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<document id="81333D1D683D4AC1DF794E95A92D5247" ID-CLB-Dataset="24885" ID-DOI="10.11646/zootaxa.4695.6.9" ID-GBIF-Dataset="e0078a85-2a99-4e9b-8add-91c9029db2b6" ID-ISSN="1175-5326" ID-Zenodo-Dep="3542761" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1573811501692" checkinUser="plazi" docAuthor="Delgado, Leonardo &amp; Mora-Aguilar, Eder F." docDate="2019" docId="03C287E25F364930D4B383E6FC4EA3C7" docLanguage="en" docName="zootaxa.4695.6.9.pdf" docOrigin="Zootaxa 4695 (6)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Onthophagus chimalapensis Delgado &amp; Mora-Aguilar 2019, new species" docType="treatment" docVersion="9" lastPageNumber="590" masterDocId="FFFBFF9A5F364934D424864EFFD1A420" masterDocTitle="A new Mexican species of Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae), with a revised key to the species of the O. dicranius species complex" masterLastPageNumber="592" masterPageNumber="586" pageNumber="586" updateTime="1698765991886" updateUser="ExternalLinkService">
<mods:mods id="9DE3638AE86D85869381E41E9754C649" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="3CF2282AE2A15C90E0C7C58741E5AC58">
<mods:title id="5ACC4B00DF17FB64C4AA987BA3A6F3E3">A new Mexican species of Onthophagus Latreille (Coleoptera: Scarabaeidae: Scarabaeinae), with a revised key to the species of the O. dicranius species complex</mods:title>
</mods:titleInfo>
<mods:name id="A5B0DCC774D7EF30182F5A651CC529AB" type="personal">
<mods:role id="22175906F1E98DF18EE235F8F6DDD315">
<mods:roleTerm id="2C15A3D52AA459F4744D64C54E518E8A">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="A1093341EA96BEA64569F7BCBDF8E4D2">Delgado, Leonardo</mods:namePart>
</mods:name>
<mods:name id="0E3AADEF864B0799F237973CD4EE9870" type="personal">
<mods:role id="FA3128C20846ECE3210AEFB8CD851E3A">
<mods:roleTerm id="7FF6D59F96E39236C2286C0D13B550B5">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="7C7F659EFC0E60519FD31311E87D5DF8">Mora-Aguilar, Eder F.</mods:namePart>
</mods:name>
<mods:typeOfResource id="7FF89938AAE9B2B05BDED9FBD7126617">text</mods:typeOfResource>
<mods:relatedItem id="8DAD79493AE2EDA0795C819ABFDC3E4C" type="host">
<mods:titleInfo id="76C17BF212CC330EA60D209CF490537A">
<mods:title id="450068B4E9F796BA74A0D1423FEAB4B1">Zootaxa</mods:title>
</mods:titleInfo>
<mods:part id="BC6B274DD86864B81DE29539BADBE4C3">
<mods:date id="735BC531EA9BC74AAC7166990031302F">2019</mods:date>
<mods:detail id="6247E5B1907BDB932B9E9924A6FAEAD4" type="pubDate">
<mods:number id="15A38118CA7E0EAB0F79234BEB382749">2019-11-13</mods:number>
</mods:detail>
<mods:detail id="DD4B4F5CAB076F061146229E13B22FB5" type="volume">
<mods:number id="6E7D7B319D77784528D2F3A372DB4696">4695</mods:number>
</mods:detail>
<mods:detail id="6E5A4289E7ABC1E12061813BD82CFBD6" type="issue">
<mods:number id="445CF9E305B0E33F7E320415E66EB0FD">6</mods:number>
</mods:detail>
<mods:extent id="655A7FE6D2F5D22623D046CE0AC5BD84" unit="page">
<mods:start id="00E38F2AEF52FA9E10ACA7C8BF83028C">586</mods:start>
<mods:end id="9C238443719C51BA2AB97DD25D10A370">592</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification id="AE9A1E48C49FAEC8CD043925FA6C0701">journal article</mods:classification>
<mods:identifier id="C640788F7EBCC8F721829660C6173B68" type="CLB-Dataset">24885</mods:identifier>
<mods:identifier id="11FDFC441F835167F8937D6ECF6F54ED" type="DOI">10.11646/zootaxa.4695.6.9</mods:identifier>
<mods:identifier id="72815113E1FDC426E5E5B2A16FC3662E" type="GBIF-Dataset">e0078a85-2a99-4e9b-8add-91c9029db2b6</mods:identifier>
<mods:identifier id="5AD964243D92B4B1E7C8545161B03E7D" type="ISSN">1175-5326</mods:identifier>
<mods:identifier id="DB6674F442AD8CD5C3153BCD1C853174" type="Zenodo-Dep">3542761</mods:identifier>
</mods:mods>
<treatment id="03C287E25F364930D4B383E6FC4EA3C7" ID-GBIF-Taxon="160406264" LSID="urn:lsid:zoobank.org:act:185F1648-2A1D-4C43-AAEA-755085B823B6" httpUri="http://treatment.plazi.org/id/03C287E25F364930D4B383E6FC4EA3C7" lastPageId="4" lastPageNumber="590" pageId="0" pageNumber="586">
<subSubSection id="C371657F5F364934D4B383E6FE41A1C6" pageId="0" pageNumber="586" type="nomenclature">
<paragraph id="8BD436F45F364934D4B383E6FC00A1E2" blockId="0.[151,977,1448,1546]" box="[151,977,1448,1475]" pageId="0" pageNumber="586">
<heading id="D09C81985F364934D4B383E6FC00A1E2" bold="true" box="[151,977,1448,1475]" fontSize="11" level="1" pageId="0" pageNumber="586" reason="1">
<emphasis id="B91FEAE65F364934D4B383E6FC00A1E2" bold="true" box="[151,977,1448,1475]" pageId="0" pageNumber="586">
<taxonomicName id="4C6B4D775F364934D4B383E6FCE5A1E2" ID-CoL="7XML7" authority="Delgado &amp; Mora-Aguilar" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[151,820,1448,1475]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="0" pageNumber="586" phylum="Arthropoda" rank="species" species="chimalapensis" status="sp. nov.">
<emphasis id="B91FEAE65F364934D4B383E6FE21A1E3" bold="true" box="[151,496,1448,1475]" italics="true" pageId="0" pageNumber="586">Onthophagus chimalapensis</emphasis>
Delgado &amp; Mora-Aguilar
</taxonomicName>
,
<taxonomicNameLabel id="A22C579D5F364934D76583E7FC00A1E2" box="[833,977,1448,1475]" pageId="0" pageNumber="586" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BD436F45F364934D4B38382FE41A1C6" blockId="0.[151,977,1448,1546]" box="[151,400,1484,1510]" pageId="0" pageNumber="586">
(
<figureCitation id="13502A715F364934D4BB8382FEDAA1C6" box="[159,267,1484,1510]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="0" pageNumber="586">Figs. 1–8</figureCitation>
,
<figureCitation id="13502A715F364934D5338382FE8EA1C6" box="[279,351,1484,1510]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="0" pageNumber="586">13, 16</figureCitation>
,
<figureCitation id="13502A715F364934D54F8382FE59A1C6" box="[363,392,1484,1510]" captionStart="FIGURE 19" captionStartId="4.[151,260,1240,1267]" captionTargetBox="[165,1421,178,1203]" captionTargetId="figure@4.[151,1436,164,1220]" captionText="FIGURE 19. Distribution of Onthophagus species. Onthophagus chimalapensis new species (circle), O. subcancer (square). Soft to dark grey lines represent elevational gradients of 1,000, 2,000, and 3,000 m, respectively." figureDoi="http://doi.org/10.5281/zenodo.3613728" httpUri="https://zenodo.org/record/3613728/files/figure.png" pageId="0" pageNumber="586">19</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C371657F5F364934D4B383A1FC17A22A" box="[151,966,1519,1546]" pageId="0" pageNumber="586" type="description">
<paragraph id="8BD436F45F364934D4B383A1FC17A22A" blockId="0.[151,977,1448,1546]" box="[151,966,1519,1546]" pageId="0" pageNumber="586">
<uri id="FFFA3AF65F364934D4B383A1FC17A22A" box="[151,966,1519,1546]" pageId="0" pageNumber="586">
urn:lsid:zoobank.org:act:
<uuid id="FFCD0C215F364934D58F83A1FC17A22A" box="[427,966,1519,1546]" pageId="0" pageNumber="586">185F1648-2A1D-4C43-AAEA-755085B823B6</uuid>
</uri>
</paragraph>
</subSubSection>
<subSubSection id="C371657F5F364935D4B38076FEA3A7CF" lastPageId="1" lastPageNumber="587" pageId="0" pageNumber="586" type="materials_examined">
<paragraph id="8BD436F45F364934D4B38076FC6DA2C2" blockId="0.[151,1437,1591,2015]" pageId="0" pageNumber="586">
