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<mods:title id="22BFE575BFF25B0AAF29AEA45C3A7A08">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="4C634D67FFD7FFE8FDC44303BCBA3B19" ID-CoL="4FSJX" authority="Bates, 1887" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[620,968,264,291]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FDC44303BC473B19" bold="true" box="[620,821,264,291]" italics="true" pageId="41" pageNumber="42">Phanaeus pyrois</emphasis>
<bibRefCitation id="EFF24B15FFD7FFE8FC954302BCBA3B19" author="Bates H. W." box="[829,968,265,291]" pageId="41" pageNumber="42" refId="ref36135" refString="Bates H. W. 1887. Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor &amp; Francis, London." type="book" year="1887">Bates, 1887</bibRefCitation>
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<subSubSection id="C379656FFFD7FFE8FDC14326BCB83B7D" box="[617,970,301,327]" pageId="41" pageNumber="42" type="description">
<paragraph id="8BDC36E4FFD7FFE8FDC14326BCB83B7D" blockId="41.[617,970,264,327]" box="[617,970,301,327]" pageId="41" pageNumber="42">
<figureCitation id="13582A61FFD7FFE8FDC14326BDBB3B7D" box="[617,713,301,327]" captionStart="Fig" captionStartId="8.[189,232,1681,1707]" captionTargetBox="[334,1251,265,1642]" captionTargetId="figure-153@8.[333,1254,265,1655]" captionTargetPageId="8" captionText="Fig. 1. Endophallite copulatrix of the Phanaeus endymion species group (except for P. arletteae Arnaud, 2018). A. Phanaeus bravoensis Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (IEXA). B. Phanaeus chiapanecus sp. nov. (TAMU). C. Phanaeus edmondsi Moctezuma, Deloya &amp; Halffter, 2019 (IEXA). D. Phanaeus endymion Harold, 1863 (VMC). E. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). F. Phanaeus halffterorum Edmonds, 1979 (TAMU). G. Phanaeus huichol Moctezuma, Sánchez- Huerta &amp; Halffter, 2017 (IEXA). H. Phanaeus jackenioi sp. nov. (UVGC). I. Phanaeus malyi Arnaud, 2002 (VMC). J. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). K. Phanaeus pacificus sp. nov. (UVGC). L. Phanaeus panamensis sp. nov. (TAMU). M. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). N. Phanaeus pyrois Bates, 1887 (VMC).O. Phanaeus rzedowskii sp. nov. (TAMU).P. Phanaeus zapotecus Edmonds, 2006 (VMC). Q. Phanaeus zoque Moctezuma &amp; Halffter, 2017 (IEXA)." figureDoi="http://doi.org/10.5281/zenodo.4836748" httpUri="https://zenodo.org/record/4836748/files/figure.png" pageId="41" pageNumber="42">Figs 1N</figureCitation>
, 
<figureCitation id="13582A61FFD7FFE8FD7E4326BD803B7D" box="[726,754,301,327]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="41" pageNumber="42">2J</figureCitation>
, 
<figureCitation id="13582A61FFD7FFE8FD564326BC6C3B7D" box="[766,798,301,327]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="41" pageNumber="42">12</figureCitation>
, 
<figureCitation id="13582A61FFD7FFE8FC824326BC383B7D" box="[810,842,301,327]" captionStart="Fig" captionStartId="51.[189,232,1233,1259]" captionTargetBox="[193,1395,270,1197]" captionTargetId="figure-241@51.[189,1398,265,1199]" captionTargetPageId="51" captionText="Fig. 16. Distribution of the Phanaeus endymion species group in Central America (in part)." figureDoi="http://doi.org/10.5281/zenodo.4836798" httpUri="https://zenodo.org/record/4836798/files/figure.png" pageId="41" pageNumber="42">16</figureCitation>
, 
<figureCitation id="13582A61FFD7FFE8FCFE4326BCF93B7D" box="[854,907,301,327]" 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="41" pageNumber="42">18O</figureCitation>
, 
<figureCitation id="13582A61FFD7FFE8FC304326BCB83B7D" box="[920,970,301,327]" 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="41" pageNumber="42">19O</figureCitation>
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<treatmentCitation id="0AC210F5FFD7FFE8FF154364BD303BB0" author="Bates H. W." box="[189,578,367,394]" page="58" pageId="41" pageNumber="42" year="1887">
<taxonomicName id="4C634D67FFD7FFE8FF154364BD303BB0" ID-CoL="4FSJX" authority="Bates, 1887: 58" authorityName="Bates, Det. W. D. Edmonds" authorityPageNumber="58" authorityYear="1887" box="[189,578,367,394]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FF154364BEF23BB3" box="[189,384,367,393]" italics="true" pageId="41" pageNumber="42">Phanaeus pyrois</emphasis>
<bibRefCitation id="EFF24B15FFD7FFE8FE204364BD303BB0" author="Bates H. W." box="[392,578,367,394]" pageId="41" pageNumber="42" refId="ref36135" refString="Bates H. W. 1887. Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor &amp; Francis, London." type="book" year="1887">Bates, 1887: 58</bibRefCitation>
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, pl. 2, table 3, figs 22–23 (in part).
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<taxonomicName id="4C634D67FFD7FFE8FF1543BABEF33BF1" ID-CoL="4FSJX" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[189,385,433,459]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FF1543BABEF33BF1" box="[189,385,433,459]" italics="true" pageId="41" pageNumber="42">Phanaeus pyrois</emphasis>
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– Nevinson 1982: 6 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FCA643BABCCD3BF6" author="Gillet J. J. E." box="[782,959,433,460]" page="85" pageId="41" pageNumber="42" year="1911">
<bibRefCitation id="EFF24B15FFD7FFE8FCA643BABCCD3BF6" author="Gillet J. J. E." box="[782,959,433,460]" pageId="41" pageNumber="42" refId="ref38289" refString="Gillet J. J. E. 1911. Coleopterorum Catalogus. Pars 38: Scarabaeidae: Coprinae. I. W. Junk, Berlin." type="book" year="1911">Gillet 1911: 85</bibRefCitation>
</treatmentCitation>
(in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FBF443BABA373BF1" author="Olsoufieff G. d'" box="[1116,1349,433,459]" page="37" pageId="41" pageNumber="42" year="1924">
<bibRefCitation id="EFF24B15FFD7FFE8FBF443BABA373BF1" author="Olsoufieff G. d'" box="[1116,1349,433,459]" pageId="41" pageNumber="42" 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: 37</bibRefCitation>
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, 93, 152 (in part). — 
<bibRefCitation id="EFF24B15FFD7FFE8FE1443DEBDE53BCA" author="Blackwelder R. E." box="[444,663,469,496]" pageId="41" pageNumber="42" pagination="1 - 1492" refId="ref36166" refString="Available from https: // www. biodiversitylibrary. org / item / 135502 # page / 7 / mode / 1 up [accessed Jun. 2020]. Blackwelder R. E. 1944 - 1957. Checklist of the coleopterous insects of Mexico, Central American, the West Indies, and South America. Parts 1 - 6. United States National Museum, Bulletin 185: 1 - 1492. https: // doi. org / 10.5479 / si. 03629236.185.927" type="journal article" year="1944">Blackwelder 1944</bibRefCitation>
–1957: 210 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FC6B43DEBBC73BD5" author="Edmonds W. D." box="[963,1205,469,495]" page="830" pageId="41" pageNumber="42" year="1972">
<bibRefCitation id="EFF24B15FFD7FFE8FC6B43DEBBC73BD5" author="Edmonds W. D." box="[963,1205,469,495]" pageId="41" pageNumber="42" pagination="731 - 874" refId="ref37422" refString="Edmonds W. D. 1972. Comparative skeletal morphology, systematics and evolution of the Phanaeine dung beetles (Coleoptera: Scarabaeidae). The University of Kansas Science Bulletin 49: 731 - 874." type="journal article" year="1972">Edmonds 1972: 830</bibRefCitation>
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(in part); 1979: 103 (in part); 1994: 3, 5, 8–9, 39, 44–46, 103 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FC1C43F2BA603829" author="Howden H. F. &amp; Young O. P." box="[948,1298,505,531]" page="134" pageId="41" pageNumber="42" year="1981">
