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<mods:title id="75D8FADD33E4208CABF68540F2269891">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="5FCCBE65265C8656EB494341EC857800">Kohlmann, Bert</mods:namePart>
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<mods:namePart id="71D4CD80CB3D6BFF57EF91F3C30D3B51">Solís, Ángel</mods:namePart>
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<treatment id="03AA23621479FF8DFF28A850FAA5AFCB" ID-DOI="http://doi.org/10.5281/zenodo.15225841" ID-GBIF-Taxon="258730473" ID-Zenodo-Dep="15225841" LSID="urn:lsid:plazi:treatment:03AA23621479FF8DFF28A850FAA5AFCB" httpUri="http://treatment.plazi.org/id/03AA23621479FF8DFF28A850FAA5AFCB" lastPageId="27" lastPageNumber="428" pageId="26" pageNumber="427" scope_family="Scarabaeidae" scope_order="Coleoptera" scope_subFamily="Scarabaeinae">
<subSubSection id="C319C1FF1479FF8CFF28A850FD0AA8A7" box="[199,667,1662,1689]" pageId="26" pageNumber="427" type="nomenclature">
<paragraph id="8BBC92741479FF8CFF28A850FD0AA8A7" blockId="26.[151,1437,151,2013]" box="[199,667,1662,1689]" pageId="26" pageNumber="427">
<emphasis id="B9774E661479FF8CFF28A850FD05A8A7" bold="true" box="[199,660,1662,1689]" pageId="26" pageNumber="427">
<taxonomicName id="4C03E9F71479FF8CFF28A850FE4DA8A7" ID-CoL="7W5H3" authorityName="Harold" authorityYear="1869" box="[199,476,1662,1689]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661479FF8CFF28A850FE4DA8A7" bold="true" box="[199,476,1662,1689]" italics="true" pageId="26" pageNumber="427">Onthophagus hoepfneri</emphasis>
</taxonomicName>
species group*
</emphasis>
.
</paragraph>
</subSubSection>
<subSubSection id="C319C1FF1479FF8CFD4DA851FA10A8DF" pageId="26" pageNumber="427" type="diagnosis">
<paragraph id="8BBC92741479FF8CFD4DA851FA10A8DF" blockId="26.[151,1437,151,2013]" pageId="26" pageNumber="427">
This group is diagnosed by no male head horns present (
<figureCitation id="13388EF11479FF8CFACAA851FA01A8A7" box="[1317,1424,1662,1689]" captionStart="FIG" captionStartId="15.[151,198,1589,1613]" captionTargetBox="[151,1436,181,1565]" captionTargetId="figure-19@15.[151,1436,181,1565]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.15213978" httpUri="https://zenodo.org/record/15213978/files/figure.png" pageId="26" pageNumber="427">Fig. 5.14</figureCitation>
); male protibial apex with some sparse short setae subequal to apical spur (
<figureCitation id="13388EF11479FF8CFC0BA88DFBC6A883" box="[996,1111,1698,1725]" 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="26" pageNumber="427">Figs. 1.59</figureCitation>
–66); pronotum with a small pointed or rounded anterior-pronotal projection (
<figureCitation id="13388EF11479FF8CFD57A8E9FCB2A8DF" box="[696,803,1734,1761]" captionStart="FIG" captionStartId="15.[151,198,1589,1613]" captionTargetBox="[151,1436,181,1565]" captionTargetId="figure-19@15.[151,1436,181,1565]" captionTargetPageId="15" captionText="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." figureDoi="http://doi.org/10.5281/zenodo.15213978" httpUri="https://zenodo.org/record/15213978/files/figure.png" pageId="26" pageNumber="427">Fig. 5.14</figureCitation>
); with coarse punctures in metafemur (
<figureCitation id="13388EF11479FF8CFB39A8E9FADBA8DF" box="[1238,1354,1734,1761]" 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="26" pageNumber="427">Figs. 2.59</figureCitation>
–66).