<materialsCitation id="3B033CA95F364934D4B38076FCFDA2BA" ID-GBIF-Occurrence="2556210901" collectingDate="2017-07-26" collectorName="E. Arriaga-Varela &amp; F. Alvarado" country="Mexico" county="San Miguel Chimalapa" elevation="2" latitude="16.664139" location="El Reten" longLatPrecision="1" longitude="-94.23338" municipality="San Antonio" pageId="0" pageNumber="586" specimenCount="2" stateProvince="Oaxaca" typeStatus="holotype">
<emphasis id="B91FEAE65F364934D4B38076FE94A272" bold="true" box="[151,325,1591,1618]" pageId="0" pageNumber="586">Type material.</emphasis>
<typeStatus id="54D088565F364934D5688076FE65A272" box="[332,436,1592,1618]" pageId="0" pageNumber="586" type="holotype">Holotype</typeStatus>
male, and one female
<typeStatus id="54D088565F364934D6948077FCC1A273" box="[688,784,1593,1619]" pageId="0" pageNumber="586" type="paratype">paratype</typeStatus>
labeled: “
<collectingCountry id="F37C76645F364934D7A58076FC26A272" box="[897,1015,1592,1618]" name="Mexico" pageId="0" pageNumber="586">MEXICO</collectingCountry>
:
<collectingRegion id="49AFF8165F364934D0258076FB89A272" box="[1025,1112,1592,1618]" country="Mexico" name="Oaxaca" pageId="0" pageNumber="586">Oaxaca</collectingRegion>
,
<collectingCounty id="62B54E785F364934D0468076FAB6A272" box="[1122,1383,1592,1618]" pageId="0" pageNumber="586">San Miguel Chimalapa</collectingCounty>
,
<collectingMunicipality id="6BB0AC8E5F364934D1568076FF27A256" pageId="0" pageNumber="586">San Antonio</collectingMunicipality>
,
<location id="8EB4602F5F364934D5278012FEBBA256" LSID="urn:lsid:plazi:treatment:03C287E25F364930D4B383E6FC4EA3C7:8EB4602F5F364934D5278012FEBBA256" box="[259,362,1628,1654]" country="Mexico" county="San Miguel Chimalapa" latitude="16.664139" longLatPrecision="1" longitude="-94.23338" municipality="San Antonio" name="El Reten" pageId="0" pageNumber="586" stateProvince="Oaxaca">El Retén</location>
,
<location id="8EB4602F5F364934D5538012FE01A256" LSID="urn:lsid:plazi:treatment:03C287E25F364930D4B383E6FC4EA3C7:8EB4602F5F364934D5538012FE01A256" box="[375,464,1628,1654]" country="Mexico" county="San Miguel Chimalapa" latitude="16.664139" longLatPrecision="1" longitude="-94.23338" municipality="San Antonio" name="Camino" pageId="0" pageNumber="586" stateProvince="Oaxaca">Camino</location>
a la Torre,
<geoCoordinate id="EE5F50335F364934D67C8012FC1BA257" box="[600,970,1627,1655]" pageId="0" pageNumber="586">16º 39′ 50.9” N, 94º 14′ 0.2” W</geoCoordinate>
,
<date id="FFD510345F364934D7F38012FBB6A256" box="[983,1127,1628,1654]" pageId="0" pageNumber="586" value="2017-07-26">
<collectingDate id="EF91E9DC5F364934D7F38012FBB6A256" box="[983,1127,1628,1654]" pageId="0" pageNumber="586" value="2017-07-26">26-VII-2017</collectingDate>
</date>
,
<location id="8EB4602F5F364934D0518012FB03A256" LSID="urn:lsid:plazi:treatment:03C287E25F364930D4B383E6FC4EA3C7:8EB4602F5F364934D0518012FB03A256" box="[1141,1234,1628,1654]" country="Mexico" county="San Miguel Chimalapa" latitude="16.664139" longLatPrecision="1" longitude="-94.23338" municipality="San Antonio" name="Winkler" pageId="0" pageNumber="586" stateProvince="Oaxaca">Winkler</location>
,
<quantity id="4C939B115F364934D0FA8012FAEEA257" box="[1246,1343,1628,1655]" metricMagnitude="0" metricUnit="m" metricValue="1.8" pageId="0" pageNumber="586" unit="m" value="1.8">
<elevation id="0046D1C75F364934D0FA8012FAEEA257" box="[1246,1343,1628,1655]" metricMagnitude="0" metricUnit="m" metricValue="1.8" pageId="0" pageNumber="586" unit="m" value="1.8">1,800 m</elevation>
</quantity>
, bosque mesófilo,
<collectorName id="269E53225F364934D52280CFFE1DA2BA" box="[262,460,1664,1691]" pageId="0" pageNumber="586">E. Arriaga-Varela</collectorName>
,
<collectorName id="269E53225F364934D5F380CFFD8AA2BA" box="[471,603,1664,1691]" pageId="0" pageNumber="586">F. Alvarado</collectorName>
, E. Mora-A. cols.”
</materialsCitation>
<materialsCitation id="3B033CA95F364934D71580CEFC6DA2C2" ID-GBIF-Occurrence="2556210902" collectingDate="2017-07-26" collectorName="L. Delgado" country="Mexico" county="San Miguel Chimalapa" elevation="2" latitude="16.664139" location="Trampa de Intercepcion" longLatPrecision="1" longitude="-94.23338" municipality="San Antonio" pageId="0" pageNumber="586" specimenCount="1" stateProvince="Oaxaca" typeStatus="paratype">
One male
<typeStatus id="54D088565F364934D78780CFFBD2A2BB" box="[931,1027,1665,1691]" pageId="0" pageNumber="586" type="paratype">paratype</typeStatus>
with the following data: “
<collectingCountry id="F37C76645F364934D10780CEFA48A2BA" box="[1315,1433,1664,1690]" name="Mexico" pageId="0" pageNumber="586">MEXICO</collectingCountry>
:
<collectingRegion id="49AFF8165F364934D4B380EAFF3FA29E" box="[151,238,1700,1726]" country="Mexico" name="Oaxaca" pageId="0" pageNumber="586">Oaxaca</collectingRegion>
, San Miguel Chimalapa, San Antonio, El Gringo,
<geoCoordinate id="EE5F50335F364934D70D80EAFBABA29F" box="[809,1146,1699,1727]" pageId="0" pageNumber="586">16° 40′ 43” N, 94° 15′ 46” W</geoCoordinate>
, VI-VII-2013,
<location id="8EB4602F5F364934D10080EBFEF7A2C2" LSID="urn:lsid:plazi:treatment:03C287E25F364930D4B383E6FC4EA3C7:8EB4602F5F364934D10080EBFEF7A2C2" country="Mexico" county="San Miguel Chimalapa" latitude="16.664139" longLatPrecision="1" longitude="-94.23338" municipality="San Antonio" name="Trampa de Intercepcion" pageId="0" pageNumber="586" stateProvince="Oaxaca">Trampa de Intercepción</location>
,
<quantity id="4C939B115F364934D5158086FE5EA2C3" box="[305,399,1736,1763]" metricMagnitude="0" metricUnit="m" metricValue="1.6" pageId="0" pageNumber="586" unit="m" value="1.6">
<elevation id="0046D1C75F364934D5158086FE5EA2C3" box="[305,399,1736,1763]" metricMagnitude="0" metricUnit="m" metricValue="1.6" pageId="0" pageNumber="586" unit="m" value="1.6">1,600 m</elevation>
</quantity>
, bosque mesófilo, E. Mora-A.,
<collectorName id="269E53225F364934D6D48087FCBFA2C2" box="[752,878,1736,1763]" pageId="0" pageNumber="586">L. Delgado</collectorName>
cols.”.
</materialsCitation>
</paragraph>
<paragraph id="8BD436F45F364934D4E380A2FAD3A326" blockId="0.[151,1437,1591,2015]" box="[199,1282,1772,1799]" pageId="0" pageNumber="586">
<materialsCitation id="3B033CA95F364934D4E380A2FAD3A326" ID-GBIF-Occurrence="2451575216" box="[199,1282,1772,1799]" collectionCode="IEXA, CNIN" pageId="0" pageNumber="586" specimenCount="3" typeStatus="holotype">
<typeStatus id="54D088565F364934D4E380A2FEFEA326" box="[199,303,1772,1798]" pageId="0" pageNumber="586" type="holotype">Holotype</typeStatus>
and one female
<typeStatus id="54D088565F364934D5CC80A3FD99A327" box="[488,584,1773,1799]" pageId="0" pageNumber="586" type="paratype">paratype</typeStatus>
are deposited in
<collectionCode id="ED7AAE315F364934D72C80A2FC9FA326" box="[776,846,1772,1798]" pageId="0" pageNumber="586">IEXA</collectionCode>
, the male
<typeStatus id="54D088565F364934D7E480A3FBF1A327" box="[960,1056,1773,1799]" pageId="0" pageNumber="586" type="paratype">paratype</typeStatus>
deposited in
<collectionCode id="ED7AAE315F364934D09280A2FB2FA326" box="[1206,1278,1772,1798]" country="Mexico" httpUri="http://grbio.org/cool/qbc0-xx4g" name="Coleccion Nacional de Insectos, Universidad Nacional Autonoma de Mexico" pageId="0" pageNumber="586">CNIN</collectionCode>
.
