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<document id="E86A04670145936A5D5A8BB08C23839C" ID-CLB-Dataset="25961" ID-DOI="10.11646/zootaxa.4658.1.1" ID-GBIF-Dataset="31d6e081-95c3-4086-a45a-c11555dda3aa" ID-ISSN="1175-5326" ID-Zenodo-Dep="3373040" ID-ZooBank="ADCBCCA2-63C1-42CD-A3C9-C14F954F4A21" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1566364910789" checkinUser="plazi" docAuthor="Ziani, Stefano, Abdel-Dayem, Mahmoud Saleh, Aldhafer, Hathal Mohammed &amp; Barbero, Enrico" docDate="2019" docId="8861532B88461837FF7609B6FDB6C5B8" docLanguage="en" docName="zootaxa.4658.1.1.pdf" docOrigin="Zootaxa 4658 (1)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Onthophagus (Onthophagus) falsus Gillet 1925" docType="treatment" docVersion="13" lastPageNumber="22" masterDocId="74582B5388531822FFE10957FFA3C74B" masterDocTitle="An overview of the Onthophagini from the Arabian Peninsula (Coleoptera: Scarabaeoidea: Scarabaeidae)" masterLastPageNumber="36" masterPageNumber="1" pageNumber="22" updateTime="1698751869822" updateUser="ExternalLinkService">
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<mods:title id="01927F1B425AB0782BEF9DB0526EE5FF">An overview of the Onthophagini from the Arabian Peninsula (Coleoptera: Scarabaeoidea: Scarabaeidae)</mods:title>
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<mods:namePart id="B607379FDED5407C79E0B7205D064AC6">Ziani, Stefano</mods:namePart>
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<mods:namePart id="72719F2D86A77C8A4CAC7C90CFBEFCD1">Abdel-Dayem, Mahmoud Saleh</mods:namePart>
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<mods:namePart id="1459D829DD4F21686F972BBFA95E0475">Aldhafer, Hathal Mohammed</mods:namePart>
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<mods:namePart id="5B7CCF2FF0766DC06473CC68BE50D136">Barbero, Enrico</mods:namePart>
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<treatment id="8861532B88461837FF7609B6FDB6C5B8" ID-DOI="http://doi.org/10.5281/zenodo.5667086" ID-GBIF-Taxon="159217244" ID-Zenodo-Dep="5667086" LSID="urn:lsid:plazi:treatment:8861532B88461837FF7609B6FDB6C5B8" httpUri="http://treatment.plazi.org/id/8861532B88461837FF7609B6FDB6C5B8" lastPageNumber="22" pageId="21" pageNumber="22">
<subSubSection id="48D2B1B688461837FF7609B6FD41C7B0" box="[151,738,225,252]" pageId="21" pageNumber="22" type="nomenclature">
<paragraph id="0077E23D88461837FF7609B6FD41C7B0" blockId="21.[151,738,225,288]" box="[151,738,225,252]" pageId="21" pageNumber="22">
<heading id="5B3F555188461837FF7609B6FD41C7B0" bold="true" box="[151,738,225,252]" fontSize="11" level="1" pageId="21" pageNumber="22" reason="1">
<taxonomicName id="C7C899BE88461837FF7609B6FD41C7B0" ID-CoL="7XMVB" authority="Gillet, 1925" authorityName="Gillet" authorityYear="1925" box="[151,738,225,252]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="falsus" subGenus="Onthophagus">
<emphasis id="32BC3E2F88461837FF7609B6FD41C7B0" bold="true" box="[151,738,225,252]" pageId="21" pageNumber="22">
<emphasis id="32BC3E2F88461837FF7609B6FE9FC7B7" bold="true" box="[151,316,225,252]" italics="true" pageId="21" pageNumber="22">Onthophagus</emphasis>
(
<emphasis id="32BC3E2F88461837FEAC09B6FE51C7B7" bold="true" box="[333,498,225,252]" italics="true" pageId="21" pageNumber="22">Onthophagus</emphasis>
)
<emphasis id="32BC3E2F88461837FDE209B6FDE8C7B7" bold="true" box="[515,587,225,252]" italics="true" pageId="21" pageNumber="22">falsus</emphasis>
Gillet, 1925
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="48D2B1B688461837FF760852FF5AC66B" box="[151,249,261,288]" pageId="21" pageNumber="22" type="description">