<bibRefCitation id="EFF24B15FFD7FFE8FC1C43F2BA603829" author="Howden H. F. &amp; Young O. P." box="[948,1298,505,531]" pageId="41" pageNumber="42" pagination="1 - 204" refId="ref39249" refString="Howden H. F. &amp; Young O. P. 1981. Panamanian Scarabaeinae: Taxonomy, distribution, and habits (Coleoptera, Scarabaeidae). Contributions of the American Entomological Institute 18: 1 - 204." type="journal article" year="1981">Howden &amp; Young 1981: 134</bibRefCitation>
</treatmentCitation>
, 136 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FEF44016BD4A380D" author="Krajcik M." box="[348,568,540,567]" page="152" pageId="41" pageNumber="42" year="2006">
<bibRefCitation id="EFF24B15FFD7FFE8FEF44016BD4A380D" author="Krajcik M." box="[348,568,540,567]" pageId="41" pageNumber="42" 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: 152</bibRefCitation>
</treatmentCitation>
(in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FD484016BCE1380D" author="Price D. L." box="[736,915,541,567]" page="17" pageId="41" pageNumber="42" year="2007">
<bibRefCitation id="EFF24B15FFD7FFE8FD484016BCE1380D" author="Price D. L." box="[736,915,541,567]" pageId="41" pageNumber="42" pagination="1 - 18" refId="ref41753" refString="Price D. L. 2007. A phylogenetic analysis of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae) based on morphological data. Insect Systematics &amp; Evolution 38: 1 - 18. https: // doi. org / 10.1163 / 187631207788784058" type="journal article" year="2007">Price 2007: 17</bibRefCitation>
</treatmentCitation>
, figs 52–53, 54 (in part); 2009: 145 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FEFF404BBDE53861" author="Solis A. &amp; Kohlmann B." box="[343,663,576,603]" page="1" pageId="41" pageNumber="42" year="2012">
<bibRefCitation id="EFF24B15FFD7FFE8FEFF404BBDE53861" author="Solis A. &amp; Kohlmann B." box="[343,663,576,603]" pageId="41" pageNumber="42" pagination="1 - 32" refId="ref42322" refString="Solis A. &amp; Kohlmann B. 2012. Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica. Zootaxa 3482: 1 - 32." type="journal article" year="2012">Solís &amp; Kohlmann 2012: 1</bibRefCitation>
</treatmentCitation>
, 8–10, 31, fig. 1 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FC5F404BBA463861" author="Edmonds W. D. &amp; Zidek J." box="[1015,1332,576,603]" page="1" pageId="41" pageNumber="42" year="2012">
<bibRefCitation id="EFF24B15FFD7FFE8FC5F404BBA463861" author="Edmonds W. D. &amp; Zidek J." box="[1015,1332,576,603]" pageId="41" pageNumber="42" 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>
, 5–6, 8, 13 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FE15406EBC473845" author="Moctezuma V. &amp; Halffter G." box="[445,821,612,639]" page="55" pageId="41" pageNumber="42" year="2017">
<bibRefCitation id="EFF24B15FFD7FFE8FE15406EBC473845" author="Moctezuma V. &amp; Halffter G." box="[445,821,612,639]" pageId="41" pageNumber="42" 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). — Moctezuma 
<emphasis id="B917EAF6FFD7FFE8FBF7406EBBE03844" box="[1119,1170,612,638]" italics="true" pageId="41" pageNumber="42">et al</emphasis>
. 2017: 114, 130 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FEFD4083BDF43898" author="Kohlmann B. &amp; Arriaga-Jimenez A. &amp; Ros M." box="[341,646,648,675]" page="69" pageId="41" pageNumber="42" year="2018">
<bibRefCitation id="EFF24B15FFD7FFE8FEFD4083BDF43898" author="Kohlmann B. &amp; Arriaga-Jimenez A. &amp; Ros M." box="[341,646,648,675]" pageId="41" pageNumber="42" 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="B917EAF6FFD7FFE8FE714082BD7E3898" box="[473,524,648,674]" italics="true" pageId="41" pageNumber="42">et al</emphasis>
. 2018: 69
</bibRefCitation>
</treatmentCitation>
, 78–79, 83, 88, 89, fig. 8a, d (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FBDA4083BE6538FC" author="Chamorro W. &amp; Marin-Armijos D. &amp; Asenjo A." httpUri="http://treatment.plazi.org/id/58071E6D2CBB4638F57CB9C8B5D2211E" page="220" pageId="41" pageNumber="42" year="2019">
<bibRefCitation id="EFF24B15FFD7FFE8FBDA4083BE6538FC" author="Chamorro W. &amp; Marin-Armijos D. &amp; Asenjo A." pageId="41" pageNumber="42" 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">
Chamorro 
<emphasis id="B917EAF6FFD7FFE8FB594082BA573898" box="[1265,1317,648,674]" italics="true" pageId="41" pageNumber="42">et al</emphasis>
. 2019: 220
</bibRefCitation>
</treatmentCitation>
(in part).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF1540DBBA053934" blockId="41.[189,1399,433,854]" pageId="41" pageNumber="42">
<treatmentCitationGroup id="AB7311CAFFD7FFE8FF1540DBBA053934" pageId="41" pageNumber="42">
<taxonomicName id="4C634D67FFD7FFE8FF1540DBBD2238D0" ID-CoL="4FSJX" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[189,592,720,747]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois" subGenus="Notiophanaeus">
<emphasis id="B917EAF6FFD7FFE8FF1540DBBE4338D0" box="[189,305,720,746]" italics="true" pageId="41" pageNumber="42">Phanaeus</emphasis>
(
<emphasis id="B917EAF6FFD7FFE8FEEB40DBBE8738D0" box="[323,501,720,746]" italics="true" pageId="41" pageNumber="42">Notiophanaeus</emphasis>
) 
<emphasis id="B917EAF6FFD7FFE8FDA040DABD2238D0" box="[520,592,721,746]" italics="true" pageId="41" pageNumber="42">pyrois</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFD7FFE8FDD840DBBC3738D0" author="Edmonds W. D." box="[624,837,720,746]" page="2" pageId="41" pageNumber="42" year="1994">
<bibRefCitation id="EFF24B15FFD7FFE8FDD840DBBC3738D0" author="Edmonds W. D." box="[624,837,720,746]" pageId="41" pageNumber="42" 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: 2</bibRefCitation>
</treatmentCitation>
, 8, 41, 44, figs 210, 214–215, 221 (in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FF4240FFBECC3934" author="Arnaud P." box="[234,446,756,782]" page="96" pageId="41" pageNumber="42" year="2002">
<bibRefCitation id="EFF24B15FFD7FFE8FF4240FFBECC3934" author="Arnaud P." box="[234,446,756,782]" pageId="41" pageNumber="42" refId="ref35859" refString="Arnaud P. 2002 b. Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury." type="book" year="2002">Arnaud 2002b: 96</bibRefCitation>
</treatmentCitation>
(in part). — 
<treatmentCitation id="0AC210F5FFD7FFE8FDFD40FFBCFE3934" author="Edmonds W. D. &amp; Zidek J." box="[597,908,756,782]" page="3" pageId="41" pageNumber="42" year="2012">
<bibRefCitation id="EFF24B15FFD7FFE8FDFD40FFBCFE3934" author="Edmonds W. D. &amp; Zidek J." box="[597,908,756,782]" pageId="41" pageNumber="42" 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: 3</bibRefCitation>
</treatmentCitation>
, 13, figs 138, 142–143, 156–159 (in part).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF154113BC8E3908" blockId="41.[189,1399,433,854]" box="[189,1020,792,818]" pageId="41" pageNumber="42">
<treatmentCitationGroup id="AB7311CAFFD7FFE8FF154113BC8E3908" box="[189,1020,792,818]" pageId="41" pageNumber="42">
<taxonomicName id="4C634D67FFD7FFE8FF154113BDEE3908" ID-CoL="4FSJX" authority="pyrois" box="[189,668,792,818]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="subSpecies" species="pyrois" subGenus="Notiophanaeus" subSpecies="pyrois">
<emphasis id="B917EAF6FFD7FFE8FF154113BE433908" box="[189,305,792,818]" italics="true" pageId="41" pageNumber="42">Phanaeus</emphasis>
(
<emphasis id="B917EAF6FFD7FFE8FEEA4113BE863908" box="[322,500,792,818]" italics="true" pageId="41" pageNumber="42">Notiophanaeus</emphasis>
) 
<emphasis id="B917EAF6FFD7FFE8FDAC4112BDEE3908" box="[516,668,793,818]" italics="true" pageId="41" pageNumber="42">pyrois pyrois</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFD7FFE8FD104113BCFF3908" author="Arnaud P." box="[696,909,792,818]" page="96" pageId="41" pageNumber="42" year="2002">
<bibRefCitation id="EFF24B15FFD7FFE8FD104113BCFF3908" author="Arnaud P." box="[696,909,792,818]" pageId="41" pageNumber="42" refId="ref35859" refString="Arnaud P. 2002 b. Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury." type="book" year="2002">Arnaud 2002b: 96</bibRefCitation>
</treatmentCitation>
(in part).