</materialsCitation>
</paragraph>
<paragraph id="8BD436F45F364935D4E3815EFF03A643" blockId="0.[151,1437,1591,2015]" lastBlockId="1.[151,1437,153,2015]" lastPageId="1" lastPageNumber="587" pageId="0" pageNumber="586">
<emphasis id="B91FEAE65F364934D4E3815EFE2DA30A" bold="true" box="[199,508,1807,1834]" pageId="0" pageNumber="586">
<typeStatus id="54D088565F364934D4E3815EFEE5A30A" box="[199,308,1808,1834]" pageId="0" pageNumber="586" type="holotype">Holotype</typeStatus>
Male. Body size.
</emphasis>
Total length: 6.0 mm; width: 3.5 mm.
<emphasis id="B91FEAE65F364934D7888141FBD5A30A" bold="true" box="[940,1028,1807,1834]" pageId="0" pageNumber="586">Colour.</emphasis>
Dorsum shiny, dark brown; margins of head, femora, and tibiae reddish brown; apex and lateral margins of elytra up to the level of posterior margin of first sternite yellowish orange. Antennae reddish brown (
<figureCitation id="13502A715F364934D7048116FC7DA352" box="[800,940,1879,1906]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="0" pageNumber="586">Figs. 1, 3, 5</figureCitation>
).
<emphasis id="B91FEAE65F364934D7E78116FBDBA352" bold="true" box="[963,1034,1880,1906]" pageId="0" pageNumber="586">Head.</emphasis>
Clypeus concave, frons and vertex slightly convex. Surface of head with small, simple, or weakly annulate fine and sparse punctures, more accentuated near the margins. Clypeus armed with an apical horn slightly recurved backward; basal portion of horn wide and flat, tapering towards the rounded apex, sides parallel. Clypeal surface adjacent at the base of the horn concave; genae lobed, narrowed to the ocular canthus; clypeo-genal junction emarginate.
<emphasis id="B91FEAE65F374935D00D86D7FB59A493" bold="true" box="[1065,1160,153,179]" pageId="1" pageNumber="587">Thorax.</emphasis>
Pronotum with a feeble anteromedian protrusion marked by 2 lateral tubercles separated by a central concavity; anterolateral pronotal declivities concave; anterolateral edges of pronotum rounded. Pronotal disc with small, simple, or weakly annulate, sparse punctures; most punctures separated by more than 2 puncture diameters; punctures towards lateral and apical margins becoming larger, annulate, and denser. Metasternum gradually descending to mesosternum, with fine and moderate-sized punctures, denser and larger near the margins. Elytral striae distinctly impressed; with small, weakly annulate punctures separated by 2
<emphasis id="B91FEAE65F374935D63D873FFDF6A5AB" bold="true" box="[537,551,369,395]" pageId="1" pageNumber="587">–</emphasis>
3 puncture diameters; elytral intervals finely rugulose, with small, weakly annulate punctures, separated by 2
<emphasis id="B91FEAE65F374935D5DB87DBFDDCA58F" bold="true" box="[511,525,405,431]" pageId="1" pageNumber="587">–</emphasis>
3 diameters.
<emphasis id="B91FEAE65F374935D6B887DAFCC9A58F" bold="true" box="[668,792,404,431]" pageId="1" pageNumber="587">Abdomen.</emphasis>
Pygidium convex, coarsely punctate; punctures annulate, confluent on disc, and each bearing a minute seta.
<emphasis id="B91FEAE65F374935D6F287F7FCC4A5F3" bold="true" box="[726,789,441,467]" pageId="1" pageNumber="587">Legs.</emphasis>
Protibiae elongate, apex with internal angle slightly protruded, anterior edge with tuft of long setae, the two apical teeth separated from their base (
<figureCitation id="13502A715F374935D0A08793FB16A5D7" box="[1156,1223,477,503]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Fig. 7</figureCitation>
); apex of metatibia with setae mostly stout and short, and with some fine and long setae between them.
<emphasis id="B91FEAE65F374935D041844EFB0BA63B" bold="true" box="[1125,1242,512,539]" pageId="1" pageNumber="587">Genitalia.</emphasis>
Parameres moderately short, with apex slightly rounded in frontal view, acuminate and directed downward in lateral view (
<figureCitation id="13502A715F374935D140846BFF15A643" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Figs. 7–8</figureCitation>
).
</paragraph>
<paragraph id="8BD436F45F374935D4E38423FEA3A7CF" blockId="1.[151,1437,153,2015]" pageId="1" pageNumber="587">
<emphasis id="B91FEAE65F374935D4E38423FD54A6A7" bold="true" box="[199,645,620,647]" pageId="1" pageNumber="587">
<typeStatus id="54D088565F374935D4E38423FEEEA6A7" box="[199,319,621,647]" pageId="1" pageNumber="587" type="paratype">Paratypes</typeStatus>
(one male and one female).
</emphasis>
Total length: 5.3–5.8 mm; width 3.1–3.5 mm. The female differs from
<typeStatus id="54D088565F374935D4B384DFFF28A68B" box="[151,249,657,683]" pageId="1" pageNumber="587" type="holotype">holotype</typeStatus>
in the following respects: clypeus unarmed, sharply bidentate, teeth slightly reflexed and separated by a V-shaped notch, lateral margins of teeth slightly oblique, lateral clypeal margin slightly curved; clypeus flat, with surface irregularly wrinkled; clypeofrontal suture with a sharp and straight carina, not reaching the lateral borders; frons with indistinct swellings in front of the eyes; pronotum nearly rounded in dorsal view, evenly convex, except for the pronotal disc which has a rounded, flat surface anteriorly; protibiae robust, lacking tuft of apical setae and with the inner angle not protruded (
<figureCitation id="13502A715F374935D608850BFD69A77F" box="[556,696,837,863]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Figs. 2, 4, 6</figureCitation>
,
<figureCitation id="13502A715F374935D6E1850BFD35A740" box="[709,740,837,864]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="1" pageNumber="587">16</figureCitation>
). The male
<typeStatus id="54D088565F374935D748850BFC1DA77F" box="[876,972,837,863]" pageId="1" pageNumber="587" type="paratype">paratype</typeStatus>
differs from the
<typeStatus id="54D088565F374935D0AB850BFB20A77F" box="[1167,1265,837,863]" pageId="1" pageNumber="587" type="holotype">holotype</typeStatus>
because of the smaller size and the following sets of characters: profemora and mesofemora, apex and lateral margins of elytra darker; clypeal horn slightly shorter, genae less expanded; pronotal protrusion weaker, with tubercles shorter, and anterolateral declivities less concave. Punctation varies little in density and depth in both sexes. Other characters are similar to
<typeStatus id="54D088565F374935D52D859BFEBEA7CF" box="[265,367,981,1007]" pageId="1" pageNumber="587" type="holotype">holotype</typeStatus>
.
</paragraph>
</subSubSection>
<subSubSection id="C371657F5F374935D4E385B7FBF1A017" pageId="1" pageNumber="587" type="etymology">
<paragraph id="8BD436F45F374935D4E385B7FBF1A017" blockId="1.[151,1437,153,2015]" pageId="1" pageNumber="587">
<emphasis id="B91FEAE65F374935D4E385B7FE9FA033" bold="true" box="[199,334,1017,1043]" pageId="1" pageNumber="587">Etymology.</emphasis>
The specific epithet is derived from Chimalapas, the region of the state of
<collectingRegion id="49AFF8165F374935D0B385B7FB3AA033" box="[1175,1259,1017,1043]" country="Mexico" name="Oaxaca" pageId="1" pageNumber="587">Oaxaca</collectingRegion>
where this species was collected, combined with the Latin suffix –
<emphasis id="B91FEAE65F374935D6FB8250FCC6A017" box="[735,791,1054,1079]" italics="true" pageId="1" pageNumber="587">ensis</emphasis>
, meaning belonging to.