<paragraph id="0077E23D88461837FF760852FF5AC66B" blockId="21.[151,738,225,288]" box="[151,249,261,288]" pageId="21" pageNumber="22">
[
<figureCitation id="98F3FEB888461837FF7F0852FF51C66B" box="[158,242,261,288]" captionStart="FIGURES 1418" captionStartId="23.[151,264,1872,1897]" captionTargetBox="[293,1306,179,1808]" captionTargetId="figure@23.[243,1344,166,1852]" captionTargetPageId="23" captionText="FIGURES 1418. Onthophagini of Arabian Peninsula. 14, Onthophagus (Micronthophagus) ochreatus dOrbigny, 1898: YE (Al Hudaydah) 10 km W Al Mansuriyah; 15, Onthophagus (Onthophagiellus) millingeni dOrbigny, 1898: YE (Sanaa) Haddah; 16, Onthophagus (s. str.) falsus Gillet, 1925: SA (Riyadh) Wadi Al Dawaser; 17, Onthophagus (Proagoderus) quadriarmatus Fairmaire, 1892, ♂: YE (Al Mahrah) Wadi N of Damqut vill.; 18, Onthophagus (Proagoderus) quadriarmatus Fairmaire, 1892, ♀: YE (Al Mahrah) Wadi N of Damqut vill. Photos by Gabriele Fiumi." httpUri="https://zenodo.org/record/3373051/files/figure.png" pageId="21" pageNumber="22">Fig. 16</figureCitation>
]
</paragraph>
</subSubSection>
<subSubSection id="48D2B1B688461837FF76081AFDB6C5B8" pageId="21" pageNumber="22" type="materials_examined">
<paragraph id="0077E23D88461837FF76081AFE19C623" blockId="21.[151,1437,333,756]" box="[151,442,333,360]" pageId="21" pageNumber="22">
<emphasis id="32BC3E2F88461837FF76081AFECBC623" bold="true" box="[151,360,333,360]" pageId="21" pageNumber="22">Literature quotes</emphasis>
. None.
</paragraph>
<paragraph id="0077E23D88461837FF260826FAA5C698" blockId="21.[151,1437,333,756]" pageId="21" pageNumber="22">
<emphasis id="32BC3E2F88461837FF260826FE0AC6C7" bold="true" box="[199,425,369,396]" pageId="21" pageNumber="22">Material examined</emphasis>
.
<materialsCitation id="B0A0E86088461837FE570826FE93C6E4" ID-GBIF-Occurrence="2350485848" collectingDate="19. xi." collectionCode="KSMA" collectorName="Alfalfa, M. &amp; M. Ur Rehman" country="Saudi Arabia" location="Al-Wasiel" pageId="21" pageNumber="22" specimenCount="1" specimenCount-male="1" stateProvince="Ar Riyad">
<collectingCountry id="78DFA2AD88461837FE570826FDECC6C2" box="[438,591,369,396]" name="Saudi Arabia" pageId="21" pageNumber="22">
<emphasis id="32BC3E2F88461837FE570826FE54C6C2" bold="true" box="[438,503,369,396]" pageId="21" pageNumber="22">SAUDI</emphasis>
<taxonomicName id="C7C899BE88461837FE1D0826FDECC6C2" box="[508,591,369,396]" class="Insecta" family="Cerambycidae" genus="Arabia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="35" phylum="Arthropoda" rank="genus">
<emphasis id="32BC3E2F88461837FE1D0826FDECC6C2" bold="true" box="[508,591,369,396]" pageId="21" pageNumber="22">ARABIA</emphasis>
</taxonomicName>
</collectingCountry>
:
<collectingRegion id="C20C2CDF88461837FDBF0826FCB8C6C0" box="[606,795,369,395]" country="Saudi Arabia" name="Ar Riyad" pageId="21" pageNumber="22">Riyadh Province</collectingRegion>
,
<location id="0517B4E688461837FCC50826FC3BC6C0" LSID="urn:lsid:plazi:treatment:8861532B88461837FF7609B6FDB6C5B8:0517B4E688461837FCC50826FC3BC6C0" box="[804,920,369,395]" country="Saudi Arabia" name="Al-Wasiel" pageId="21" pageNumber="22" stateProvince="Ar Riyad">Al-Wasiel</location>
,
<date id="7476C4FD88461837FC430826FC7CC6C7" box="[930,991,369,396]" pageId="21" pageNumber="22">
<collectingDate id="64323D1588461837FC430826FC7CC6C7" box="[930,991,369,396]" pageId="21" pageNumber="22">19.xi.</collectingDate>
</date>
205, SW on
<collectorName id="AD3D87EB88461837FB880826FB45C6C7" box="[1129,1254,369,396]" pageId="21" pageNumber="22">Alfalfa, M.</collectorName>
Ur Rehman leg.
