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<treatmentCitation id="345110B00772FF9EFEE7A643024CFBEA" author="Ferreira" box="[359,547,1034,1056]" page="779" pageId="13" pageNumber="14" year="1972">
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: 779
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<emphasis id="8784EAB30772FF9EFF17A6600121FBF5" box="[151,334,1065,1087]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
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<treatmentCitation id="345110B00772FF9EFEE7A660024EFBF5" author="Kabakov" box="[359,545,1065,1087]" page="94" pageId="13" pageNumber="14" year="1979">
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: 94
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(diagnosis)
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: 413
</treatmentCitation>
(new combination, comment taxonomy)
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: 17
</treatmentCitation>
(comment taxonomy)
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: 260
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(diagnosis, comment)
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Monaghan
<emphasis id="8784EAB30772FF9EFE59A6EC0262FB70" box="[473,525,1188,1210]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
2007
</bibRefCitation>
: 678
</treatmentCitation>
(phylogeny)
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<emphasis id="8784EAB30772FF9EFF17A68B0121FB12" box="[151,334,1218,1240]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
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: 24
</treatmentCitation>
(comment)
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<treatmentCitation id="345110B00772FF9EFEE7A6AB02D9FB3D" author="Tarasov" box="[359,694,1249,1271]" page="2" pageId="13" pageNumber="14" year="2011">
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: 2
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(phylogeny)
<taxonomicName id="72F04D220772FF9EFF17A7490121FADC" ID-CoL="45DS" box="[151,334,1280,1302]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFF17A7490121FADC" box="[151,334,1280,1302]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
<treatmentCitation id="345110B00772FF9EFEE7A7490191FADC" author="Genier" box="[359,510,1280,1302]" page="2" pageId="13" pageNumber="14" year="2013">
<bibRefCitation id="D1614B500772FF9EFEE7A7490187FADC" author="Genier" box="[359,488,1280,1302]" pageId="13" pageNumber="14" refString="Genier, F. (2013) Transfert d' Onthophagus bubalus Harold, 1867, dans le genre Phalops Erichson, 1847, et notes sur sa position phylogenetique (Coleoptera, Scarabaeidae, Scarabaeinae). Catharsius, La Revue, 7, 1 - 4." type="journal article" year="2013">Génier 2013</bibRefCitation>
: 2
</treatmentCitation>
(comment taxonomy)
</treatmentCitationGroup>
</paragraph>
<paragraph id="B54F36A10772FF9EFF17A757028BFAFE" blockId="13.[151,953,942,1424]" box="[151,740,1310,1333]" pageId="13" pageNumber="14">
<treatmentCitationGroup id="95E0118F0772FF9EFF17A757028BFAFE" box="[151,740,1310,1333]" pageId="13" pageNumber="14">
<taxonomicName id="72F04D220772FF9EFF17A7570121FAFE" ID-CoL="45DS" box="[151,334,1310,1332]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFF17A7570121FAFE" box="[151,334,1310,1332]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
<treatmentCitation id="345110B00772FF9EFEE7A757020DFAFE" author="Mlambo" box="[359,610,1310,1333]" page="318" pageId="13" pageNumber="14" year="2015">
<bibRefCitation id="D1614B500772FF9EFEE7A757025CFAFE" author="Mlambo" box="[359,563,1310,1333]" pageId="13" pageNumber="14" refString="Mlambo, S., Sole, C. L. &amp; Scholtz, C. H. (2015) A molecular phylogeny of the African Scarabaeinae (Coleoptera: Scarabaeidae). Arthropod Systematics &amp; Phylogeny, 73, 303 - 321." type="journal article" year="2015">
Mlambo
<emphasis id="8784EAB30772FF9EFE43A7690199FAFE" box="[451,502,1310,1333]" italics="true" pageId="13" pageNumber="14">et al.</emphasis>