</paragraph>
</subSubSection>
<subSubSection id="C371657F5F374930D4E3820EFC4EA3C7" lastPageId="4" lastPageNumber="590" pageId="1" pageNumber="587" type="distribution">
<paragraph id="8BD436F45F374935D4E3820EFC65A0CB" blockId="1.[151,1437,153,2015]" pageId="1" pageNumber="587">
<emphasis id="B91FEAE65F374935D4E3820EFEB0A07B" bold="true" box="[199,353,1088,1115]" pageId="1" pageNumber="587">Distribution.</emphasis>
<taxonomicName id="4C6B4D775F374935D543820FFD74A07B" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[359,677,1089,1115]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F374935D543820FFD74A07B" box="[359,677,1089,1115]" italics="true" pageId="1" pageNumber="587">Onthophagus chimalapensis</emphasis>
</taxonomicName>
is known from two neighbouring localities, in the highly conserved mountainous areas of El Retén in the Chimalapas region, in the State of
<collectingRegion id="49AFF8165F374935D7EC822BFBCEA05F" box="[968,1055,1125,1151]" country="Mexico" name="Oaxaca" pageId="1" pageNumber="587">Oaxaca</collectingRegion>
,
<collectingCountry id="F37C76645F374935D00F822BFB50A05F" box="[1067,1153,1125,1151]" name="Mexico" pageId="1" pageNumber="587">Mexico</collectingCountry>
(
<figureCitation id="13502A715F374935D0B4822BFB35A05F" box="[1168,1252,1125,1151]" captionStart="FIGURE 19" captionStartId="4.[151,260,1240,1267]" captionTargetBox="[165,1421,178,1203]" captionTargetId="figure@4.[151,1436,164,1220]" captionText="FIGURE 19. Distribution of Onthophagus species. Onthophagus chimalapensis new species (circle), O. subcancer (square). Soft to dark grey lines represent elevational gradients of 1,000, 2,000, and 3,000 m, respectively." figureDoi="http://doi.org/10.5281/zenodo.3613728" httpUri="https://zenodo.org/record/3613728/files/figure.png" pageId="1" pageNumber="587">Fig. 19</figureCitation>
). Both localities and surrounding areas have a rugged topography with ravines and hills with moist cloud forests ranging from about
<quantity id="4C939B115F374935D4B382E2FF22A0E7" box="[151,243,1196,1223]" metricMagnitude="0" metricUnit="m" metricValue="1.45" pageId="1" pageNumber="587" unit="m" value="1.45">1,450 m</quantity>
to
<quantity id="4C939B115F374935D53382E3FEA7A0E7" box="[279,374,1197,1223]" metricMagnitude="0" metricUnit="m" metricValue="1.8" pageId="1" pageNumber="587" unit="m" value="1.8">1,800 m</quantity>
. The cloud forests of the Chimalapas region are separated from the cloud forests of the Meseta Central de
<collectingRegion id="49AFF8165F374935D536829FFEBDA0CB" box="[274,364,1233,1259]" country="Mexico" name="Chiapas" pageId="1" pageNumber="587">Chiapas</collectingRegion>
and the Sierra Madre de
<collectingRegion id="49AFF8165F374935D6A3829FFD30A0CB" box="[647,737,1233,1259]" country="Mexico" name="Chiapas" pageId="1" pageNumber="587">Chiapas</collectingRegion>
by drier lowlands.
</paragraph>
<paragraph id="8BD436F45F374935D4E382BAFC22A1C7" blockId="1.[151,1437,153,2015]" pageId="1" pageNumber="587">
<emphasis id="B91FEAE65F374935D4E382BAFE6DA12F" bold="true" box="[199,444,1268,1295]" pageId="1" pageNumber="587">Taxonomic remarks.</emphasis>
<taxonomicName id="4C6B4D775F374935D5E682BBFD2EA12F" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[450,767,1269,1295]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F374935D5E682BBFD2EA12F" box="[450,767,1269,1295]" italics="true" pageId="1" pageNumber="587">Onthophagus chimalapensis</emphasis>
</taxonomicName>
(
<figureCitation id="13502A715F374935D72A82BBFCA7A12F" box="[782,886,1269,1295]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Figs. 1–8</figureCitation>
) is similar to
<taxonomicName id="4C6B4D775F374935D02A82BBFAD6A12F" authority="Howden" authorityName="Howden" authorityYear="1973" box="[1038,1287,1269,1295]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F374935D02A82BBFB73A12F" box="[1038,1186,1269,1295]" italics="true" pageId="1" pageNumber="587">O. subcancer</emphasis>
Howden
</taxonomicName>
(
<figureCitation id="13502A715F374935D13282BBFA41A12F" box="[1302,1424,1269,1295]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="1" pageNumber="587">Figs. 9–12</figureCitation>
), as both species share the following characters: length less than
<quantity id="4C939B115F374935D74E8357FC61A113" box="[874,944,1305,1331]" metricMagnitude="-3" metricUnit="m" metricValue="7.0" pageId="1" pageNumber="587" unit="mm" value="7.0">7 mm</quantity>
; colour dark brown lacking metallic iridescence; elytra with apex yellowish red; mesofemora and metafemora light brown or yellowish red (
<figureCitation id="13502A715F374935D0CF8373FA88A177" box="[1259,1369,1341,1367]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Figs. 1–4</figureCitation>
); dorsum glabrous, punctures of the pronotal disc simple or weakly annulate, small, and sparse; pygidium coarsely punctate; protibiae with the two apical teeth widely separated (joined only at the base); clypeal horn of male recurved; head lacking of carinae, tubercles, or horns between the eyes; clypeus of female bidentate at middle; clypeofrontal suture with a central carina; frons without carina or tubercles behind of eyes.
</paragraph>
<paragraph id="8BD436F45F374935D4E383BFFD7CA29F" blockId="1.[151,1437,153,2015]" pageId="1" pageNumber="587">
The new species differs from
<taxonomicName id="4C6B4D775F374935D60583BFFD69A22B" authorityName="Howden" authorityYear="1973" box="[545,696,1521,1547]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F374935D60583BFFD69A22B" box="[545,696,1521,1547]" italics="true" pageId="1" pageNumber="587">O. subcancer</emphasis>
</taxonomicName>
by the following characters: male with frons and vertex convex and finely punctate (not flat and with distinct, moderate-sized punctures) (
<figureCitation id="13502A715F374935D7C8805BFBEFA20F" box="[1004,1086,1556,1583]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Figs. 5</figureCitation>
,
<figureCitation id="13502A715F374935D068805BFB8DA20F" box="[1100,1116,1557,1583]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="1" pageNumber="587">9</figureCitation>
), female with frons without tubercles (not with two tubercles at level of the anterior border of eyes), and pronotal disc with a rounded, slightly concave surface anteriorly (not evenly convex) (
<figureCitation id="13502A715F374935D68B8013FD2CA257" box="[687,765,1629,1655]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="1" pageNumber="587">Figs. 6</figureCitation>
,
<figureCitation id="13502A715F374935D72C8013FCF6A257" box="[776,807,1629,1655]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="1" pageNumber="587">10</figureCitation>
). This combination of characters is diagnostic to the new species. The size and shape of the pronotal protrusion observed on the
<typeStatus id="54D088565F374935D78A80CFFBC1A2BB" box="[942,1040,1665,1691]" pageId="1" pageNumber="587" type="holotype">holotype</typeStatus>
and male
<typeStatus id="54D088565F374935D0A780CFFB32A2BB" box="[1155,1251,1665,1691]" pageId="1" pageNumber="587" type="paratype">paratype</typeStatus>
of
<taxonomicName id="4C6B4D775F374935D12D80CFFF1EA29F" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F374935D12D80CFFF1EA29F" italics="true" pageId="1" pageNumber="587">O. chimalapensis</emphasis>
</taxonomicName>
show seemingly a moderate development.
