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<mods:title id="7BA6B40464B9E7CAE145AF08353450E9">The Phanaeus tridens species group (Coleoptera: Scarabaeoidea): a dung beetle group with genital morphological stasis but a changing ecological niche</mods:title>
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<mods:affiliation id="8AB3314BFE91AD25D2295B53A778823A">Centro Tlaxcala de Biología de la Conducta, Universidad Autónoma de Tlaxcala, Tlaxcala de Xicohténcatl, Tlaxcala 90070, Mexico, Km 1.5 Carretera Tlaxcala-Puebla S / N, La Loma Xicohtencatl.</mods:affiliation>
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<mods:namePart id="692C906B2ACD5C1C89609AA8C1AB8DE3">Halffter, Gonzalo</mods:namePart>
<mods:affiliation id="3277FC2CE1B083E200DBBC89E4BDD11E">Emeritus Researcher, Instituto de Ecología A. C., Xalapa, Veracruz 91073, Mexico, Carretera antigua a Coatepec 351, El Haya.</mods:affiliation>
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<mods:namePart id="A3D47F50DC8C8EFB68438D37CEC6D2C2">Lizardo, Viridiana</mods:namePart>
<mods:affiliation id="00B51783CBE2A237DA95122FD4E899AD">Facultad de Ciencias, Universidad Nacional Autónoma de México, Mexico City 04510, Mexico, Apartado Postal 70 - 399.</mods:affiliation>
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<taxonomicName id="54824D1F8A32FFA5FECBFF14ED56FF67" ID-CoL="4FSKJ" authority="Bates, 1887" authorityName="Bates" authorityYear="1887" box="[286,624,190,213]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="eximius">
<emphasis id="A1F6EA8E8A32FFA5FECBFF14ED56FF67" bold="true" box="[286,624,190,213]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FECBFF14EECFFF67" bold="true" box="[286,489,190,213]" italics="true" pageId="24" pageNumber="471">Phanaeus eximius</emphasis>
<bibRefCitation id="F7134B6D8A32FFA5FE3AFF14ED56FF67" author="Bates" box="[495,624,190,213]" firstAuthor="Bates" pageId="24" pageNumber="471" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">Bates, 1887</bibRefCitation>
</emphasis>
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<paragraph id="933D369C8A32FFA5FEA4FF76ED38FF5D" blockId="24.[151,677,220,273]" box="[369,542,220,239]" pageId="24" pageNumber="471">
(
<figureCitation id="0BB92A198A32FFA5FEA2FF76EEFFFF5D" box="[375,473,220,239]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29–31</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FE37FF76EEDFFF5D" box="[482,505,220,239]" captionStart="Figs 32–42" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="24" pageNumber="471">42</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FDD7FF76ED3EFF5D" box="[514,536,220,239]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">54</figureCitation>
)
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<paragraph id="933D369C8A32FFA5FF42FF54ED83FEA3" blockId="24.[151,677,220,273]" box="[151,677,254,273]" pageId="24" pageNumber="471">
<treatmentCitationGroup id="B39211B28A32FFA5FF42FF54ED83FEA3" box="[151,677,254,273]" pageId="24" pageNumber="471">
<treatmentCitation id="1223108D8A32FFA5FF42FF54EE99FEA3" author="BATES H. W." box="[151,447,254,273]" page="62" pageId="24" pageNumber="471" year="1887">
<taxonomicName id="54824D1F8A32FFA5FF42FF54EE99FEA3" ID-CoL="4FSKJ" authority="Bates, 1887: 62" authorityName="Bates" authorityPageNumber="62" authorityYear="1887" box="[151,447,254,273]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="eximius">
<emphasis id="A1F6EA8E8A32FFA5FF42FF54EE14FEA3" box="[151,306,254,273]" italics="true" pageId="24" pageNumber="471">Phanaeus eximius</emphasis>
<bibRefCitation id="F7134B6D8A32FFA5FEE2FF54EE99FEA3" author="Bates" box="[311,447,254,273]" firstAuthor="Bates" page="62" pageId="24" pageNumber="471" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">Bates, 1887: 62</bibRefCitation>
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. 
<typeStatus id="4C39883E8A32FFA5FE12FF54EED7FEA3" box="[455,497,254,273]" pageId="24" pageNumber="471">Type</typeStatus>
locality: 
<collectingCountry id="EB95760C8A32FFA5FD97FF54ED87FEA3" box="[578,673,254,273]" name="Guatemala" pageId="24" pageNumber="471">Guatemala</collectingCountry>
.