</treatmentCitationGroup>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF154137BC5D396C" blockId="41.[189,1399,433,854]" box="[189,815,828,854]" pageId="41" pageNumber="42">
<treatmentCitationGroup id="AB7311CAFFD7FFE8FF154137BC5D396C" box="[189,815,828,854]" pageId="41" pageNumber="42">
<taxonomicName id="4C634D67FFD7FFE8FF154137BEA2396F" ID-CoL="4FSJX" authority="pyrois" box="[189,464,828,854]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="subSpecies" species="pyrois" subSpecies="pyrois">
<emphasis id="B917EAF6FFD7FFE8FF154137BEA2396F" box="[189,464,828,854]" italics="true" pageId="41" pageNumber="42">Phanaeus pyrois pyrois</emphasis>
</taxonomicName>
– 
<treatmentCitation id="0AC210F5FFD7FFE8FE444137BDB3396C" author="Arnaud P." box="[492,705,828,854]" page="97" pageId="41" pageNumber="42" year="2002">
<bibRefCitation id="EFF24B15FFD7FFE8FE444137BDB3396C" author="Arnaud P." box="[492,705,828,854]" pageId="41" pageNumber="42" refId="ref35859" refString="Arnaud P. 2002 b. Phanaeini. Dendropaemon, Tetramereia, Homalotarsus, Megatharsis, Diabroctis, Coprophanaeus, Oxysternon, Phanaeus, Sulcophanaeus. Hillside Books, Canterbury." type="book" year="2002">Arnaud 2002b: 97</bibRefCitation>
</treatmentCitation>
(in part).
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFD7FFE8FF154176BEEA3E40" pageId="41" pageNumber="42" type="diagnosis">
<paragraph id="8BDC36E4FFD7FFE8FF154176BE4439AD" blockId="41.[189,310,893,919]" box="[189,310,893,919]" pageId="41" pageNumber="42">
<heading id="D0948188FFD7FFE8FF154176BE4439AD" bold="true" box="[189,310,893,919]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="3">
<emphasis id="B917EAF6FFD7FFE8FF154176BE4439AD" bold="true" box="[189,310,893,919]" pageId="41" pageNumber="42">Diagnosis</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF1541A6BEEA3E40" blockId="41.[187,1399,941,1146]" pageId="41" pageNumber="42">
Easily diagnosed species by the pronotum bright metallic red (
<figureCitation id="13582A61FFD7FFE8FC1A41A6BB3E39FD" box="[946,1100,941,967]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="41" pageNumber="42">Figs 12A, D</figureCitation>
), green (
<figureCitation id="13582A61FFD7FFE8FB1041A6BA7F39FD" box="[1208,1293,941,967]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="41" pageNumber="42">Fig. 2J</figureCitation>
) or dark metallic blue (12B), with elytral striae not strongly impressed basally (
<figureCitation id="13582A61FFD7FFE8FBAF41DABB1339D1" box="[1031,1121,977,1003]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="41" pageNumber="42">Fig. 12</figureCitation>
). The rest of the green/ blue species of the 
<taxonomicName id="4C634D67FFD7FFE8FE0141FEBD4B3E35" authorityName="Harold. Ex-Mus" authorityYear="1863" box="[425,569,1013,1039]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="endymion">
<emphasis id="B917EAF6FFD7FFE8FE0141FEBD4B3E35" box="[425,569,1013,1039]" italics="true" pageId="41" pageNumber="42">P. endymion</emphasis>
</taxonomicName>
species group are recognized by the elytral striae strongly impressed basally as a distinct fossa. A black dorsal colour is never found in 
<taxonomicName id="4C634D67FFD7FFE8FC624612BB423E08" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[970,1072,1049,1074]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FC624612BB423E08" box="[970,1072,1049,1074]" italics="true" pageId="41" pageNumber="42">P. pyrois</emphasis>
</taxonomicName>
specimens. Minor males of 
<taxonomicName id="4C634D67FFD7FFE8FF164636BE113E6C" authority="Moctezuma &amp; Halffter, 2021" authorityName="Moctezuma &amp; Halffter" authorityYear="2021" box="[190,355,1085,1110]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="panamensis" status="sp. nov.">
<emphasis id="B917EAF6FFD7FFE8FF164636BE113E6C" box="[190,355,1085,1110]" italics="true" pageId="41" pageNumber="42">P. panamensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A224578DFFD7FFE8FECF4636BEB23E6D" box="[359,448,1085,1111]" pageId="41" pageNumber="42" rank="species">sp. nov.</taxonomicNameLabel>
and red 
<taxonomicName id="4C634D67FFD7FFE8FDB54636BDF03E6C" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[541,642,1085,1110]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FDB54636BDF03E6C" box="[541,642,1085,1110]" italics="true" pageId="41" pageNumber="42">P. pyrois</emphasis>
</taxonomicName>
may be strongly mimetic, but easily separated by the endophallite copulatrix (
<figureCitation id="13582A61FFD7FFE8FEEA466BBEF53E40" box="[322,391,1120,1146]" captionStart="Fig" captionStartId="8.[189,232,1681,1707]" captionTargetBox="[334,1251,265,1642]" captionTargetId="figure-153@8.[333,1254,265,1655]" captionTargetPageId="8" captionText="Fig. 1. Endophallite copulatrix of the Phanaeus endymion species group (except for P. arletteae Arnaud, 2018). A. Phanaeus bravoensis Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (IEXA). B. Phanaeus chiapanecus sp. nov. (TAMU). C. Phanaeus edmondsi Moctezuma, Deloya &amp; Halffter, 2019 (IEXA). D. Phanaeus endymion Harold, 1863 (VMC). E. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). F. Phanaeus halffterorum Edmonds, 1979 (TAMU). G. Phanaeus huichol Moctezuma, Sánchez- Huerta &amp; Halffter, 2017 (IEXA). H. Phanaeus jackenioi sp. nov. (UVGC). I. Phanaeus malyi Arnaud, 2002 (VMC). J. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). K. Phanaeus pacificus sp. nov. (UVGC). L. Phanaeus panamensis sp. nov. (TAMU). M. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). N. Phanaeus pyrois Bates, 1887 (VMC).O. Phanaeus rzedowskii sp. nov. (TAMU).P. Phanaeus zapotecus Edmonds, 2006 (VMC). Q. Phanaeus zoque Moctezuma &amp; Halffter, 2017 (IEXA)." figureDoi="http://doi.org/10.5281/zenodo.4836748" httpUri="https://zenodo.org/record/4836748/files/figure.png" pageId="41" pageNumber="42">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFD7FFE8FF1546A9BEB33D15" pageId="41" pageNumber="42" type="materials_examined">