2015
</bibRefCitation>
: 318
</treatmentCitation>
(phylogeny)
</treatmentCitationGroup>
</paragraph>
<paragraph id="B54F36A10772FF9EFF17A7740287FAB8" blockId="13.[151,953,942,1424]" pageId="13" pageNumber="14">
<treatmentCitationGroup id="95E0118F0772FF9EFF17A7740287FAB8" pageId="13" pageNumber="14">
<taxonomicName id="72F04D220772FF9EFF17A7740121FA99" ID-CoL="45DS" box="[151,334,1341,1363]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFF17A7740121FA99" box="[151,334,1341,1363]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
<treatmentCitation id="345110B00772FF9EFEE7A7740213FA99" author="Breeschoten" box="[359,636,1341,1363]" page="87" pageId="13" pageNumber="14" year="2016">
<bibRefCitation id="D1614B500772FF9EFEE7A7740236FA99" author="Breeschoten" box="[359,601,1341,1363]" pageId="13" pageNumber="14" refString="Breeschoten, T., Doorenweerd, C., Tarasov, S. &amp; Vogler, A. P. (2016) Phylogenetics and biogeography of the dung beetle genus Onthophagus inferred from mitochondrial genomes. Molecular Phylogenetics and Evolution, 105, 86 - 95. https: // doi. org / 10.1016 / j. ympev. 2016.08.016" type="journal article" year="2016">
Breeschoten
<emphasis id="8784EAB30772FF9EFE6AA7770278FA99" box="[490,535,1341,1363]" italics="true" pageId="13" pageNumber="14">et al</emphasis>
, 2016
</bibRefCitation>
: 87
</treatmentCitation>
(phylogeny)
<taxonomicName id="72F04D220772FF9EFF17A7150121FAB8" ID-CoL="45DS" box="[151,334,1372,1394]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFF17A7150121FAB8" box="[151,334,1372,1394]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
<bibRefCitation id="D1614B500772FF9EFEE7A7140250FAB8" author="Roggero" box="[359,575,1372,1394]" pageId="13" pageNumber="14" refString="Roggero, A., Barbero, E. &amp; Palestrini, C. (2016 a) Revised classification and phylogeny of an Afrotropical species group based on molecular and morphological data, with the description of a new genus (Coleoptera: Scarabaeidae: Onthophagini). Organisms Diversity &amp; Evolution, 2016, [1 - 18]." type="journal article" year="2016" yearSuffix="a">
Roggero
<emphasis id="8784EAB30772FF9EFE44A714019EFAB8" box="[452,497,1372,1394]" italics="true" pageId="13" pageNumber="14">et al</emphasis>
. 2016a
</bibRefCitation>
: [1] (phylogeny)
</treatmentCitationGroup>
</paragraph>
<paragraph id="B54F36A10772FF9EFF17A73302B7FA5A" blockId="13.[151,953,942,1424]" box="[151,728,1402,1424]" pageId="13" pageNumber="14">
<treatmentCitationGroup id="95E0118F0772FF9EFF17A73302B7FA5A" box="[151,728,1402,1424]" pageId="13" pageNumber="14">
<taxonomicName id="72F04D220772FF9EFF17A7330121FA5A" ID-CoL="45DS" box="[151,334,1402,1424]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFF17A7330121FA5A" box="[151,334,1402,1424]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
<treatmentCitation id="345110B00772FF9EFEE7A7320238FA5A" author="Roggero" box="[359,599,1402,1424]" page="2" pageId="13" pageNumber="14" year="2016">
<bibRefCitation id="D1614B500772FF9EFEE7A732022FFA5A" author="Roggero" box="[359,576,1402,1424]" pageId="13" pageNumber="14" refString="Roggero, A., Dierkens, M., Barbero, E. &amp; Palestrini, C. (2016 b) Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society, 2016, 1 - 23. https: // doi. org / 10.1111 / zoj. 12498" type="journal article" year="2016" yearSuffix="b">
Roggero
<emphasis id="8784EAB30772FF9EFE44A732019EFA5A" box="[452,497,1402,1424]" italics="true" pageId="13" pageNumber="14">et al</emphasis>
. 2016b
</bibRefCitation>
: 2
</treatmentCitation>
(phylogeny)
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="FDEA652A0772FF9EFF17A7F403A3FA1C" box="[151,972,1469,1494]" pageId="13" pageNumber="14" type="materials_examined">
<paragraph id="B54F36A10772FF9EFF17A7F403A3FA1C" blockId="13.[151,1437,1469,2034]" box="[151,972,1469,1494]" pageId="13" pageNumber="14">
<emphasis id="8784EAB30772FF9EFF17A7F40144FA1C" bold="true" box="[151,299,1469,1494]" pageId="13" pageNumber="14">Type species</emphasis>
.