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</paragraph>
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<subSubSection id="DB9865178A32FFA5FF42FE82EFCBFE72" pageId="24" pageNumber="471" type="materials_examined">
<paragraph id="933D369C8A32FFA5FF42FE82EE8BFEC2" blockId="24.[151,760,296,448]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FF42FE82EE5DFE89" bold="true" box="[151,379,296,315]" pageId="24" pageNumber="471">
<typeStatus id="4C39883E8A32FFA5FF42FE82EFE2FE89" box="[151,196,296,315]" pageId="24" pageNumber="471">Type</typeStatus>
material examined.
</emphasis>
L ECTOTYPE (designated by 
<bibRefCitation id="F7134B6D8A32FFA5FDB8FE82EDD3FE89" author="EDMONDS" box="[621,757,296,315]" firstAuthor="EDMONDS" pageId="24" pageNumber="471" pagination="1 - 105" refId="ref25450" refString="EDMONDS W. D. 1994: Revision of Phanaeus MacLeay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science of the Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">EDMONDS 1994</bibRefCitation>
, examined from photographs; 
<figureCitation id="0BB92A198A32FFA5FE73FEE8EECDFEE7" box="[422,491,322,341]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Fig. 31</figureCitation>
): ♂, without specific locality (
<collectionCode id="F593AE598A32FFA5FF48FEF7EFC7FEC2" box="[157,225,349,368]" pageId="24" pageNumber="471">BMNH</collectionCode>
: NHMUK 013903535).
</paragraph>
<paragraph id="933D369C8A32FFA5FF42FED2EFCBFE72" blockId="24.[151,760,296,448]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FF42FED2EFD7FE17" bold="true" pageId="24" pageNumber="471">
Non-type material examined. 
<collectingCountry id="EB95760C8A32FFA5FE64FED2ED67FE39" box="[433,577,376,395]" name="Guatemala" pageId="24" pageNumber="471">GUATEMALA</collectingCountry>
: W I THOUT SPEC I F I C LOCAL I TY:
</emphasis>
<specimenCount id="8584FD158A32FFA5FF2DFE38EE3CFE17" box="[248,282,396,421]" pageId="24" pageNumber="471" type="female">1 ♀</specimenCount>
(
<collectionCode id="F593AE598A32FFA5FEF2FE38EE4BFE17" box="[295,365,402,421]" pageId="24" pageNumber="471">GHVM</collectionCode>
). 
<emphasis id="A1F6EA8E8A32FFA5FEA8FE38EDE9FE17" bold="true" box="[381,719,402,421]" pageId="24" pageNumber="471">
<collectingCountry id="EB95760C8A32FFA5FEA8FE38ED23FE17" box="[381,517,402,421]" name="Nicaragua" pageId="24" pageNumber="471">NICARAGUA</collectingCountry>
: U NKNOWN LOCAL I TY:
</emphasis>
1 ♂ (
<collectionCode id="F593AE598A32FFA5FF48FE07EFC5FE72" box="[157,227,429,448]" pageId="24" pageNumber="471">GHVM</collectionCode>
).