<paragraph id="8BDC36E4FFD7FFE8FF1546A9BE1E3E86" blockId="41.[189,364,1186,1212]" box="[189,364,1186,1212]" pageId="41" pageNumber="42">
<heading id="D0948188FFD7FFE8FF1546A9BE1E3E86" bold="true" box="[189,364,1186,1212]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="3">
<emphasis id="B917EAF6FFD7FFE8FF1546A9BE1E3E86" bold="true" box="[189,364,1186,1212]" pageId="41" pageNumber="42">Type material</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF4246D9BD9B3ED6" blockId="41.[189,1399,1233,1368]" box="[234,745,1233,1260]" pageId="41" pageNumber="42">
<emphasis id="B917EAF6FFD7FFE8FF4246D9BE143ED6" bold="true" box="[234,358,1234,1260]" pageId="41" pageNumber="42">
<typeStatus id="54D88846FFD7FFE8FF4246D9BE143ED6" box="[234,358,1234,1260]" pageId="41" pageNumber="42" type="lectotype">Lectotype</typeStatus>
</emphasis>
(studied from photographs, 
<specimenCount id="9D65FD6DFFD7FFE8FD1B46D9BD923ED6" box="[691,736,1234,1260]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
)
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF1546FDBCAD3F62" blockId="41.[189,1399,1233,1368]" pageId="41" pageNumber="42">
<materialsCitation id="3B0B3CB9FFD7FFE8FF1546FDBCA93F62" ID-GBIF-Occurrence="3093980511" collectionCode="BMNH" collectorName="T. Belt" country="Nicaragua" location="Chontales" pageId="41" pageNumber="42" specimenCount="2" specimenCount-male="2" stateProvince="Chontales" typeStatus="lectotype">
<collectingCountry id="F3747674FFD7FFE8FF1546FDBE193F2A" box="[189,363,1270,1296]" name="Nicaragua" pageId="41" pageNumber="42">NICARAGUA</collectingCountry>
– 
<emphasis id="B917EAF6FFD7FFE8FE3C46FEBD633F35" bold="true" box="[404,529,1269,1295]" pageId="41" pageNumber="42">
<collectingRegion id="49A7F806FFD7FFE8FE3C46FEBD633F35" box="[404,529,1269,1295]" country="Nicaragua" name="Chontales" pageId="41" pageNumber="42">Chontales</collectingRegion>
</emphasis>
• 
<specimenCount id="9D65FD6DFFD7FFE8FD9F46FDBD3D3F2A" box="[567,591,1270,1296]" pageId="41" pageNumber="42" type="male">♂</specimenCount>
, 
<bibRefCitation id="EFF24B15FFD7FFE8FDC946FEBC383F2A" author="Edmonds W. D." box="[609,842,1269,1296]" pageId="41" pageNumber="42" 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: 45</bibRefCitation>
(
<figureCitation id="13582A61FFD7FFE8FCC846FEBCA63F2A" box="[864,980,1269,1296]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="41" pageNumber="42">Fig. 12D</figureCitation>
); “NHMUK 013678267/ B. C. A. p. 58, sp.8. / LECTO-TYPE/ 
<taxonomicName id="4C634D67FFD7FFE8FDBF4712BC4E3F0E" authority="Bates." authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[535,828,1305,1332]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FDBF4712BD973F09" box="[535,741,1305,1331]" italics="true" pageId="41" pageNumber="42">Phanaeus pyrois</emphasis>
Bates.
</taxonomicName>
LECTOTYPE 
<specimenCount id="9D65FD6DFFD7FFE8FC574711BB673F0E" box="[1023,1045,1306,1332]" pageId="41" pageNumber="42" type="male">♂</specimenCount>
P. ARNAUD DET 1980/T. Belt /Type / 
<taxonomicName id="4C634D67FFD7FFE8FEEB4735BEDA3F6D" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[323,424,1342,1367]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="41" pageNumber="42" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD7FFE8FEEB4735BEDA3F6D" box="[323,424,1342,1367]" italics="true" pageId="41" pageNumber="42">P. pyrois</emphasis>
</taxonomicName>
♂/ Sp. figured”; NHMUK 013678267; 
<collectionCode id="ED72AE21FFD7FFE8FCD74735BCA93F62" box="[895,987,1342,1368]" collectionName="United Kingdom, London, The Natural History Museum [formerly British Museum (Natural History)]" pageId="41" pageNumber="42">BMNH</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF154774BDC23FA0" blockId="41.[189,688,1407,1434]" box="[189,688,1407,1434]" pageId="41" pageNumber="42">
<emphasis id="B917EAF6FFD7FFE8FF154774BD703FA3" bold="true" box="[189,514,1407,1433]" pageId="41" pageNumber="42">Non-type material revised</emphasis>
(
<specimenCount id="9D65FD6DFFD7FFE8FDBB4774BD253FA0" box="[531,599,1407,1434]" pageId="41" pageNumber="42" type="male">7 ♂♂</specimenCount>
, 
<specimenCount id="9D65FD6DFFD7FFE8FDCC4774BDD63FA0" box="[612,676,1407,1434]" pageId="41" pageNumber="42" type="female">4 ♀♀</specimenCount>
)
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF1547A4BB273C46" blockId="41.[189,1399,1454,1660]" pageId="41" pageNumber="42">
<materialsCitation id="3B0B3CB9FFD7FFE8FF1547A4BA113FF3" ID-GBIF-Occurrence="3093980353" box="[189,1379,1454,1481]" collectingDate="1998-06-30" collectionCode="TAMU" collectorName="JM. Mars" country="Nicaragua" location="Volcan Mombacho" pageId="41" pageNumber="42" specimenCount="1" specimenCount-male="1" stateProvince="Granada">
<collectingCountry id="F3747674FFD7FFE8FF1547A4BE193FF3" box="[189,363,1455,1481]" name="Nicaragua" pageId="41" pageNumber="42">NICARAGUA</collectingCountry>
– 
<emphasis id="B917EAF6FFD7FFE8FE2447A5BE8E3FF2" bold="true" box="[396,508,1454,1480]" pageId="41" pageNumber="42">
<collectingRegion id="49A7F806FFD7FFE8FE2447A5BE8E3FF2" box="[396,508,1454,1480]" country="Nicaragua" name="Granada" pageId="41" pageNumber="42">Granada</collectingRegion>
</emphasis>
• 
<specimenCount id="9D65FD6DFFD7FFE8FDB247A4BD393FF3" box="[538,587,1455,1481]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
; “ 
<location id="8EBC603FFFD7FFE8FDCB47A5BC383FF2" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FDCB47A5BC383FF2" box="[611,842,1454,1480]" country="Nicaragua" name="Volcan Mombacho" pageId="41" pageNumber="42" stateProvince="Granada">Volcán Mombacho</location>
. 