</paragraph>
<paragraph id="8BD436F45F374935D4E38087FADDA3FF" blockId="1.[151,1437,153,2015]" pageId="1" pageNumber="587">
Many of the characters shared between
<taxonomicName id="4C6B4D775F374935D6A28087FC99A2C3" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[646,840,1737,1763]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F374935D6A28087FC99A2C3" box="[646,840,1737,1763]" italics="true" pageId="1" pageNumber="587">O. chimalapensis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6B4D775F374935D75B8087FBC4A2C3" authorityName="Howden" authorityYear="1973" box="[895,1045,1737,1763]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F374935D75B8087FBC4A2C3" box="[895,1045,1737,1763]" italics="true" pageId="1" pageNumber="587">O. subcancer</emphasis>
</taxonomicName>
are also shared with the species of the
<taxonomicName id="4C6B4D775F374935D4E580A3FE9CA327" authorityName="Bates" authorityYear="1887" box="[193,333,1773,1799]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F374935D4E580A3FE9CA327" box="[193,333,1773,1799]" italics="true" pageId="1" pageNumber="587">O. dicranius</emphasis>
</taxonomicName>
species group (
<emphasis id="B91FEAE65F374935D5DA80A0FDEDA327" box="[510,572,1774,1799]" italics="true" pageId="1" pageNumber="587">sensu</emphasis>
<bibRefCitation id="EFFA4B055F374935D66780A3FC88A327" author="Kohlmann, B. &amp; Solis, A." box="[579,857,1773,1799]" pageId="1" pageNumber="587" pagination="159 - 261" refId="ref4401" refString="Kohlmann, B. &amp; Solis, A. (2001) El genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, 49, 159 - 261." type="journal article" year="2001">Kohlmann &amp; Solís 2001</bibRefCitation>
). In fact,
<bibRefCitation id="EFFA4B055F374935D7E780A3FB33A327" author="Zunino, M. &amp; Halffter, G." box="[963,1250,1773,1799]" pageId="1" pageNumber="587" pagination="157 - 178" refId="ref4529" refString="Zunino, M. &amp; Halffter, G. (1997) Sobre Onthophagus Latreille, 1802 americanos (Coleoptera: Scarabaeidae: Scarabaeinae). Elytron, 11, 157 - 178." type="journal article" year="1997">Zunino &amp; Halffter (1997)</bibRefCitation>
had included
<taxonomicName id="4C6B4D775F374935D1A580A3FEDBA30B" authorityName="Howden" authorityYear="1973" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F374935D1A580A3FEDBA30B" italics="true" pageId="1" pageNumber="587">O. subcancer</emphasis>
</taxonomicName>
into the
<taxonomicName id="4C6B4D775F374935D554815FFE29A30B" authorityName="Bates" authorityYear="1887" box="[368,504,1809,1835]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="mirabilis">
<emphasis id="B91FEAE65F374935D554815FFE29A30B" box="[368,504,1809,1835]" italics="true" pageId="1" pageNumber="587">O. mirabilis</emphasis>
</taxonomicName>
species set of the
<taxonomicName id="4C6B4D775F374935D6ED815FFC87A30B" box="[713,854,1809,1835]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="clypeatus">
<emphasis id="B91FEAE65F374935D6ED815FFC87A30B" box="[713,854,1809,1835]" italics="true" pageId="1" pageNumber="587">O. clypeatus</emphasis>
</taxonomicName>
group. Likewise,
<bibRefCitation id="EFFA4B055F374935D007815FFAF7A30B" author="Howden, H &amp; Gill, B. D." box="[1059,1318,1809,1835]" pageId="1" pageNumber="587" pagination="201 - 224" refId="ref4305" refString="Howden, H .. &amp; Gill, B. D. (1987) New species and new records of Panamanian and Costa Rican Scarabaeinae (Coleoptera: Scarabaeidae). The Coleopterists Bulletin, 41, 201 - 224." type="journal article" year="1987">Howden &amp; Gill (1987)</bibRefCitation>
also mentioned the similarity of
<taxonomicName id="4C6B4D775F374935D5BE817BFDF4A36F" authorityName="Bates" authorityYear="1887" box="[410,549,1845,1871]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F374935D5BE817BFDF4A36F" box="[410,549,1845,1871]" italics="true" pageId="1" pageNumber="587">O. dicranius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6B4D775F374935D67D817BFC63A36F" authority="Howden &amp; Gill" authorityName="Howden &amp; Gill" authorityYear="1987" box="[601,946,1845,1871]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="dorsipilulus">
<emphasis id="B91FEAE65F374935D67D817BFCD1A36F" box="[601,768,1845,1871]" italics="true" pageId="1" pageNumber="587">O. dorsipilulus</emphasis>
Howden &amp; Gill
</taxonomicName>
to
<taxonomicName id="4C6B4D775F374935D7F7817BFBB6A36F" authorityName="Howden" authorityYear="1973" box="[979,1127,1845,1871]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F374935D7F7817BFBB6A36F" box="[979,1127,1845,1871]" italics="true" pageId="1" pageNumber="587">O. subcancer</emphasis>
</taxonomicName>
. Morphological differences of both
<taxonomicName id="4C6B4D775F374935D4C88117FE51A353" authorityName="Howden" authorityYear="1973" box="[236,384,1881,1907]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F374935D4C88117FE51A353" box="[236,384,1881,1907]" italics="true" pageId="1" pageNumber="587">O. subcancer</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6B4D775F374935D5908117FDA4A353" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[436,629,1881,1907]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F374935D5908117FDA4A353" box="[436,629,1881,1907]" italics="true" pageId="1" pageNumber="587">O. chimalapensis</emphasis>
</taxonomicName>
with the species of the
<taxonomicName id="4C6B4D775F374935D7518117FBD1A353" authorityName="Bates" authorityYear="1887" box="[885,1024,1881,1907]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F374935D7518117FBD1A353" box="[885,1024,1881,1907]" italics="true" pageId="1" pageNumber="587">O. dicranius</emphasis>
</taxonomicName>
species group are basically related to punctation: the punctures of the pronotal disc are sparse, simple, or weakly annulate and fine (punctures not annulate and/or large, and dense in the species of the
<taxonomicName id="4C6B4D775F374935D6AE81EFFCC7A39B" authorityName="Bates" authorityYear="1887" box="[650,790,1953,1979]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F374935D6AE81EFFCC7A39B" box="[650,790,1953,1979]" italics="true" pageId="1" pageNumber="587">O. dicranius</emphasis>
</taxonomicName>
species group); while the elytral striae have annulate and small punctures (not annulate and/or large punctures in the species of the
<taxonomicName id="4C6B4D775F374935D7EB818BFB8AA3FF" authorityName="Bates" authorityYear="1887" box="[975,1115,1989,2015]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="587" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F374935D7EB818BFB8AA3FF" box="[975,1115,1989,2015]" italics="true" pageId="1" pageNumber="587">O. dicranius</emphasis>
</taxonomicName>
species group).
</paragraph>
<caption id="DF14667C5F344936D4B381DAFB6CA3D7" ID-DOI="http://doi.org/10.5281/zenodo.3613724" ID-Zenodo-Dep="3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="2" pageNumber="588" startId="2.[151,275,1940,1967]" targetBox="[183,1411,197,1903]" targetPageId="2">
<paragraph id="8BD436F45F344936D4B381DAFB6CA3D7" blockId="2.[151,1436,1940,2039]" pageId="2" pageNumber="588">
<emphasis id="B91FEAE65F344936D4B381DAFE9CA38F" bold="true" box="[151,333,1940,1967]" pageId="2" pageNumber="588">FIGURES 1–8.</emphasis>
<taxonomicName id="4C6B4D775F344936D57181DBFD45A38F" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[341,660,1941,1967]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="588" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F344936D57181DBFD45A38F" box="[341,660,1941,1967]" italics="true" pageId="2" pageNumber="588">Onthophagus chimalapensis</emphasis>
</taxonomicName>
.
<emphasis id="B91FEAE65F344936D68081DBFD63A38F" bold="true" box="[676,690,1941,1967]" pageId="2" pageNumber="588">1</emphasis>
, Habitus,
<typeStatus id="54D088565F344936D70381DBFC5CA38F" box="[807,909,1941,1967]" pageId="2" pageNumber="588" type="holotype">holotype</typeStatus>
;
<emphasis id="B91FEAE65F344936D7BD81DBFC76A38F" bold="true" box="[921,935,1941,1967]" pageId="2" pageNumber="588">2</emphasis>
, habitus, female
<typeStatus id="54D088565F344936D04F81DBFB1EA38F" box="[1131,1231,1941,1967]" pageId="2" pageNumber="588" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F344936D0FF81DBFB38A38F" bold="true" box="[1243,1257,1941,1967]" pageId="2" pageNumber="588">3</emphasis>
, lateral view of male
<typeStatus id="54D088565F344936D4F181F7FEE8A3F3" box="[213,313,1977,2003]" pageId="2" pageNumber="588" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F344936D56181F7FE82A3F3" bold="true" box="[325,339,1977,2003]" pageId="2" pageNumber="588">4</emphasis>
, lateral view of female
<typeStatus id="54D088565F344936D64481F7FD15A3F3" box="[608,708,1977,2003]" pageId="2" pageNumber="588" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F344936D6F481F7FD0FA3F3" bold="true" box="[720,734,1977,2003]" pageId="2" pageNumber="588">5</emphasis>
, ¾ profile view of male
<typeStatus id="54D088565F344936D7D281F7FB8DA3F3" box="[1014,1116,1977,2003]" pageId="2" pageNumber="588" type="holotype">holotype</typeStatus>
;
<emphasis id="B91FEAE65F344936D04C81F7FBA7A3F3" bold="true" box="[1128,1142,1977,2003]" pageId="2" pageNumber="588">6</emphasis>
, ¾ profile view of female
<typeStatus id="54D088565F344936D4B38193FF2AA3D7" box="[151,251,2013,2039]" pageId="2" pageNumber="588" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F344936D5228193FEC5A3D7" bold="true" box="[262,276,2013,2039]" pageId="2" pageNumber="588">7</emphasis>
, lateral view of genitalia, male
<typeStatus id="54D088565F344936D6538193FD0AA3D7" box="[631,731,2013,2039]" pageId="2" pageNumber="588" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F344936D6C28193FD25A3D7" bold="true" box="[742,756,2013,2039]" pageId="2" pageNumber="588">8</emphasis>
, dorsal view of genitalia, male
<typeStatus id="54D088565F344936D0718193FB68A3D7" box="[1109,1209,2013,2039]" pageId="2" pageNumber="588" type="paratype">paratype</typeStatus>
.
</paragraph>
</caption>
<caption id="DF14667C5F354937D4B38173FDF4A3C7" ID-DOI="http://doi.org/10.5281/zenodo.3613726" ID-Zenodo-Dep="3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="3" pageNumber="589" startId="3.[151,275,1853,1880]" targetBox="[192,1415,190,1785]" targetPageId="3">
<paragraph id="8BD436F45F354937D4B38173FDF4A3C7" blockId="3.[151,1436,1853,2024]" pageId="3" pageNumber="589">
<emphasis id="B91FEAE65F354937D4B38173FE8BA377" bold="true" box="[151,346,1853,1880]" pageId="3" pageNumber="589">FIGURES 9–18.</emphasis>
<taxonomicName id="4C6B4D775F354937D5468170FE28A378" authorityName="Latreille" authorityYear="1802" box="[354,505,1854,1880]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="genus">
<emphasis id="B91FEAE65F354937D5468170FE28A378" box="[354,505,1854,1880]" italics="true" pageId="3" pageNumber="589">Onthophagus</emphasis>
</taxonomicName>
species.
