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<mods:title id="F442A070A3AF8308675D65902B16FABF">Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species</mods:title>
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<taxonomicName id="4C634D67FFFBFFC4FDFA47E1BC903C3F" authority="Arnaud, 2018" authorityName="Arnaud" authorityYear="2018" box="[594,994,1514,1541]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="arletteae">
<emphasis id="B917EAF6FFFBFFC4FDFA47E1BC483C3F" bold="true" box="[594,826,1514,1541]" italics="true" pageId="5" pageNumber="6">Phanaeus arletteae</emphasis>
<bibRefCitation id="EFF24B15FFFBFFC4FCE847E0BC903C3F" author="Arnaud" box="[832,994,1515,1541]" firstAuthor="Arnaud" pageId="5" pageNumber="6" pagination="3 - 7" refId="ref35890" refString="Arnaud P. 2018. Description d'une nouvelle espece de Phanaeus d'Equateur et revalidation de l'espece Coprophanaeus (C.) edmondsi Arnaud (Coleoptera: Scarabaeidae, Scarabaeinae). Besoiro 26: 3 - 7." type="journal article" year="2018">Arnaud, 2018</bibRefCitation>
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<paragraph id="8BDC36E4FFFBFFC4FD064405BCFA3C12" blockId="5.[594,994,1514,1576]" box="[686,904,1550,1576]" pageId="5" pageNumber="6">
<figureCitation id="13582A61FFFBFFC4FD064405BC753C12" box="[686,775,1550,1576]" captionStart="Fig" captionStartId="54.[191,234,1247,1273]" captionTargetBox="[193,1393,268,1195]" captionTargetId="figure-243@54.[189,1396,265,1198]" captionTargetPageId="54" captionText="Fig. 17. Distribution of the Phanaeus endymion species group in South America. Phanaeus malyi Arnaud, 2001 was reported from Colombia without accurate locality. Phanaeus arletteae Arnaud, 2018 was modified from Arnaud (2018)." figureDoi="http://doi.org/10.5281/zenodo.4836800" httpUri="https://zenodo.org/record/4836800/files/figure.png" pageId="5" pageNumber="6">Figs 17</figureCitation>
, 
<figureCitation id="13582A61FFFBFFC4FCBC4405BC3B3C12" box="[788,841,1550,1576]" captionStart="Fig" captionStartId="55.[189,232,1681,1707]" captionTargetBox="[258,1329,264,1646]" captionTargetId="figure-157@55.[258,1329,264,1646]" captionTargetPageId="55" captionText="Fig. 18. Pronotum of females within the Phanaeus endymion species group. A. Phanaeus arletteae Arnaud, 2018 (modified from Arnaud 2018). B. Phanaeus bravoensis Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (by Jose Luis Sánchez-Huerta).C. Phanaeus chiapanecus sp. nov. D. Phanaeus edmondsi Moctezuma, Deloya &amp; Halffter, 2019 (by Jose Luis Sánchez-Huerta). E. Phanaeus endymion Harold, 1863. F. Phanaeus funereus Balthasar, 1939 stat. rev. G. Phanaeus halffterorum Edmonds, 1979 (by Jose Luis Sánchez-Huerta).H. Phanaeus huichol Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (by Jose Luis Sánchez-Huerta). I. Phanaeus jackenioi sp. nov. J. Phanaeus malyi Arnaud, 2002. K. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. L. Phanaeus pacificus sp. nov. M. Phanaeus panamensis sp. nov. N. Phanaeus porioni Arnaud, 2001 stat. rev. O. Phanaeus pyrois Bates, 1887. P. Phanaeus rzedowskii sp. nov. Q. Phanaeus zapotecus Edmonds, 2006. R. Phanaeus zoque Moctezuma &amp; Halffter, 2017." figureDoi="http://doi.org/10.5281/zenodo.4836802" httpUri="https://zenodo.org/record/4836802/files/figure.png" pageId="5" pageNumber="6">18A</figureCitation>
, 
<figureCitation id="13582A61FFFBFFC4FCFD4405BCFA3C12" box="[853,904,1550,1576]" captionStart="Fig" captionStartId="56.[189,231,1678,1704]" captionTargetBox="[306,1282,265,1646]" captionTargetId="figure-158@56.[306,1282,265,1646]" captionTargetPageId="56" captionText="Fig. 19. Elytral integument of species within the Phanaeus endymion species group. A. Phanaeus arletteae Arnaud, 2018 (modified from Arnaud 2018). B. Phanaeus bravoensis Moctezuma, Sánchez- Huerta &amp; Halffter, 2017 (by Jose Luis Sánchez-Huerta). C. Phanaeus chiapanecus sp. nov. D. Phanaeus edmondsi Moctezuma, Deloya &amp; Halffter, 2019 (by Jose Luis Sánchez-Huerta). E. Phanaeus endymion Harold, 1863. F. Phanaeus funereus Balthasar, 1939 stat. rev. G. Phanaeus halffterorum Edmonds, 1979 (by Jose Luis Sánchez-Huerta). H. Phanaeus huichol Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (by Jose Luis Sánchez-Huerta). I. Phanaeus jackenioi sp. nov. J. Phanaeus malyi Arnaud, 2002.K. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. L. Phanaeus pacificus sp. nov. M. Phanaeus panamensis sp. nov. N. Phanaeus porioni Arnaud, 2001 stat. rev. O. Phanaeus pyrois Bates, 1887. P. Phanaeus rzedowskii sp. nov. Q. Phanaeus zapotecus Edmonds, 2006. R. Phanaeus zoque Moctezuma &amp; Halffter, 2017." figureDoi="http://doi.org/10.5281/zenodo.4836804" httpUri="https://zenodo.org/record/4836804/files/figure.png" pageId="5" pageNumber="6">19A</figureCitation>