<location id="8EBC603FFFD7FFE8FCFE47A4BC863FF3" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FCFE47A4BC863FF3" box="[854,1012,1455,1481]" country="Nicaragua" name="Bosque Seco" pageId="41" pageNumber="42" stateProvince="Granada">Bosque Seco</location>
. 
<date id="FFDD1024FFD7FFE8FBA947A4BB033FF3" box="[1025,1137,1455,1481]" pageId="41" pageNumber="42" value="1998-06-30">
<collectingDate id="EF99E9CCFFD7FFE8FBA947A4BB033FF3" box="[1025,1137,1455,1481]" pageId="41" pageNumber="42" value="1998-06-30">30-VI-98</collectingDate>
</date>
. 
<collectorName id="26965332FFD7FFE8FBD747A4BB853FF3" box="[1151,1271,1455,1481]" pageId="41" pageNumber="42">JM. Mars</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FAA747A4BA113FF3" box="[1295,1379,1455,1481]" collectionName="USA, Texas, College Station, Texas A &amp; M University" pageId="41" pageNumber="42">TAMU</collectionCode>
</materialsCitation>
• 
<materialsCitation id="3B0B3CB9FFD7FFE8FF1547D9BB5E3FD6" ID-GBIF-Occurrence="3093980488" box="[189,1068,1490,1517]" collectingDate="1998-05-21" collectingMethod="Malaise" collectionCode="VMC" collectorName="JM. Mars" country="Nicaragua" location="Volcan Mombacho" pageId="41" pageNumber="42" specimenCount="1" specimenCount-male="1" stateProvince="Granada">
<specimenCount id="9D65FD6DFFD7FFE8FF1547D9BF9F3FD7" box="[189,237,1490,1517]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
; “ 
<location id="8EBC603FFFD7FFE8FEAC47D9BE983FD6" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FEAC47D9BE983FD6" box="[260,490,1490,1516]" country="Nicaragua" name="Volcan Mombacho" pageId="41" pageNumber="42" stateProvince="Granada">Volcán Mombacho</location>
. 
<location id="8EBC603FFFD7FFE8FE5E47D9BD003FD7" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FE5E47D9BD003FD7" box="[502,626,1490,1517]" country="Nicaragua" name="Santa Ana" pageId="41" pageNumber="42" stateProvince="Granada">Santa Ana</location>
. 
<date id="FFDD1024FFD7FFE8FDD647D9BD903FD6" box="[638,738,1490,1516]" pageId="41" pageNumber="42" value="1998-05-21">
<collectingDate id="EF99E9CCFFD7FFE8FDD647D9BD903FD6" box="[638,738,1490,1516]" pageId="41" pageNumber="42" value="1998-05-21">21-V-98</collectingDate>
</date>
. 
<collectingMethod id="52224EF3FFD7FFE8FD4647D9BC3D3FD6" box="[750,847,1490,1516]" pageId="41" pageNumber="42">Malaise</collectingMethod>
. 
<collectorName id="26965332FFD7FFE8FCF247D8BCA33FD6" box="[858,977,1490,1517]" pageId="41" pageNumber="42">JM. Mars</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FC4147D9BB5E3FD6" box="[1001,1068,1490,1516]" pageId="41" pageNumber="42">VMC</collectionCode>
</materialsCitation>
• 
<materialsCitation id="3B0B3CB9FFD7FFE8FBEE47D9BDC23C2A" ID-GBIF-Occurrence="3093980448" collectingDate="1998-06-30" collectionCode="IEXA" collectorName="JM. Mars" country="Nicaragua" location="Volcan Mombacho" pageId="41" pageNumber="42" specimenCount="1" specimenCount-male="1" stateProvince="Granada">
<specimenCount id="9D65FD6DFFD7FFE8FBEE47D9BB043FD7" box="[1094,1142,1490,1517]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
; “ 
<location id="8EBC603FFFD7FFE8FB2547D9BA013FD6" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FB2547D9BA013FD6" box="[1165,1395,1490,1516]" country="Nicaragua" name="Volcan Mombacho" pageId="41" pageNumber="42" stateProvince="Granada">Volcán Mombacho</location>
. 
<location id="8EBC603FFFD7FFE8FF1547FDBE3E3C2B" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FF1547FDBE3E3C2B" box="[189,332,1526,1553]" country="Nicaragua" name="El Progreso" pageId="41" pageNumber="42" stateProvince="Granada">El Progreso</location>
. 
<date id="FFDD1024FFD7FFE8FEF047FDBEBB3C2A" box="[344,457,1526,1552]" pageId="41" pageNumber="42" value="1998-06-30">
<collectingDate id="EF99E9CCFFD7FFE8FEF047FDBEBB3C2A" box="[344,457,1526,1552]" pageId="41" pageNumber="42" value="1998-06-30">30-VI-98</collectingDate>
</date>
. 
<collectorName id="26965332FFD7FFE8FE7E47FCBD393C2A" box="[470,587,1526,1553]" pageId="41" pageNumber="42">JM. Mars</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FDCE47FDBDC23C2A" box="[614,688,1526,1552]" pageId="41" pageNumber="42">IEXA</collectionCode>
</materialsCitation>
. – 
<materialsCitation id="3B0B3CB9FFD7FFE8FD7D47FDBD473C0E" ID-GBIF-Occurrence="3093980450" collectingDate="2005-12-03" collectingDateMax="2005-12-08" collectingDateMin="2005-12-03" collectionCode="TAMU" collectorName="D. G. Marqua" country="Nicaragua" location="El Jaguar Coffee Finca" pageId="41" pageNumber="42" specimenCount="1" specimenCount-male="1" stateProvince="Jinotega">
<emphasis id="B917EAF6FFD7FFE8FD7D47FDBC4D3C2A" bold="true" box="[725,831,1526,1552]" pageId="41" pageNumber="42">
<collectingRegion id="49A7F806FFD7FFE8FD7D47FDBC4D3C2A" box="[725,831,1526,1552]" country="Nicaragua" name="Jinotega" pageId="41" pageNumber="42">Jinotega</collectingRegion>
</emphasis>
• 
<specimenCount id="9D65FD6DFFD7FFE8FCF247FDBCF83C2B" box="[858,906,1526,1553]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
; “ 
<location id="8EBC603FFFD7FFE8FC0B47FDBBC93C2A" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FC0B47FDBBC93C2A" box="[931,1211,1526,1553]" country="Nicaragua" name="El Jaguar Coffee Finca" pageId="41" pageNumber="42" stateProvince="Jinotega">El Jaguar Coffee Finca</location>
. 
<date id="FFDD1024FFD7FFE8FB6F47FDBA003C2B" box="[1223,1394,1526,1553]" pageId="41" pageNumber="42" value="2005-12-03" valueMax="2005-12-08" valueMin="2005-12-03">
<collectingDate id="EF99E9CCFFD7FFE8FB6F47FDBA003C2B" box="[1223,1394,1526,1553]" pageId="41" pageNumber="42" value="2005-12-03" valueMax="2005-12-08" valueMin="2005-12-03">
XII-3–8– 
<quantity id="4C9B9B01FFD7FFE8FA9D47FDBA003C2B" box="[1333,1394,1526,1553]" metricMagnitude="2" metricUnit="m" metricValue="6.1125677087999994" pageId="41" pageNumber="42" unit="ft" value="2005.4356">2005</quantity>
</collectingDate>
</date>
. 4356 ft. 