</paragraph>
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<subSubSection id="DB9865178A32FFA5FF42FE73EEC5FC22" pageId="24" pageNumber="471" type="diagnosis">
<paragraph id="933D369C8A32FFA5FF42FE73EEC5FC22" blockId="24.[151,760,473,1584]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FF42FE73EE2AFE42" bold="true" box="[151,268,473,496]" pageId="24" pageNumber="471">Diagnosis.</emphasis>
Metallic green (
<figureCitation id="0BB92A198A32FFA5FE6AFE73ED1EFE42" box="[447,568,473,496]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Fig. 30–31</figureCitation>
), deep blue-green (
<figureCitation id="0BB92A198A32FFA5FF4AFE53EFD6FDA2" box="[159,240,505,528]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Fig. 29</figureCitation>
), dark blue (
<figureCitation id="0BB92A198A32FFA5FEADFE53EEEEFDA2" box="[376,456,505,528]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">Fig. 54</figureCitation>
), green-yellow or red. Sides of pronotal disc finely granulate (
<figureCitation id="0BB92A198A32FFA5FE3AFDB3ED40FD82" box="[495,614,537,560]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29, 31</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FDBAFDB3EDABFD82" box="[623,653,537,560]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">54</figureCitation>
). Pronotal disc coarsely granulorugose (
<figureCitation id="0BB92A198A32FFA5FE19FD93ED62FDE2" box="[460,580,569,592]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29, 31</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FD9BFD93ED4AFDE2" box="[590,620,569,592]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">54</figureCitation>
). Posteromedial process of pronotum produced into triangular denticle, widened laterally, not bifurcated apically (
<figureCitation id="0BB92A198A32FFA5FDBCFDD3ED99FD22" box="[617,703,633,656]" captionStart="Figs 32–42" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="24" pageNumber="471">Figs 42</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FD18FDD3EDCDFD22" box="[717,747,633,656]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">54</figureCitation>
). Anteromedial portion of pronotal disc with two strongly developed denticles (
<figureCitation id="0BB92A198A32FFA5FE54FD13EEF0FD62" box="[385,470,697,720]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">Fig. 54</figureCitation>
). Anterolateral margins of pronotal disc with inconspicuous line of tubercles (
<figureCitation id="0BB92A198A32FFA5FD19FD73EF93FCA2" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">Fig. 54</figureCitation>
). Posterolateral angles of pronotum more developed than posteromedial process (
<figureCitation id="0BB92A198A32FFA5FE07FCB3ED0EFC82" box="[466,552,793,816]" captionStart="Figs 32–42" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="24" pageNumber="471">Figs 42</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FDE2FCB3ED73FC82" box="[567,597,793,816]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">54</figureCitation>
). Elytral striae scabriculous, coarsely impressed, coarsely punctate (
<figureCitation id="0BB92A198A32FFA5FD1DFC93EFFBFCC2" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29–31</figureCitation>
). Elytral interstriae highly vermiculate, superficially punctate, convex (
<figureCitation id="0BB92A198A32FFA5FE8FFCD3EEF0FC22" box="[346,470,889,912]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29–31</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="C7FD66148A32FFA5FCE9FD8BEB4EFDFD" ID-DOI="http://doi.org/10.5281/zenodo.5821192" ID-Zenodo-Dep="5821192" httpUri="https://zenodo.org/record/5821192/files/figure.png" pageId="24" pageNumber="471" startId="24.[828,858,545,564]" targetBox="[841,1417,205,525]" targetPageId="24">
<paragraph id="933D369C8A32FFA5FCE9FD8BEB4EFDFD" blockId="24.[828,1436,545,591]" pageId="24" pageNumber="471">
Fig.63. Phallobase, parameres and endophallite copulatrix of the 
<taxonomicName id="54824D1F8A32FFA5FA85FD8BEABAFD86" authorityName="Castelnau" authorityYear="1840" box="[1360,1436,545,564]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A32FFA5FA85FD8BEABAFD86" box="[1360,1436,545,564]" italics="true" pageId="24" pageNumber="471">P.tridens</emphasis>
</taxonomicName>
species group. Scale bar = 1.0 mm.
</paragraph>
</caption>
<caption id="C7FD66148A32FFA5FCE9FDDAEBDAFD2C" ID-Table-UUID="C7FD66148A32FFA5FCE9FDDAEBDAFD2C" httpUri="http://table.plazi.org/id/C7FD66148A32FFA5FCE9FDDAEBDAFD2C" pageId="24" pageNumber="471" startId="24.[828,874,624,643]" targetBox="[838,1375,703,1038]" targetIsTable="true" targetPageId="24">
<paragraph id="933D369C8A32FFA5FCE9FDDAEBDAFD2C" blockId="24.[828,1436,624,670]" pageId="24" pageNumber="471">
Table 2. Number of single points used for modelling and model evaluation of the 
<taxonomicName id="54824D1F8A32FFA5FCA7FD21ECE4FD2C" authorityName="Castelnau" authorityYear="1840" box="[882,962,651,670]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A32FFA5FCA7FD21ECE4FD2C" box="[882,962,651,670]" italics="true" pageId="24" pageNumber="471">P. tridens</emphasis>
</taxonomicName>
species group’s distribution models.
