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<document id="2C78A5A116C04F240D90BB952B91DD2B" ID-CLB-Dataset="46432" ID-DOI="10.5281/zenodo.200834" ID-GBIF-Dataset="0d504835-4a24-4134-ae33-9448d9fb5884" ID-ISSN="1175-5326" ID-Zenodo-Dep="200834" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460384737241" checkinUser="plazi" docAuthor="Fonseca, Claudio Ruy Vasconcelos Da, Barbosa, Márcio Luís Leitão &amp; Fernandez, Maria Fernanda Souza" docDate="2011" docId="E922B809FFFDFF9466CBCAA8FB06F937" docLanguage="en" docName="zt03012p020.pdf" docOrigin="Zootaxa 3012" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Aulacocyclinae Kaup 1868" docType="treatment" docVersion="8" lastPageNumber="6" masterDocId="151BC071FFF9FF92665CCA3FFF92FFC5" masterDocTitle="A hypothetical evolutionary history of passalid beetles narrated by the comparative anatomy of the hindgut (Coleoptera: Passalidae)" masterLastPageNumber="20" masterPageNumber="1" pageNumber="5" updateTime="1698297348626" updateUser="ExternalLinkService">
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<mods:title id="7D8AF539492D8ADA4700D64A0FE75CDD">A hypothetical evolutionary history of passalid beetles narrated by the comparative anatomy of the hindgut (Coleoptera: Passalidae)</mods:title>
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<mods:namePart id="8164AF91BB95607FCCB5E753BA76056D">Fonseca, Claudio Ruy Vasconcelos Da</mods:namePart>
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<mods:namePart id="EA63CB83E52AB83C3D26CB0B194CF774">Barbosa, Márcio Luís Leitão</mods:namePart>
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<mods:namePart id="4B81B66A8FEEDBFD83D26E00C7FD5EC5">Fernandez, Maria Fernanda Souza</mods:namePart>
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<mods:date id="586BA380E0FD1EF9E0F8D94E2000F61B">2011</mods:date>
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<treatment id="E922B809FFFDFF9466CBCAA8FB06F937" ID-DOI="http://doi.org/10.5281/zenodo.5658203" ID-GBIF-Taxon="119415003" ID-Zenodo-Dep="5658203" LSID="urn:lsid:plazi:treatment:E922B809FFFDFF9466CBCAA8FB06F937" httpUri="http://treatment.plazi.org/id/E922B809FFFDFF9466CBCAA8FB06F937" lastPageId="6" lastPageNumber="6" pageId="4" pageNumber="5">
<subSubSection id="29915A94FFFDFF9466CBCAA8FB06F937" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5" type="nomenclature">
<paragraph id="6134091FFFFDFF9666CBCAA8FDE1FF74" blockId="4.[151,627,151,210]" box="[151,627,151,177]" pageId="4" pageNumber="5">
<heading id="3A7CBE73FFFDFF9666CBCAA8FDE1FF74" bold="true" box="[151,627,151,177]" fontSize="11" level="1" pageId="4" pageNumber="5" reason="1">
<emphasis id="53FFD50DFFFDFF9666CBCAA8FDE1FF74" bold="true" box="[151,627,151,177]" pageId="4" pageNumber="5">
Subfamily
<taxonomicName id="A68B729CFFFDFF966743CAA8FDE1FF74" ID-CoL="J78" authority="Kaup, 1868" authorityName="Kaup" authorityYear="1868" box="[287,627,151,177]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="subFamily" subFamily="Aulacocyclinae">
Aulacocyclinae
<bibRefCitation id="051A74EEFFFDFF9667BFCAA8FDE1FF74" author="Kaup" box="[483,627,151,177]" pageId="4" pageNumber="5" refString="Kaup, J. J. (1868) Podromus zu einer monographie der Passaliden. Coleopterologishe Hefte, 3, 4 - 32." type="journal article" year="1868">Kaup, 1868</bibRefCitation>
</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="6134091FFFFDFF9666CBCA85FF79FF17" blockId="4.[151,627,151,210]" box="[151,235,186,210]" pageId="4" pageNumber="5">
(
<figureCitation id="F9B0159AFFFDFF9666C3CA85FF72FF17" box="[159,224,186,210]" captionStart="FIGURE 1" captionStartId="4.[151,250,1840,1863]" captionTargetBox="[185,1422,522,1793]" captionTargetId="figure@4.[151,1436,518,1819]" captionTargetPageId="4" captionText="FIGURE 1. Aulacocyclinae hindgut. A— Aulacocyclus edentulus (MacLeay), Australia; B— Comacupes cylindraceus (Perty), Philippines; C— Taeniocerus bicanthatus (Percheron), Borneo; D— Cylindrocaulus bucerus Fairmaire, Japan; E— Ceracupes fronticornis (Westwood), Taiwan." httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5">Fig. 1</figureCitation>
)
</paragraph>
<paragraph id="6134091FFFFDFF9666CBCB3DFBE1FE0B" blockId="4.[151,1436,258,463]" pageId="4" pageNumber="5">
The genera
<taxonomicName id="A68B729CFFFDFF966741CB3CFE65FEDF" authority="Kaup" authorityName="Kaup" box="[285,503,259,283]" class="Insecta" family="Passallidae" genus="Aulacocyclus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF966741CB3CFE21FEDE" box="[285,435,259,283]" italics="true" pageId="4" pageNumber="5">Aulacocyclus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFDFF96645BCB3DFDD4FEDF" box="[519,582,258,282]" captionStart="FIGURE 1" captionStartId="4.[151,250,1840,1863]" captionTargetBox="[185,1422,522,1793]" captionTargetId="figure@4.[151,1436,518,1819]" captionTargetPageId="4" captionText="FIGURE 1. Aulacocyclinae hindgut. A— Aulacocyclus edentulus (MacLeay), Australia; B— Comacupes cylindraceus (Perty), Philippines; C— Taeniocerus bicanthatus (Percheron), Borneo; D— Cylindrocaulus bucerus Fairmaire, Japan; E— Ceracupes fronticornis (Westwood), Taiwan." httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5">Fig. 1</figureCitation>
A),