<collectorName id="26965332FFD7FFE8FE894410BEBE3C0E" box="[289,460,1562,1589]" pageId="41" pageNumber="42">D. G. Marqua</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FE494411BD473C0E" box="[481,565,1562,1588]" collectionName="USA, Texas, College Station, Texas A &amp; M University" pageId="41" pageNumber="42">TAMU</collectionCode>
</materialsCitation>
• 
<materialsCitation id="3B0B3CB9FFD7FFE8FDE74411BEE83C62" ID-GBIF-Occurrence="3093980310" collectingDate="2005-06-05" collectingDateMax="2005-06-10" collectingDateMin="2005-06-05" collectionCode="VMC" collectorName="D. G. Marqua" country="Nicaragua" elevation="1328" location="El Jaguar Coffee Finca" pageId="41" pageNumber="42" specimenCount="2" specimenCount-female="2" stateProvince="Jinotega">
<specimenCount id="9D65FD6DFFD7FFE8FDE74411BDE13C0F" box="[591,659,1562,1589]" pageId="41" pageNumber="42" type="female">2 ♀♀</specimenCount>
; “ 
<location id="8EBC603FFFD7FFE8FD054411BCB03C0E" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FD054411BCB03C0E" box="[685,962,1562,1589]" country="Nicaragua" name="El Jaguar Coffee Finca" pageId="41" pageNumber="42" stateProvince="Jinotega">El Jaguar Coffee Finca</location>
. 
<date id="FFDD1024FFD7FFE8FC654411BB0E3C0F" box="[973,1148,1562,1589]" pageId="41" pageNumber="42" value="2005-06-05" valueMax="2005-06-10" valueMin="2005-06-05">
<collectingDate id="EF99E9CCFFD7FFE8FC654411BB0E3C0F" box="[973,1148,1562,1589]" pageId="41" pageNumber="42" value="2005-06-05" valueMax="2005-06-10" valueMin="2005-06-05">VI-5–10–2005</collectingDate>
</date>
, el. 
<quantity id="4C9B9B01FFD7FFE8FB054411BA633C0E" box="[1197,1297,1562,1589]" metricMagnitude="3" metricUnit="m" metricValue="1.3277087999999997" pageId="41" pageNumber="42" unit="ft" value="4356.0">
<elevation id="004ED1D7FFD7FFE8FB054411BA633C0E" box="[1197,1297,1562,1589]" metricMagnitude="3" metricUnit="m" metricValue="1.3277087999999997" pageId="41" pageNumber="42" unit="ft" value="4356.0">4,356 ft.</elevation>
</quantity>
Coll. 
<collectorName id="26965332FFD7FFE8FAF24410BE303C62" pageId="41" pageNumber="42">D. G. Marqua</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FEFF4435BEE83C62" box="[343,410,1598,1624]" pageId="41" pageNumber="42">VMC</collectionCode>
</materialsCitation>
• 
<materialsCitation id="3B0B3CB9FFD7FFE8FE1A4435BE863C46" ID-GBIF-Occurrence="3093980525" collectingDate="2005-12" collectionCode="TAMU" collectorName="D. G. Marqua" country="Nicaragua" elevation="1340" latitude="13.241112" location="Finca El Jaguar" longLatPrecision="21" longitude="-86.05445" pageId="41" pageNumber="42" specimenCount="3" specimenCount-female="2" specimenCount-male="1" stateProvince="Jinotega">
<specimenCount id="9D65FD6DFFD7FFE8FE1A4435BE923C63" box="[434,480,1598,1625]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
, 
<specimenCount id="9D65FD6DFFD7FFE8FE444435BD5D3C63" box="[492,559,1598,1625]" pageId="41" pageNumber="42" type="female">2 ♀♀</specimenCount>
; “ 
<location id="8EBC603FFFD7FFE8FDEF4435BC703C63" LSID="urn:lsid:plazi:treatment:03CA87F2FFD7FFEDFDC44303BE123858:8EBC603FFFD7FFE8FDEF4435BC703C63" box="[583,770,1598,1625]" country="Nicaragua" latitude="13.241112" longLatPrecision="21" longitude="-86.05445" name="Finca El Jaguar" pageId="41" pageNumber="42" stateProvince="Jinotega">Finca El Jaguar</location>
, 
<locationDeviation id="9A6E2FC6FFD7FFE8FCA54435BCF03C62" box="[781,898,1598,1624]" pageId="41" pageNumber="42">32kmNW</locationDeviation>
. 
<quantity id="4C9B9B01FFD7FFE8FC254435BC903C62" box="[909,994,1598,1624]" metricMagnitude="3" metricUnit="m" metricValue="1.34" pageId="41" pageNumber="42" unit="m" value="1340.0">
<elevation id="004ED1D7FFD7FFE8FC254435BC903C62" box="[909,994,1598,1624]" metricMagnitude="3" metricUnit="m" metricValue="1.34" pageId="41" pageNumber="42" unit="m" value="1340.0">1340m</elevation>
</quantity>
. 13°14’28’’N-86°03’16’’W. 
<date id="FFDD1024FFD7FFE8FA984435BA053C63" box="[1328,1399,1598,1625]" pageId="41" pageNumber="42" value="2005-12">
<collectingDate id="EF99E9CCFFD7FFE8FA984435BA053C63" box="[1328,1399,1598,1625]" pageId="41" pageNumber="42" value="2005-12">xii-05</collectingDate>
</date>
col 
<collectorName id="26965332FFD7FFE8FF404469BEF93C46" box="[232,395,1634,1660]" pageId="41" pageNumber="42">D.G. Marqua</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FE084469BE863C46" box="[416,500,1634,1660]" collectionName="USA, Texas, College Station, Texas A &amp; M University" pageId="41" pageNumber="42">TAMU</collectionCode>
</materialsCitation>
• 
<materialsCitation id="3B0B3CB9FFD7FFE8FDA54469BB203C46" ID-GBIF-Occurrence="3093980517" box="[525,1106,1634,1660]" collectingDate="2005-12" collectionCode="VMC" collectorName="D. G. Marqua" country="Nicaragua" elevation="1340" latitude="13.241112" location="Finca El Jaguar" longLatPrecision="21" longitude="-86.05445" pageId="41" pageNumber="42" specimenCount="1" specimenCount-male="1" stateProvince="Jinotega">
<specimenCount id="9D65FD6DFFD7FFE8FDA54469BD4E3C46" box="[525,572,1634,1660]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
; same collection data as for preceding; 
<collectionCode id="ED72AE21FFD7FFE8FBA34469BB203C46" box="[1035,1106,1634,1660]" pageId="41" pageNumber="42">VMC</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF1544AFBB953C84" blockId="41.[189,1255,1700,1727]" box="[189,1255,1700,1727]" pageId="41" pageNumber="42">
<materialsCitation id="3B0B3CB9FFD7FFE8FF1544AFBB913C84" ID-GBIF-Occurrence="3093980530" box="[189,1251,1700,1727]" collectingDate="1980-02-26" collectionCode="TAMU" collectorName="H &amp; A Howden" country="Costa Rica" elevation="650" location="Cartago" pageId="41" pageNumber="42" specimenCount="651" specimenCount-male="1" stateProvince="Cartago">
<collectingCountry id="F3747674FFD7FFE8FF1544AFBE153C84" box="[189,359,1700,1726]" name="Costa Rica" pageId="41" pageNumber="42">COSTA RICA</collectingCountry>
– 
<emphasis id="B917EAF6FFD7FFE8FE2A44AFBE9B3C84" bold="true" box="[386,489,1700,1726]" pageId="41" pageNumber="42">
<collectingRegion id="49A7F806FFD7FFE8FE2A44AFBE9B3C84" box="[386,489,1700,1726]" country="Costa Rica" name="Cartago" pageId="41" pageNumber="42">Cartago</collectingRegion>
</emphasis>
• 
<specimenCount id="9D65FD6DFFD7FFE8FDAA44AFBD433C84" box="[514,561,1700,1726]" pageId="41" pageNumber="42" type="male">1 ♂</specimenCount>
; “Turrialba. 