<specimenCount id="16CE29B488461837FF7608C2FF61C6FB" box="[151,194,405,432]" pageId="21" pageNumber="22" type="male">1 ♂</specimenCount>
(
<collectionCode id="66D97AF888461837FF3008C2FE8BC6E4" box="[209,296,405,431]" pageId="21" pageNumber="22">KSMA</collectionCode>
)
</materialsCitation>
;
<materialsCitation id="B0A0E86088461837FEDD08C2FE10C698" ID-GBIF-Occurrence="2350486019" collectingDate="2018-05-09" collectionCode="KSMA" collectorName="H. Al Dhafer" country="Saudi Arabia" elevation="700" location="Al Muhmmadiya Farm" municipality="Wadi Al Dawaser" pageId="21" pageNumber="22" specimenCount="2" specimenCount-male="2" stateProvince="Riyadh Province">
<collectingMunicipality id="E013784788461837FEDD08C2FDA7C6FB" box="[316,516,405,432]" pageId="21" pageNumber="22">Wadi Al Dawaser</collectingMunicipality>
,
<location id="0517B4E688461837FDEF08C2FCB7C6FB" LSID="urn:lsid:plazi:treatment:8861532B88461837FF7609B6FDB6C5B8:0517B4E688461837FDEF08C2FCB7C6FB" box="[526,788,405,432]" country="Saudi Arabia" municipality="Wadi Al Dawaser" name="Al Muhmmadiya Farm" pageId="21" pageNumber="22" stateProvince="Riyadh Province">Al Muhmmadiya Farm</location>
,
<quantity id="C7304FD888461837FCC108C2FCC4C6FB" box="[800,871,405,432]" metricMagnitude="2" metricUnit="m" metricValue="7.0" pageId="21" pageNumber="22" unit="m" value="700.0">
<elevation id="8BE5050E88461837FCC108C2FCC4C6FB" box="[800,871,405,432]" metricMagnitude="2" metricUnit="m" metricValue="7.0" pageId="21" pageNumber="22" unit="m" value="700.0">700 m</elevation>
</quantity>
(2025.497N 4443.563E),
<date id="7476C4FD88461837FB7A08C2FB5EC6E4" box="[1179,1277,405,431]" pageId="21" pageNumber="22" value="2018-05-09">
<collectingDate id="64323D1588461837FB7A08C2FB5EC6E4" box="[1179,1277,405,431]" pageId="21" pageNumber="22" value="2018-05-09">9.v.2018</collectingDate>
</date>
,
<collectorName id="AD3D87EB88461837FAEB08C1FA3EC6E4" box="[1290,1437,405,432]" pageId="21" pageNumber="22">H. Al Dhafer</collectorName>
<emphasis id="32BC3E2F88461837FF7608EDFF6BC698" box="[151,200,441,467]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
. leg.