<taxonomicName id="72F04D220772FF9EFEB9A7F402BBFA1C" authority="Fabricius, 1775" authorityName="Fabricius" authorityYear="1775" box="[313,724,1469,1494]" class="Insecta" family="Scarabaeidae" genus="Scarabaeus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="species" species="bonasus">
<emphasis id="8784EAB30772FF9EFEB9A7F40270FA1F" box="[313,543,1469,1493]" italics="true" pageId="13" pageNumber="14">Scarabaeus bonasus</emphasis>
<bibRefCitation id="D1614B500772FF9EFDA5A7F402BBFA1C" author="Fabricius" box="[549,724,1469,1494]" pageId="13" pageNumber="14" refString="Fabricius, J. C. (1775) Systema Entomologiae sistens Insectorum Classes, Ordines, Genera, Species adiectis Synonymis, Locis Descriptionibus, Observationibus. Officina Libraria Kortii, Flensburg &amp; Leipzig, Germany, xxxii + 832 pp." type="book" year="1775">Fabricius, 1775</bibRefCitation>
</taxonomicName>
(original designation)
</paragraph>
</subSubSection>
<subSubSection id="FDEA652A0772FF9EFF47A7A8035BF863" pageId="13" pageNumber="14" type="diagnosis">
<paragraph id="B54F36A10772FF9EFF47A7A8035BF863" blockId="13.[151,1437,1469,2034]" pageId="13" pageNumber="14">
<emphasis id="8784EAB30772FF9EFF47A7A80155FA30" bold="true" box="[199,314,1505,1530]" pageId="13" pageNumber="14">Diagnosis</emphasis>
. Externally, the genus
<taxonomicName id="72F04D220772FF9EFDE8A7A8035DFA33" box="[616,818,1505,1529]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFDE8A7A8035DFA33" box="[616,818,1505,1529]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
is almost indistinguishable from the rest of the
<taxonomicName id="72F04D220772FF9EFF17A44C015AF9D4" box="[151,309,1541,1566]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="tribe" tribe="Onthophagini">Onthophagini</taxonomicName>
. However, all
<taxonomicName id="72F04D220772FF9EFE68A44C02DDF9D7" box="[488,690,1541,1565]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFE68A44C02DDF9D7" box="[488,690,1541,1565]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
species have the anterior portion of the hypomeron produced anteriorly and visible from above; therefore, setae that would be located along the anterior edge in ventral portion are moved to a dorsal position. The only other external characters found separating the genus
<taxonomicName id="72F04D220772FF9EFB52A40405F3F9AF" box="[1234,1436,1613,1637]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFB52A40405F3F9AF" box="[1234,1436,1613,1637]" italics="true" pageId="13" pageNumber="14">Digitonthophagus</emphasis>
</taxonomicName>
and the other closely related genera, is the presence of a distinctly shaped internoapical tooth of the male protibia (
<figureCitation id="2DCB2A240772FF9EFF1FA4DC0172F964" box="[159,285,1685,1710]" captionStart-0="FIGURES 7 10" captionStart-1="FIGURES 11 14" captionStart-2="FIGURES 15 18" captionStart-3="FIGURES 19 22" captionStartId-0="87.[151,264,1949,1971]" captionStartId-1="88.[151,264,1977,1999]" captionStartId-2="89.[151,264,1942,1964]" captionStartId-3="90.[151,264,1959,1981]" captionTargetBox-0="[228,1347,259,1889]" captionTargetBox-1="[202,1376,213,1923]" captionTargetBox-2="[218,1372,259,1904]" captionTargetBox-3="[200,1342,226,1900]" captionTargetId-0="figure@87.[201,1365,193,1905]" captionTargetId-1="figure@88.[196,1383,193,1938]" captionTargetId-2="figure@89.[206,1381,193,1921]" captionTargetId-3="figure@90.[195,1367,193,1916]" captionTargetPageId-0="87" captionTargetPageId-1="88" captionTargetPageId-2="89" captionTargetPageId-3="90" captionText-0="FIGURES 7 10. Male dorsal habitus. 7. Digitonthophagus bonasus, alloreferent (Tamil Nadu State, India); 8. D. uks, holotype; 9. D. sahelicus, holotype; 10. D. catta, alloreferent (Tamil Nadu State, India)." captionText-1="FIGURES 11 14. Male dorsal habitus. 