</paragraph>
</caption>
<paragraph id="933D369C8A32FFA5FC93FD15EA1AFBBC" pageId="24" pageNumber="471">
<table id="E182C43C8A320042FC93FD15EA79FBBC" box="[838,1375,703,1038]" gridcols="3" gridrows="9" pageId="24" pageNumber="471">
<tr id="2DB234DE8A320042FC93FD15EA79FD5F" box="[838,1375,703,749]" gridrow="0" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FD15ECEFFD5F" box="[838,969,703,749]" gridcol="0" gridrow="0" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FC93FD15ECAEFD60" bold="true" box="[838,904,703,722]" pageId="24" pageNumber="471">Species</emphasis>
</th>
<th id="6E635DA28A320042FC3EFD15EBB6FD5F" box="[1003,1168,703,749]" gridcol="1" gridrow="0" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FC3EFD15EBB6FD5F" bold="true" pageId="24" pageNumber="471">Unique Points (training / testing)</emphasis>
</th>
<th id="6E635DA28A320042FB6CFD15EA79FD5F" box="[1209,1375,703,749]" gridcol="2" gridrow="0" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FB6CFD15EA00FD5F" bold="true" pageId="24" pageNumber="471">Continuous Boyce Index (CBI)</emphasis>
</th>
</tr>
<tr id="2DB234DE8A320042FC93FD54EA79FCA3" box="[838,1375,766,785]" gridrow="1" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FD54ECEFFCA3" box="[838,969,766,785]" gridcol="0" gridrow="1" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FD54EC95FCA3" authorityName="Arnaud" authorityYear="2001" baseAuthorityName="Arnaud" baseAuthorityYear="2001" box="[838,947,766,785]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="balthasari">
<emphasis id="A1F6EA8E8A32FFA5FC93FD54EC95FCA3" box="[838,947,766,785]" italics="true" pageId="24" pageNumber="471">P. balthasari</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFD54EBB6FCA3" box="[1003,1168,766,785]" gridcol="1" gridrow="1" pageId="24" pageNumber="471">18 (13 / 5)</td>
<td id="6E635DA28A320042FB6CFD54EA79FCA3" box="[1209,1375,766,785]" gridcol="2" gridrow="1" pageId="24" pageNumber="471">0.836</td>
</tr>
<tr id="2DB234DE8A320042FC93FC88EA79FC87" box="[838,1375,802,821]" gridrow="2" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FC88ECEFFC87" box="[838,969,802,821]" gridcol="0" gridrow="2" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FC88ECEFFC87" authorityName="Balthasar" authorityYear="1939" box="[838,969,802,821]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="substriolatus">
<emphasis id="A1F6EA8E8A32FFA5FC93FC88ECEFFC87" box="[838,969,802,821]" italics="true" pageId="24" pageNumber="471">P. substriolatus</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFC88EBB6FC87" box="[1003,1168,802,821]" gridcol="1" gridrow="2" pageId="24" pageNumber="471">6 (4 / 2)</td>
<td id="6E635DA28A320042FB6CFC88EA79FC87" box="[1209,1375,802,821]" gridcol="2" gridrow="2" pageId="24" pageNumber="471">0.7</td>
</tr>
<tr id="2DB234DE8A320042FC93FCECEA79FCEB" box="[838,1375,838,857]" gridrow="3" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FCECECEFFCEB" box="[838,969,838,857]" gridcol="0" gridrow="3" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FCECECB9FCEB" authorityName="Harold" authorityYear="1863" box="[838,927,838,857]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="daphnis">
<emphasis id="A1F6EA8E8A32FFA5FC93FCECECB9FCEB" box="[838,927,838,857]" italics="true" pageId="24" pageNumber="471">P. daphnis</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFCECEBB6FCEB" box="[1003,1168,838,857]" gridcol="1" gridrow="3" pageId="24" pageNumber="471">37 (30 / 7)</td>
<td id="6E635DA28A320042FB6CFCECEA79FCEB" box="[1209,1375,838,857]" gridcol="2" gridrow="3" pageId="24" pageNumber="471">0.876</td>
</tr>
<tr id="2DB234DE8A320042FC93FCC1EA79FCCC" box="[838,1375,875,894]" gridrow="4" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FCC1ECEFFCCC" box="[838,969,875,894]" gridcol="0" gridrow="4" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FCC1ECBAFCCC" authorityName="Bates" authorityYear="1887" box="[838,924,875,894]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="eximius">