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<treatmentCitation id="0AC210F5FFFBFFC4FF154444BC473C53" author="Arnaud P." box="[189,821,1615,1642]" page="4" pageId="5" pageNumber="6" year="2018">
<taxonomicName id="4C634D67FFFBFFC4FF154444BC473C53" authority="Arnaud, 2018: 4" authorityName="Arnaud" authorityPageNumber="4" authorityYear="2018" box="[189,821,1615,1642]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="arletteae" subGenus="Notiophanaeus">
<emphasis id="B917EAF6FFFBFFC4FF154444BE433C53" box="[189,305,1615,1641]" italics="true" pageId="5" pageNumber="6">Phanaeus</emphasis>
(
<emphasis id="B917EAF6FFFBFFC4FEEA4444BE863C53" box="[322,500,1615,1641]" italics="true" pageId="5" pageNumber="6">Notiophanaeus</emphasis>
) 
<emphasis id="B917EAF6FFFBFFC4FDAC4444BD1E3C53" box="[516,620,1615,1641]" italics="true" pageId="5" pageNumber="6">arletteae</emphasis>
<bibRefCitation id="EFF24B15FFFBFFC4FDDA4444BC473C53" author="Arnaud" box="[626,821,1615,1641]" firstAuthor="Arnaud" page="4" pageId="5" pageNumber="6" pagination="3 - 7" refId="ref35890" refString="Arnaud P. 2018. Description d'une nouvelle espece de Phanaeus d'Equateur et revalidation de l'espece Coprophanaeus (C.) edmondsi Arnaud (Coleoptera: Scarabaeidae, Scarabaeinae). Besoiro 26: 3 - 7." type="journal article" year="2018">Arnaud, 2018: 4</bibRefCitation>
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.
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<treatmentCitationGroup id="AB7311CAFFFBFFC4FF15449ABD843CC8" pageId="5" pageNumber="6">
<taxonomicName id="4C634D67FFFBFFC4FF15449ABD013C91" ID-CoL="856BJ" authorityName="Harold" authorityYear="1871" box="[189,627,1681,1707]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="blanchardi">
<emphasis id="B917EAF6FFFBFFC4FF15449ABECE3C91" box="[189,444,1681,1707]" italics="true" pageId="5" pageNumber="6">Phanaeus blanchardi</emphasis>
ab. 
<emphasis id="B917EAF6FFFBFFC4FE5C449ABD013C91" box="[500,627,1681,1707]" italics="true" pageId="5" pageNumber="6">viridicollis</emphasis>
</taxonomicName>
(unavailable infrasubspecific name) – 
<treatmentCitation id="0AC210F5FFFBFFC4FBE4449ABA383C91" author="Olsoufieff G. d'" box="[1100,1354,1681,1707]" page="149" pageId="5" pageNumber="6" year="1924">
<bibRefCitation id="EFF24B15FFFBFFC4FBE4449ABA383C91" author="Olsoufieff" box="[1100,1354,1681,1707]" firstAuthor="Olsoufieff" page="149" pageId="5" pageNumber="6" pagination="4 - 172" refId="ref41334" refString="Olsoufieff G. d'. 1924. Les Phanaeides (Coleoptera-Lamellicornia). Famille Scarabaeidae - Tr. Coprini. Insecta, Revue illustree d'Entomologique 13: 4 - 172." type="journal article" year="1924">Olsoufieff 1924: 149</bibRefCitation>
</treatmentCitation>
. — 
<treatmentCitation id="0AC210F5FFFBFFC4FF4244BFBE963CF4" author="Edmonds W. D." box="[234,484,1716,1742]" page="443" pageId="5" pageNumber="6" year="1994">
<bibRefCitation id="EFF24B15FFFBFFC4FF4244BFBE963CF4" author="Edmonds" box="[234,484,1716,1742]" firstAuthor="Edmonds" page="443" pageId="5" pageNumber="6" pagination="1 - 105" refId="ref37487" refString="Edmonds W. D. 1994. Revision of Phanaeus Macleay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science, Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">Edmonds 1994: 443</bibRefCitation>
</treatmentCitation>
. — 
<treatmentCitation id="0AC210F5FFFBFFC4FDB244BFBD8E3CF4" author="Arnaud P." box="[538,764,1716,1742]" page="98" pageId="5" pageNumber="6" year="2002">
<bibRefCitation id="EFF24B15FFFBFFC4FDB244BFBD8E3CF4" author="Arnaud" box="[538,764,1716,1742]" firstAuthor="Arnaud" page="98" pageId="5" pageNumber="6" refId="ref35859" refString="Arnaud P. 2002 b. Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury." type="book" year="2002" yearSuffix="b">Arnaud 2002b: 98</bibRefCitation>
</treatmentCitation>
. — 
<treatmentCitation id="0AC210F5FFFBFFC4FC9D44BFBBFC3CF4" author="Edmonds W. D. &amp; Zidek J." box="[821,1166,1716,1742]" page="13" pageId="5" pageNumber="6" year="2012">