<taxonomicName id="A68B729CFFFDFF96642BCB3CFCACFEDF" authority="Kaup" authorityName="Kaup" box="[631,830,259,282]" class="Insecta" family="Passallidae" genus="Comacupes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF96642BCB3CFD68FEDF" box="[631,762,259,282]" italics="true" pageId="4" pageNumber="5">Comacupes</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFDFF966512CB3DFC1CFEDF" box="[846,910,258,282]" captionStart="FIGURE 1" captionStartId="4.[151,250,1840,1863]" captionTargetBox="[185,1422,522,1793]" captionTargetId="figure@4.[151,1436,518,1819]" captionTargetPageId="4" captionText="FIGURE 1. Aulacocyclinae hindgut. A— Aulacocyclus edentulus (MacLeay), Australia; B— Comacupes cylindraceus (Perty), Philippines; C— Taeniocerus bicanthatus (Percheron), Borneo; D— Cylindrocaulus bucerus Fairmaire, Japan; E— Ceracupes fronticornis (Westwood), Taiwan." httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5">Fig. 1</figureCitation>
B),
<taxonomicName id="A68B729CFFFDFF9665E0CB3BFB1AFEDF" authority="Kaup" authorityName="Kaup" box="[956,1160,259,283]" class="Insecta" family="Passallidae" genus="Taeniocerus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF9665E0CB3BFBD6FEDE" box="[956,1092,260,283]" italics="true" pageId="4" pageNumber="5">Taeniocerus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFDFF9662C5CB3DFB48FEDF" box="[1177,1242,258,282]" captionStart="FIGURE 1" captionStartId="4.[151,250,1840,1863]" captionTargetBox="[185,1422,522,1793]" captionTargetId="figure@4.[151,1436,518,1819]" captionTargetPageId="4" captionText="FIGURE 1. Aulacocyclinae hindgut. A— Aulacocyclus edentulus (MacLeay), Australia; B— Comacupes cylindraceus (Perty), Philippines; C— Taeniocerus bicanthatus (Percheron), Borneo; D— Cylindrocaulus bucerus Fairmaire, Japan; E— Ceracupes fronticornis (Westwood), Taiwan." httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5">Fig. 1</figureCitation>
C),
<taxonomicName id="A68B729CFFFDFF966354CB3CFEBEFEFB" authority="Fairmaire" authorityName="Fairmaire" class="Insecta" family="Passallidae" genus="Cylindrocaulus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF966354CB3CFF2AFEFB" italics="true" pageId="4" pageNumber="5">Cylindrocaulus</emphasis>
Fairmaire
</taxonomicName>
(
<figureCitation id="F9B0159AFFFDFF966760CB19FEE8FEFB" box="[316,378,294,318]" captionStart="FIGURE 1" captionStartId="4.[151,250,1840,1863]" captionTargetBox="[185,1422,522,1793]" captionTargetId="figure@4.[151,1436,518,1819]" captionTargetPageId="4" captionText="FIGURE 1. Aulacocyclinae hindgut. A— Aulacocyclus edentulus (MacLeay), Australia; B— Comacupes cylindraceus (Perty), Philippines; C— Taeniocerus bicanthatus (Percheron), Borneo; D— Cylindrocaulus bucerus Fairmaire, Japan; E— Ceracupes fronticornis (Westwood), Taiwan." httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5">Fig. 1</figureCitation>
D) and
<taxonomicName id="A68B729CFFFDFF96678FCB18FD1DFEFB" authority="Kaup" authorityName="Kaup" box="[467,655,295,318]" class="Insecta" family="Passallidae" genus="Ceracupes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF96678FCB18FDD9FEFB" box="[467,587,295,318]" italics="true" pageId="4" pageNumber="5">Ceracupes</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFDFF9664C3CB19FD4CFEFB" box="[671,734,294,318]" captionStart="FIGURE 1" captionStartId="4.[151,250,1840,1863]" captionTargetBox="[185,1422,522,1793]" captionTargetId="figure@4.[151,1436,518,1819]" captionTargetPageId="4" captionText="FIGURE 1. Aulacocyclinae hindgut. A— Aulacocyclus edentulus (MacLeay), Australia; B— Comacupes cylindraceus (Perty), Philippines; C— Taeniocerus bicanthatus (Percheron), Borneo; D— Cylindrocaulus bucerus Fairmaire, Japan; E— Ceracupes fronticornis (Westwood), Taiwan." httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5">Fig. 1</figureCitation>
E) are included in this subfamily. The ileum in these genera is similar to
<taxonomicName id="A68B729CFFFDFF96674DCB75FE37FEA7" box="[273,421,330,354]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="family">Scarabaeidae</taxonomicName>
(
<bibRefCitation id="051A74EEFFFDFF9667EBCB75FDC2FEA7" author="Umeya" box="[439,592,330,354]" pageId="4" pageNumber="5" refString="Umeya, K. (1960) A comparative morphology of the alimentary tract in the adults of lamellicorn-beetles (Coleoptera). Memoir of Faculty of the Agricultural. Hokkaido University, 3, 60 - 113." type="journal article" year="1960">Umeya 1960</bibRefCitation>
;
<bibRefCitation id="051A74EEFFFDFF96643CCB75FCCEFEA7" author="Lopez-Guerrero" box="[608,860,330,354]" pageId="4" pageNumber="5" refString="Lopez-Guerrero, Y. (2002) Anatomy and histology of the digestive system of Cephalodesmus armiger Westwood (Coleoptera, Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 56, 97 - 106." type="journal article" year="2002">López-Guerrero 2002</bibRefCitation>
), where the diverticula are assembled around the proximal edge and vary in shape and number. There are about six diverticula in
<taxonomicName id="A68B729CFFFDFF966247CB50FB0CFE43" box="[1051,1182,367,390]" class="Insecta" family="Passallidae" genus="Comacupes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF966247CB50FB0CFE43" box="[1051,1182,367,390]" italics="true" pageId="4" pageNumber="5">Comacupes</emphasis>
</taxonomicName>
and none in
<taxonomicName id="A68B729CFFFDFF96636CCB51FF72FE6E" class="Insecta" family="Passallidae" genus="Cylindrocaulus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF96636CCB51FF72FE6E" italics="true" pageId="4" pageNumber="5">Cylindrocaulus</emphasis>
</taxonomicName>
, but the average is three to four for the subfamily.