</paragraph>
</subSubSection>
<subSubSection id="C319C1FF1479FF8DFF28A8C5FAA5AFCB" lastPageId="27" lastPageNumber="428" pageId="26" pageNumber="427" type="multiple">
<paragraph id="8BBC92741479FF8DFF28A8C5FE53AF23" blockId="26.[151,1437,151,2013]" lastBlockId="27.[151,1437,151,2013]" lastPageId="27" lastPageNumber="428" pageId="26" pageNumber="427">
We propose, based on the barcode and morphological analyses, that this species group includes, at present, besides 
<taxonomicName id="4C03E9F71479FF8CFEB3A921FDDEA917" authority="Harold" authorityName="Harold" authorityYear="1869" box="[348,591,1807,1833]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661479FF8CFEB3A921FE67A917" box="[348,502,1807,1833]" italics="true" pageId="26" pageNumber="427">O. hoepfneri</emphasis>
Harold
</taxonomicName>
, four more species, 
<taxonomicName id="4C03E9F71479FF8CFCBAA921FBDCA917" authority=", Bates" authorityName="Bates" box="[853,1101,1807,1833]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="coscineus">
<emphasis id="B9774E661479FF8CFCBAA921FC60A916" box="[853,1009,1807,1833]" italics="true" pageId="26" pageNumber="427">O. coscineus</emphasis>
, Bates
</taxonomicName>
, 
<taxonomicName id="4C03E9F71479FF8CFBB7A921FAE7A917" authority="Brown" authorityName="Brown" authorityYear="1927" box="[1112,1398,1807,1833]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="oklahomensis">
<emphasis id="B9774E661479FF8CFBB7A921FA8EA917" box="[1112,1311,1807,1833]" italics="true" pageId="26" pageNumber="427">O. oklahomensis</emphasis>
Brown
</taxonomicName>
, 
<taxonomicName id="4C03E9F71479FF8CFA6EA921FE0BA973" authority="Harold" authorityName="Harold" authorityYear="1871" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="pennsylvanicus">
<emphasis id="B9774E661479FF8CFA6EA921FED3A973" italics="true" pageId="26" pageNumber="427">O. pennsylvanicus</emphasis>
Harold
</taxonomicName>
, and 
<taxonomicName id="4C03E9F71479FF8CFE37A91DFCD4A973" authority="Harold. Using" authorityName="Harold. Using" box="[472,837,1843,1869]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="tuberculifrons">
<emphasis id="B9774E661479FF8CFE37A91DFD0DA973" box="[472,668,1843,1869]" italics="true" pageId="26" pageNumber="427">O. tuberculifrons</emphasis>
Harold. Using
</taxonomicName>
a phylogenetic analysis, 
<bibRefCitation id="EF92EF851479FF8CFB80A91DFAD6A973" author="Emlen, D. J. &amp; Marangelo, J. &amp; Ball, B. &amp; Cunningham, C. W." box="[1135,1351,1842,1869]" pageId="26" pageNumber="427" pagination="1060 - 1084" refId="ref34152" refString="Emlen, D. J., Marangelo, J., Ball, B. &amp; Cunningham, C. W. (2005) Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution, 59, 1060-1084. https://doi.org/10.1111/j.0014-3820.2005.tb01044.x" type="journal article" year="2005">
Emlen 
<emphasis id="B9774E661479FF8CFB50A91DFB6AA973" box="[1215,1275,1843,1869]" italics="true" pageId="26" pageNumber="427">et al.</emphasis>
(2005
</bibRefCitation>
: fig. 2, bootstrap support of 79) had already obtained a branch formed by 
<taxonomicName id="4C03E9F71479FF8CFC48A979FBAAA94E" box="[935,1083,1879,1905]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="coscineus">
<emphasis id="B9774E661479FF8CFC48A979FBAAA94E" box="[935,1083,1879,1905]" italics="true" pageId="26" pageNumber="427">O. coscineus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C03E9F71479FF8CFB9EA979FAD3A94F" authorityName="Harold" authorityYear="1871" box="[1137,1346,1879,1905]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="pennsylvanicus">
<emphasis id="B9774E661479FF8CFB9EA979FAD3A94F" box="[1137,1346,1879,1905]" italics="true" pageId="26" pageNumber="427">O. pennsylvanicus</emphasis>
</taxonomicName>
. At the same time, because of the low support values indicated by the maximum-likelihood tree (
<figureCitation id="13388EF11479FF8CFB47A955FB7FA9AB" box="[1192,1262,1915,1941]" captionStart="FIG" captionStartId="19.[151,198,1784,1808]" captionTargetBox="[152,1427,189,1751]" captionTargetId="figure-19@19.[151,1436,181,1760]" captionTargetPageId="19" captionText="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="http://doi.org/10.5281/zenodo.15213984" httpUri="https://zenodo.org/record/15213984/files/figure.png" pageId="26" pageNumber="427">Fig. 8</figureCitation>
), more species groups might emerge from the 
<taxonomicName id="4C03E9F71479FF8CFE11A9B1FD1CA987" authorityName="Harold" authorityYear="1869" box="[510,653,1951,1977]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661479FF8CFE11A9B1FD1CA987" box="[510,653,1951,1977]" italics="true" pageId="26" pageNumber="427">O. hoepfneri</emphasis>