<bibRefCitation id="EFF24B15FFFBFFC4FC9D44BFBBFC3CF4" author="Edmonds &amp; Zidek" box="[821,1166,1716,1742]" firstAuthor="Edmonds" page="13" pageId="5" pageNumber="6" pagination="1 - 108" refId="ref37762" 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: 13</bibRefCitation>
</treatmentCitation>
(as an unavailable infrasubspecific name referable to 
<taxonomicName id="4C634D67FFFBFFC4FD2844D3BD973CCB" ID-CoL="4FSJX" baseAuthorityName="Edmonds &amp; Zidek" baseAuthorityYear="2012" box="[640,741,1752,1777]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFFBFFC4FD2844D3BD973CCB" box="[640,741,1752,1777]" italics="true" pageId="5" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFFBFFC4FF1544F1BC553D02" blockId="5.[189,1399,1681,2024]" pageId="5" pageNumber="6">
<treatmentCitationGroup id="AB7311CAFFFBFFC4FF1544F1BC553D02" pageId="5" pageNumber="6">
<taxonomicName id="4C634D67FFFBFFC4FF1544F1BC633D2E" ID-CoL="856BJ" authorityName="Harold" authorityYear="1871" box="[189,785,1786,1813]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="blanchardi" subGenus="Phanaeus">
<emphasis id="B917EAF6FFFBFFC4FF1544F1BE433D2E" box="[189,305,1786,1812]" italics="true" pageId="5" pageNumber="6">Phanaeus</emphasis>
(
<emphasis id="B917EAF6FFFBFFC4FEE044F1BECE3D2E" box="[328,444,1786,1812]" italics="true" pageId="5" pageNumber="6">Phanaeus</emphasis>
) 
<emphasis id="B917EAF6FFFBFFC4FE7C44F1BD263D2E" box="[468,596,1786,1812]" italics="true" pageId="5" pageNumber="6">blanchardi</emphasis>
ab. 
<emphasis id="B917EAF6FFFBFFC4FD3A44F1BC633D2E" box="[658,785,1786,1812]" italics="true" pageId="5" pageNumber="6">viridicollis</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFFBFFC4FC9344F1BBD43D2E" author="Martinez A &amp; Pereira F. S." box="[827,1190,1786,1812]" page="68" pageId="5" pageNumber="6" year="1967">
<bibRefCitation id="EFF24B15FFFBFFC4FC9344F1BBD43D2E" author="Martinez &amp; Pereira" box="[827,1190,1786,1812]" firstAuthor="Martinez" page="68" pageId="5" pageNumber="6" pagination="53 - 69" refId="ref40118" refString="Martinez A &amp; Pereira F. S. 1967. Notas escarabeidologicas III (Col. Scarabaeidae-Scarabaeinae). Revista de la Sociedad Entomologica Argentina 29: 53 - 69." type="journal article" year="1967">Martínez &amp; Pereira 1967: 68</bibRefCitation>
</treatmentCitation>
(as synonym of 
<taxonomicName id="4C634D67FFFBFFC4FF424515BE193D02" ID-CoL="874G8" authorityName="Balthasar" authorityYear="1939" baseAuthorityName="Balthasar" baseAuthorityYear="1939" box="[234,363,1822,1848]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="funereus">
<emphasis id="B917EAF6FFFBFFC4FF424515BE193D02" box="[234,363,1822,1848]" italics="true" pageId="5" pageNumber="6">P. funereus</emphasis>
</taxonomicName>
) (unavailable infrasubspecific name).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFFBFFC4FF15454ABD843D98" blockId="5.[189,1399,1681,2024]" pageId="5" pageNumber="6">
<treatmentCitationGroup id="AB7311CAFFFBFFC4FF15454ABD843D98" pageId="5" pageNumber="6">
<taxonomicName id="4C634D67FFFBFFC4FF15454ABDF03D61" ID-CoL="7HQWR" authorityName="Olsoufieff" authorityYear="1924" box="[189,642,1857,1883]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="viridicollis" subGenus="Notiophanaeus">
<emphasis id="B917EAF6FFFBFFC4FF15454ABE433D61" box="[189,305,1857,1883]" italics="true" pageId="5" pageNumber="6">Phanaeus</emphasis>
(
<emphasis id="B917EAF6FFFBFFC4FEE9454ABE813D61" box="[321,499,1857,1883]" italics="true" pageId="5" pageNumber="6">Notiophanaeus</emphasis>
) 
<emphasis id="B917EAF6FFFBFFC4FDAB454ABDF03D61" box="[515,642,1857,1883]" italics="true" pageId="5" pageNumber="6">viridicollis</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFFBFFC4FD34454ABCE93D61" author="Arnaud P." box="[668,923,1857,1883]" page="98" pageId="5" pageNumber="6" year="2002">
<bibRefCitation id="EFF24B15FFFBFFC4FD34454ABCE93D61" author="Arnaud" box="[668,923,1857,1883]" firstAuthor="Arnaud" page="98 - 99" pageId="5" pageNumber="6" refId="ref35859" refString="Arnaud P. 2002 b. Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury." type="book" year="2002" yearSuffix="b">Arnaud 2002b: 98–99</bibRefCitation>
</treatmentCitation>
(as a new combination for 
<taxonomicName id="4C634D67FFFBFFC4FB7E4549BA053D61" ID-CoL="856BJ" authorityName="Harold" authorityYear="1871" box="[1238,1399,1857,1883]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="blanchardi">
<emphasis id="B917EAF6FFFBFFC4FB7E4549BB9A3D61" box="[1238,1256,1858,1883]" italics="true" pageId="5" pageNumber="6">P</emphasis>
. 
