
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<paragraph id="8BBC92741478FF8DFF28AADCFD77AB33" blockId="27.[151,1437,151,2013]" box="[199,742,1266,1293]" pageId="27" pageNumber="428">
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<emphasis id="B9774E661478FF8DFF28AADCFD3CAB33" bold="true" box="[199,685,1266,1293]" pageId="27" pageNumber="428">
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<taxonomicName id="4C03E9F71478FF8DFF28AADCFE61AB33" ID-CoL="7XNH7" box="[199,496,1266,1293]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="marginatus">
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<emphasis id="B9774E661478FF8DFF28AADCFE61AB33" bold="true" box="[199,496,1266,1293]" italics="true" pageId="27" pageNumber="428">Onthophagus marginatus</emphasis>
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</taxonomicName>
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species group*.
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</emphasis>
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This
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</paragraph>
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</subSubSection>
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<subSubSection id="C319C1FF1478FF8DFD02AADDFB7AAB47" pageId="27" pageNumber="428" type="diagnosis">
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<paragraph id="8BBC92741478FF8DFD02AADDFB7AAB47" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="428">
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group is diagnosed by a pale antennal club; head frons with a strong carina; elytral intervals having a median row of tubercles each bearing a strong seta; distal edges of the elytra have contrasting pale markings (brown in
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<taxonomicName id="4C03E9F71478FF8DFD9BAB15FD66AB6B" box="[628,759,1339,1365]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="fragosus">
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<emphasis id="B9774E661478FF8DFD9BAB15FD66AB6B" box="[628,759,1339,1365]" italics="true" pageId="27" pageNumber="428">O. fragosus</emphasis>
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</taxonomicName>
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); the pygidium has distinct setiferous umbilicate punctures; males with a brush of fused long hair at the distal end of the protibiae. Its two species live in
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<collectingCountry id="F314D2E41478FF8DFB46AB71FB79AB47" box="[1193,1256,1375,1401]" name="Cuba" pageId="27" pageNumber="428">Cuba</collectingCountry>
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.
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</paragraph>
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</subSubSection>
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<subSubSection id="C319C1FF1478FF8DFF28ABADFC71A917" pageId="27" pageNumber="428" type="discussion">
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<paragraph id="8BBC92741478FF8DFF28ABADFE27A813" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="428">
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Although no species of this group could be included in the mtDNA analysis, this aggregation is established based on a characteristic set of morphological characters mentioned in the diagnosis. The group is composed at present by
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<taxonomicName id="4C03E9F71478FF8DFEFDABE5FD80ABDB" authority="Laporte" authorityName="Laporte" box="[274,529,1483,1509]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="423" phylum="Arthropoda" rank="species" species="marginatus">
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O.
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<emphasis id="B9774E661478FF8DFEDBABE5FE22ABDA" box="[308,435,1483,1508]" italics="true" pageId="27" pageNumber="428">marginatus</emphasis>
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Laporte
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</taxonomicName>
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and
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<taxonomicName id="4C03E9F71478FF8DFDA8ABE5FC36ABDB" authority="Genier & Howden." authorityName="Genier & Howden." box="[583,935,1483,1509]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="fragosus">
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<emphasis id="B9774E661478FF8DFDA8ABE5FD58ABDB" box="[583,713,1483,1509]" italics="true" pageId="27" pageNumber="428">O. fragosus</emphasis>
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Génier & Howden.
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</taxonomicName>
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The first species is widely distributed on the island; the second has been collected in litter on an isolated patch of moist forest (
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<quantity id="4CFB3F911478FF8DFBF4ABC0FB07A837" box="[1051,1174,1518,1545]" metricMagnitude="2" metricUnit="m" metricValue="7.925" metricValueMax="9.35" metricValueMin="6.5" pageId="27" pageNumber="428" unit="m" value="792.5" valueMax="935.0" valueMin="650.0">650-935 m</quantity>
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a.s.l.) (
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<bibRefCitation id="EF92EF851478FF8DFB09ABC1FA06A837" author="Matthews, E." box="[1254,1431,1519,1545]" pageId="27" pageNumber="428" pagination="1 - 134" refId="ref36431" refString="Matthews, E. (1966) A taxonomic and zoogeographic survey of the Scarabaeinae of the Antilles (Coleoptera: Scarabaeidae). Memoirs of the American Entomological Society, 21, 1-134." type="journal article" year="1966">Matthews 1966</bibRefCitation>
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;
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<bibRefCitation id="EF92EF851478FF8DFF78A83DFE3BA813" author="Genier, F. & Howden, H. F." box="[151,426,1555,1581]" pageId="27" pageNumber="428" pagination="597 - 599" refId="ref34552" refString="Genier, F. & Howden, H. F. (2014) Onthophagus fragosus n. sp. a second endemic species of Onthophagus Latreille from Cuba (Coleoptera: Scarabaeidae, Scarabaeinae). Zootaxa, 3795 (5), 597-599. https://doi.org/10.11646/zootaxa.3795.5.10" type="journal article" year="2014">Génier & Howden 2014</bibRefCitation>
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).