<emphasis id="A1F6EA8E8A32FFA5FC93FCC1ECBAFCCC" box="[838,924,875,894]" italics="true" pageId="24" pageNumber="471">P. eximius</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFCC1EBB6FCCC" box="[1003,1168,875,894]" gridcol="1" gridrow="4" pageId="24" pageNumber="471">16 (13 / 3)</td>
<td id="6E635DA28A320042FB6CFCC1EA79FCCC" box="[1209,1375,875,894]" gridcol="2" gridrow="4" pageId="24" pageNumber="471">0.909</td>
</tr>
<tr id="2DB234DE8A320042FC93FC25EA79FC10" box="[838,1375,911,930]" gridrow="5" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FC25ECEFFC10" box="[838,969,911,930]" gridcol="0" gridrow="5" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FC25EC86FC10" authorityName="Bates" authorityYear="1887" box="[838,928,911,930]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="furiosus">
<emphasis id="A1F6EA8E8A32FFA5FC93FC25EC86FC10" box="[838,928,911,930]" italics="true" pageId="24" pageNumber="471">P. furiosus</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFC25EBB6FC10" box="[1003,1168,911,930]" gridcol="1" gridrow="5" pageId="24" pageNumber="471">79 (63 / 16)</td>
<td id="6E635DA28A320042FB6CFC25EA79FC10" box="[1209,1375,911,930]" gridcol="2" gridrow="5" pageId="24" pageNumber="471">0.847</td>
</tr>
<tr id="2DB234DE8A320042FC93FC19EA79FC74" box="[838,1375,947,966]" gridrow="6" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FC19ECEFFC74" box="[838,969,947,966]" gridcol="0" gridrow="6" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FC19ECB9FC74" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[838,927,947,966]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="herbeus">
<emphasis id="A1F6EA8E8A32FFA5FC93FC19ECB9FC74" box="[838,927,947,966]" italics="true" pageId="24" pageNumber="471">P. herbeus</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFC19EBB6FC74" box="[1003,1168,947,966]" gridcol="1" gridrow="6" pageId="24" pageNumber="471">8 (5 / 3)</td>
<td id="6E635DA28A320042FB6CFC19EA79FC74" box="[1209,1375,947,966]" gridcol="2" gridrow="6" pageId="24" pageNumber="471">0.909</td>
</tr>
<tr id="2DB234DE8A320042FC93FC7DEA79FC58" box="[838,1375,983,1002]" gridrow="7" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FC7DECEFFC58" box="[838,969,983,1002]" gridcol="0" gridrow="7" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FC7DECBEFC58" authorityName="Arnaud" authorityYear="2001" baseAuthorityName="Arnaud" baseAuthorityYear="2001" box="[838,920,983,1002]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="moroni">
<emphasis id="A1F6EA8E8A32FFA5FC93FC7DECBEFC58" box="[838,920,983,1002]" italics="true" pageId="24" pageNumber="471">P. moroni</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFC7DEBB6FC58" box="[1003,1168,983,1002]" gridcol="1" gridrow="7" pageId="24" pageNumber="471">6 (5 / 1)</td>
<td id="6E635DA28A320042FB6CFC7DEA79FC58" box="[1209,1375,983,1002]" gridcol="2" gridrow="7" pageId="24" pageNumber="471">0.972</td>
</tr>
<tr id="2DB234DE8A320042FC93FC51EA79FBBC" box="[838,1375,1019,1038]" gridrow="8" pageId="24" pageNumber="471">
<th id="6E635DA28A320042FC93FC51ECEFFBBC" box="[838,969,1019,1038]" gridcol="0" gridrow="8" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FC93FC51ECBEFBBC" authorityName="Harold" authorityYear="1863" box="[838,920,1019,1038]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="nimrod">
<emphasis id="A1F6EA8E8A32FFA5FC93FC51ECBEFBBC" box="[838,920,1019,1038]" italics="true" pageId="24" pageNumber="471">P. nimrod</emphasis>
</taxonomicName>
</th>
<td id="6E635DA28A320042FC3EFC51EBB6FBBC" box="[1003,1168,1019,1038]" gridcol="1" gridrow="8" pageId="24" pageNumber="471">8 (5 / 3)</td>
<td id="6E635DA28A320042FB6CFC51EA79FBBC" box="[1209,1375,1019,1038]" gridcol="2" gridrow="8" pageId="24" pageNumber="471">0.906</td>
</tr>
</table>
</paragraph>
<subSubSection id="DB9865178A32FFA5FF42FC33EED7FB22" pageId="24" pageNumber="471" type="description">
<paragraph id="933D369C8A32FFA5FF42FC33EED7FB22" blockId="24.[151,760,473,1584]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FF42FC33EEBBFC02" bold="true" box="[151,413,921,944]" pageId="24" pageNumber="471">
Variation. 