<collectingCountry id="199C498FFFFDFF966544CBACFCA5FE6E" box="[792,823,403,427]" name="American Samoa" pageId="4" pageNumber="5">As</collectingCountry>
to segmentation, there is variation from broad (
<taxonomicName id="A68B729CFFFDFF96630CCBACFF45FE0A" class="Insecta" family="Passallidae" genus="Comacupes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF96630CCBACFF45FE0A" italics="true" pageId="4" pageNumber="5">Comacupes</emphasis>
</taxonomicName>
) to close clustering, which makes the ileum look bellow-shaped (
<taxonomicName id="A68B729CFFFDFF9665EBCB89FBF1FE0B" box="[951,1123,438,462]" class="Insecta" family="Passallidae" genus="Cylindrocaulus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFDFF9665EBCB89FBF1FE0B" box="[951,1123,438,462]" italics="true" pageId="4" pageNumber="5">Cylindrocaulus</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="35F45997FFFDFF9666CBCD0FFE7FF841" httpUri="https://zenodo.org/record/200835/files/figure.png" pageId="4" pageNumber="5" targetBox="[185,1422,522,1793]" targetPageId="4">
<paragraph id="6134091FFFFDFF9666CBCD0FFE7FF841" blockId="4.[151,1436,1840,1924]" pageId="4" pageNumber="5">
<emphasis id="53FFD50DFFFDFF9666CBCD0FFE81F883" bold="true" box="[151,275,1840,1863]" pageId="4" pageNumber="5">FIGURE 1.</emphasis>
<taxonomicName id="A68B729CFFFDFF966745CD0EFE24F882" box="[281,438,1841,1863]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="subFamily" subFamily="Aulacocyclinae">Aulacocyclinae</taxonomicName>
hindgut. A—
<taxonomicName id="A68B729CFFFDFF96641FCD0EFC3EF882" authority="MacLeay" authorityName="MacLeay" box="[579,940,1841,1863]" class="Insecta" family="Passallidae" genus="Aulacocyclus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="edentulus">
<emphasis id="53FFD50DFFFDFF96641FCD0EFCA0F882" box="[579,818,1841,1863]" italics="true" pageId="4" pageNumber="5">Aulacocyclus edentulus</emphasis>
(MacLeay)
</taxonomicName>
, Australia; B—
<taxonomicName id="A68B729CFFFDFF966217CD0DFA0AF882" authority="Perty" authorityName="Perty" box="[1099,1432,1841,1863]" class="Insecta" family="Passallidae" genus="Comacupes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="cylindraceus">
<emphasis id="53FFD50DFFFDFF966217CD0DFADAF882" box="[1099,1352,1841,1863]" italics="true" pageId="4" pageNumber="5">Comacupes cylindraceus</emphasis>
(Perty)
</taxonomicName>
, Philippines; C—
<taxonomicName id="A68B729CFFFDFF96671ECD6EFD2DF8A0" authority="Percheron" authorityName="Percheron" box="[322,703,1871,1894]" class="Insecta" family="Passallidae" genus="Taeniocerus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="bicanthatus">
<emphasis id="53FFD50DFFFDFF96671ECD6EFDA9F8A0" box="[322,571,1871,1894]" italics="true" pageId="4" pageNumber="5">Taeniocerus bicanthatus</emphasis>
(Percheron)
</taxonomicName>
, Borneo; D—
<taxonomicName id="A68B729CFFFDFF96650CCD70FB23F8A0" authority="Fairmaire" authorityName="Fairmaire" box="[848,1201,1871,1893]" class="Insecta" family="Passallidae" genus="Cylindrocaulus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="bucerus">
<emphasis id="53FFD50DFFFDFF96650CCD70FBD6F8A0" box="[848,1092,1871,1893]" italics="true" pageId="4" pageNumber="5">Cylindrocaulus bucerus</emphasis>
Fairmaire
</taxonomicName>
, Japan; E—
<taxonomicName id="A68B729CFFFDFF96636CCD6FFE00F841" authority="Westwood" authorityName="Westwood" class="Insecta" family="Passallidae" genus="Ceracupes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="species" species="fronticornis">
<emphasis id="53FFD50DFFFDFF96636CCD6FFE9CF841" italics="true" pageId="4" pageNumber="5">Ceracupes fronticornis</emphasis>
(Westwood)
</taxonomicName>
, Taiwan.
</paragraph>
</caption>
<paragraph id="6134091FFFFCFF9766CBCAA5FF6AFF10" blockId="5.[151,444,151,213]" pageId="5" pageNumber="6">
<emphasis id="53FFD50DFFFCFF9766CBCAA5FE2EFF60" bold="true" box="[151,444,151,180]" pageId="5" pageNumber="6">Branch Solenocyclinae+</emphasis>
<figureCitation id="F9B0159AFFFCFF9766CBCA82FF6AFF10" box="[151,248,189,213]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStart-2="FIGURE 4" captionStart-3="FIGURE 5" captionStart-4="FIGURE 6" captionStart-5="FIGURE 7" captionStart-6="FIGURE 8" captionStartId-0="5.[151,250,1915,1938]" captionStartId-1="6.[151,250,1301,1324]" captionStartId-2="7.[151,250,1857,1880]" captionStartId-3="8.[151,250,1790,1813]" captionStartId-4="9.[151,250,1347,1370]" captionStartId-5="10.[151,250,1899,1922]" captionStartId-6="11.[151,250,1410,1433]" captionTargetBox-0="[170,1422,687,1884]" captionTargetBox-1="[156,1431,205,1278]" captionTargetBox-2="[160,1418,198,1805]" captionTargetBox-3="[153,1429,196,1752]" captionTargetBox-4="[159,1432,198,1314]" captionTargetBox-5="[202,1411,197,1885]" captionTargetBox-6="[156,1431,196,1375]" captionTargetId-0="figure@5.[165,1424,687,1925]" captionTargetId-1="figure@6.[151,1436,193,1280]" captionTargetId-2="figure@7.[151,1436,193,1843]" captionTargetId-3="figure@8.[151,1436,193,1769]" captionTargetId-4="figure@9.[151,1436,193,1326]" captionTargetId-5="figure@10.[199,1420,193,1891]" captionTargetId-6="figure@11.[151,1436,193,1389]" captionTargetPageId-0="5" captionTargetPageId-1="6" captionTargetPageId-2="7" captionTargetPageId-3="8" captionTargetPageId-4="9" captionTargetPageId-5="10" captionTargetPageId-6="11" captionText-0="FIGURE 2. Solenocyclinae hindgut. A— Trichostigmus thoreyi (Kaup), Philippines; B— Leptaulax dentatus (Fabricius), India; C— Stephanocephalus hostilis (Percheron), Madagascar; D— Didimus aberrans (Kuwert), Democratic Republic of the Congo; E— Pentalobus barbatus (Fabricius), Ivory Coast; F— Semicyclus grayi Kaup, Madagascar." captionText-1="FIGURE 3. Solenocyclinae hindgut. A— Solenocyclus exaratus Klug, Madagascar; B— Vitellinus approximatus (Klug), Madagascar; C— Ciceronius morbillosus (Klug), Madagascar; D— Erionomus planiceps (Eschscholtz), Democratic Republic of the Congo; E— Flaminius nonfriedi Kuwert, Madagascar." captionText-2="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." captionText-3="FIGURE 5. Proculinae hindgut. A— Verres corticicola (Truqui), Mexico; B— Veturius heydeni (Kaup), Venezuela; C— Proculejus truquii Kaup, Mexico; D— Ogyges laevissimus (Kaup), Guatemala; E— Undulifer incisus (Truqui), Mexico; F— Oileus rimator (Truqui), Mexico; G— Proculus goryi (Melly), Guatemala; H— Xylopassaloides pterocavis Reyes-Castillo, Fonseca &amp; Castillo, Mexico." captionText-4="FIGURE 6. Passalinae hindgut. A— Ptichopus angulatus (Percheron), Costa Rica; B— Passalus interruptus (Linnaeus), Brazil; C— Spasalus crenatus (MacLeay), Puerto Rico; D— Paxillus leachi MacLeay, Brazil; E— Passipassalus buhrnheimi Fonseca &amp; Reyes-Castillo, Brazil (Fig. E from Fonseca &amp; Reyes – Castillo 1993)." captionText-5="FIGURE 7. Macrolininae hindgut. A— Pleurarius pilipes Kaup, India; B— Ophrygonius inaequalis (Burmeister), India; C— Episphenus moorei Kaup, India; D— Aceraius grandis (Burmeister), Taiwan; E— Macrolinus latipennis (Percheron), Java; F— Pseudepisphenus perplexus Gravely, New Guinea; G— Protomocoelus australis (Boisduval), Solomon Islands; H— Labienus ptox (Kaup), New Guinea; I— Analaches puberilis Kuwert, New Guinea." captionText-6="FIGURE 8. Macrolininae hindgut. A— Mastochilus polyphyllus (MacLeay), Australia; B— Pharochilus dilatatus (Dalman), Australia; C— Cetejus virginalis (Kaup), New Guinea; D— Gonatas naviculator (Percheron), New Guinea; E— Plesthenus quadricornis (Kaup), Sulawesi; F— Pelopides gravidus (Kuwert), Sumatra." httpUri-0="https://zenodo.org/record/200836/files/figure.png" httpUri-1="https://zenodo.org/record/200837/files/figure.png" httpUri-2="https://zenodo.org/record/200838/files/figure.png" httpUri-3="https://zenodo.org/record/200839/files/figure.png" httpUri-4="https://zenodo.org/record/200840/files/figure.png" httpUri-5="https://zenodo.org/record/200841/files/figure.png" httpUri-6="https://zenodo.org/record/200842/files/figure.png" pageId="5" pageNumber="6">Figs 2–8</figureCitation>
</paragraph>
<paragraph id="6134091FFFFCFF9466CBCB3AFB06F937" blockId="5.[151,1436,261,654]" lastBlockId="6.[151,1436,1430,1778]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
This branch embodies all genera that are part of
<taxonomicName id="A68B729CFFFCFF97649CCB3AFCA6FED8" box="[704,820,261,285]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subFamily" subFamily="Passalinae">Passalinae</taxonomicName>
in Reyes-Castillo’s (1970) classification. The features of the ileum confirm this observation, as it is obvious that Solenocyclinae+ is the sister group of
<taxonomicName id="A68B729CFFFCFF9762BBCB13FA0BFE81" box="[1255,1433,300,324]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subFamily" subFamily="Aulacocyclinae">Aulacocyclinae</taxonomicName>
. Concerning the ileum architecture, Solenocyclinae+ show transformations that start to become noticeable on the African genera
<taxonomicName id="A68B729CFFFCFF976714CB48FDCBFE4A" authority="Kaup" authorityName="Kaup" box="[328,601,375,399]" class="Insecta" family="Passallidae" genus="Stephanocephalus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF976714CB48FD81FE4A" box="[328,531,375,399]" italics="true" pageId="5" pageNumber="6">Stephanocephalus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF976436CB48FD3DFE4A" box="[618,687,375,399]" captionStart="FIGURE 2" captionStartId="5.[151,250,1915,1938]" captionTargetBox="[170,1422,687,1884]" captionTargetId="figure@5.[165,1424,687,1925]" captionTargetPageId="5" captionText="FIGURE 2. Solenocyclinae hindgut. A— Trichostigmus thoreyi (Kaup), Philippines; B— Leptaulax dentatus (Fabricius), India; C— Stephanocephalus hostilis (Percheron), Madagascar; D— Didimus aberrans (Kuwert), Democratic Republic of the Congo; E— Pentalobus barbatus (Fabricius), Ivory Coast; F— Semicyclus grayi Kaup, Madagascar." httpUri="https://zenodo.org/record/200836/files/figure.png" pageId="5" pageNumber="6">Fig. 2</figureCitation>
C),
<taxonomicName id="A68B729CFFFCFF976486CB48FCECFE4A" authority="Kaup" authorityName="Kaup" box="[730,894,375,399]" class="Insecta" family="Passallidae" genus="Didimus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF976486CB48FCABFE4A" box="[730,825,375,399]" italics="true" pageId="5" pageNumber="6">Didimus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF9765D3CB48FC46FE4A" box="[911,980,375,399]" captionStart="FIGURE 2" captionStartId="5.[151,250,1915,1938]" captionTargetBox="[170,1422,687,1884]" captionTargetId="figure@5.[165,1424,687,1925]" captionTargetPageId="5" captionText="FIGURE 2. Solenocyclinae hindgut. A— Trichostigmus thoreyi (Kaup), Philippines; B— Leptaulax dentatus (Fabricius), India; C— Stephanocephalus hostilis (Percheron), Madagascar; D— Didimus aberrans (Kuwert), Democratic Republic of the Congo; E— Pentalobus barbatus (Fabricius), Ivory Coast; F— Semicyclus grayi Kaup, Madagascar." httpUri="https://zenodo.org/record/200836/files/figure.png" pageId="5" pageNumber="6">Fig. 2</figureCitation>
D),
<taxonomicName id="A68B729CFFFCFF97625CCB48FB51FE4A" authority="Kaup" authorityName="Kaup" box="[1024,1219,375,399]" class="Insecta" family="Passallidae" genus="Pentalobus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF97625CCB48FBECFE4A" box="[1024,1150,375,399]" italics="true" pageId="5" pageNumber="6">Pentalobus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF976289CB48FA88FE4A" box="[1237,1306,375,399]" captionStart="FIGURE 2" captionStartId="5.[151,250,1915,1938]" captionTargetBox="[170,1422,687,1884]" captionTargetId="figure@5.[165,1424,687,1925]" captionTargetPageId="5" captionText="FIGURE 2. Solenocyclinae hindgut. A— Trichostigmus thoreyi (Kaup), Philippines; B— Leptaulax dentatus (Fabricius), India; C— Stephanocephalus hostilis (Percheron), Madagascar; D— Didimus aberrans (Kuwert), Democratic Republic of the Congo; E— Pentalobus barbatus (Fabricius), Ivory Coast; F— Semicyclus grayi Kaup, Madagascar." httpUri="https://zenodo.org/record/200836/files/figure.png" pageId="5" pageNumber="6">Fig. 2</figureCitation>
E),