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</paragraph>
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<paragraph id="8BBC92741478FF8DFF28A819FC71A917" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="428">
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There is also a third species in the Greater Antilles living in Hispaniola,
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<taxonomicName id="4C03E9F71478FF8DFC06A819FA0DA86F" authority="Palisot. Matthews (1966)" authorityName="Palisot. Matthews" authorityYear="1966" box="[1001,1436,1591,1617]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="albicornis">
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<emphasis id="B9774E661478FF8DFC06A819FBECA86F" box="[1001,1149,1591,1617]" italics="true" pageId="27" pageNumber="428">O. albicornis</emphasis>
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Palisot.
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<bibRefCitation id="EF92EF851478FF8DFB32A819FA0DA86F" author="Matthews, E." box="[1245,1436,1591,1617]" pageId="27" pageNumber="428" pagination="1 - 134" refId="ref36431" refString="Matthews, E. (1966) A taxonomic and zoogeographic survey of the Scarabaeinae of the Antilles (Coleoptera: Scarabaeidae). Memoirs of the American Entomological Society, 21, 1-134." type="journal article" year="1966">Matthews (1966)</bibRefCitation>
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</taxonomicName>
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is of the opinion that the species of the Greater Antilles are derived from a common ancestor. This might be so because
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<taxonomicName id="4C03E9F71478FF8DFF17A851FE1FA8A7" authorityName="Palisot. Matthews" authorityYear="1966" box="[248,398,1663,1689]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="albicornis">
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<emphasis id="B9774E661478FF8DFF17A851FE1FA8A7" box="[248,398,1663,1689]" italics="true" pageId="27" pageNumber="428">O. albicornis</emphasis>
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</taxonomicName>
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shares with the two Cuban species the following characters: distal edges of the elytra have contrasting pale markings (brown in
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<taxonomicName id="4C03E9F71478FF8DFDDCA88DFD24A883" box="[563,693,1699,1725]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="fragosus">
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<emphasis id="B9774E661478FF8DFDDCA88DFD24A883" box="[563,693,1699,1725]" italics="true" pageId="27" pageNumber="428">O. fragosus</emphasis>
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</taxonomicName>
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); the antennal clubs are white while alive, turning yellowish when dead; prominent brush of fused setae emerging from the distal end of the male foretibia.
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<bibRefCitation id="EF92EF851478FF8DFB84A8E9FA0DA8DF" author="Genier, F. & Howden, H. F." box="[1131,1436,1735,1761]" pageId="27" pageNumber="428" pagination="597 - 599" refId="ref34552" refString="Genier, F. & Howden, H. F. (2014) Onthophagus fragosus n. sp. a second endemic species of Onthophagus Latreille from Cuba (Coleoptera: Scarabaeidae, Scarabaeinae). Zootaxa, 3795 (5), 597-599. https://doi.org/10.11646/zootaxa.3795.5.10" type="journal article" year="2014">Génier and Howden (2014)</bibRefCitation>
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indicated that the protibiae of
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<taxonomicName id="4C03E9F71478FF8DFE0AA8C5FDF7A93B" box="[485,614,1771,1797]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="fragosus">
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<emphasis id="B9774E661478FF8DFE0AA8C5FDF7A93B" box="[485,614,1771,1797]" italics="true" pageId="27" pageNumber="428">O. fragosus</emphasis>
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</taxonomicName>
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lacked the apical setal brush. However, photographs provided by Andrew Smith of the male
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<taxonomicName id="4C03E9F71478FF8DFE89A921FE79A917" box="[358,488,1807,1833]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="fragosus">
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<emphasis id="B9774E661478FF8DFE89A921FE79A917" box="[358,488,1807,1833]" italics="true" pageId="27" pageNumber="428">O. fragosus</emphasis>
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</taxonomicName>
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clearly shows the existence of this structure.