<specimenCount id="16CE29B488461837FEE708EEFEE6C69F" box="[262,325,441,468]" pageId="21" pageNumber="22" type="male">2 ♂♂</specimenCount>
(
<collectionCode id="66D97AF888461837FEB508EEFE08C698" box="[340,427,441,467]" pageId="21" pageNumber="22">KSMA</collectionCode>
)
</materialsCitation>
;
<materialsCitation id="B0A0E86088461837FE5F08EEFAA1C698" ID-GBIF-Occurrence="2350485859" box="[446,1282,441,468]" collectingDate="2015-11-14" collectionCode="KSMA" collectorName="E. M. Khan" country="Saudi Arabia" location="Wadi Hanifa" pageId="21" pageNumber="22" specimenCount="2" specimenCount-male="2" stateProvince="Riyadh Province">
<location id="0517B4E688461837FE5F08EEFDECC698" LSID="urn:lsid:plazi:treatment:8861532B88461837FF7609B6FDB6C5B8:0517B4E688461837FE5F08EEFDECC698" box="[446,591,441,467]" country="Saudi Arabia" name="Wadi Hanifa" pageId="21" pageNumber="22" stateProvince="Riyadh Province">Wadi Hanifa</location>
,
<date id="7476C4FD88461837FDBB08EEFD77C69F" box="[602,724,441,468]" pageId="21" pageNumber="22" value="2015-11-14">
<collectingDate id="64323D1588461837FDBB08EEFD77C69F" box="[602,724,441,468]" pageId="21" pageNumber="22" value="2015-11-14">14.xi.2015</collectingDate>
</date>
, HP in date palm,
<collectorName id="AD3D87EB88461837FC4308EDFBBDC698" box="[930,1054,441,468]" pageId="21" pageNumber="22">E.M. Khan</collectorName>
leg.
<specimenCount id="16CE29B488461837FBB408EEFB37C69F" box="[1109,1172,441,468]" pageId="21" pageNumber="22" type="male">2 ♂♂</specimenCount>
(
<collectionCode id="66D97AF888461837FB4208EEFB59C698" box="[1187,1274,441,467]" pageId="21" pageNumber="22">KSMA</collectionCode>
)
</materialsCitation>
.
</paragraph>
<paragraph id="0077E23D88461837FF26088AFBCFC550" blockId="21.[151,1437,333,756]" pageId="21" pageNumber="22">
<emphasis id="32BC3E2F88461837FF26088AFEFAC6B3" bold="true" box="[199,345,477,504]" pageId="21" pageNumber="22">Distribution</emphasis>
.
<emphasis id="32BC3E2F88461837FE87088AFE38C6B3" bold="true" box="[358,411,477,504]" pageId="21" pageNumber="22">Asia</emphasis>
:
<collectingCountry id="78DFA2AD88461837FE46088AFD93C6BC" box="[423,560,477,503]" name="Afghanistan" pageId="21" pageNumber="22">Afghanistan</collectingCountry>
(
<bibRefCitation id="64599FCC88461837FDDC088AFD4AC6B3" author="Balthasar, V." box="[573,745,477,504]" pageId="21" pageNumber="22" refId="ref36564" refString="Balthasar, V. (1963) Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band 2. Coprinae. Tschechoslowakischen Akademie der Wissenschaften, Prag, 628 pp." type="book" year="1963">Balthasar 1963</bibRefCitation>
), central and northern
<collectingCountry id="78DFA2AD88461837FC01088AFBB9C6BC" box="[992,1050,477,503]" name="India" pageId="21" pageNumber="22">India</collectingCountry>
(