<emphasis id="A1F6EA8E8A32FFA5FEC7FC33EEBBFC02" bold="true" box="[274,413,921,944]" italics="true" pageId="24" pageNumber="471">
<taxonomicName id="54824D1F8A32FFA5FEC7FC33EE7EFC02" authorityName="Greenslade P &amp; Greenslade Pjm" authorityYear="1987" box="[274,344,921,944]" class="Insecta" family="Scarabaeidae" genus="Minor" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="genus">Minor</taxonomicName>
male.
</emphasis>
</emphasis>
Similar to major males, except for reduction of secondary sexual characters (i.e., cephalic horn, pronotal processes and posterolateral angles). 
<emphasis id="A1F6EA8E8A32FFA5FD65FC73EF97FBA2" bold="true" italics="true" pageId="24" pageNumber="471">Female.</emphasis>
Similar to male, except for head showing trituberculate carina; pronotal sculpture granulate to superficially punctate, smooth or rugose posteriorly; pronotum with anteromedial black macula, and anteromedial carina followed by posterior concavity (
<figureCitation id="0BB92A198A32FFA5FE41FBD3EEC2FB22" box="[404,484,1145,1168]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Fig. 30</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A32FFA5FF42FB33EDDEFA02" pageId="24" pageNumber="471" type="discussion">
<paragraph id="933D369C8A32FFA5FF42FB33EDDEFA02" blockId="24.[151,760,473,1584]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FF42FB33EE3FFB02" bold="true" box="[151,281,1177,1200]" pageId="24" pageNumber="471">Comments.</emphasis>
Apparently, this species is well defined and is not involved in taxonomical controversies (
<bibRefCitation id="F7134B6D8A32FFA5FD5BFB13EFF7FB42" author="EDMONDS" firstAuthor="EDMONDS" pageId="24" pageNumber="471" pagination="1 - 105" refId="ref25450" refString="EDMONDS W. D. 1994: Revision of Phanaeus MacLeay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science of the Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">EDMONDS 1994</bibRefCitation>
, 
<bibRefCitation id="F7134B6D8A32FFA5FF0AFB73EEA2FB42" author="ARNAUD" box="[223,388,1241,1264]" firstAuthor="ARNAUD" pageId="24" pageNumber="471" refId="ref24193" refString="ARNAUD P. 2002: Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury, 151 pp." type="book" year="2002">ARNAUD 2002</bibRefCitation>
, 
<bibRefCitation id="F7134B6D8A32FFA5FE41FB73ED90FB42" author="EDMONDS &amp; ZIDEK" box="[404,694,1241,1264]" firstAuthor="EDMONDS" pageId="24" pageNumber="471" pagination="1 - 108" refId="ref25590" refString="EDMONDS W. D. &amp; ZIDEK J. 2012: Taxonomy of Phanaeus revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Insecta Mundi 274: 1 - 108." type="journal article" year="2012">EDMONDS &amp; ZÍDEK 2012</bibRefCitation>
). The only variation that was reported by previous studies is its several chromatic phases (
<figureCitation id="0BB92A198A32FFA5FE7BFAB3ED0EFA82" box="[430,552,1305,1328]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29–31</figureCitation>
, 
<figureCitation id="0BB92A198A32FFA5FDE7FAB3ED76FA82" box="[562,592,1305,1328]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="24" pageNumber="471">54</figureCitation>
). All specimens here revised show the distinctly vermiculate elytral striae (
<figureCitation id="0BB92A198A32FFA5FF4AFAF3EE3AFAC2" box="[159,284,1369,1392]" captionStart="Figs 29–31" captionStartId="20.[760,797,2004,2023]" captionTargetBox="[162,1438,503,2076]" captionTargetId="figure-184@20.[155,1432,503,2079]" captionTargetPageId="20" captionText="Figs 29–31. Phanaeus eximius Bates, 1887. 29 – male blue phase; 30 – female green phase (typical); 31 – lectotype and labels (by Keita Matsumoto, BMNH). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821178" httpUri="https://zenodo.org/record/5821178/files/figure.png" pageId="24" pageNumber="471">Figs 29–31</figureCitation>
). Nevertheless, further studies are needed to understand the intraspecific variation in 
<taxonomicName id="54824D1F8A32FFA5FDEAFAD3ED8FFA22" authorityName="Bates" authorityYear="1887" box="[575,681,1401,1424]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="eximius">
<emphasis id="A1F6EA8E8A32FFA5FDEAFAD3ED8FFA22" box="[575,681,1401,1424]" italics="true" pageId="24" pageNumber="471">P. eximius</emphasis>
</taxonomicName>
since a limited number of specimens was available for our work.