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</paragraph>
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</subSubSection>
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<mods:title id="84DC0EC87ECAB7BD3A8D463809E81546">A review of the species groups of the Western Hemisphere Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) using COI barcoding and gene trees</mods:title>
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<mods:namePart id="B995B55ADAF33FF8FD83144B55144047">Kohlmann, Bert</mods:namePart>
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<mods:namePart id="090C275215D6C9BAA1BF18D4690EFA89">Solís, Ángel</mods:namePart>
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<subSubSection id="C319C1FF147AFF8FFF28A850FD08A8A7" box="[199,665,1662,1689]" pageId="25" pageNumber="426" type="nomenclature">
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<paragraph id="8BBC9274147AFF8FFF28A850FD08A8A7" blockId="25.[151,1437,151,2013]" box="[199,665,1662,1689]" pageId="25" pageNumber="426">
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<taxonomicName id="4C03E9F7147AFF8FFF28A850FE75A8A7" ID-CoL="7XMYF" authorityName="Bates" authorityYear="1887" box="[199,484,1662,1689]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="gazellinus">
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<emphasis id="B9774E66147AFF8FFF28A850FE75A8A7" bold="true" box="[199,484,1662,1689]" italics="true" pageId="25" pageNumber="426">Onthophagus gazellinus</emphasis>
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species group
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.
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</paragraph>
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<subSubSection id="C319C1FF147AFF8FFD4DA851FD5EA973" pageId="25" pageNumber="426" type="diagnosis">
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<paragraph id="8BBC9274147AFF8FFD4DA851FD5EA973" blockId="25.[151,1437,151,2013]" pageId="25" pageNumber="426">
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||||
This group is diagnosed by the male having no head horns (
|
||||
<figureCitation id="13388EF1147AFF8FFA8BA851FF4AA883" captionStart-0="FIG" captionStart-1="FIG" captionStart-2="FIG" captionStart-3="FIG" captionStartId-0="15.[151,198,1589,1613]" captionStartId-1="17.[151,198,1790,1814]" captionStartId-2="18.[151,198,395,419]" captionStartId-3="19.[151,198,1784,1808]" captionTargetBox-0="[151,1436,181,1565]" captionTargetBox-1="[167,1430,190,1757]" captionTargetBox-2="[158,1435,188,365]" captionTargetBox-3="[152,1427,189,1751]" captionTargetId-0="figure-19@15.[151,1436,181,1565]" captionTargetId-1="figure-19@17.[158,1430,181,1766]" captionTargetId-2="figure-22@18.[152,1435,181,371]" captionTargetId-3="figure-19@19.[151,1436,181,1760]" captionTargetPageId-0="15" captionTargetPageId-1="17" captionTargetPageId-2="18" captionTargetPageId-3="19" captionText-0="FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed." captionText-1="FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes." captionText-2="FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale." captionText-3="FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes." figureDoi-0="http://doi.org/10.5281/zenodo.15213978" figureDoi-1="http://doi.org/10.5281/zenodo.15213980" figureDoi-2="http://doi.org/10.5281/zenodo.15213982" figureDoi-3="http://doi.org/10.5281/zenodo.15213984" httpUri-0="https://zenodo.org/record/15213978/files/figure.png" httpUri-1="https://zenodo.org/record/15213980/files/figure.png" httpUri-2="https://zenodo.org/record/15213982/files/figure.png" httpUri-3="https://zenodo.org/record/15213984/files/figure.png" pageId="25" pageNumber="426">Figs. 5.7–8</figureCitation>
|
||||
); male protibial apex with a tuft of long setae resembling a fine-pointed paintbrush (
|
||||
<figureCitation id="13388EF1147AFF8FFB90A88DFB61A883" box="[1151,1264,1698,1725]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.57</figureCitation>
|
||||
–58); pronotum with four transversely aligned tubercles along a straight carina (
|
||||