<emphasis id="B917EAF6FFFBFFC4FB5F454ABA053D61" box="[1271,1399,1857,1883]" italics="true" pageId="5" pageNumber="6">blanchardi</emphasis>
</taxonomicName>
ab. 
<taxonomicName id="4C634D67FFFBFFC4FEBA456FBEE33D44" ID-CoL="7HQWR" authorityName="Olsoufieff" authorityYear="1924" box="[274,401,1892,1918]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="viridicollis">
<emphasis id="B917EAF6FFFBFFC4FEBA456FBEE33D44" box="[274,401,1892,1918]" italics="true" pageId="5" pageNumber="6">viridicollis</emphasis>
</taxonomicName>
) (unavailable infrasubspecific name). — 
<treatmentCitation id="0AC210F5FFFBFFC4FCC1456FBBD63D44" author="Edmonds W. D. &amp; Zidek J." box="[873,1188,1892,1918]" page="13" pageId="5" pageNumber="6" year="2012">
<bibRefCitation id="EFF24B15FFFBFFC4FCC1456FBBD63D44" author="Edmonds &amp; Zidek" box="[873,1188,1892,1918]" firstAuthor="Edmonds" page="13" pageId="5" pageNumber="6" pagination="1 - 108" refId="ref37762" 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: 13</bibRefCitation>
</treatmentCitation>
(as an unavailable infrasubspecific name referable to 
<taxonomicName id="4C634D67FFFBFFC4FD284583BD973D9B" ID-CoL="4FSJX" baseAuthorityName="Edmonds &amp; Zidek" baseAuthorityYear="2012" box="[640,741,1928,1953]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFFBFFC4FD284583BD973D9B" box="[640,741,1928,1953]" italics="true" pageId="5" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFFBFFC4FF1545A1BD843DD2" blockId="5.[189,1399,1681,2024]" pageId="5" pageNumber="6">
<treatmentCitationGroup id="AB7311CAFFFBFFC4FF1545A1BD843DD2" pageId="5" pageNumber="6">
<taxonomicName id="4C634D67FFFBFFC4FF1545A1BECC3DFE" ID-CoL="7HQWR" authorityName="Olsoufieff" authorityYear="1924" box="[189,446,1962,1988]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="viridicollis">
<emphasis id="B917EAF6FFFBFFC4FF1545A1BECC3DFE" box="[189,446,1962,1988]" italics="true" pageId="5" pageNumber="6">Phanaeus viridicollis</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFFBFFC4FE4145A1BDA23DFF" author="Krajcik M." box="[489,720,1962,1989]" page="153" pageId="5" pageNumber="6" year="2006">
<bibRefCitation id="EFF24B15FFFBFFC4FE4145A1BDA23DFF" author="Krajcik" box="[489,720,1962,1989]" firstAuthor="Krajcik" page="153" pageId="5" pageNumber="6" pagination="1 - 189" refId="ref39668" refString="Krajcik M. 2006. Checklist of Scarabaeoidea of the World. 1. Scarabaeinae (Coleoptera: Scarabaeidae: Scarabaeinae). Animma. X. Suplement 3: 1 - 189." type="journal article" year="2006">Krajcik 2006: 153</bibRefCitation>
</treatmentCitation>
. — 
<treatmentCitation id="0AC210F5FFFBFFC4FCA645A1BB113DFE" author="Edmonds W. D. &amp; Zidek J." box="[782,1123,1962,1988]" page="1" pageId="5" pageNumber="6" year="2012">
<bibRefCitation id="EFF24B15FFFBFFC4FCA645A1BB113DFE" author="Edmonds &amp; Zidek" box="[782,1123,1962,1988]" firstAuthor="Edmonds" page="1" pageId="5" pageNumber="6" pagination="1 - 108" refId="ref37762" 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: 1</bibRefCitation>
</treatmentCitation>
, 6 (as an unavailable infrasubspecific name referable to 
<taxonomicName id="4C634D67FFFBFFC4FD2845C5BD973DDD" ID-CoL="4FSJX" baseAuthorityName="Edmonds &amp; Zidek" baseAuthorityYear="2012" box="[640,741,1998,2023]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFFBFFC4FD2845C5BD973DDD" box="[640,741,1998,2023]" italics="true" pageId="5" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF154301BEB23B7D" blockId="6.[189,1399,265,399]" pageId="6" pageNumber="7">
<treatmentCitationGroup id="AB7311CAFFF8FFC7FF154301BEB23B7D" pageId="6" pageNumber="7">
Non 
<taxonomicName id="4C634D67FFF8FFC7FF514302BECC3B19" ID-CoL="4FSJX" baseAuthorityName="Edmonds &amp; Zidek" baseAuthorityYear="2012" box="[249,446,265,291]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFF8FFC7FF514302BECC3B19" box="[249,446,265,291]" italics="true" pageId="6" pageNumber="7">Phanaeus pyrois</emphasis>