<taxonomicName id="A68B729CFFFCFF97631FCB47FE9AFE76" authority="Kaup" authorityName="Kaup" class="Insecta" family="Passallidae" genus="Semicyclus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF97631FCB47FF56FE76" italics="true" pageId="5" pageNumber="6">Semicyclus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF976744CBA4FEC9FE76" box="[280,347,411,435]" captionStart="FIGURE 2" captionStartId="5.[151,250,1915,1938]" captionTargetBox="[170,1422,687,1884]" captionTargetId="figure@5.[165,1424,687,1925]" captionTargetPageId="5" captionText="FIGURE 2. Solenocyclinae hindgut. A— Trichostigmus thoreyi (Kaup), Philippines; B— Leptaulax dentatus (Fabricius), India; C— Stephanocephalus hostilis (Percheron), Madagascar; D— Didimus aberrans (Kuwert), Democratic Republic of the Congo; E— Pentalobus barbatus (Fabricius), Ivory Coast; F— Semicyclus grayi Kaup, Madagascar." httpUri="https://zenodo.org/record/200836/files/figure.png" pageId="5" pageNumber="6">Fig. 2</figureCitation>
F) and
<taxonomicName id="A68B729CFFFCFF9767F7CBA4FDEAFE76" authority="Kuwert" authorityName="Kuwert" box="[427,632,411,435]" class="Insecta" family="Passallidae" genus="Flaminius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF9767F7CBA4FD8EFE76" box="[427,540,411,435]" italics="true" pageId="5" pageNumber="6">Flaminius</emphasis>
Kuwert
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF9764DBCBA4FD5BFE76" box="[647,713,411,435]" captionStart="FIGURE 3" captionStartId="6.[151,250,1301,1324]" captionTargetBox="[156,1431,205,1278]" captionTargetId="figure@6.[151,1436,193,1280]" captionTargetPageId="6" captionText="FIGURE 3. Solenocyclinae hindgut. A— Solenocyclus exaratus Klug, Madagascar; B— Vitellinus approximatus (Klug), Madagascar; C— Ciceronius morbillosus (Klug), Madagascar; D— Erionomus planiceps (Eschscholtz), Democratic Republic of the Congo; E— Flaminius nonfriedi Kuwert, Madagascar." httpUri="https://zenodo.org/record/200837/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
E), possessing a collection of protuberances in the distal half. In
<taxonomicName id="A68B729CFFFCFF9766CBCB80FEFCFE12" authority="Kaup" authorityName="Kaup" box="[151,366,447,471]" class="Insecta" family="Passallidae" genus="Solenocyclus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF9766CBCB80FEBBFE12" box="[151,297,447,471]" italics="true" pageId="5" pageNumber="6">Solenocyclus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF976723CB80FE53FE12" box="[383,449,447,471]" captionStart="FIGURE 3" captionStartId="6.[151,250,1301,1324]" captionTargetBox="[156,1431,205,1278]" captionTargetId="figure@6.[151,1436,193,1280]" captionTargetPageId="6" captionText="FIGURE 3. Solenocyclinae hindgut. A— Solenocyclus exaratus Klug, Madagascar; B— Vitellinus approximatus (Klug), Madagascar; C— Ciceronius morbillosus (Klug), Madagascar; D— Erionomus planiceps (Eschscholtz), Democratic Republic of the Congo; E— Flaminius nonfriedi Kuwert, Madagascar." httpUri="https://zenodo.org/record/200837/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
A),
<taxonomicName id="A68B729CFFFCFF9767B3CB80FD27FE12" authority="Kuwert" authorityName="Kuwert" box="[495,693,447,471]" class="Insecta" family="Passallidae" genus="Vitellinus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF9767B3CB80FDCAFE12" box="[495,600,447,471]" italics="true" pageId="5" pageNumber="6">Vitellinus</emphasis>
Kuwert
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF97649ACB80FC98FE12" box="[710,778,447,471]" captionStart="FIGURE 3" captionStartId="6.[151,250,1301,1324]" captionTargetBox="[156,1431,205,1278]" captionTargetId="figure@6.[151,1436,193,1280]" captionTargetPageId="6" captionText="FIGURE 3. Solenocyclinae hindgut. A— Solenocyclus exaratus Klug, Madagascar; B— Vitellinus approximatus (Klug), Madagascar; C— Ciceronius morbillosus (Klug), Madagascar; D— Erionomus planiceps (Eschscholtz), Democratic Republic of the Congo; E— Flaminius nonfriedi Kuwert, Madagascar." httpUri="https://zenodo.org/record/200837/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
B),
<taxonomicName id="A68B729CFFFCFF976569CBFFFC66FE12" authority="Kaup" authorityName="Kaup" box="[821,1012,448,471]" class="Insecta" family="Passallidae" genus="Ciceronius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF976569CBFFFC3DFE12" box="[821,943,448,471]" italics="true" pageId="5" pageNumber="6">Ciceronius</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF976259CB80FBDAFE12" box="[1029,1096,447,471]" captionStart="FIGURE 3" captionStartId="6.[151,250,1301,1324]" captionTargetBox="[156,1431,205,1278]" captionTargetId="figure@6.[151,1436,193,1280]" captionTargetPageId="6" captionText="FIGURE 3. Solenocyclinae hindgut. A— Solenocyclus exaratus Klug, Madagascar; B— Vitellinus approximatus (Klug), Madagascar; C— Ciceronius morbillosus (Klug), Madagascar; D— Erionomus planiceps (Eschscholtz), Democratic Republic of the Congo; E— Flaminius nonfriedi Kuwert, Madagascar." httpUri="https://zenodo.org/record/200837/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
C) and
<taxonomicName id="A68B729CFFFCFF9762C2CBFFFACFFE12" authority="Kaup" authorityName="Kaup" box="[1182,1373,448,471]" class="Insecta" family="Passallidae" genus="Erionomus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF9762C2CBFFFA8BFE12" box="[1182,1305,448,471]" italics="true" pageId="5" pageNumber="6">Erionomus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF976332CB80FF37FE3E" captionStart="FIGURE 3" captionStartId="6.[151,250,1301,1324]" captionTargetBox="[156,1431,205,1278]" captionTargetId="figure@6.[151,1436,193,1280]" captionTargetPageId="6" captionText="FIGURE 3. Solenocyclinae hindgut. A— Solenocyclus exaratus Klug, Madagascar; B— Vitellinus approximatus (Klug), Madagascar; C— Ciceronius morbillosus (Klug), Madagascar; D— Erionomus planiceps (Eschscholtz), Democratic Republic of the Congo; E— Flaminius nonfriedi Kuwert, Madagascar." httpUri="https://zenodo.org/record/200837/files/figure.png" pageId="5" pageNumber="6">Fig. 3</figureCitation>
D), such protuberances extend their range to cover the whole surface of the ileum. The number and size of the protuberances are not constant, as sometimes they can be dense in