<figureCitation id="13388EF1147AFF8FFC6BA8E9FB9AA8DF" box="[900,1035,1734,1761]" captionStart-0="FIG" captionStart-1="FIG" captionStart-2="FIG" captionStart-3="FIG" captionStartId-0="15.[151,198,1589,1613]" captionStartId-1="17.[151,198,1790,1814]" captionStartId-2="18.[151,198,395,419]" captionStartId-3="19.[151,198,1784,1808]" captionTargetBox-0="[151,1436,181,1565]" captionTargetBox-1="[167,1430,190,1757]" captionTargetBox-2="[158,1435,188,365]" captionTargetBox-3="[152,1427,189,1751]" captionTargetId-0="figure-19@15.[151,1436,181,1565]" captionTargetId-1="figure-19@17.[158,1430,181,1766]" captionTargetId-2="figure-22@18.[152,1435,181,371]" captionTargetId-3="figure-19@19.[151,1436,181,1760]" captionTargetPageId-0="15" captionTargetPageId-1="17" captionTargetPageId-2="18" captionTargetPageId-3="19" captionText-0="FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed." captionText-1="FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes." captionText-2="FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale." captionText-3="FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes." figureDoi-0="http://doi.org/10.5281/zenodo.15213978" figureDoi-1="http://doi.org/10.5281/zenodo.15213980" figureDoi-2="http://doi.org/10.5281/zenodo.15213982" figureDoi-3="http://doi.org/10.5281/zenodo.15213984" httpUri-0="https://zenodo.org/record/15213978/files/figure.png" httpUri-1="https://zenodo.org/record/15213980/files/figure.png" httpUri-2="https://zenodo.org/record/15213982/files/figure.png" httpUri-3="https://zenodo.org/record/15213984/files/figure.png" pageId="25" pageNumber="426">Figs. 5.7–8</figureCitation>
|
||||
), base unrimmed; metafemur light brown with fine punctures (
|
||||
<figureCitation id="13388EF1147AFF8FFE3FA8C5FDD5A93B" box="[464,580,1770,1797]" captionStart="FIG" captionStartId="8.[151,198,1395,1419]" captionTargetBox="[319,1275,188,1362]" captionTargetId="figure-22@8.[312,1275,181,1371]" captionTargetPageId="8" captionText="FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213972" httpUri="https://zenodo.org/record/15213972/files/figure.png" pageId="25" pageNumber="426">Figs. 2.57</figureCitation>
|
||||
–58); at the excavated posterior pronotal margin a line of 5–6 large, ocellate punctures with one short, stiff, upward pointing seta, punctures disappearing towards middle of the pronotal margin; apex of elytra and pygidium with long, erect setae.
|
||||
</paragraph>
|
||||
</subSubSection>
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<subSubSection id="C319C1FF147AFF8CFF28A979FEEBAEEB" lastPageId="26" lastPageNumber="427" pageId="25" pageNumber="426" type="discussion">
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||||
<bibRefCitation id="EF92EF85147AFF8FFF28A979FE0AA94F" author="Boucomont, A." box="[199,411,1879,1905]" pageId="25" pageNumber="426" pagination="293 - 332" refId="ref33043" refString="Boucomont, A. (1932) Synopsis des Onthophagus d'Amerique du Sud (Col. Scarab.). Annales de la Societe Entomologique de France, 101, 293-332. https://doi.org/10.1080/21686351.1932.12280217" type="journal article" year="1932">Boucomont (1932)</bibRefCitation>
|
||||
initially established this species group as his “10
|
||||
<superScript id="7C763F3C147AFF8FFC54A979FC53A95B" attach="left" box="[955,962,1879,1893]" fontSize="6" pageId="25" pageNumber="426">e</superScript>
|
||||
Groupe” and was supported by
|
||||
<bibRefCitation id="EF92EF85147AFF8FFAC9A979FEACA9AB" author="Kohlmann, B. & Solis, A." pageId="25" pageNumber="426" pagination="159 - 261" refId="ref35740" refString="Kohlmann, B. & Solis, A. (2001) El genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, 9, 159-261." type="journal article" year="2001">Kohlmann & Solís (2001)</bibRefCitation>
|
||||
under the name of
|
||||
<taxonomicName id="4C03E9F7147AFF8FFDFBA955FD39A9AB" authorityName="Bates" authorityYear="1887" box="[532,680,1915,1941]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="gazellinus">
|
||||
<emphasis id="B9774E66147AFF8FFDFBA955FD39A9AB" box="[532,680,1915,1941]" italics="true" pageId="25" pageNumber="426">O. gazellinus</emphasis>
|
||||
</taxonomicName>
|
||||
species group. This group contains, at present, only two species,
|
||||
<taxonomicName id="4C03E9F7147AFF8FFA6EA955FEDFA987" authority="Bates" authorityName="Bates" authorityYear="1887" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="gazellinus">
|
||||
<emphasis id="B9774E66147AFF8FFA6EA955FE98A987" italics="true" pageId="25" pageNumber="426">O. gazellinus</emphasis>
|
||||
Bates
|
||||
</taxonomicName>
|
||||
and
|
||||