</taxonomicName>
(error) – 
<treatmentCitation id="0AC210F5FFF8FFC7FD9D4301BCCA3B1E" author="Moctezuma V. &amp; Halffter G." box="[565,952,265,292]" page="55" pageId="6" pageNumber="7" year="2017">
<bibRefCitation id="EFF24B15FFF8FFC7FD9D4301BCCA3B1E" author="Moctezuma &amp; Halffter" box="[565,952,265,292]" firstAuthor="Moctezuma" page="55" pageId="6" pageNumber="7" pagination="47 - 56" refId="ref40548" refString="Moctezuma V. &amp; Halffter G. 2017. A new species of Phanaeus Macleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71: 47 - 56. https: // doi. org / 10.1649 / 0010 - 065 X- 71.1.47" type="journal article" year="2017">Moctezuma &amp; Halffter 2017: 55</bibRefCitation>
</treatmentCitation>
(in part). — 
<bibRefCitation id="EFF24B15FFF8FFC7FBF24301BE683B7D" author="Moctezuma V. &amp; Halffter G." journalOrPublisher="The Coleopterists Bulletin" pageId="6" pageNumber="7" pagination="47 - 56" part="71" refId="ref40548" refString="Moctezuma V. &amp; Halffter G. 2017. A new species of Phanaeus Macleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico. The Coleopterists Bulletin 71: 47 - 56. https: // doi. org / 10.1649 / 0010 - 065 X- 71.1.47" title="A new species of Phanaeus Macleay (Coleoptera: Scarabaeidae: Scarabaeinae) from Los Chimalapas, Oaxaca, Mexico" type="journal article" year="2017">
Moctezuma 
<emphasis id="B917EAF6FFF8FFC7FB584301BA513B19" box="[1264,1315,265,291]" italics="true" pageId="6" pageNumber="7">et al</emphasis>
. 2017: 114
</bibRefCitation>
, 130 (in part).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF15435BBBEC3B50" blockId="6.[189,1399,265,399]" box="[189,1182,336,363]" pageId="6" pageNumber="7">
<treatmentCitationGroup id="AB7311CAFFF8FFC7FF15435BBBEC3B50" box="[189,1182,336,363]" pageId="6" pageNumber="7">
<taxonomicName id="4C634D67FFF8FFC7FF15435BBD1E3B50" authorityName="Arnaud" authorityYear="2018" box="[189,620,336,363]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="arletteae" subGenus="Notiophanaeus">
<emphasis id="B917EAF6FFF8FFC7FF15435BBE433B50" box="[189,305,336,362]" italics="true" pageId="6" pageNumber="7">Phanaeus</emphasis>
(
<emphasis id="B917EAF6FFF8FFC7FEEA435BBE863B50" box="[322,500,336,362]" italics="true" pageId="6" pageNumber="7">Notiophanaeus</emphasis>
) 
<emphasis id="B917EAF6FFF8FFC7FDAC435BBD1E3B50" box="[516,620,336,362]" italics="true" pageId="6" pageNumber="7">arletteae</emphasis>
</taxonomicName>
<emphasis id="B917EAF6FFF8FFC7FDDC435ABDF13B50" box="[628,643,337,362]" italics="true" pageId="6" pageNumber="7">–</emphasis>
<treatmentCitation id="0AC210F5FFF8FFC7FD22435BBCC73B51" author="Chamorro W. &amp; Marin-Armijos D. &amp; Granda V. &amp; Vaz-de-Mello F. Z." box="[650,949,336,363]" page="97" pageId="6" pageNumber="7" year="2018">
<bibRefCitation id="EFF24B15FFF8FFC7FD22435BBCC73B51" author="Chamorro" box="[650,949,336,363]" etAl="et al." firstAuthor="Chamorro" page="97" pageId="6" pageNumber="7" pagination="72 - 100" refId="ref36557" refString="Chamorro W., Marin-Armijos D., Granda V. &amp; Vaz-de-Mello F. Z. 2018. Listado de especies y clave de generos y subgeneros de escarabajos estercoleros (Coleoptera: Scarabaeidae: Scarabaeinae) presentes y presuntos para Ecuador. Revista Colombiana de Entomologia 44: 72 - 100. https: // doi. org / 10.25100 / socolen. v 44 i 1.6545" type="journal article" year="2018">
Chamorro 
<emphasis id="B917EAF6FFF8FFC7FCA0435ABC4E3B50" box="[776,828,336,362]" italics="true" pageId="6" pageNumber="7">et al</emphasis>
. 2018: 97
</bibRefCitation>
</treatmentCitation>
; 
<treatmentCitation id="0AC210F5FFF8FFC7FC69435ABB483B51" author="Chamorro W. &amp; Marin-Armijos D. &amp; Asenjo A." box="[961,1082,337,363]" httpUri="http://treatment.plazi.org/id/451442EF53FBDB52171039B944A5395A" page="213" pageId="6" pageNumber="7" year="2019">