11. D. eucatta, holotype; 12. D. falciger, holotype; 13. Digitonthophagus biflagellatus, holotype; 14. D. gazella, non type (Central Province, Zambia)." captionText-2="FIGURES 15 18. Male dorsal habitus. 15. D. petilus, holotype; 16. D. viridicollis, holotype; 17. D. aksumensis, holotype; 18. D. dilatatus, holotype." captionText-3="FIGURES 19 22. Male dorsal habitus. 19. Digitonthophagus fimator, holotype; 20. D. lusinganus, lectotype; 21. D. ulcerosus, holotype; 22. D. namaquensis, holotype." httpUri-0="https://zenodo.org/record/439451/files/figure.png" httpUri-1="https://zenodo.org/record/439452/files/figure.png" httpUri-2="https://zenodo.org/record/439453/files/figure.png" httpUri-3="https://zenodo.org/record/439454/files/figure.png" pageId="13" pageNumber="14">Figs. 722</figureCitation>
). This tooth is usually elongate, ventrally flat, and obliquely oriented downward. This character must be combined with females completely lacking this protibial internoapical tooth. In
<taxonomicName id="72F04D220772FF9EFBC9A4F0048FF91B" box="[1097,1248,1721,1745]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="14" phylum="Arthropoda" rank="genus">
<emphasis id="8784EAB30772FF9EFBC9A4F0048FF91B" box="[1097,1248,1721,1745]" italics="true" pageId="13" pageNumber="14">Onthophagus</emphasis>
</taxonomicName>
species where a similar tooth is present, the shape of this tooth is spiniform and forwardly directed in males and present in females as well but reduced in size (in African species examined). Internally, the parameres are rather short and the basolateral paramerites always present with the apex of the later adjacent to the lateral portion of the apical carinae. The internal sac of the aedeagus is lacking the lamella copulatrix and associated small sclerites and bristle pads. The frontolateral peripheral sclerite (
<figureCitation id="2DCB2A240772FF9EFDB4A52402F0F84C" box="[564,671,1901,1926]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStart-2="FIGURE 4" captionStart-3="FIGURE 5" captionStartId-0="6.[151,250,836,858]" captionStartId-1="6.[151,250,1897,1919]" captionStartId-2="7.[151,250,1857,1879]" captionStartId-3="8.[151,250,1240,1262]" captionTargetBox-0="[345,1247,205,812]" captionTargetBox-1="[151,1436,910,1872]" captionTargetBox-2="[151,1436,193,1835]" captionTargetBox-3="[151,1436,193,1219]" captionTargetId-0="figure@6.[340,1247,195,812]" captionTargetId-1="figure@6.[151,1436,910,1872]" captionTargetId-2="figure@7.[151,1436,193,1835]" captionTargetId-3="figure@8.[151,1436,193,1219]" captionTargetPageId-0="6" captionTargetPageId-1="6" captionTargetPageId-2="7" captionTargetPageId-3="8" captionText-0="FIGURE 2. Apical portion of internal sac of Digitonthophagus bonasus (membranous portion colored red)." captionText-1="FIGURE 3. Detailed terminology of the frontolateral peripheral and associated sclerites of Digitonthophagus gazella." captionText-2="FIGURE 4. Detailed terminology of the frontolateral peripheral and associated sclerites of Digitonthophagus gazella, right lateral view." captionText-3="FIGURE 5. Detailed terminology of the frontolateral peripheral and associated sclerites of Digitonthophagus eucatta, ventral view." httpUri-0="https://zenodo.org/record/439446/files/figure.png" httpUri-1="https://zenodo.org/record/439447/files/figure.png" httpUri-2="https://zenodo.org/record/439448/files/figure.png" httpUri-3="https://zenodo.org/record/439449/files/figure.png" pageId="13" pageNumber="14">Figs. 25</figureCitation>
) is located at the apex of the internal sac and a subapicoventral lobe oriented perpendicularly to the main plan of FLP is present.