<taxonomicName id="4C03E9F7147AFF8FFE66A9B1FD0FA987" authority="Arrow" authorityName="Arrow" authorityYear="1913" box="[393,670,1951,1977]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="onthochromus">
|
||||
<emphasis id="B9774E66147AFF8FFE66A9B1FDC1A987" box="[393,592,1951,1977]" italics="true" pageId="25" pageNumber="426">O. onthochromus</emphasis>
|
||||
Arrow
|
||||
</taxonomicName>
|
||||
(
|
||||
<tableCitation id="C681A7CF147AFF8FFD5DA9B1FC9FA987" box="[690,782,1951,1977]" captionStart="TABLE 7" captionStartId="21.[152,245,728,752]" captionTargetBox="[163,1421,809,1957]" captionText="TABLE 7. Species groups and their proposed constituent species. Total number of taxa per group in parentheses. Species group Species and subspecies" pageId="25" pageNumber="426">Table 7</tableCitation>
|
||||
).
|
||||
<bibRefCitation id="EF92EF85147AFF8FFCC7A9B1FB90A987" author="Boucomont, A." box="[808,1025,1951,1977]" pageId="25" pageNumber="426" pagination="293 - 332" refId="ref33043" refString="Boucomont, A. (1932) Synopsis des Onthophagus d'Amerique du Sud (Col. Scarab.). Annales de la Societe Entomologique de France, 101, 293-332. https://doi.org/10.1080/21686351.1932.12280217" type="journal article" year="1932">Boucomont (1932)</bibRefCitation>
|
||||
defined the group as having large species with long erect setae at the end of the elytra and on the pygidium; male head unarmed and female with two carinae; thorax base not rimmed; coloration light, elytral points brown. This species group is distributed from
|
||||
<collectingCountry id="F314D2E41479FF8CFF78AE95FE9BAEEB" box="[151,266,187,213]" name="Nicaragua" pageId="26" pageNumber="427">Nicaragua</collectingCountry>
|
||||
to
|
||||
<collectingCountry id="F314D2E41479FF8CFEC1AE95FEE7AEEB" box="[302,374,187,213]" name="Brazil" pageId="26" pageNumber="427">Brazil</collectingCountry>
|
||||
.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
123
data/03/AA/23/03AA2362147AFF8FFF28AAA8FA83A84B.xml
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123
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Normal file
|
|
@ -0,0 +1,123 @@
|
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<mods:title id="828E54B6B1610CC995DB31611DF7DE44">A review of the species groups of the Western Hemisphere Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae) using COI barcoding and gene trees</mods:title>
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<mods:affiliation id="B33A3CEC543FD659396B2B30FD3452AD">BioAlfa Barcoding Project Santo Domingo de Heredia, Costa Rica.</mods:affiliation>
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<mods:nameIdentifier id="8805005DAED7873F79BA78274BBD6396" type="email">bkohlmann64@gmail.com</mods:nameIdentifier>
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<mods:namePart id="32D96F0040A340C9B920733BCE7EF4B4">Solís, Ángel</mods:namePart>
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<subSubSection id="C319C1FF147AFF8FFF28AAA8FD02AA9F" box="[199,659,1158,1185]" pageId="25" pageNumber="426" type="nomenclature">
|
||||
<paragraph id="8BBC9274147AFF8FFF28AAA8FD02AA9F" blockId="25.[151,1437,151,2013]" box="[199,659,1158,1185]" pageId="25" pageNumber="426">
|
||||
<emphasis id="B9774E66147AFF8FFF28AAA8FD1DAA9F" bold="true" box="[199,652,1158,1185]" pageId="25" pageNumber="426">
|
||||
<taxonomicName id="4C03E9F7147AFF8FFF28AAA8FE47AA9F" ID-CoL="7XMTN" authorityName="Bates" authorityYear="1887" box="[199,470,1158,1185]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="eulophus">
|
||||
<emphasis id="B9774E66147AFF8FFF28AAA8FE47AA9F" bold="true" box="[199,470,1158,1185]" italics="true" pageId="25" pageNumber="426">Onthophagus eulophus</emphasis>
|
||||
</taxonomicName>
|
||||
species group*
|
||||
</emphasis>
|
||||
.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
<subSubSection id="C319C1FF147AFF8FFD75AAA9FD25AB33" pageId="25" pageNumber="426" type="diagnosis">
|
||||
<paragraph id="8BBC9274147AFF8FFD75AAA9FD25AB33" blockId="25.[151,1437,151,2013]" pageId="25" pageNumber="426">
|
||||
This group is diagnosed by a slightly emarginate clypeus; apical and internal male protibial margins projected (
|
||||
<figureCitation id="13388EF1147AFF8FFD85AA85FD4BAAFB" box="[618,730,1194,1221]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.23</figureCitation>
|
||||