</taxonomicName>
branch once more species are included. These maximum-likelihood analyses suggest the existence of at least two clades within this species group, one formed by 
<taxonomicName id="4C03E9F71479FF8CFB36A9EDFAFBA9E2" box="[1241,1386,1987,2013]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="26" pageNumber="427" phylum="Arthropoda" rank="species" species="coscineus">
<emphasis id="B9774E661479FF8CFB36A9EDFAFBA9E2" box="[1241,1386,1987,2013]" italics="true" pageId="26" pageNumber="427">O. coscineus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C03E9F71478FF8DFF78AEB9FEC9AE8F" box="[151,344,151,177]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="tuberculifrons">
<emphasis id="B9774E661478FF8DFF78AEB9FEC9AE8F" box="[151,344,151,177]" italics="true" pageId="27" pageNumber="428">O. tuberculifrons</emphasis>
</taxonomicName>
, another by 
<taxonomicName id="4C03E9F71478FF8DFE07AEB9FDE7AE8F" authorityName="Harold" authorityYear="1869" box="[488,630,151,177]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661478FF8DFE07AEB9FDE7AE8F" box="[488,630,151,177]" italics="true" pageId="27" pageNumber="428">O. hoepfneri</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C03E9F71478FF8DFD69AEB9FCD3AE8F" authorityName="Brown" authorityYear="1927" box="[646,834,151,177]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="oklahomensis">
<emphasis id="B9774E661478FF8DFD69AEB9FCD3AE8F" box="[646,834,151,177]" italics="true" pageId="27" pageNumber="428">O. oklahomensis</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C03E9F71478FF8DFC6DAEB9FBC0AE8F" authorityName="Harold" authorityYear="1871" box="[898,1105,151,177]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="pennsylvanicus">
<emphasis id="B9774E661478FF8DFC6DAEB9FBC0AE8F" box="[898,1105,151,177]" italics="true" pageId="27" pageNumber="428">O. pennsylvanicus</emphasis>
</taxonomicName>
. More specimens are needed to clarify this situation. 
<bibRefCitation id="EF92EF851478FF8DFE44AE95FD13AEEB" author="Boucomont, A." box="[427,642,187,213]" pageId="27" pageNumber="428" 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>
placed 
<taxonomicName id="4C03E9F71478FF8DFD33AE95FCFAAEEB" authorityName="Harold" authorityYear="1869" box="[732,875,187,213]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661478FF8DFD33AE95FCFAAEEB" box="[732,875,187,213]" italics="true" pageId="27" pageNumber="428">O. hoepfneri</emphasis>
</taxonomicName>
in his “8 
<superScript id="7C763F3C1478FF8DFC39AE95FC4CAEF7" attach="left" box="[982,989,187,201]" fontSize="6" pageId="27" pageNumber="428">e</superScript>
Groupe”, which corresponds to the 
<taxonomicName id="4C03E9F71478FF8DFA6EAE95FF7FAEC7" authorityName="Harold" authorityYear="1880" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="landolti">
<emphasis id="B9774E661478FF8DFA6EAE95FF7FAEC7" italics="true" pageId="27" pageNumber="428">O. landolti</emphasis>
</taxonomicName>
species group, and 
<taxonomicName id="4C03E9F71478FF8DFE34AEF1FD0BAEC7" authorityName="Brown" authorityYear="1927" box="[475,666,223,249]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="oklahomensis">
<emphasis id="B9774E661478FF8DFE34AEF1FD0BAEC7" box="[475,666,223,249]" italics="true" pageId="27" pageNumber="428">O. oklahomensis</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C03E9F71478FF8DFD47AEF1FCEAAEC7" authorityName="Harold" authorityYear="1871" box="[680,891,223,249]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="pennsylvanicus">
<emphasis id="B9774E661478FF8DFD47AEF1FCEAAEC7" box="[680,891,223,249]" italics="true" pageId="27" pageNumber="428">O. pennsylvanicus</emphasis>
</taxonomicName>
, and 
<taxonomicName id="4C03E9F71478FF8DFC52AEF1FB10AEC7" box="[957,1153,223,249]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="tuberculifrons">
<emphasis id="B9774E661478FF8DFC52AEF1FB10AEC7" box="[957,1153,223,249]" italics="true" pageId="27" pageNumber="428">O. tuberculifrons</emphasis>
</taxonomicName>
in his “12 
<superScript id="7C763F3C1478FF8DFB10AEF1FA97AED3" attach="left" box="[1279,1286,223,237]" fontSize="6" pageId="27" pageNumber="428">e</superScript>
Groupe”; 
<taxonomicName id="4C03E9F71478FF8DFA6EAEF1FE95AF22" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="coscineus">
<emphasis id="B9774E661478FF8DFA6EAEF1FE95AF22" italics="true" pageId="27" pageNumber="428">O. coscineus</emphasis>
</taxonomicName>
was unassigned.