<taxonomicName id="A68B729CFFFCFF976522C838FBA0FDDA" authority="Kaup" authorityName="Kaup" box="[894,1074,519,543]" class="Insecta" family="Passallidae" genus="Leptaulax" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF976522C838FC7CFDDA" box="[894,1006,519,543]" italics="true" pageId="5" pageNumber="6">Leptaulax</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF97621DC838FB16FDDA" box="[1089,1156,519,543]" captionStart="FIGURE 2" captionStartId="5.[151,250,1915,1938]" captionTargetBox="[170,1422,687,1884]" captionTargetId="figure@5.[165,1424,687,1925]" captionTargetPageId="5" captionText="FIGURE 2. Solenocyclinae hindgut. A— Trichostigmus thoreyi (Kaup), Philippines; B— Leptaulax dentatus (Fabricius), India; C— Stephanocephalus hostilis (Percheron), Madagascar; D— Didimus aberrans (Kuwert), Democratic Republic of the Congo; E— Pentalobus barbatus (Fabricius), Ivory Coast; F— Semicyclus grayi Kaup, Madagascar." httpUri="https://zenodo.org/record/200836/files/figure.png" pageId="5" pageNumber="6">Fig. 2</figureCitation>
B), giving a very striking differentiation to the ileum. In addition to the protuberances, the long diverticula, which arise from the proximal region in
<taxonomicName id="A68B729CFFFCFF976759C86DFE24FDAF" box="[261,438,594,618]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="subFamily" subFamily="Aulacocyclinae">Aulacocyclinae</taxonomicName>
, undergo shortening in the Solenocyclinae+, and in some instances turning into just one diverticulum. Protuberance clustering increases in taxa from Indo-Australian
<taxonomicName id="A68B729CFFFCFF9765A4C849FB21FD4B" authority="Kaup" authorityName="Kaup" box="[1016,1203,630,654]" class="Insecta" family="Passallidae" genus="Pleurarius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFCFF9765A4C849FBE2FD4B" box="[1016,1136,630,654]" italics="true" pageId="5" pageNumber="6">Pleurarius</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFCFF97629EC849FA96FD4B" box="[1218,1284,630,654]" captionStart="FIGURE 7" captionStartId="10.[151,250,1899,1922]" captionTargetBox="[202,1411,197,1885]" captionTargetId="figure@10.[199,1420,193,1891]" captionTargetPageId="10" captionText="FIGURE 7. Macrolininae hindgut. A— Pleurarius pilipes Kaup, India; B— Ophrygonius inaequalis (Burmeister), India; C— Episphenus moorei Kaup, India; D— Aceraius grandis (Burmeister), Taiwan; E— Macrolinus latipennis (Percheron), Java; F— Pseudepisphenus perplexus Gravely, New Guinea; G— Protomocoelus australis (Boisduval), Solomon Islands; H— Labienus ptox (Kaup), New Guinea; I— Analaches puberilis Kuwert, New Guinea." httpUri="https://zenodo.org/record/200841/files/figure.png" pageId="5" pageNumber="6">Fig. 7</figureCitation>
A) to Neotro- The Neotropical
<emphasis id="53FFD50DFFFFFF9467DBCFA8FE25FA6B" box="[391,439,1431,1454]" italics="true" pageId="6" pageNumber="7">taxa</emphasis>
are included in two tribes: Passalini and
<taxonomicName id="A68B729CFFFFFF9465C9CFA9FB2EFA6B" authority="(Boucher 2006)" baseAuthorityName="Boucher" baseAuthorityYear="2006" box="[917,1212,1430,1454]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="tribe" tribe="Proculini">
Proculini (
<bibRefCitation id="051A74EEFFFFFF946251CFA9FB26FA6B" author="Boucher" box="[1037,1204,1430,1454]" pageId="6" pageNumber="7" refString="Boucher, S. (2006) Evolution et phylogenie des Coleopteres Passalidae (Scarabaeoidea): Les taxons du groupe famille, la tribu neotropicale des Proculini et son complexe Ve t u r i us. Annales de la Societe Entomologique de France (n. s.), 41, 239 - 604." type="journal article" year="2006">Boucher 2006</bibRefCitation>
)
</taxonomicName>
. The ileum in most genera of
<taxonomicName id="A68B729CFFFFFF946757CF85FEE0FA17" box="[267,370,1466,1490]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="tribe" tribe="Proculini">Proculini</taxonomicName>
(exposed clypeus);
<taxonomicName id="A68B729CFFFFFF946404CF85FCCBFA17" authority="Kuwert" authorityName="Kuwert" box="[600,857,1466,1490]" class="Insecta" family="Passallidae" genus="Odontotaenius" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF946404CF85FD6FFA17" box="[600,765,1466,1490]" italics="true" pageId="6" pageNumber="7">Odontotaenius</emphasis>
Kuwert
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF946537CF85FC22FA17" box="[875,944,1466,1490]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
A),
<taxonomicName id="A68B729CFFFFFF946580CF84FBEAFA17" authority="Kaup" authorityName="Kaup" box="[988,1144,1467,1490]" class="Insecta" family="Passallidae" genus="Spurius" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF946580CF84FBA0FA17" box="[988,1074,1467,1490]" italics="true" pageId="6" pageNumber="7">Spurius</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9462D5CF85FB5DFA17" box="[1161,1231,1466,1490]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
B),
<taxonomicName id="A68B729CFFFFFF9462A5CF85FA09FA17" authority="Kaup" authorityName="Kaup" box="[1273,1435,1466,1490]" class="Insecta" family="Passallidae" genus="Popilius" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9462A5CF85FAC5FA17" box="[1273,1367,1466,1490]" italics="true" pageId="6" pageNumber="7">Popilius</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9466C3CFE1FF77FA33" box="[159,229,1502,1526]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
C),
<taxonomicName id="A68B729CFFFFFF94674DCFE1FD93FA33" authority="Kaup" authorityName="Kaup" box="[273,513,1502,1526]" class="Insecta" family="Passallidae" genus="Pseudacanthus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF94674DCFE1FE29FA33" box="[273,443,1502,1526]" italics="true" pageId="6" pageNumber="7">Pseudacanthus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF94644FCFE1FDCBFA33" box="[531,601,1502,1526]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
D),
<taxonomicName id="A68B729CFFFFFF9464DACFE1FC8BFA33" authority="Kaup" authorityName="Kaup" box="[646,793,1502,1526]" class="Insecta" family="Passallidae" genus="Vindex" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9464DACFE1FD42FA33" box="[646,720,1502,1526]" italics="true" pageId="6" pageNumber="7">Vindex</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF946577CFE1FCE3FA33" box="[811,881,1502,1526]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
E),
<taxonomicName id="A68B729CFFFFFF9465C7CFE1FBE5FA33" authority="Kuwert" authorityName="Kuwert" box="[923,1143,1502,1526]" class="Insecta" family="Passallidae" genus="Petrejoides" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9465C7CFE1FB88FA33" box="[923,1050,1502,1526]" italics="true" pageId="6" pageNumber="7">Petrejoides</emphasis>
Kuwert