<emphasis id="B91FEAE65F354937D6448173FDBFA377" bold="true" box="[608,622,1853,1879]" pageId="3" pageNumber="589">9</emphasis>
, ¾ profile view, male of
<taxonomicName id="4C6B4D775F354937D7A88170FBF3A378" authorityName="Howden" authorityYear="1973" box="[908,1058,1854,1880]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F354937D7A88170FBF3A378" box="[908,1058,1854,1880]" italics="true" pageId="3" pageNumber="589">O. subcancer</emphasis>
</taxonomicName>
;
<emphasis id="B91FEAE65F354937D0158173FB9CA377" bold="true" box="[1073,1101,1853,1879]" pageId="3" pageNumber="589">10</emphasis>
, ¾ profile view, female of
<taxonomicName id="4C6B4D775F354937D1A58170FEDBA35C" authorityName="Howden" authorityYear="1973" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F354937D1A58170FEDBA35C" italics="true" pageId="3" pageNumber="589">O. subcancer</emphasis>
</taxonomicName>
;
<emphasis id="B91FEAE65F354937D53E812FFEE5A35B" bold="true" box="[282,308,1889,1915]" pageId="3" pageNumber="589">11</emphasis>
, lateral view of genitalia, male of
<taxonomicName id="4C6B4D775F354937D69D812CFC9EA35C" authorityName="Howden" authorityYear="1973" box="[697,847,1890,1916]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F354937D69D812CFC9EA35C" box="[697,847,1890,1916]" italics="true" pageId="3" pageNumber="589">O. subcancer</emphasis>
</taxonomicName>
;
<emphasis id="B91FEAE65F354937D77B812FFCAAA35B" bold="true" box="[863,891,1889,1915]" pageId="3" pageNumber="589">12</emphasis>
, dorsal view of genitalia, male of
<taxonomicName id="4C6B4D775F354937D0DB812CFA45A35C" authorityName="Howden" authorityYear="1973" box="[1279,1428,1890,1916]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F354937D0DB812CFA45A35C" box="[1279,1428,1890,1916]" italics="true" pageId="3" pageNumber="589">O. subcancer</emphasis>
</taxonomicName>
;
<emphasis id="B91FEAE65F354937D4B381CBFF62A3BF" bold="true" box="[151,179,1925,1951]" pageId="3" pageNumber="589">13</emphasis>
, protibia of
<taxonomicName id="4C6B4D775F354937D56481C8FDD2A380" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[320,515,1926,1952]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F354937D56481C8FDD2A380" box="[320,515,1926,1952]" italics="true" pageId="3" pageNumber="589">O. chimalapensis</emphasis>
</taxonomicName>
,
<typeStatus id="54D088565F354937D63681CBFDA9A3BF" box="[530,632,1925,1951]" pageId="3" pageNumber="589" type="holotype">holotype</typeStatus>
;
<emphasis id="B91FEAE65F354937D6A781CBFD4EA3BF" bold="true" box="[643,671,1925,1951]" pageId="3" pageNumber="589">14</emphasis>
, protibia of
<taxonomicName id="4C6B4D775F354937D70881C8FC21A380" box="[812,1008,1926,1952]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="aperodorsatus">
<emphasis id="B91FEAE65F354937D70881C8FC21A380" box="[812,1008,1926,1952]" italics="true" pageId="3" pageNumber="589">O. aperodorsatus</emphasis>
</taxonomicName>
, male
<typeStatus id="54D088565F354937D01981C8FB70A380" box="[1085,1185,1926,1952]" pageId="3" pageNumber="589" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F354937D08981CBFB18A3BF" bold="true" box="[1197,1225,1925,1951]" pageId="3" pageNumber="589">15</emphasis>
, protibia of
<taxonomicName id="4C6B4D775F354937D17181C8FEDBA3E4" authorityName="Howden" authorityYear="1973" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="neomirabilis">
<emphasis id="B91FEAE65F354937D17181C8FEDBA3E4" italics="true" pageId="3" pageNumber="589">O. neomirabilis</emphasis>
</taxonomicName>
, male;
<emphasis id="B91FEAE65F354937D57A81E7FEABA3E3" bold="true" box="[350,378,1961,1987]" pageId="3" pageNumber="589">16</emphasis>
, protibia of
<taxonomicName id="4C6B4D775F354937D62281E4FD19A3E4" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[518,712,1962,1988]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F354937D62281E4FD19A3E4" box="[518,712,1962,1988]" italics="true" pageId="3" pageNumber="589">O. chimalapensis</emphasis>
</taxonomicName>
, female
<typeStatus id="54D088565F354937D70E81E4FC5FA3E4" box="[810,910,1962,1988]" pageId="3" pageNumber="589" type="paratype">paratype</typeStatus>
;
<emphasis id="B91FEAE65F354937D7BE81E7FC67A3E3" bold="true" box="[922,950,1961,1987]" pageId="3" pageNumber="589">17</emphasis>
, protibia of
<taxonomicName id="4C6B4D775F354937D06581E4FAC1A3E4" authorityName="Howden &amp; Gill" authorityYear="1993" box="[1089,1296,1962,1988]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="asperodorsatus">
<emphasis id="B91FEAE65F354937D06581E4FAC1A3E4" box="[1089,1296,1962,1988]" italics="true" pageId="3" pageNumber="589">O. asperodorsatus</emphasis>
</taxonomicName>
, female;
<emphasis id="B91FEAE65F354937D15D81E7FA44A3E3" bold="true" box="[1401,1429,1961,1987]" pageId="3" pageNumber="589">18</emphasis>
, protibia of
<taxonomicName id="4C6B4D775F354937D5308180FE15A3C8" authorityName="Howden" authorityYear="1973" box="[276,452,1998,2024]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="589" phylum="Arthropoda" rank="species" species="neomirabilis">
<emphasis id="B91FEAE65F354937D5308180FE15A3C8" box="[276,452,1998,2024]" italics="true" pageId="3" pageNumber="589">O. neomirabilis</emphasis>
</taxonomicName>
, female.
</paragraph>
</caption>
<caption id="DF14667C5F324930D4B38296FAEDA137" ID-DOI="http://doi.org/10.5281/zenodo.3613728" ID-Zenodo-Dep="3613728" httpUri="https://zenodo.org/record/3613728/files/figure.png" pageId="4" pageNumber="590" startId="4.[151,260,1240,1267]" targetBox="[165,1421,178,1203]" targetPageId="4">
<paragraph id="8BD436F45F324930D4B38296FAEDA137" blockId="4.[151,1436,1240,1303]" pageId="4" pageNumber="590">
<emphasis id="B91FEAE65F324930D4B38296FEFCA0D3" bold="true" box="[151,301,1240,1267]" pageId="4" pageNumber="590">FIGURE 19.</emphasis>
Distribution of
<taxonomicName id="4C6B4D775F324930D5FB8297FDA7A0D3" authorityName="Latreille" authorityYear="1802" box="[479,630,1241,1267]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="genus">
<emphasis id="B91FEAE65F324930D5FB8297FDA7A0D3" box="[479,630,1241,1267]" italics="true" pageId="4" pageNumber="590">Onthophagus</emphasis>
</taxonomicName>
species.
<taxonomicName id="4C6B4D775F324930D6FE8297FBC6A0D3" authority="Delgado &amp; Mora-Aguilar, 2019" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[730,1047,1241,1267]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="chimalapensis" status="sp. nov.">
<emphasis id="B91FEAE65F324930D6FE8297FBC6A0D3" box="[730,1047,1241,1267]" italics="true" pageId="4" pageNumber="590">Onthophagus chimalapensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A22C579D5F324930D03A8297FB72A0D3" box="[1054,1187,1241,1267]" pageId="4" pageNumber="590" rank="species">new species</taxonomicNameLabel>
(circle),
<taxonomicName id="4C6B4D775F324930D12C8297FA4DA0D3" authorityName="Howden" authorityYear="1973" box="[1288,1436,1241,1267]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F324930D12C8297FA4DA0D3" box="[1288,1436,1241,1267]" italics="true" pageId="4" pageNumber="590">O. subcancer</emphasis>
</taxonomicName>
(square). Soft to dark grey lines represent elevational gradients of
<geoCoordinate id="EE5F50335F324930D75882B3FC6FA137" box="[892,958,1277,1303]" degrees="1,000" direction="north" orientation="latitude" pageId="4" pageNumber="590" precision="55" value="1.0">1,000</geoCoordinate>
,
<geoCoordinate id="EE5F50335F324930D7ED82B3FBDAA137" box="[969,1035,1277,1303]" degrees="2,000" direction="east" orientation="longitude" pageId="4" pageNumber="590" precision="55" value="2.0">2,000</geoCoordinate>
, and
<quantity id="4C939B115F324930D06282B3FB75A137" box="[1094,1188,1277,1303]" metricMagnitude="0" metricUnit="m" metricValue="3.0" pageId="4" pageNumber="590" unit="m" value="3.0">3,000 m</quantity>
, respectively.