</paragraph>
<paragraph id="8BBC92741478FF8DFF28AF09FAA5AFCB" blockId="27.[151,1437,151,2013]" pageId="27" pageNumber="428">
The relationship of the 
<taxonomicName id="4C03E9F71478FF8DFE3CAF09FDF3AF7F" authorityName="Harold" authorityYear="1869" box="[467,610,295,321]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661478FF8DFE3CAF09FDF3AF7F" box="[467,610,295,321]" italics="true" pageId="27" pageNumber="428">O. hoepfneri</emphasis>
</taxonomicName>
species group with the 
<taxonomicName id="4C03E9F71478FF8DFC9CAF09FC7CAF7E" authorityName="Harold" authorityYear="1869" box="[883,1005,295,321]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="crinitus">
<emphasis id="B9774E661478FF8DFC9CAF09FC7CAF7E" box="[883,1005,295,321]" italics="true" pageId="27" pageNumber="428">O. crinitus</emphasis>
</taxonomicName>
species group suggested in this study (
<figureCitation id="13388EF11478FF8DFF70AF65FE9DAF5B" box="[159,268,331,357]" captionStart-0="FIG" captionStart-1="FIG" captionStartId-0="7.[151,198,1393,1417]" captionStartId-1="8.[151,198,1395,1419]" captionTargetBox-0="[332,1263,188,1360]" captionTargetBox-1="[319,1275,188,1362]" captionTargetId-0="figure-19@7.[325,1263,181,1369]" captionTargetId-1="figure-22@8.[312,1275,181,1371]" captionTargetPageId-0="7" captionTargetPageId-1="8" captionText-0="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." captionText-1="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-0="http://doi.org/10.5281/zenodo.15213969" figureDoi-1="http://doi.org/10.5281/zenodo.15213972" httpUri-0="https://zenodo.org/record/15213969/files/figure.png" httpUri-1="https://zenodo.org/record/15213972/files/figure.png" pageId="27" pageNumber="428">Figs. 1–2</figureCitation>
) was expected. In another maximum-likelihood genetic analysis, 
<bibRefCitation id="EF92EF851478FF8DFBEDAF65FB48AF5B" author="Emlen, D. J. &amp; Marangelo, J. &amp; Ball, B. &amp; Cunningham, C. W." box="[1026,1241,330,357]" pageId="27" pageNumber="428" pagination="1060 - 1084" refId="ref34152" refString="Emlen, D. J., Marangelo, J., Ball, B. &amp; Cunningham, C. W. (2005) Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution, 59, 1060-1084. https://doi.org/10.1111/j.0014-3820.2005.tb01044.x" type="journal article" year="2005">
Emlen 
<emphasis id="B9774E661478FF8DFBBDAF65FB1FAF5B" box="[1106,1166,331,357]" italics="true" pageId="27" pageNumber="428">et al.</emphasis>
(2005
</bibRefCitation>
: fig. 4, Bayesian posterior probability of 99) obtained a branch formed by 
<taxonomicName id="4C03E9F71478FF8DFCE1AF41FC14AFB6" authorityName="Harold" authorityYear="1869" box="[782,901,367,393]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="crinitus">
<emphasis id="B9774E661478FF8DFCE1AF41FC14AFB6" box="[782,901,367,393]" italics="true" pageId="27" pageNumber="428">O. crinitus</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C03E9F71478FF8DFC54AF41FB16AFB7" authorityName="Harold" authorityYear="1871" box="[955,1159,367,393]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="pennsylvanicus">
<emphasis id="B9774E661478FF8DFC54AF41FB16AFB7" box="[955,1159,367,393]" italics="true" pageId="27" pageNumber="428">O. pennsylvanicus</emphasis>
</taxonomicName>
. The 
<bibRefCitation id="EF92EF851478FF8DFB2AAF41FA0DAFB7" author="Emlen, D. J. &amp; Marangelo, J. &amp; Ball, B. &amp; Cunningham, C. W." box="[1221,1436,366,393]" pageId="27" pageNumber="428" pagination="1060 - 1084" refId="ref34152" refString="Emlen, D. J., Marangelo, J., Ball, B. &amp; Cunningham, C. W. (2005) Diversity in the weapons of sexual selection: horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution, 59, 1060-1084. https://doi.org/10.1111/j.0014-3820.2005.tb01044.x" type="journal article" year="2005">