–25); male protibial apex with some short setae (
|
||||
<figureCitation id="13388EF1147AFF8FFB1DAA85FAF3AAFB" box="[1266,1378,1194,1221]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.23</figureCitation>
|
||||
–25); male protibial spur strongly bent inwards (
|
||||
<figureCitation id="13388EF1147AFF8FFD9DAAE1FD75AAD7" box="[626,740,1230,1257]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.23</figureCitation>
|
||||
–25); male pronotum with a horn or a plate-like horn, which is level or turned upwards, never bent downwards.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
<subSubSection id="C319C1FF147AFF8FFF28AB39FA83A84B" pageId="25" pageNumber="426" type="discussion">
|
||||
<paragraph id="8BBC9274147AFF8FFF28AB39FA83A84B" blockId="25.[151,1437,151,2013]" pageId="25" pageNumber="426">
|
||||
Although no species of this group could be included in the mtDNA analysis, this aggregation is established based on a characteristic set of morphological characters, like having apical and internal male protibial margins projected and a male protibial spur strongly bent inwards (
|
||||
<figureCitation id="13388EF1147AFF8FFCC5AB71FC0AAB47" box="[810,923,1374,1401]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.23</figureCitation>
|
||||
–25). Moreover, contrary to the
|
||||
<taxonomicName id="4C03E9F7147AFF8FFAECAB71FA0DAB46" authorityName="Bates" authorityYear="1887" box="[1283,1436,1375,1401]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="mexicanus">
|
||||
<emphasis id="B9774E66147AFF8FFAECAB71FA0DAB46" box="[1283,1436,1375,1401]" italics="true" pageId="25" pageNumber="426">O. mexicanus</emphasis>
|
||||
</taxonomicName>
|
||||
species group that have long and slender male protibiae (
|
||||
<figureCitation id="13388EF1147AFF8FFCE8ABADFCE6ABA3" box="[775,887,1411,1437]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.26</figureCitation>
|
||||
–28), this group has short and stout male protibiae (
|
||||
<figureCitation id="13388EF1147AFF8FFF70AB89FE80ABFF" box="[159,273,1446,1473]" captionStart="FIG" captionStartId="7.[151,198,1393,1417]" captionTargetBox="[332,1263,188,1360]" captionTargetId="figure-19@7.[325,1263,181,1369]" captionTargetPageId="7" captionText="FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed." figureDoi="http://doi.org/10.5281/zenodo.15213969" httpUri="https://zenodo.org/record/15213969/files/figure.png" pageId="25" pageNumber="426">Figs. 1.23</figureCitation>
|
||||
–25). All species of this group seem to be associated with rodent nests; this behavior is supported by the shortened male protibiae and bent male protibial spurs (Moctezuma &
|
||||
<bibRefCitation id="EF92EF85147AFF8FFC59ABE5FBC0ABDB" author="Halffter, G." box="[950,1105,1483,1509]" pageId="25" pageNumber="426" pagination="1 - 30" refId="ref36735" refString="Moctezuma. V. & Halffter, G. (2021) Species redescriptions and new species of the Onthophagus mexicanus species group (Coleoptera: Scarabaeidae), with notes on distribution and rodent-dung beetle associations. Zoological Studies, 60, 1-30." type="journal article" year="2021">Halffter 2021</bibRefCitation>
|
||||
;
|
||||
<bibRefCitation id="EF92EF85147AFF8FFBB2ABE5FAF4ABDB" author="Moctezuma, V. & Halffter, G. & Mora-Aguilar, E." box="[1117,1381,1483,1509]" pageId="25" pageNumber="426" pagination="1 - 9" refId="ref36871" refString="Moctezuma, V., Halffter, G. & Mora-Aguilar, E. (2023) A new species of the Onthophagus mirabilis species complex (Coleoptera: Scarabaeidae: Scarabaeinae) from the Chimalapas region, Oaxaca, Mexico. Revista Mexicana de Biodiversidad, 94 (e 945081), 1-9. https://doi.org/10.22201/ib.20078706e.2023.94.5081" type="journal article" year="2023">
|
||||
Moctezuma
|
||||
<emphasis id="B9774E66147AFF8FFB06ABE5FAB0ABDB" box="[1257,1313,1483,1509]" italics="true" pageId="25" pageNumber="426">et al.</emphasis>
|
||||
2023
|
||||
</bibRefCitation>
|
||||
). We consider that this group includes eight species (
|
||||
<tableCitation id="C681A7CF147AFF8FFD50ABC1FC88A837" box="[703,793,1519,1545]" captionStart="TABLE 7" captionStartId="21.[152,245,728,752]" captionTargetBox="[163,1421,809,1957]" captionText="TABLE 7. Species groups and their proposed constituent species. Total number of taxa per group in parentheses. Species group Species and subspecies" pageId="25" pageNumber="426">Table 7</tableCitation>
|
||||
).