</paragraph>
</subSubSection>
<subSubSection id="FDEA652A0772FF9DFF47A5FC00B0FD62" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14" type="description">
<paragraph id="B54F36A10772FF9DFF47A5FC00B0FD62" blockId="13.[151,1437,1469,2034]" lastBlockId="14.[151,1437,151,968]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
<emphasis id="8784EAB30772FF9EFF47A5FC013EF804" bold="true" box="[199,337,1973,1998]" pageId="13" pageNumber="14">Description</emphasis>
.
<emphasis id="8784EAB30772FF9EFEDFA5FC01E0F804" bold="true" box="[351,399,1973,1998]" pageId="13" pageNumber="14">Size</emphasis>
. Small to moderately large (6.517.0 mm).
<emphasis id="8784EAB30772FF9EFCFCA5FC03D0F804" bold="true" box="[892,959,1973,1998]" pageId="13" pageNumber="14">Color</emphasis>
. Overall color fulvous to light brown with structure edges and sutures more-or-less darker; pronotal disc and head darker and with distinct metallic sheen; elytral margins brown to green, disc with a more-or-less distinct darker irregular macula; metasternum and femoral ventral surface usually with much darker maculae.
<emphasis id="8784EAB30771FF9DFD70A2F50340FF1F" bold="true" box="[752,815,188,213]" pageId="14" pageNumber="15">Head</emphasis>
. With a sharply defined clypeofrontal carina in both sexes; antenna with 9 antennomeres, with deep pits on distal surface of antennomeres 78.
<emphasis id="8784EAB30771FF9DFB59A296053EFF32" bold="true" box="[1241,1361,223,248]" pageId="14" pageNumber="15">Pronotum</emphasis>
. With distinct squamiform granules anteriorly posterior to the anterior declivity that is always simply punctate, granules larger anteriorly and extending more-or-less posteriorly; anterior discal punctures weakly-defined and becoming well-defined posteriorly, when present.
<emphasis id="8784EAB30771FF9DFDE6A30502DDFEAF" bold="true" box="[614,690,332,357]" pageId="14" pageNumber="15">Elytra</emphasis>
. Juxtasutural interval with strong metallic sheen along sutures; usually with a dark oval macula on the apical declivity on intervals 45.
<emphasis id="8784EAB30771FF9DFC60A326043CFE42" bold="true" box="[992,1107,367,392]" pageId="14" pageNumber="15">Pygidium</emphasis>
. Surface with scattered, fine granulate punctures; each puncture with an erect, yellow seta.
<emphasis id="8784EAB30771FF9DFCF4A3DD038FFE67" bold="true" box="[884,992,404,429]" pageId="14" pageNumber="15">Ventrites</emphasis>
. Hypomeron slightly convex; anterior hypomeral carina well-defined throughout; mesosternum with a bisinuate, glossy surface anteriorly and a more-orless developed longitudinal glossy, keel-shaped structure medially; posterior portion completely rugopunctate and setose; metasternal median lobe usually with a distinct longitudinal ridge behind the anterior suture.