</paragraph>
</caption>
<paragraph id="8BD436F45F324930D4E3830BFC4EA3C7" blockId="4.[151,1437,1349,2023]" pageId="4" pageNumber="590">
Since the separation of the
<taxonomicName id="4C6B4D775F324930D5D0830BFDAEA17F" authorityName="Bates" authorityYear="1887" box="[500,639,1349,1375]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D5D0830BFDAEA17F" box="[500,639,1349,1375]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6B4D775F324930D690830BFCEAA17F" authorityName="Bates" authorityYear="1887" box="[692,827,1349,1375]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="mirabilis">
<emphasis id="B91FEAE65F324930D690830BFCEAA17F" box="[692,827,1349,1375]" italics="true" pageId="4" pageNumber="590">O. mirabilis</emphasis>
</taxonomicName>
species groups from the
<taxonomicName id="4C6B4D775F324930D074830BFB0AA17F" box="[1104,1243,1349,1375]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="clypeatus">
<emphasis id="B91FEAE65F324930D074830BFB0AA17F" box="[1104,1243,1349,1375]" italics="true" pageId="4" pageNumber="590">O. clypeatus</emphasis>
</taxonomicName>
species group by
<bibRefCitation id="EFFA4B055F324930D4B38327FE4AA1A3" author="Howden, H. F. &amp; Gill, B. D." box="[151,411,1385,1411]" pageId="4" pageNumber="590" pagination="1091 - 1114" refId="ref4346" refString="Howden, H. F. &amp; Gill, B. D. (1993) Mesoamerican Onthophagus Latreille in the dicranius and mirabilis species group (Coleoptera: Scarabaeidae). The Canadian Entomologist, 125, 1091 - 1114. https: // doi. org / 10.4039 / Ent 1251091 - 6" type="journal article" year="1993">Howden &amp; Gill (1993)</bibRefCitation>
, there have been different views in considering only one or two groups.
<bibRefCitation id="EFFA4B055F324930D0E88327FF33A187" author="Genier, F. &amp; Howden, H. F." pageId="4" pageNumber="590" pagination="130 - 144" refId="ref4262" refString="Genier, F. &amp; Howden, H. F. (1999) Two new Central American Onthophagus Latreille of the mirabilis species group (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 53, 130 - 144." type="journal article" year="1999">Génier &amp; Howden (1999)</bibRefCitation>
redefined these groupings by using the colour of the body, structure of protibiae, and morphological differences observed on the head and pronotum of females. As suggested by
<bibRefCitation id="EFFA4B055F324930D78A83FFFB1CA1EB" author="Genier, F. &amp; Howden, H. F." box="[942,1229,1457,1483]" pageId="4" pageNumber="590" pagination="130 - 144" refId="ref4262" refString="Genier, F. &amp; Howden, H. F. (1999) Two new Central American Onthophagus Latreille of the mirabilis species group (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 53, 130 - 144." type="journal article" year="1999">Génier &amp; Howden (1999)</bibRefCitation>
and more recently by
<bibRefCitation id="EFFA4B055F324930D49E839BFE8BA1CF" author="Genier, F." box="[186,346,1493,1519]" pageId="4" pageNumber="590" pagination="574 - 580" refId="ref4215" refString="Genier, F. (2017) A new Guatemalan cloud forest endemic Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae). The Canadian Entomologist, 149, 574 - 580. https: // doi. org / 10.4039 / tce. 2017.32" type="journal article" year="2017">Génier (2017)</bibRefCitation>
, we consider the
<taxonomicName id="4C6B4D775F324930D639839BFD78A1CF" authorityName="Bates" authorityYear="1887" box="[541,681,1493,1519]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D639839BFD78A1CF" box="[541,681,1493,1519]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6B4D775F324930D6FB839BFCB6A1CF" authorityName="Bates" authorityYear="1887" box="[735,871,1493,1519]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="mirabilis">
<emphasis id="B91FEAE65F324930D6FB839BFCB6A1CF" box="[735,871,1493,1519]" italics="true" pageId="4" pageNumber="590">O. mirabilis</emphasis>
</taxonomicName>
as species complexes within the
<taxonomicName id="4C6B4D775F324930D0FF839BFAB6A1CF" authorityName="Bates" authorityYear="1887" box="[1243,1383,1493,1519]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D0FF839BFAB6A1CF" box="[1243,1383,1493,1519]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
species group. In this regard,
<taxonomicName id="4C6B4D775F324930D59A83B7FD82A233" authorityName="Howden" authorityYear="1973" box="[446,595,1529,1555]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F324930D59A83B7FD82A233" box="[446,595,1529,1555]" italics="true" pageId="4" pageNumber="590">O. subcancer</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6B4D775F324930D6AE83B7FC9CA233" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[650,845,1529,1555]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F324930D6AE83B7FC9CA233" box="[650,845,1529,1555]" italics="true" pageId="4" pageNumber="590">O. chimalapensis</emphasis>
</taxonomicName>
are placed into the
<taxonomicName id="4C6B4D775F324930D00E83B7FB66A233" authorityName="Bates" authorityYear="1887" box="[1066,1207,1529,1555]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D00E83B7FB66A233" box="[1066,1207,1529,1555]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
species complex, as both species exhibit the body bicolored, and the two apical teeth of protibia widely separated (joined only at the base); not the body concolor, and the two apical teeth of protibia joined beyond the base (
<figureCitation id="13502A715F324930D0B5800FFACDA27B" box="[1169,1308,1601,1627]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="4" pageNumber="590">Figs. 13–18</figureCitation>
). However, females of these two species, like the species of the
<taxonomicName id="4C6B4D775F324930D6C4802BFCB6A25F" authorityName="Bates" authorityYear="1887" box="[736,871,1637,1663]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="mirabilis">
<emphasis id="B91FEAE65F324930D6C4802BFCB6A25F" box="[736,871,1637,1663]" italics="true" pageId="4" pageNumber="590">O. mirabilis</emphasis>
</taxonomicName>
species complex, show remarkable differences on head and pronotum (
<figureCitation id="13502A715F324930D5A480C7FDD9A283" box="[384,520,1673,1699]" captionStart="FIGURES 1–8" captionStartId="2.[151,275,1940,1967]" captionTargetBox="[183,1411,197,1903]" captionTargetId="figure@2.[151,1436,164,1920]" captionText="FIGURES 1–8. Onthophagus chimalapensis. 1, Habitus, holotype; 2, habitus, female paratype; 3, lateral view of male paratype; 4, lateral view of female paratype; 5, ¾ profile view of male holotype; 6, ¾ profile view of female paratype; 7, lateral view of genitalia, male paratype; 8, dorsal view of genitalia, male paratype." figureDoi="http://doi.org/10.5281/zenodo.3613724" httpUri="https://zenodo.org/record/3613724/files/figure.png" pageId="4" pageNumber="590">Figs. 2, 4, 6</figureCitation>
,
<figureCitation id="13502A715F324930D63080C7FDE2A283" box="[532,563,1673,1699]" captionStart="FIGURES 9–18" captionStartId="3.[151,275,1853,1880]" captionTargetBox="[192,1415,190,1785]" captionTargetId="figure@3.[151,1435,164,1833]" captionText="FIGURES 9–18. Onthophagus species. 9, ¾ profile view, male of O. subcancer; 10, ¾ profile view, female of O. subcancer; 11, lateral view of genitalia, male of O. subcancer; 12, dorsal view of genitalia, male of O. subcancer; 13, protibia of O. chimalapensis, holotype; 14, protibia of O. aperodorsatus, male paratype; 15, protibia of O. neomirabilis, male; 16, protibia of O. chimalapensis, female paratype; 17, protibia of O. asperodorsatus, female; 18, protibia of O. neomirabilis, female." figureDoi="http://doi.org/10.5281/zenodo.3613726" httpUri="https://zenodo.org/record/3613726/files/figure.png" pageId="4" pageNumber="590">10</figureCitation>
). Thus, the
<taxonomicName id="4C6B4D775F324930D69380C7FC95A283" authorityName="Bates" authorityYear="1887" box="[695,836,1673,1699]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D69380C7FC95A283" box="[695,836,1673,1699]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
species complex comprises the following species:
<taxonomicName id="4C6B4D775F324930D1A580C7FDD0A2E7" authority="Howden &amp; Gill" authorityName="Howden &amp; Gill" authorityYear="1993" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="asperodorsatus">
<emphasis id="B91FEAE65F324930D1A580C7FE92A2E7" italics="true" pageId="4" pageNumber="590">O. asperodorsatus</emphasis>
Howden &amp; Gill
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D62B80E3FD02A2E7" authorityName="Delgado &amp; Mora-Aguilar" authorityYear="2019" box="[527,723,1709,1735]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="chimalapensis">
<emphasis id="B91FEAE65F324930D62B80E3FD02A2E7" box="[527,723,1709,1735]" italics="true" pageId="4" pageNumber="590">O. chimalapensis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D6C080E3FB4DA2E7" authority="Kohlmann &amp; Solis" authorityName="Kohlmann &amp; Solis" authorityYear="2001" box="[740,1180,1709,1735]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="cryptodicranius">