<bibRefCitation id="EFF24B15FFF8FFC7FC69435ABB483B51" author="Chamorro" box="[961,1082,337,363]" firstAuthor="Chamorro" page="213" pageId="6" pageNumber="7" pagination="1 - 343" refId="ref36626" refString="Chamorro W., Marin-Armijos D. &amp; Asenjo A. 2019. Scarabaeinae dung beetles from Ecuador: a catalog, nomenclatural acts, and distribution records. ZooKeys 826: 1 - 343. https: // doi. org / 10.3897 / zookeys. 826.26488" type="journal article" year="2019">2019: 213</bibRefCitation>
</treatmentCitation>
, pl. 46a.
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF15437FBC6F3BB4" blockId="6.[189,1399,265,399]" box="[189,797,372,399]" pageId="6" pageNumber="7">
<treatmentCitationGroup id="AB7311CAFFF8FFC7FF15437FBC6F3BB4" box="[189,797,372,399]" pageId="6" pageNumber="7">
<taxonomicName id="4C634D67FFF8FFC7FF15437FBED23BB4" authorityName="Arnaud" authorityYear="2018" box="[189,416,372,398]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="arletteae">
<emphasis id="B917EAF6FFF8FFC7FF15437FBED23BB4" box="[189,416,372,398]" italics="true" pageId="6" pageNumber="7">Phanaeus arletteae</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFF8FFC7FE16437FBD9F3BB4" author="Kohlmann B. &amp; Arriaga-Jimenez A. &amp; Ros M." box="[446,749,372,399]" page="83" pageId="6" pageNumber="7" year="2018">
<bibRefCitation id="EFF24B15FFF8FFC7FE16437FBD9F3BB4" author="Kohlmann" box="[446,749,372,399]" etAl="et al." firstAuthor="Kohlmann" page="83" pageId="6" pageNumber="7" pagination="67 - 93" refId="ref39574" refString="Kohlmann B., Arriaga-Jimenez A. &amp; Ros M. 2018. Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae). ZooKeys 743: 67 - 93. https: // doi. org / 10.3897 / zookeys. 743.23029" type="journal article" year="2018">
Kohlmann 
<emphasis id="B917EAF6FFF8FFC7FDE9437EBD063BB4" box="[577,628,372,398]" italics="true" pageId="6" pageNumber="7">et al</emphasis>
. 2018: 83
</bibRefCitation>
</treatmentCitation>
, 88.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFF8FFC7FF1543BDBCFC3882" pageId="6" pageNumber="7" type="materials_examined">
<paragraph id="8BDC36E4FFF8FFC7FF1543BDBE1E3BEA" blockId="6.[189,364,438,464]" box="[189,364,438,464]" pageId="6" pageNumber="7">
<heading id="D0948188FFF8FFC7FF1543BDBE1E3BEA" bold="true" box="[189,364,438,464]" fontSize="11" level="3" pageId="6" pageNumber="7" reason="3">
<emphasis id="B917EAF6FFF8FFC7FF1543BDBE1E3BEA" bold="true" box="[189,364,438,464]" pageId="6" pageNumber="7">Type material</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF4243EEBC83383A" blockId="6.[189,1398,485,583]" box="[234,1009,485,512]" pageId="6" pageNumber="7">
<emphasis id="B917EAF6FFF8FFC7FF4243EEBE2E3BC5" bold="true" box="[234,348,485,511]" pageId="6" pageNumber="7">
<typeStatus id="54D88846FFF8FFC7FF4243EEBE2E3BC5" box="[234,348,485,511]" pageId="6" pageNumber="7" type="holotype">Holotype</typeStatus>
</emphasis>
(revised from photograph from the original description)
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF154002BE7E387D" blockId="6.[189,1398,485,583]" pageId="6" pageNumber="7">
<materialsCitation id="3B0B3CB9FFF8FFC7FF154002BE7A387D" ID-GBIF-Occurrence="3093980301" collectionCode="CPFA" country="Ecuador" location="Guayas" municipality="Kilometric" pageId="6" pageNumber="7" specimenCount="1" specimenCount-male="1" stateProvince="Guayas">
<collectingCountry id="F3747674FFF8FFC7FF154002BE393819" box="[189,331,521,547]" name="Ecuador" pageId="6" pageNumber="7">ECUADOR</collectingCountry>
• 
<specimenCount id="9D65FD6DFFF8FFC7FEC34001BEF1381E" box="[363,387,522,548]" pageId="6" pageNumber="7" type="male">♂</specimenCount>
, 
<bibRefCitation id="EFF24B15FFF8FFC7FE3A4002BD263819" author="Arnaud" box="[402,596,521,547]" firstAuthor="Arnaud" page="4" pageId="6" pageNumber="7" pagination="3 - 7" refId="ref35890" refString="Arnaud P. 2018. Description d'une nouvelle espece de Phanaeus d'Equateur et revalidation de l'espece Coprophanaeus (C.) edmondsi Arnaud (Coleoptera: Scarabaeidae, Scarabaeinae). Besoiro 26: 3 - 7." type="journal article" year="2018">Arnaud 2018: 4</bibRefCitation>