<emphasis id="8784EAB30771FF9DFAA2A3B605FAFDD2" bold="true" box="[1314,1429,511,536]" pageId="14" pageNumber="15">Aedeagus</emphasis>
. Parameres rather short and basolateral paramerites always present, with the apex of the later adjacent to the lateral portion of the apical carinae.
<emphasis id="8784EAB30771FF9DFD81A00E02A2FDAA" bold="true" box="[513,717,583,608]" pageId="14" pageNumber="15">Female genitalia</emphasis>
. Infundibulum thin, simple with a median portion reduced; receptaculum seminis elongate, simple, lacking annulate ridges internally (female genitalia characters from
<bibRefCitation id="D1614B500771FF9DFACBA02500BCFD62" author="Zunino" pageId="14" pageNumber="15" refString="Zunino, M. (1981) Insect of Saudi Arabia, Coleoptera: Fam. Scarabaeidae, Tribe Onthophagini. Fauna of Saudi Arabia, 3, 408 - 416." type="journal article" year="1981">Zunino 1981</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="FDEA652A0771FF9DFF47A0FD01CEFA57" pageId="14" pageNumber="15" type="discussion">
<paragraph id="B54F36A10771FF9DFF47A0FD0503FC02" blockId="14.[151,1437,151,968]" pageId="14" pageNumber="15">
<emphasis id="8784EAB30771FF9DFF47A0FD015BFD07" bold="true" box="[199,308,692,717]" pageId="14" pageNumber="15">Remarks</emphasis>
. Balthasars (1959) original description was restricted to external morphology and included a number of unrelated species (
<taxonomicName id="72F04D220771FF9DFE67A09102B7FD3A" authority="Balthasar" authorityName="Balthasar" box="[487,728,727,752]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="actaeon">
<emphasis id="8784EAB30771FF9DFE67A091020BFD3A" box="[487,612,728,752]" italics="true" pageId="14" pageNumber="15">O. actaeon</emphasis>
Balthasar
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFD63A0910461FD25" authority="Boucomont" authorityName="Boucomont" box="[739,1038,728,752]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="anguliceps">
<emphasis id="8784EAB30771FF9DFD63A0910310FD3A" box="[739,895,728,752]" italics="true" pageId="14" pageNumber="15">O. anguliceps</emphasis>
Boucomont
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFB9AA0910480FD3A" authority="Arrow" authorityName="Arrow" box="[1050,1263,728,752]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="avocetta">
<emphasis id="8784EAB30771FF9DFB9AA09104F2FD3A" box="[1050,1181,728,752]" italics="true" pageId="14" pageNumber="15">O. avocetta</emphasis>
Arrow
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFB7CA0910161FCDF" authority="van Lansberge" authorityName="van Lansberge" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="blumei">
<emphasis id="8784EAB30771FF9DFB7CA0910504FD3A" box="[1276,1387,728,752]" italics="true" pageId="14" pageNumber="15">O. blumei</emphasis>
van Lansberge
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFE9BA0B50261FCDF" authority="Fabricius" authorityName="Fabricius" box="[283,526,764,789]" class="Insecta" family="Scarabaeidae" genus="Scarabaeus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="bonasus">
<emphasis id="8784EAB30771FF9DFE9BA0B501F5FCDE" box="[283,410,764,788]" italics="true" pageId="14" pageNumber="15">O. bonasus</emphasis>
Fabricius
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFD9BA0B50359FCDF" authority="d'Orbigny" authorityName="d'Orbigny" box="[539,822,764,789]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="cameloides">
<emphasis id="8784EAB30771FF9DFD9BA0B502D2FCDE" box="[539,701,764,788]" italics="true" pageId="14" pageNumber="15">O. cameloides</emphasis>
d'Orbigny
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFCC4A0B504F5FCDE" authority="Boucomont" authorityName="Boucomont" box="[836,1178,764,788]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="curvicarinatus">
<emphasis id="8784EAB30771FF9DFCC4A0B50464FCDE" box="[836,1035,764,788]" italics="true" pageId="14" pageNumber="15">O. curvicarinatus</emphasis>
Boucomont