<emphasis id="B91FEAE65F324930D6C080E3FC68A2E7" box="[740,953,1709,1735]" italics="true" pageId="4" pageNumber="590">O. cryptodicranius</emphasis>
Kohlmann &amp; Solís
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D08E80E3FAE9A2E7" authorityName="Bates" authorityYear="1887" box="[1194,1336,1709,1735]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D08E80E3FAE9A2E7" box="[1194,1336,1709,1735]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D16C80E3FF27A2CB" authorityName="Howden &amp; Gill" authorityYear="1987" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dorsipilulus">
<emphasis id="B91FEAE65F324930D16C80E3FF27A2CB" italics="true" pageId="4" pageNumber="590">O. dorsipilulus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D522809FFD47A2CB" authority="Kohlmann &amp; Solis" authorityName="Kohlmann &amp; Solis" authorityYear="2001" box="[262,662,1745,1771]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="inediapterus">
<emphasis id="B91FEAE65F324930D522809FFE64A2CB" box="[262,437,1745,1771]" italics="true" pageId="4" pageNumber="590">O. inediapterus</emphasis>
Kohlmann &amp; Solís
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D687809FFC42A2CB" authority="Genier" authorityName="Genier" authorityYear="2017" box="[675,915,1745,1771]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="kohlmanni">
<emphasis id="B91FEAE65F324930D687809FFCECA2CB" box="[675,829,1745,1771]" italics="true" pageId="4" pageNumber="590">O. kohlmanni</emphasis>
Génier
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D784809FFB26A2CB" authority="Kohlmann &amp; Solis" authorityName="Kohlmann &amp; Solis" authorityYear="2001" box="[928,1271,1745,1771]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="nubilus">
<emphasis id="B91FEAE65F324930D784809FFBC7A2CB" box="[928,1046,1745,1771]" italics="true" pageId="4" pageNumber="590">O. nubilus</emphasis>
Kohlmann &amp; Solís
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D120809FFE9AA32F" authority="Howden &amp; Gill" authorityName="Howden &amp; Gill" authorityYear="1993" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="petenensis">
<emphasis id="B91FEAE65F324930D120809FFA4CA2CB" box="[1284,1437,1745,1771]" italics="true" pageId="4" pageNumber="590">O. petenensis</emphasis>
Howden &amp; Gill
</taxonomicName>
,
<taxonomicName id="4C6B4D775F324930D57C80BBFD4AA32F" authority="Howden &amp; Gill" authorityName="Howden &amp; Gill" authorityYear="1993" box="[344,667,1781,1807]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="quetzalis">
<emphasis id="B91FEAE65F324930D57C80BBFE0EA32F" box="[344,479,1781,1807]" italics="true" pageId="4" pageNumber="590">O. quetzalis</emphasis>
Howden &amp; Gill
</taxonomicName>
, and
<taxonomicName id="4C6B4D775F324930D6FC80BBFCBEA32F" authorityName="Howden" authorityYear="1973" box="[728,879,1781,1807]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="subcancer">
<emphasis id="B91FEAE65F324930D6FC80BBFCBEA32F" box="[728,879,1781,1807]" italics="true" pageId="4" pageNumber="590">O. subcancer</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFFA4B055F324930D75B80BBFBA4A32F" author="Howden, H. F. &amp; Gill, B. D." box="[895,1141,1781,1807]" pageId="4" pageNumber="590" pagination="1091 - 1114" refId="ref4346" refString="Howden, H. F. &amp; Gill, B. D. (1993) Mesoamerican Onthophagus Latreille in the dicranius and mirabilis species group (Coleoptera: Scarabaeidae). The Canadian Entomologist, 125, 1091 - 1114. https: // doi. org / 10.4039 / Ent 1251091 - 6" type="journal article" year="1993">Howden &amp; Gill 1993</bibRefCitation>
;
<bibRefCitation id="EFFA4B055F324930D0A680BBFA46A32F" author="Genier, F. &amp; Howden, H. F." box="[1154,1431,1781,1807]" pageId="4" pageNumber="590" pagination="130 - 144" refId="ref4262" refString="Genier, F. &amp; Howden, H. F. (1999) Two new Central American Onthophagus Latreille of the mirabilis species group (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 53, 130 - 144." type="journal article" year="1999">Génier &amp; Howden 1999</bibRefCitation>
;
<bibRefCitation id="EFFA4B055F324930D4B38157FE61A313" author="Kohlmann, B. &amp; Solis, A." box="[151,432,1817,1843]" pageId="4" pageNumber="590" pagination="159 - 261" refId="ref4401" refString="Kohlmann, B. &amp; Solis, A. (2001) El genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, 49, 159 - 261." type="journal article" year="2001">Kohlmann &amp; Solís 2001</bibRefCitation>
;
<bibRefCitation id="EFFA4B055F324930D5998157FD9EA313" author="Genier, F." box="[445,591,1817,1843]" pageId="4" pageNumber="590" pagination="574 - 580" refId="ref4215" refString="Genier, F. (2017) A new Guatemalan cloud forest endemic Onthophagus Latreille, 1802 (Coleoptera: Scarabaeidae: Scarabaeinae). The Canadian Entomologist, 149, 574 - 580. https: // doi. org / 10.4039 / tce. 2017.32" type="journal article" year="2017">Génier 2017</bibRefCitation>
). Although
<taxonomicName id="4C6B4D775F324930D6F78157FC06A313" authorityName="Howden &amp; Gill" authorityYear="1993" box="[723,983,1817,1843]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="quetzalis">
<emphasis id="B91FEAE65F324930D6F78157FC06A313" box="[723,983,1817,1843]" italics="true" pageId="4" pageNumber="590">Onthophagus quetzalis</emphasis>
</taxonomicName>
was initially considered into the
<taxonomicName id="4C6B4D775F324930D1778157FF31A377" authorityName="Bates" authorityYear="1887" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="mirabilis">
<emphasis id="B91FEAE65F324930D1778157FF31A377" italics="true" pageId="4" pageNumber="590">O. mirabilis</emphasis>
</taxonomicName>
species complex (
<bibRefCitation id="EFFA4B055F324930D5948173FD75A377" author="Howden, H. F. &amp; Gill, B. D." box="[432,676,1853,1879]" pageId="4" pageNumber="590" pagination="1091 - 1114" refId="ref4346" refString="Howden, H. F. &amp; Gill, B. D. (1993) Mesoamerican Onthophagus Latreille in the dicranius and mirabilis species group (Coleoptera: Scarabaeidae). The Canadian Entomologist, 125, 1091 - 1114. https: // doi. org / 10.4039 / Ent 1251091 - 6" type="journal article" year="1993">Howden &amp; Gill 1993</bibRefCitation>
), after was included in the
<taxonomicName id="4C6B4D775F324930D7F38173FBB5A377" authorityName="Bates" authorityYear="1887" box="[983,1124,1853,1879]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D7F38173FBB5A377" box="[983,1124,1853,1879]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
species complex by
<bibRefCitation id="EFFA4B055F324930D1748173FEB5A35B" author="Genier, F. &amp; Howden, H. F." pageId="4" pageNumber="590" pagination="130 - 144" refId="ref4262" refString="Genier, F. &amp; Howden, H. F. (1999) Two new Central American Onthophagus Latreille of the mirabilis species group (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 53, 130 - 144." type="journal article" year="1999">Génier &amp; Howden (1999)</bibRefCitation>
this change was supported by the morphological characters previously mentioned. Unlike
<bibRefCitation id="EFFA4B055F324930D17F812FFE5AA3BF" author="Kohlmann, B. &amp; Solis, A." pageId="4" pageNumber="590" pagination="159 - 261" refId="ref4401" refString="Kohlmann, B. &amp; Solis, A. (2001) El genero Onthophagus (Coleoptera: Scarabaeidae) en Costa Rica. Giornale Italiano di Entomologia, 49, 159 - 261." type="journal article" year="2001">Kohlmann &amp; Solís (2001)</bibRefCitation>
, we remove from this complex
<taxonomicName id="4C6B4D775F324930D6DE81CBFB2DA3BF" authority="Zunino &amp; Halffter" authorityName="Zunino &amp; Halffter" authorityYear="1981" box="[762,1276,1925,1951]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="micropterus">
<emphasis id="B91FEAE65F324930D6DE81CBFBF1A3BF" box="[762,1056,1925,1951]" italics="true" pageId="4" pageNumber="590">Onthophagus micropterus</emphasis>
Zunino &amp; Halffter
</taxonomicName>
, as males and females have conspicuous tubercles aligned with the midline of the eyes, not head destitute of tubercles or horns at this region, as in the others species of the
<taxonomicName id="4C6B4D775F324930D64F8183FD29A3C7" authorityName="Bates" authorityYear="1887" box="[619,760,1997,2023]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="590" phylum="Arthropoda" rank="species" species="dicranius">
<emphasis id="B91FEAE65F324930D64F8183FD29A3C7" box="[619,760,1997,2023]" italics="true" pageId="4" pageNumber="590">O. dicranius</emphasis>
</taxonomicName>
species group.
</paragraph>
</subSubSection>
</treatment>
</document>
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