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9462D5CFE1FB5DFA33" box="[1161,1231,1502,1526]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
F),
<taxonomicName id="A68B729CFFFFFF9462A4CFE1FA0EFA32" authority="Zang" authorityName="Zang" box="[1272,1436,1502,1527]" class="Insecta" family="Passallidae" genus="Heliscus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9462A4CFE1FACBFA33" box="[1272,1369,1502,1526]" italics="true" pageId="6" pageNumber="7">Heliscus</emphasis>
Zang
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9466C3CC3DFF77F9DF" box="[159,229,1538,1562]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
G),
<taxonomicName id="A68B729CFFFFFF94674FCC3DFDA5F9DF" authority="Kuwert" authorityName="Kuwert" box="[275,567,1538,1562]" class="Insecta" family="Passallidae" genus="Chondrocephalus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF94674FCC3DFE4BF9DF" box="[275,473,1538,1562]" italics="true" pageId="6" pageNumber="7">Chondrocephalus</emphasis>
Kuwert
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF946416CC3DFD02F9DF" box="[586,656,1538,1562]" captionStart="FIGURE 4" captionStartId="7.[151,250,1857,1880]" captionTargetBox="[160,1418,198,1805]" captionTargetId="figure@7.[151,1436,193,1843]" captionTargetPageId="7" captionText="FIGURE 4. Proculinae hindgut. A— Odontotaenius striatopunctatus (Percheron), Mexico; B— Spurius bicornis (Truqui), Costa Rica; C— Popilius marginatus (Percheron), Brazil; D— Pseudacanthus mexicanus (Truqui), Mexico; E— Vindex agnoscendus (Percheron), Mexico; F— Petrejoides tenuis Kuwert, Costa Rica; G— Heliscus tropicus (Percheron), Mexico; H— Chondrocephalus granulifrons (Bates), Mexico." httpUri="https://zenodo.org/record/200838/files/figure.png" pageId="6" pageNumber="7">Fig. 4</figureCitation>
H),
<taxonomicName id="A68B729CFFFFFF9464E1CC3CFCD8F9DF" authority="Kaup" authorityName="Kaup" box="[701,842,1539,1562]" class="Insecta" family="Passallidae" genus="Verres" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9464E1CC3CFC92F9DF" box="[701,768,1539,1562]" italics="true" pageId="6" pageNumber="7">Verres</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF946501CC3DFC30F9DF" box="[861,930,1538,1562]" captionStart="FIGURE 5" captionStartId="8.[151,250,1790,1813]" captionTargetBox="[153,1429,196,1752]" captionTargetId="figure@8.[151,1436,193,1769]" captionTargetPageId="8" captionText="FIGURE 5. Proculinae hindgut. A— Verres corticicola (Truqui), Mexico; B— Veturius heydeni (Kaup), Venezuela; C— Proculejus truquii Kaup, Mexico; D— Ogyges laevissimus (Kaup), Guatemala; E— Undulifer incisus (Truqui), Mexico; F— Oileus rimator (Truqui), Mexico; G— Proculus goryi (Melly), Guatemala; H— Xylopassaloides pterocavis Reyes-Castillo, Fonseca &amp; Castillo, Mexico." httpUri="https://zenodo.org/record/200839/files/figure.png" pageId="6" pageNumber="7">Fig. 5</figureCitation>
A),
<taxonomicName id="A68B729CFFFFFF94658DCC3CFBE1F9DF" authority="Kaup" authorityName="Kaup" box="[977,1139,1539,1562]" class="Insecta" family="Passallidae" genus="Veturius" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF94658DCC3CFBB8F9DF" box="[977,1066,1539,1562]" italics="true" pageId="6" pageNumber="7">Veturius</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9462DACC3DFB59F9DF" box="[1158,1227,1538,1562]" captionStart="FIGURE 5" captionStartId="8.[151,250,1790,1813]" captionTargetBox="[153,1429,196,1752]" captionTargetId="figure@8.[151,1436,193,1769]" captionTargetPageId="8" captionText="FIGURE 5. Proculinae hindgut. A— Verres corticicola (Truqui), Mexico; B— Veturius heydeni (Kaup), Venezuela; C— Proculejus truquii Kaup, Mexico; D— Ogyges laevissimus (Kaup), Guatemala; E— Undulifer incisus (Truqui), Mexico; F— Oileus rimator (Truqui), Mexico; G— Proculus goryi (Melly), Guatemala; H— Xylopassaloides pterocavis Reyes-Castillo, Fonseca &amp; Castillo, Mexico." httpUri="https://zenodo.org/record/200839/files/figure.png" pageId="6" pageNumber="7">Fig. 5</figureCitation>
B) and
<taxonomicName id="A68B729CFFFFFF946378CC3DFF46F9FB" authority="Kaup" authorityName="Kaup" class="Insecta" family="Passallidae" genus="Proculejus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF946378CC3DFA0EF9DF" box="[1316,1436,1538,1562]" italics="true" pageId="6" pageNumber="7">Proculejus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9466B9CC19FEBBF9FB" box="[229,297,1574,1598]" captionStart="FIGURE 5" captionStartId="8.[151,250,1790,1813]" captionTargetBox="[153,1429,196,1752]" captionTargetId="figure@8.[151,1436,193,1769]" captionTargetPageId="8" captionText="FIGURE 5. Proculinae hindgut. A— Verres corticicola (Truqui), Mexico; B— Veturius heydeni (Kaup), Venezuela; C— Proculejus truquii Kaup, Mexico; D— Ogyges laevissimus (Kaup), Guatemala; E— Undulifer incisus (Truqui), Mexico; F— Oileus rimator (Truqui), Mexico; G— Proculus goryi (Melly), Guatemala; H— Xylopassaloides pterocavis Reyes-Castillo, Fonseca &amp; Castillo, Mexico." httpUri="https://zenodo.org/record/200839/files/figure.png" pageId="6" pageNumber="7">Fig. 5</figureCitation>
C); has the long diverticulum facing down. The long diverticulum faces up for
<taxonomicName id="A68B729CFFFFFF9462E8CC19FACFF9FB" authority="Kaup" authorityName="Kaup" box="[1204,1373,1574,1598]" class="Insecta" family="Passallidae" genus="Proculus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9462E8CC19FA8AF9FB" box="[1204,1304,1574,1598]" italics="true" pageId="6" pageNumber="7">Proculus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF946332CC19FF37F9A7" captionStart="FIGURE 5" captionStartId="8.[151,250,1790,1813]" captionTargetBox="[153,1429,196,1752]" captionTargetId="figure@8.[151,1436,193,1769]" captionTargetPageId="8" captionText="FIGURE 5. Proculinae hindgut. A— Verres corticicola (Truqui), Mexico; B— Veturius heydeni (Kaup), Venezuela; C— Proculejus truquii Kaup, Mexico; D— Ogyges laevissimus (Kaup), Guatemala; E— Undulifer incisus (Truqui), Mexico; F— Oileus rimator (Truqui), Mexico; G— Proculus goryi (Melly), Guatemala; H— Xylopassaloides pterocavis Reyes-Castillo, Fonseca &amp; Castillo, Mexico." httpUri="https://zenodo.org/record/200839/files/figure.png" pageId="6" pageNumber="7">Fig. 5</figureCitation>