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFB26A0B505F7FCDF" authority="Harold" authorityName="Harold" box="[1190,1432,764,789]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="diabolicus">
<emphasis id="8784EAB30771FF9DFB26A0B50550FCDE" box="[1190,1343,764,788]" italics="true" pageId="14" pageNumber="15">O. diabolicus</emphasis>
Harold
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFF17A169011DFCF2" authority="Arrow" authorityName="Arrow" box="[151,370,800,824]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="digitatus">
<emphasis id="8784EAB30771FF9DFF17A1690171FCF2" box="[151,286,800,824]" italics="true" pageId="14" pageNumber="15">O. digitatus</emphasis>
Arrow
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFE00A16902A6FCFD" authority="Boucomont" authorityName="Boucomont" box="[384,713,800,824]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="diversiformis">
<emphasis id="8784EAB30771FF9DFE00A1690257FCF2" box="[384,568,800,824]" italics="true" pageId="14" pageNumber="15">O. diversiformis</emphasis>
Boucomont
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFD56A16903FBFCF2" authority="Arrow" authorityName="Arrow" box="[726,916,800,824]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="fossor">
<emphasis id="8784EAB30771FF9DFD56A1690350FCF2" box="[726,831,800,824]" italics="true" pageId="14" pageNumber="15">O. fossor</emphasis>
Arrow
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFC22A16904E3FCF2" authority="Fabricius" authorityName="Fabricius" box="[930,1164,799,824]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="gazella">
<emphasis id="8784EAB30771FF9DFC22A1690477FCF2" box="[930,1048,800,824]" italics="true" pageId="14" pageNumber="15">O. gazella</emphasis>
Fabricius
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFB1BA169051CFCFD" authority="Bates" authorityName="Bates" box="[1179,1395,800,824]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="kuluensis">
<emphasis id="8784EAB30771FF9DFB1BA1690547FCF2" box="[1179,1320,800,824]" italics="true" pageId="14" pageNumber="15">O. kuluensis</emphasis>
Bates
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFA01A1690110FC96" authority="Arrow" authorityName="Arrow" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="manipurensis">
<emphasis id="8784EAB30771FF9DFA01A1690142FC96" italics="true" pageId="14" pageNumber="15">O. manipurensis</emphasis>
Arrow
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFE0CA10D02FDFC96" authority="Nomura" authorityName="Nomura" box="[396,658,836,860]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="ohbayashii">
<emphasis id="8784EAB30771FF9DFE0CA10D0245FC96" box="[396,554,836,860]" italics="true" pageId="14" pageNumber="15">O. ohbayashii</emphasis>
Nomura
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFD1EA10D03FCFC96" authority="Boucomont" authorityName="Boucomont" box="[670,915,836,860]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="ribbei">
<emphasis id="8784EAB30771FF9DFD1EA10D036AFC96" box="[670,773,836,860]" italics="true" pageId="14" pageNumber="15">O. ribbei</emphasis>
Boucomont
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFC1FA10D04EEFC96" authority="Hope" authorityName="Hope" box="[927,1153,836,860]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="rubricollis">
<emphasis id="8784EAB30771FF9DFC1FA10D0456FC96" box="[927,1081,836,860]" italics="true" pageId="14" pageNumber="15">O. rubricollis</emphasis>
Hope
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFB0DA10D051AFC97" authority="Harold" authorityName="Harold" box="[1165,1397,836,861]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="solivagus">
<emphasis id="8784EAB30771FF9DFB0DA10D0574FC96" box="[1165,1307,836,860]" italics="true" pageId="14" pageNumber="15">O. solivagus</emphasis>
Harold
</taxonomicName>
,
<taxonomicName id="72F04D220771FF9DFA01A10D0103FC4A" authority="Balthasar" authorityName="Balthasar" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="troniceki">
<emphasis id="8784EAB30771FF9DFA01A10D0095FC4A" italics="true" pageId="14" pageNumber="15">O. troniceki</emphasis>