<specimenCount id="9D65FD6DFFD7FFE8FD6D44AFBC623C85" box="[709,784,1700,1727]" pageId="41" pageNumber="42" type="generic">
<quantity id="4C9B9B01FFD7FFE8FD6D44AFBC793C85" box="[709,779,1700,1727]" metricMagnitude="2" metricUnit="m" metricValue="6.5" pageId="41" pageNumber="42" unit="m" value="650.0">650m</quantity>
.
</specimenCount>
<date id="FFDD1024FFD7FFE8FCBF44AFBCDF3C84" box="[791,941,1700,1726]" pageId="41" pageNumber="42" value="1980-02-26">
<collectingDate id="EF99E9CCFFD7FFE8FCBF44AFBCDF3C84" box="[791,941,1700,1726]" pageId="41" pageNumber="42" value="1980-02-26">26.Feb.1980</collectingDate>
</date>
. 
<collectorName id="26965332FFD7FFE8FC1044AFBB003C84" box="[952,1138,1700,1726]" pageId="41" pageNumber="42">H &amp; A Howden</collectorName>
”; 
<collectionCode id="ED72AE21FFD7FFE8FB2344AFBB913C84" box="[1163,1251,1700,1726]" collectionName="USA, Texas, College Station, Texas A &amp; M University" pageId="41" pageNumber="42">TAMU</collectionCode>
</materialsCitation>
.
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF1544EDBE2F3D3A" blockId="41.[189,349,1766,1792]" box="[189,349,1766,1792]" pageId="41" pageNumber="42">
<heading id="D0948188FFD7FFE8FF1544EDBE2F3D3A" bold="true" box="[189,349,1766,1792]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="3">
<emphasis id="B917EAF6FFD7FFE8FF1544EDBE2F3D3A" bold="true" box="[189,349,1766,1792]" pageId="41" pageNumber="42">Type locality</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF15451EBEB33D15" blockId="41.[189,449,1813,1839]" box="[189,449,1813,1839]" pageId="41" pageNumber="42">
<collectingCountry id="F3747674FFD7FFE8FF15451EBE4B3D15" box="[189,313,1813,1839]" name="Nicaragua" pageId="41" pageNumber="42">Nicaragua</collectingCountry>
, 
<collectingRegion id="49A7F806FFD7FFE8FEEC451EBECF3D15" box="[324,445,1813,1839]" country="Nicaragua" name="Chontales" pageId="41" pageNumber="42">Chontales</collectingRegion>
.
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFD7FFEBFF15455CBC813CF8" lastPageId="42" lastPageNumber="43" pageId="41" pageNumber="42" type="description">
<paragraph id="8BDC36E4FFD7FFE8FF15455CBE193D4B" blockId="41.[189,363,1879,1905]" box="[189,363,1879,1905]" pageId="41" pageNumber="42">
<heading id="D0948188FFD7FFE8FF15455CBE193D4B" bold="true" box="[189,363,1879,1905]" fontSize="11" level="3" pageId="41" pageNumber="42" reason="3">
<emphasis id="B917EAF6FFD7FFE8FF15455CBE193D4B" bold="true" box="[189,363,1879,1905]" pageId="41" pageNumber="42">Redescription</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFD7FFE8FF42458DBE0F3D9B" blockId="41.[189,1400,1926,2025]" box="[234,381,1926,1953]" pageId="41" pageNumber="42">
<emphasis id="B917EAF6FFD7FFE8FF42458DBE0F3D9B" bold="true" box="[234,381,1926,1953]" pageId="41" pageNumber="42">Major male</emphasis>
</paragraph>
<paragraph id="8BDC36E4FFD7FFEBFF1545A0BCA03B7D" blockId="41.[189,1400,1926,2025]" lastBlockId="42.[189,1398,265,327]" lastPageId="42" lastPageNumber="43" pageId="41" pageNumber="42">
HEAD. Clypeus bidentate, black anteriorly, bright metallic red, green, or dark metallic blue posteriorly; roughened sculpture. Genae bright metallic red, green, or dark metallic blue; roughened sculpture. Front black, bright metallic red, green, or dark metallic blue on portions adjacent to cephalic horn. Cephalic horn black, curved posteriorly over pronotum (
<figureCitation id="13582A61FFD4FFEBFD4C4327BC4E3B7C" box="[740,828,300,326]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="42" pageNumber="43">Figs 2J</figureCitation>
, 
<figureCitation id="13582A61FFD4FFEBFCE04327BCB73B7D" box="[840,965,300,327]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="42" pageNumber="43">12A–B, D</figureCitation>
).
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF154365BC543845" blockId="42.[189,1399,366,639]" pageId="42" pageNumber="43">
PRONOTUM. Keel absent in the middle of anterior pronotal margin. Disc triangular, flat, with two distinctly developed tubercles on anterior portion. Triangle bright metallic red, green, or dark metallic blue; becoming black on posterior margin of posterolateral angles; lightly granulate, scabriculous, impunctate. Sides bright metallic red, green, or dark metallic blue; smooth sculpture, scabriculous, with superficially impressed punctures. Lateral lines of pronotal triangle straight. Posterolateral angles widened or slightly acute; projected posteriorly or posterolaterally. Lateral fossae distinctly impressed. Basal fossae obtusely oval, distinctly impressed. Posterior margin sometimes black, with superficially impressed to effaced punctures (
<figureCitation id="13582A61FFD4FFEBFD90406FBDE23844" box="[568,656,612,638]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="42" pageNumber="43">Figs 2J</figureCitation>
, 
<figureCitation id="13582A61FFD4FFEBFD34406FBC6B3845" box="[668,793,612,639]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="42" pageNumber="43">12A–B, D</figureCitation>
).
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF1540ADBB7E393C" blockId="42.[189,1399,677,774]" pageId="42" pageNumber="43">
ELYTRA. Striae fine, smooth, scabriculous, not strongly impressed basally; bright red, green, or dark blue; with superficially impressed punctation. Interstriae black, smooth, scabriculous, with almost effaced to effaced punctures. Sutural margin without apical tooth (
<figureCitation id="13582A61FFD4FFEBFCE340E7BC8D393C" box="[843,1023,748,774]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="42" pageNumber="43">Fig. 12A–B, D</figureCitation>
).