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A32FFA5FF42FA13ED57F982" pageId="24" pageNumber="471" type="distribution">
<paragraph id="933D369C8A32FFA5FF42FA13ED57F982" blockId="24.[151,760,473,1584]" pageId="24" pageNumber="471">
<emphasis id="A1F6EA8E8A32FFA5FF42FA13EE0CFA62" bold="true" box="[151,298,1465,1488]" pageId="24" pageNumber="471">Distribution.</emphasis>
<collectingCountry id="EB95760C8A32FFA5FEE4FA13EE81FA62" box="[305,423,1465,1488]" name="Guatemala" pageId="24" pageNumber="471">Guatemala</collectingCountry>
, 
<collectingCountry id="EB95760C8A32FFA5FE66FA13ED15FA62" box="[435,563,1465,1488]" name="El Salvador" pageId="24" pageNumber="471">El Salvador</collectingCountry>
, 
<collectingCountry id="EB95760C8A32FFA5FDEAFA13ED89FA62" box="[575,687,1465,1488]" name="Nicaragua" pageId="24" pageNumber="471">Nicaragua</collectingCountry>
, 
<collectingCountry id="EB95760C8A32FFA5FD6EFA13EFEEFA42" name="Costa Rica" pageId="24" pageNumber="471">Costa Rica</collectingCountry>
and probably 
<collectingCountry id="EB95760C8A32FFA5FEB6FA73EEE8FA42" box="[355,462,1497,1520]" name="Honduras" pageId="24" pageNumber="471">Honduras</collectingCountry>
. Following 
<taxonomicName id="54824D1F8A32FFA5FD98FA73ED9BFA42" authorityName="Bates" authorityYear="1887" box="[589,701,1497,1520]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="furiosus">
<emphasis id="A1F6EA8E8A32FFA5FD98FA73ED9BFA42" box="[589,701,1497,1520]" italics="true" pageId="24" pageNumber="471">P. furiosus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A32FFA5FD19FA73EF93F9A2" captionStart="Fig" captionStartId="29.[151,182,946,965]" captionTargetBox="[162,1425,196,930]" captionTargetId="figure-13@29.[162,1425,196,930]" captionTargetPageId="29" captionText="Fig. 66. Predicted distribution of P. furiosus Bates, 1887 and P. pseudofurcosus Balthasar, 1939." figureDoi="http://doi.org/10.5281/zenodo.5821201" httpUri="https://zenodo.org/record/5821201/files/figure.png" pageId="24" pageNumber="471">Fig. 66</figureCitation>
), this is the second most widely distributed species within the 
<taxonomicName id="54824D1F8A32FFA5FEDCF9B3EE4BF982" authorityName="Castelnau" authorityYear="1840" box="[265,365,1561,1584]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="24" pageNumber="471" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A32FFA5FEDCF9B3EE4BF982" box="[265,365,1561,1584]" italics="true" pageId="24" pageNumber="471">P. tridens</emphasis>
</taxonomicName>
species group (
<figureCitation id="0BB92A198A32FFA5FDC0F9B3ED42F982" box="[533,612,1561,1584]" captionStart="Fig" captionStartId="30.[151,182,940,959]" captionTargetBox="[162,1425,190,924]" captionTargetId="figure-13@30.[162,1425,190,924]" captionTargetPageId="30" captionText="Fig. 68. Predicted distribution of P. eximius Bates, 1887." figureDoi="http://doi.org/10.5281/zenodo.5821205" httpUri="https://zenodo.org/record/5821205/files/figure.png" pageId="24" pageNumber="471">Fig. 68</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>