, pl. 1, fig. a; 
<collectingRegion id="49A7F806FFF8FFC7FCA94002BC2E3819" box="[769,860,521,547]" country="Ecuador" name="Guayas" pageId="6" pageNumber="7">Guayas</collectingRegion>
, 
<locationDeviation id="9A6E2FC6FFF8FFC7FCC34002BA003819" box="[875,1394,521,547]" pageId="6" pageNumber="7">Kilometric point 18 Balzar-Quevedo Road</locationDeviation>
; 
<collectionCode id="ED72AE21FFF8FFC7FF154026BE7A387D" box="[189,264,557,583]" collectionName="P and F Arnaud" pageId="6" pageNumber="7">CPFA</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF154064BE2F38B3" blockId="6.[189,349,623,649]" box="[189,349,623,649]" pageId="6" pageNumber="7">
<heading id="D0948188FFF8FFC7FF154064BE2F38B3" bold="true" box="[189,349,623,649]" fontSize="11" level="3" pageId="6" pageNumber="7" reason="3">
<emphasis id="B917EAF6FFF8FFC7FF154064BE2F38B3" bold="true" box="[189,349,623,649]" pageId="6" pageNumber="7">Type locality</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF154095BCFC3882" blockId="6.[189,910,670,696]" box="[189,910,670,696]" pageId="6" pageNumber="7">
<collectingCountry id="F3747674FFF8FFC7FF154095BE533882" box="[189,289,670,696]" name="Ecuador" pageId="6" pageNumber="7">Ecuador</collectingCountry>
, 
<collectingRegion id="49A7F806FFF8FFC7FE844095BEF53882" box="[300,391,670,696]" country="Ecuador" name="Guayas" pageId="6" pageNumber="7">Guayas</collectingRegion>
, Kilometric point 18 Balzar-Quevedo Road.
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFF8FFC7FF1540EBBC063913" box="[189,884,736,809]" pageId="6" pageNumber="7" type="distribution">
<paragraph id="8BDC36E4FFF8FFC7FF1540EBBE2438C0" blockId="6.[189,342,736,762]" box="[189,342,736,762]" pageId="6" pageNumber="7">
<heading id="D0948188FFF8FFC7FF1540EBBE2438C0" bold="true" box="[189,342,736,762]" fontSize="11" level="3" pageId="6" pageNumber="7" reason="3">
<emphasis id="B917EAF6FFF8FFC7FF1540EBBE2438C0" bold="true" box="[189,342,736,762]" pageId="6" pageNumber="7">Distribution</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF154104BC063913" blockId="6.[189,884,783,809]" box="[189,884,783,809]" pageId="6" pageNumber="7">
Pacific slope of the Andes, south-central 
<collectingCountry id="F3747674FFF8FFC7FD354104BD8D3913" box="[669,767,783,809]" name="Ecuador" pageId="6" pageNumber="7">Ecuador</collectingCountry>
(
<figureCitation id="13582A61FFF8FFC7FCA74104BC173913" box="[783,869,783,809]" captionStart="Fig" captionStartId="54.[191,234,1247,1273]" captionTargetBox="[193,1393,268,1195]" captionTargetId="figure-243@54.[189,1396,265,1198]" captionTargetPageId="54" captionText="Fig. 17. Distribution of the Phanaeus endymion species group in South America. Phanaeus malyi Arnaud, 2001 was reported from Colombia without accurate locality. Phanaeus arletteae Arnaud, 2018 was modified from Arnaud (2018)." figureDoi="http://doi.org/10.5281/zenodo.4836800" httpUri="https://zenodo.org/record/4836800/files/figure.png" pageId="6" pageNumber="7">Fig. 17</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFF8FFC7FF15415ABDA23E4B" pageId="6" pageNumber="7" type="discussion">
<paragraph id="8BDC36E4FFF8FFC7FF15415ABE5D3951" blockId="6.[189,303,849,875]" box="[189,303,849,875]" pageId="6" pageNumber="7">
<heading id="D0948188FFF8FFC7FF15415ABE5D3951" bold="true" box="[189,303,849,875]" fontSize="11" level="3" pageId="6" pageNumber="7" reason="3">
<emphasis id="B917EAF6FFF8FFC7FF15415ABE5D3951" bold="true" box="[189,303,849,875]" pageId="6" pageNumber="7">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFF8FFC7FF15418BBDA23E4B" blockId="6.[189,1399,896,1137]" pageId="6" pageNumber="7">