G), in which the ileum is formed by large diverticula on the proximal region that becomes smaller towards the distal apex. On the other hand, the large, closely clustered protuberances distinguish
<taxonomicName id="A68B729CFFFFFF946278CC51FB1AF943" box="[1060,1160,1646,1670]" class="Insecta" family="Passallidae" genus="Proculus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF946278CC51FB1AF943" box="[1060,1160,1646,1670]" italics="true" pageId="6" pageNumber="7">Proculus</emphasis>
</taxonomicName>
from all other
<taxonomicName id="A68B729CFFFFFF946372CC51FA0AF943" box="[1326,1432,1646,1670]" class="Insecta" family="Passalidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="tribe" tribe="Proculini">Proculini</taxonomicName>
. In the Passalini (hidden clypeus);
<taxonomicName id="A68B729CFFFFFF94644DCCADFD57F96F" authority="Kaup" authorityName="Kaup" box="[529,709,1682,1706]" class="Insecta" family="Passallidae" genus="Ptichopus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF94644DCCADFD13F96F" box="[529,641,1682,1706]" italics="true" pageId="6" pageNumber="7">Ptichopus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF94648FCCADFC87F96F" box="[723,789,1682,1706]" captionStart="FIGURE 6" captionStartId="9.[151,250,1347,1370]" captionTargetBox="[159,1432,198,1314]" captionTargetId="figure@9.[151,1436,193,1326]" captionTargetPageId="9" captionText="FIGURE 6. Passalinae hindgut. A— Ptichopus angulatus (Percheron), Costa Rica; B— Passalus interruptus (Linnaeus), Brazil; C— Spasalus crenatus (MacLeay), Puerto Rico; D— Paxillus leachi MacLeay, Brazil; E— Passipassalus buhrnheimi Fonseca &amp; Reyes-Castillo, Brazil (Fig. E from Fonseca &amp; Reyes – Castillo 1993)." httpUri="https://zenodo.org/record/200840/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
A),
<taxonomicName id="A68B729CFFFFFF94651DCCADFC37F96F" box="[833,933,1682,1706]" class="Insecta" family="Passallidae" genus="Passalus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF94651DCCADFC37F96F" box="[833,933,1682,1706]" italics="true" pageId="6" pageNumber="7">Passalus</emphasis>
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9465EFCCADFC64F96F" box="[947,1014,1682,1706]" captionStart="FIGURE 6" captionStartId="9.[151,250,1347,1370]" captionTargetBox="[159,1432,198,1314]" captionTargetId="figure@9.[151,1436,193,1326]" captionTargetPageId="9" captionText="FIGURE 6. Passalinae hindgut. A— Ptichopus angulatus (Percheron), Costa Rica; B— Passalus interruptus (Linnaeus), Brazil; C— Spasalus crenatus (MacLeay), Puerto Rico; D— Paxillus leachi MacLeay, Brazil; E— Passipassalus buhrnheimi Fonseca &amp; Reyes-Castillo, Brazil (Fig. E from Fonseca &amp; Reyes – Castillo 1993)." httpUri="https://zenodo.org/record/200840/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
B),
<taxonomicName id="A68B729CFFFFFF946243CCADFB57F96F" authority="Kaup" authorityName="Kaup" box="[1055,1221,1682,1706]" class="Insecta" family="Passallidae" genus="Spasalus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF946243CCADFB11F96F" box="[1055,1155,1682,1706]" italics="true" pageId="6" pageNumber="7">Spasalus</emphasis>
Kaup
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF946288CCADFA85F96F" box="[1236,1303,1682,1706]" captionStart="FIGURE 6" captionStartId="9.[151,250,1347,1370]" captionTargetBox="[159,1432,198,1314]" captionTargetId="figure@9.[151,1436,193,1326]" captionTargetPageId="9" captionText="FIGURE 6. Passalinae hindgut. A— Ptichopus angulatus (Percheron), Costa Rica; B— Passalus interruptus (Linnaeus), Brazil; C— Spasalus crenatus (MacLeay), Puerto Rico; D— Paxillus leachi MacLeay, Brazil; E— Passipassalus buhrnheimi Fonseca &amp; Reyes-Castillo, Brazil (Fig. E from Fonseca &amp; Reyes – Castillo 1993)." httpUri="https://zenodo.org/record/200840/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
C),
<taxonomicName id="A68B729CFFFFFF94631CCCADFE93F90A" authority="MacLeay" authorityName="MacLeay" class="Insecta" family="Passallidae" genus="Paxillus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF94631CCCADFA0EF96F" box="[1344,1436,1682,1706]" italics="true" pageId="6" pageNumber="7">Paxillus</emphasis>
MacLeay
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF94674ECC89FEC4F90B" box="[274,342,1718,1742]" captionStart="FIGURE 6" captionStartId="9.[151,250,1347,1370]" captionTargetBox="[159,1432,198,1314]" captionTargetId="figure@9.[151,1436,193,1326]" captionTargetPageId="9" captionText="FIGURE 6. Passalinae hindgut. A— Ptichopus angulatus (Percheron), Costa Rica; B— Passalus interruptus (Linnaeus), Brazil; C— Spasalus crenatus (MacLeay), Puerto Rico; D— Paxillus leachi MacLeay, Brazil; E— Passipassalus buhrnheimi Fonseca &amp; Reyes-Castillo, Brazil (Fig. E from Fonseca &amp; Reyes – Castillo 1993)." httpUri="https://zenodo.org/record/200840/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
D) and
<taxonomicName id="A68B729CFFFFFF9467F2CC89FCEFF90B" authority="Fonseca &amp; Reyes-Castillo" authorityName="Fonseca &amp; Reyes-Castillo" box="[430,893,1718,1742]" class="Insecta" family="Passallidae" genus="Passipassalus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="genus">
<emphasis id="53FFD50DFFFFFF9467F2CC89FDD9F90B" box="[430,587,1718,1742]" italics="true" pageId="6" pageNumber="7">Passipassalus</emphasis>
Fonseca &amp; Reyes-Castillo
</taxonomicName>
(
<figureCitation id="F9B0159AFFFFFF9465D2CC89FC46F90B" box="[910,980,1718,1742]" captionStart="FIGURE 6" captionStartId="9.[151,250,1347,1370]" captionTargetBox="[159,1432,198,1314]" captionTargetId="figure@9.[151,1436,193,1326]" captionTargetPageId="9" captionText="FIGURE 6. Passalinae hindgut. A— Ptichopus angulatus (Percheron), Costa Rica; B— Passalus interruptus (Linnaeus), Brazil; C— Spasalus crenatus (MacLeay), Puerto Rico; D— Paxillus leachi MacLeay, Brazil; E— Passipassalus buhrnheimi Fonseca &amp; Reyes-Castillo, Brazil (Fig. E from Fonseca &amp; Reyes – Castillo 1993)." httpUri="https://zenodo.org/record/200840/files/figure.png" pageId="6" pageNumber="7">Fig. 6</figureCitation>
E); the ileum becomes flattened and the protuberances increase from four to six rows, becoming noticeably denser in Macrolininae.
</paragraph>
</subSubSection>
</treatment>
</document>
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