Balthasar
</taxonomicName>
, and
<taxonomicName id="72F04D220771FF9DFE26A12102EEFC4A" authority="Harold" authorityName="Harold" box="[422,641,871,896]" class="Insecta" family="Scarabaeidae" genus="Onthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="wallacei">
<emphasis id="8784EAB30771FF9DFE26A1210247FC4A" box="[422,552,872,896]" italics="true" pageId="14" pageNumber="15">O. wallacei</emphasis>
Harold
</taxonomicName>
). The main characteristic used by Balthasar to define his subgenus was the elongate and thin protibia of the male. This character state is highly homoplasious in
<taxonomicName id="72F04D220771FF9DFBFAA1C50561FC6F" box="[1146,1294,908,933]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="subFamily" subFamily="Scarabaeinae">Scarabaeinae</taxonomicName>
and evolved many times in the
<taxonomicName id="72F04D220771FF9DFEE6A1E6026BFC02" box="[358,516,943,968]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="tribe" tribe="Onthophagini">Onthophagini</taxonomicName>
. The subgenus, as originally defined, was clearly paraphyletic (
<bibRefCitation id="D1614B500771FF9DFB49A1E60530FC02" author="Zunino" box="[1225,1375,943,968]" pageId="14" pageNumber="15" refString="Zunino, M. (1981) Insect of Saudi Arabia, Coleoptera: Fam. Scarabaeidae, Tribe Onthophagini. Fauna of Saudi Arabia, 3, 408 - 416." type="journal article" year="1981">Zunino 1981</bibRefCitation>
).
</paragraph>
<paragraph id="B54F36A10771FF9DFF17A1BE0246FAC7" blockId="14.[151,1436,1015,1437]" pageId="14" pageNumber="15">
<emphasis id="8784EAB30771FF9DFF17A1BE0348FBDA" bold="true" box="[151,807,1015,1040]" pageId="14" pageNumber="15">Remarks for species descriptions and identification key</emphasis>
. Diagnoses based on external morphology refer to welldeveloped individuals. As in many scarabaeine species, strong allometric variation in the development of external secondary sexual characters is present. Smallest males will show completely feminized head and thoracic ornamentation, while largest males present two horns on the vertex. Pronotal and elytral granulations and punctation are usually diagnostic but variable, and extremes of one species can be similar to individuals of another species, which makes identification difficult with external characters for some specimens. Examination of the FLP sclerite shape will always provide a definite identification for males. Diagnoses for species groups must be used in conjunction with species diagnoses.
</paragraph>
<paragraph id="B54F36A10771FF9DFF47A75E01CEFA57" blockId="14.[151,1436,1015,1437]" pageId="14" pageNumber="15">
Species are arranged into six groups based on the current phylogeny in order to facilitate comparison of closely-related species. Species are listed in alphabetical order within each group except for the
<taxonomicName id="72F04D220771FF9DFB52A775009CFAB2" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="bonasus">
<emphasis id="8784EAB30771FF9DFB52A775009CFAB2" italics="true" pageId="14" pageNumber="15">Digitonthophagus bonasus</emphasis>
</taxonomicName>
group, where
<taxonomicName id="72F04D220771FF9DFE14A72801B5FAB2" box="[404,474,1376,1400]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="uks">
<emphasis id="8784EAB30771FF9DFE14A72801B5FAB2" box="[404,474,1376,1400]" italics="true" pageId="14" pageNumber="15">D. uks</emphasis>
</taxonomicName>
is treated next to
<taxonomicName id="72F04D220771FF9DFD21A7280370FAB2" box="[673,799,1376,1400]" class="Insecta" family="Scarabaeidae" genus="Digitonthophagus" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="15" phylum="Arthropoda" rank="species" species="bonasus">
<emphasis id="8784EAB30771FF9DFD21A7280370FAB2" box="[673,799,1376,1400]" italics="true" pageId="14" pageNumber="15">D. bonasus</emphasis>
</taxonomicName>
since it is the most closely related species from the same biogeographical region.
</paragraph>
</subSubSection>
</treatment>
</document>