The genital morphology of this species remains unknown. We were not able to personally revise specimens of 
<taxonomicName id="4C634D67FFF8FFC7FEC841AFBD313984" authorityName="Arnaud" authorityYear="2018" box="[352,579,932,958]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="arletteae">
<emphasis id="B917EAF6FFF8FFC7FEC841AFBD313984" box="[352,579,932,958]" italics="true" pageId="6" pageNumber="7">Phanaeus arletteae</emphasis>
</taxonomicName>
. Nevertheless, the external morphology of the 
<typeStatus id="54D88846FFF8FFC7FBD841AFBBA53984" box="[1136,1239,932,958]" pageId="6" pageNumber="7" type="holotype">holotype</typeStatus>
and a female 
<typeStatus id="54D88846FFF8FFC7FF1641C3BE5139D8" box="[190,291,968,994]" pageId="6" pageNumber="7" type="paratype">paratype</typeStatus>
were illustrated by 
<bibRefCitation id="EFF24B15FFF8FFC7FDB141C3BDBC39D8" author="Arnaud" box="[537,718,968,994]" firstAuthor="Arnaud" pageId="6" pageNumber="7" pagination="3 - 7" refId="ref35890" refString="Arnaud P. 2018. Description d'une nouvelle espece de Phanaeus d'Equateur et revalidation de l'espece Coprophanaeus (C.) edmondsi Arnaud (Coleoptera: Scarabaeidae, Scarabaeinae). Besoiro 26: 3 - 7." type="journal article" year="2018">Arnaud (2018)</bibRefCitation>
. We consider that the diagnostic characters provided by 
<bibRefCitation id="EFF24B15FFF8FFC7FF1541E0BE1F3E3C" author="Arnaud" box="[189,365,1003,1030]" firstAuthor="Arnaud" pageId="6" pageNumber="7" pagination="3 - 7" refId="ref35890" refString="Arnaud P. 2018. Description d'une nouvelle espece de Phanaeus d'Equateur et revalidation de l'espece Coprophanaeus (C.) edmondsi Arnaud (Coleoptera: Scarabaeidae, Scarabaeinae). Besoiro 26: 3 - 7." type="journal article" year="2018">Arnaud (2018)</bibRefCitation>
are adequate to separate this species from closely related taxa. The commentaries of previous authors confirmed the validity of this species (
<bibRefCitation id="EFF24B15FFF8FFC7FCC74604BBD53E13" author="Martinez &amp; Pereira" box="[879,1191,1039,1065]" firstAuthor="Martinez" pageId="6" pageNumber="7" pagination="53 - 69" refId="ref40118" refString="Martinez A &amp; Pereira F. S. 1967. Notas escarabeidologicas III (Col. Scarabaeidae-Scarabaeinae). Revista de la Sociedad Entomologica Argentina 29: 53 - 69." type="journal article" year="1967">Martínez &amp; Pereira 1967</bibRefCitation>
; 
<bibRefCitation id="EFF24B15FFF8FFC7FB104604BA003E13" author="Edmonds" box="[1208,1394,1039,1065]" firstAuthor="Edmonds" pageId="6" pageNumber="7" pagination="1 - 105" refId="ref37487" refString="Edmonds W. D. 1994. Revision of Phanaeus Macleay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science, Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">Edmonds 1994</bibRefCitation>
; 
<bibRefCitation id="EFF24B15FFF8FFC7FF154638BE1B3E77" author="Arnaud" box="[189,361,1075,1101]" firstAuthor="Arnaud" pageId="6" pageNumber="7" refId="ref35859" refString="Arnaud P. 2002 b. Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury." type="book" year="2002" yearSuffix="b">Arnaud 2002b</bibRefCitation>
; 
<bibRefCitation id="EFF24B15FFF8FFC7FEDD4638BDE63E77" author="Edmonds &amp; Zidek" box="[373,660,1075,1101]" firstAuthor="Edmonds" pageId="6" pageNumber="7" pagination="1 - 108" refId="ref37762" 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>
). Particularly, the morphology of the pronotum of the female is unique within the 
<taxonomicName id="4C634D67FFF8FFC7FE3A4653BD523E4B" baseAuthorityName="Moctezuma &amp; Halffter" baseAuthorityYear="2017" box="[402,544,1111,1137]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="7" phylum="Arthropoda" rank="species" species="endymion">
<emphasis id="B917EAF6FFF8FFC7FE3A4653BD523E4B" box="[402,544,1111,1137]" italics="true" pageId="6" pageNumber="7">P. endymion</emphasis>
</taxonomicName>
species group.
</paragraph>
</subSubSection>
</treatment>
</document>