<bibRefCitation id="64599FCC88461837FBC7088AFB11C6BC" author="Arrow, G. J." box="[1062,1202,477,503]" pageId="21" pageNumber="22" refId="ref36526" refString="Arrow, G. J. (1931) The Fauna of British India, including Ceylon and Burma. Coleoptera Lamellicornia. Part III (Coprinae). Taylor &amp; Francis, London, 428 pp." type="book" year="1931">Arrow 1931</bibRefCitation>
;
<bibRefCitation id="64599FCC88461837FB5D088AFA33C6B3" author="Mittal, I. C. &amp; Jain, R." box="[1212,1424,477,504]" pageId="21" pageNumber="22" pagination="383 - 404" refId="ref38496" refString="Mittal, I. C. &amp; Jain, R. (2015) A Checklist Of Indian Dung Beetles (Coleoptera: Scarabaeidae). Indian Journal of Entomology, 77 (4), 383 - 404. https: // doi. org / 10.5958 / 0974 - 8172.2015.00076. 0" type="journal article" year="2015">Mittal &amp; Jain 2015</bibRefCitation>
), south-western
<collectingCountry id="78DFA2AD88461837FEDA0B56FED6C550" box="[315,373,513,539]" name="India" pageId="21" pageNumber="22">India</collectingCountry>
(
<bibRefCitation id="64599FCC88461837FE650B56FDE7C550" author="Sabu, T. K. &amp; Nithya, S. &amp; Vinod, K. V." box="[388,580,513,540]" pageId="21" pageNumber="22" pagination="29 - 38" refId="ref39367" refString="Sabu, T. K., Nithya, S. &amp; Vinod, K. V. (2011) Faunal survey, endemism and possible species loss of Scarabaeinae (Coleoptera: Scarabaeidae) in the western slopes of the moist South Western Ghats, South India. Zootaxa, 2830, 29 - 38. https: // doi. org / 10.11646 / zootaxa. 2830.1. 3" type="journal article" year="2011">
Sabu
<emphasis id="32BC3E2F88461837FE250B55FE56C550" box="[452,501,513,539]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
., 2011
</bibRefCitation>
;
<bibRefCitation id="64599FCC88461837FDAE0B56FCF2C550" author="Venugopal, K. S. &amp; Sabu, K. T. &amp; Flemming, A. T." box="[591,849,513,540]" pageId="21" pageNumber="22" pagination="2685 - 2692" refId="ref39747" refString="Venugopal, K. S., Sabu, K. T. &amp; Flemming, A. T. (2012) Diversity and community structure of dung beetles (Coleoptera: Scarabaeinae) associated with semi-urban fragmented agricultural land in the Malabar coast in southern India. Journal of Threatened Taxa, 4 (7), 2685 - 2692. https: // doi. org / 10.11609 / JoTT. o 3074.2685 - 9 2" type="journal article" year="2012">
Venugopal
<emphasis id="32BC3E2F88461837FD2F0B55FD5CC550" box="[718,767,513,539]" italics="true" pageId="21" pageNumber="22">et al</emphasis>
., 2012
</bibRefCitation>
);
<collectingCountry id="78DFA2AD88461837FC850B56FC60C550" box="[868,963,513,539]" name="Pakistan" pageId="21" pageNumber="22">Pakistan</collectingCountry>
(
<bibRefCitation id="64599FCC88461837FC330B56FBFCC550" author="Arrow, G. J." box="[978,1119,513,539]" pageId="21" pageNumber="22" refId="ref36526" refString="Arrow, G. J. (1931) The Fauna of British India, including Ceylon and Burma. Coleoptera Lamellicornia. Part III (Coprinae). Taylor &amp; Francis, London, 428 pp." type="book" year="1931">Arrow 1931</bibRefCitation>
).