|
||||
<bibRefCitation id="EF92EF85147AFF8FFCDEABC1FBC8A837" author="Zunino, M. & Halffter, G." box="[817,1113,1519,1545]" pageId="25" pageNumber="426" pagination="157 - 178" refId="ref39685" refString="Zunino, M. & Halffter, G. (1997) Sobre Onthophagus Latreille, 1802 americanos (Coleoptera: Scarabaeidae: Scarabaeinae). Elytron, 11, 157-178." type="journal article" year="1997">Zunino & Halffter (1997)</bibRefCitation>
|
||||
and Moctezuma &
|
||||
<bibRefCitation id="EF92EF85147AFF8FFAABABC1FF73A813" author="Halffter, G." pageId="25" pageNumber="426" pagination="1 - 30" refId="ref36735" refString="Moctezuma. V. & Halffter, G. (2021) Species redescriptions and new species of the Onthophagus mexicanus species group (Coleoptera: Scarabaeidae), with notes on distribution and rodent-dung beetle associations. Zoological Studies, 60, 1-30." type="journal article" year="2021">Halffter (2021)</bibRefCitation>
|
||||
consider these species part of the
|
||||
<taxonomicName id="4C03E9F7147AFF8FFD86A83DFC93A812" authorityName="Bates" authorityYear="1887" box="[617,770,1555,1581]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="mexicanus">
|
||||
<emphasis id="B9774E66147AFF8FFD86A83DFC93A812" box="[617,770,1555,1581]" italics="true" pageId="25" pageNumber="426">O. mexicanus</emphasis>
|
||||
</taxonomicName>
|
||||
species group. The
|
||||
<taxonomicName id="4C03E9F7147AFF8FFC08A83DFBFEA813" authorityName="Bates" authorityYear="1887" box="[999,1135,1555,1581]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="eulophus">
|
||||
<emphasis id="B9774E66147AFF8FFC08A83DFBFEA813" box="[999,1135,1555,1581]" italics="true" pageId="25" pageNumber="426">O. eulophus</emphasis>
|
||||
</taxonomicName>
|
||||
species group is primarily distributed in the mountains of the southwestern
|
||||
<collectingCountry id="F314D2E4147AFF8FFD57A819FC40A86F" box="[696,977,1591,1617]" name="United States of America" pageId="25" pageNumber="426">United States of America</collectingCountry>
|
||||
and along the Mexican Pacific mountain slopes, except for
|
||||
<taxonomicName id="4C03E9F7147AFF8FFE8CA875FD68A84B" authority="Moctezuma & Halffter" authorityName="Moctezuma & Halffter" authorityYear="2021" box="[355,761,1627,1653]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="25" pageNumber="426" phylum="Arthropoda" rank="species" species="totonacus">
|
||||
<emphasis id="B9774E66147AFF8FFE8CA875FE63A84A" box="[355,498,1627,1653]" italics="true" pageId="25" pageNumber="426">O. totonacus</emphasis>
|
||||
Moctezuma & Halffter
|
||||
</taxonomicName>
|
||||
, which lives along the Mexican Gulf piedmont.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
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Reference in a new issue