Emlen 
<emphasis id="B9774E661478FF8DFAFAAF41FADDAFB7" box="[1301,1356,367,393]" italics="true" pageId="27" pageNumber="428">et al.</emphasis>
(2005)
</bibRefCitation>
analysis and the present barcode (
<figureCitation id="13388EF11478FF8DFDE5AFBDFDFDAF93" box="[522,620,403,429]" captionStart-0="FIG" captionStart-1="FIG" captionStartId-0="7.[151,198,1393,1417]" captionStartId-1="8.[151,198,1395,1419]" captionTargetBox-0="[332,1263,188,1360]" captionTargetBox-1="[319,1275,188,1362]" captionTargetId-0="figure-19@7.[325,1263,181,1369]" captionTargetId-1="figure-22@8.[312,1275,181,1371]" captionTargetPageId-0="7" captionTargetPageId-1="8" captionText-0="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." captionText-1="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-0="http://doi.org/10.5281/zenodo.15213969" figureDoi-1="http://doi.org/10.5281/zenodo.15213972" httpUri-0="https://zenodo.org/record/15213969/files/figure.png" httpUri-1="https://zenodo.org/record/15213972/files/figure.png" pageId="27" pageNumber="428">Figs 1–2</figureCitation>
) and partition (
<figureCitation id="13388EF11478FF8DFCF9AFBDFCC9AF93" box="[790,856,403,429]" captionStart="FIG" captionStartId="20.[151,198,1943,1967]" captionTargetBox="[319,1263,188,1912]" captionTargetId="figure-22@20.[318,1270,181,1919]" captionTargetPageId="20" captionText="FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes." figureDoi="http://doi.org/10.5281/zenodo.15213986" httpUri="https://zenodo.org/record/15213986/files/figure.png" pageId="27" pageNumber="428">Fig. 9</figureCitation>
) analyses suggest a close relationship between these two groups. 
<bibRefCitation id="EF92EF851478FF8DFEC9AF99FDF4AFEF" author="Schwery, O. &amp; O'Meara, B. C." box="[294,613,439,465]" pageId="27" pageNumber="428" refId="ref38339" refString="Schwery, O. &amp; O'Meara, B. C. (2021) Age, origin, and biogeography: unveiling the factors behind the diversification of dung beetles. Available from: https://doi.org/10.1101/2021.01.26.428346 (accessed 19 November 2024)" type="book" year="2021">Schwery &amp; O'Meara (2021)</bibRefCitation>
also recovered a mixed 
<emphasis id="B9774E661478FF8DFC90AF99FB06AFEE" box="[895,1175,439,465]" italics="true" pageId="27" pageNumber="428">
<taxonomicName id="4C03E9F71478FF8DFC90AF99FB82AFEF" authorityName="Harold" authorityYear="1869" box="[895,1043,439,465]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="hoepfneri">O. hoepfneri</taxonomicName>
– 
<taxonomicName id="4C03E9F71478FF8DFBCDAF99FB06AFEE" authorityName="Harold" authorityYear="1869" box="[1058,1175,439,465]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="crinitus">O. crinitus</taxonomicName>
</emphasis>
clade with a bootstrap support of 9. The 
<taxonomicName id="4C03E9F71478FF8DFEB0AFF5FE7DAFCB" authorityName="Harold" authorityYear="1869" box="[351,492,475,501]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="27" pageNumber="428" phylum="Arthropoda" rank="species" species="hoepfneri">
<emphasis id="B9774E661478FF8DFEB0AFF5FE7DAFCB" box="[351,492,475,501]" italics="true" pageId="27" pageNumber="428">O. hoepfneri</emphasis>
</taxonomicName>
species group is distributed from the 
<collectingCountry id="F314D2E41478FF8DFC7DAFF5FB3DAFCB" box="[914,1196,475,501]" name="United States of America" pageId="27" pageNumber="428">United States of America</collectingCountry>
to 
<collectingCountry id="F314D2E41478FF8DFB3FAFF5FABEAFCB" box="[1232,1327,475,501]" name="Ecuador" pageId="27" pageNumber="428">Ecuador</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>