</paragraph>
<paragraph id="0077E23D88461837FF260B71FDB6C5B8" blockId="21.[151,1437,333,756]" pageId="21" pageNumber="22">
<emphasis id="32BC3E2F88461837FF260B71FE97C50B" bold="true" box="[199,308,550,576]" pageId="21" pageNumber="22">Remarks</emphasis>
. First record for the Arabian Peninsula, notably
<collectingCountry id="78DFA2AD88461837FC830B72FC5AC574" box="[866,1017,549,575]" name="Saudi Arabia" pageId="21" pageNumber="22">
Saudi
<taxonomicName id="C7C899BE88461837FC480B72FC5AC574" box="[937,1017,549,575]" class="Insecta" family="Cerambycidae" genus="Arabia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="35" phylum="Arthropoda" rank="genus">Arabia</taxonomicName>
</collectingCountry>
. The taxon was placed in the subgenus
<taxonomicName id="C7C899BE88461837FF260B1EFDF7C52F" authority="Balathasar, 1935" authorityName="Balathasar" authorityYear="1935" box="[199,596,585,612]" class="Insecta" family="Scarabaeidae" genus="Gibbonthophagus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">
<emphasis id="32BC3E2F88461837FF260B1EFE33C528" box="[199,400,585,611]" italics="true" pageId="21" pageNumber="22">Gibbonthophagus</emphasis>
Balathasar, 1935
</taxonomicName>
by
<bibRefCitation id="64599FCC88461837FD600B1EFC9FC52F" author="Kabakov, O. N." box="[641,828,585,612]" pageId="21" pageNumber="22" refId="ref38133" refString="Kabakov, O. N. (2006) Plastinchatousye zhuki podsemeystva Scarabaeinae (Insecta: Coleoptera: Scarabaeidae) fauny Rossii i sopredel'nykh stran. [The lamellicorn beetle subfamily Scarabaeinae (Insecta: Coleoptera: Scarabaeidae) in the fauna of Russia and adjacent countries]. Tovarishchestvo Nauchnykh Izdanii KMK, Moskva, 374 pp." type="book" year="2006">Kabakov (2006)</bibRefCitation>
, then in
<emphasis id="32BC3E2F88461837FC7E0B1EFA97C528" box="[927,1332,585,611]" italics="true" pageId="21" pageNumber="22">
<taxonomicName id="C7C899BE88461837FC7E0B1EFB95C528" box="[927,1078,585,611]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="genus">Onthophagus</taxonomicName>
species incertae sedis
</emphasis>
by
<bibRefCitation id="64599FCC88461837FA800B1EFEC2C5C3" author="Ziani, S. &amp; Bezdek, A." pageId="21" pageNumber="22" pagination="180 - 204" refId="ref39999" refString="Ziani, S. &amp; Bezdek, A. (2016) Scarabaeidae, subfamily Scarabaeinae, tribe Onthophagini. In: Lobl, I. &amp; Lobl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Volume 3. Scarabaeoidea - Scirtoidea - Dascilloidea - Buprestoidea - Byrrhoidea. Revised and updated edition. Brill, Leiden-Boston, pp. 180 - 204." type="book chapter" year="2016">Ziani &amp; Bezděk (2016)</bibRefCitation>
. We have herein inserted
<taxonomicName id="C7C899BE88461837FD6B0B39FD4CC5CC" authorityName="Gillet" authorityYear="1925" box="[650,751,621,648]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="falsus">
<emphasis id="32BC3E2F88461837FD6B0B39FD4CC5CC" box="[650,751,621,648]" italics="true" pageId="21" pageNumber="22">O. falsus</emphasis>
</taxonomicName>
in the nominotypical subgenus because, in our opinion, the species is phylogenetically more closely related to
<taxonomicName id="C7C899BE88461837FD3E0BC6FB2DC5E7" authority="(Schreber, 1759)" baseAuthorityName="Schreber" baseAuthorityYear="1759" box="[735,1166,657,684]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="taurus">
<emphasis id="32BC3E2F88461837FD3E0BC6FC65C5E0" box="[735,966,657,683]" italics="true" pageId="21" pageNumber="22">Onthophagus taurus</emphasis>
(Schreber, 1759)
</taxonomicName>
, the
<typeStatus id="DF735C9F88461837FB260BC5FB54C5E7" box="[1223,1271,658,684]" pageId="21" pageNumber="22">type</typeStatus>
species of the subgenus, than to any
<typeStatus id="DF735C9F88461837FE780BE1FE6AC59B" box="[409,457,694,720]" pageId="21" pageNumber="22">type</typeStatus>
species of any other genus or subgenus of
<taxonomicName id="C7C899BE88461837FC5C0BE2FBF8C584" authorityName="Burmeister" authorityYear="1846" box="[957,1115,693,719]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="tribe" tribe="Onthophagini">Onthophagini</taxonomicName>
. Paramera apex and lamella copulatrix support this placement.
</paragraph>
</subSubSection>
</treatment>
</document>