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF154125BDCA397D" blockId="42.[189,696,813,840]" box="[189,696,813,840]" pageId="42" pageNumber="43">PROTIBIAE. Quadridentate with apical spine.</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF154164BCE73996" blockId="42.[189,1398,879,941]" pageId="42" pageNumber="43">TERGITE VIII. Bright metallic red, green, or dark metallic blue; scabriculous; with rough, superficially impressed punctures. Basal margin with setae variable in size.</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF1541DFBDC03E0E" blockId="42.[189,1399,979,1076]" pageId="42" pageNumber="43">
GENITALIA. Right and left lobes of endophallite copulatrix similar in size. Right lobe strongly reduced, obtusely triangular in shape; rounded superiorly. Left lobe obtusely lobed, strongly developed. Central ridge and column similar in size (
<figureCitation id="13582A61FFD4FFEBFDEF4611BDD63E0E" box="[583,676,1050,1076]" captionStart="Fig" captionStartId="8.[189,232,1681,1707]" captionTargetBox="[334,1251,265,1642]" captionTargetId="figure-153@8.[333,1254,265,1655]" captionTargetPageId="8" captionText="Fig. 1. Endophallite copulatrix of the Phanaeus endymion species group (except for P. arletteae Arnaud, 2018). A. Phanaeus bravoensis Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (IEXA). B. Phanaeus chiapanecus sp. nov. (TAMU). C. Phanaeus edmondsi Moctezuma, Deloya &amp; Halffter, 2019 (IEXA). D. Phanaeus endymion Harold, 1863 (VMC). E. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). F. Phanaeus halffterorum Edmonds, 1979 (TAMU). G. Phanaeus huichol Moctezuma, Sánchez- Huerta &amp; Halffter, 2017 (IEXA). H. Phanaeus jackenioi sp. nov. (UVGC). I. Phanaeus malyi Arnaud, 2002 (VMC). J. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). K. Phanaeus pacificus sp. nov. (UVGC). L. Phanaeus panamensis sp. nov. (TAMU). M. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). N. Phanaeus pyrois Bates, 1887 (VMC).O. Phanaeus rzedowskii sp. nov. (TAMU).P. Phanaeus zapotecus Edmonds, 2006 (VMC). Q. Phanaeus zoque Moctezuma &amp; Halffter, 2017 (IEXA)." figureDoi="http://doi.org/10.5281/zenodo.4836748" httpUri="https://zenodo.org/record/4836748/files/figure.png" pageId="42" pageNumber="43">Fig. 1N</figureCitation>
).
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF424650BE0F3E4F" blockId="42.[187,1399,1115,1213]" box="[234,381,1115,1141]" pageId="42" pageNumber="43">
<emphasis id="B917EAF6FFD4FFEBFF424650BE0F3E4F" bold="true" box="[234,381,1115,1141]" pageId="42" pageNumber="43">Minor male</emphasis>
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF154675BC173E87" blockId="42.[187,1399,1115,1213]" pageId="42" pageNumber="43">Like the major male, except for the reduction of secondary sexual characters (i.e., cephalic horn, pronotal triangle and tubercles, and pronotal posterolateral angles).</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF4246E8BE373EC7" blockId="42.[186,1400,1251,1489]" box="[234,325,1251,1277]" pageId="42" pageNumber="43">
<emphasis id="B917EAF6FFD4FFEBFF4246E8BE373EC7" bold="true" box="[234,325,1251,1277]" pageId="42" pageNumber="43">Female</emphasis>
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF15470DBA263FEB" blockId="42.[186,1400,1251,1489]" pageId="42" pageNumber="43">
Similar to the male, except for the head showing a cephalic trituberculate carina; with conical, nearly aligned tubercles; middle tubercle slightly more developed than lateral tubercles; frons with distinctly impressed punctures; pronotal sculpture smooth, with almost effaced punctures; pronotum almost completely black, becoming posteriorly and laterally bright metallic red, green, or dark metallic blue; pronotal process trituberculate, lacking concavity; pronotal tubercles nearly aligned; with middle tubercle more developed than lateral tubercles; posterior pronotal midline superficially impressed (
<figureCitation id="13582A61FFD4FFEBFB7347BDBA353FEB" box="[1243,1351,1462,1489]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="42" pageNumber="43">Fig. 12C</figureCitation>
).
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF4247F3BE123C28" blockId="42.[188,1399,1528,1730]" box="[234,352,1528,1554]" pageId="42" pageNumber="43">
<emphasis id="B917EAF6FFD4FFEBFF4247F3BE123C28" bold="true" box="[234,352,1528,1554]" pageId="42" pageNumber="43">Variation</emphasis>
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF154417BC813CF8" blockId="42.[188,1399,1528,1730]" pageId="42" pageNumber="43">
Mean length 
<quantity id="4C9B9B01FFD4FFEBFEF34410BEB43C0C" box="[347,454,1563,1590]" metricMagnitude="-2" metricUnit="m" metricValue="1.78" pageId="42" pageNumber="43" unit="mm" value="17.8">17.8 mm</quantity>
(
<quantity id="4C9B9B01FFD4FFEBFE7F4410BDFB3C0C" box="[471,649,1563,1590]" metricMagnitude="-2" metricUnit="m" metricValue="1.7399999999999998" metricValueMax="2.01" metricValueMin="1.47" pageId="42" pageNumber="43" unit="mm" value="17.4" valueMax="20.1" valueMin="14.7">14.7–20.1 mm</quantity>
). 
<taxonomicName id="4C634D67FFD4FFEBFD084410BC173C0F" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[672,869,1563,1589]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD4FFEBFD084410BC173C0F" box="[672,869,1563,1589]" italics="true" pageId="42" pageNumber="43">Phanaeus pyrois</emphasis>
</taxonomicName>
is the most variable in colour species of the 
<taxonomicName id="4C634D67FFD4FFEBFF174434BE3C3C62" authorityName="Harold. Ex-Mus" authorityYear="1863" box="[191,334,1598,1624]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="endymion">
<emphasis id="B917EAF6FFD4FFEBFF174434BE3C3C62" box="[191,334,1598,1624]" italics="true" pageId="42" pageNumber="43">P. endymion</emphasis>
</taxonomicName>
species group. The outspoken colour variability of this species was previously outlined by Bates (1886–1889), particularly for the specimens from 
<collectingCountry id="F3747674FFD4FFEBFCE4446ABCBA3C41" box="[844,968,1633,1659]" name="Nicaragua" pageId="42" pageNumber="43">Nicaragua</collectingCountry>
. Tree typical chromatic morphs were found by us (bright metallic red, 
<figureCitation id="13582A61FFD4FFEBFD93448EBDBB3CA5" box="[571,713,1669,1695]" captionStart="Fig" captionStartId="48.[189,232,1877,1903]" captionTargetBox="[228,1363,265,1833]" captionTargetId="figure-40@48.[189,1399,265,1833]" captionTargetPageId="48" captionText="Fig. 14. A–D. Phanaeus zapotecus Edmonds, 2006. A. ♂ (TAMU). B. ♀ (TAMU). C. Minor ♂ (VMC). D. Tergite VIII (TAMU). E. Phanaeus dionysius Kohlmann, Arriaga-Jiménez &amp; Rös, 2018 syn. nov., tergite VIII (IEXA)." figureDoi="http://doi.org/10.5281/zenodo.4836794" httpUri="https://zenodo.org/record/4836794/files/figure.png" pageId="42" pageNumber="43">Fig. 14A, D</figureCitation>
; green, 
<figureCitation id="13582A61FFD4FFEBFC8C448EBC043CA5" box="[804,886,1669,1695]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="42" pageNumber="43">Fig. 2J</figureCitation>
; or dark metallic blue, 
<figureCitation id="13582A61FFD4FFEBFB29448EBB983CA5" box="[1153,1258,1669,1695]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="42" pageNumber="43">Fig. 12B</figureCitation>
), but colour combinations are found and rare specimens has a bright golden sheen.
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFD4FFEBFF1544E2BB6A3D6F" pageId="42" pageNumber="43" type="distribution">
<paragraph id="8BDC36E4FFD4FFEBFF1544E2BE243D39" blockId="42.[189,342,1769,1795]" box="[189,342,1769,1795]" pageId="42" pageNumber="43">
<heading id="D0948188FFD4FFEBFF1544E2BE243D39" bold="true" box="[189,342,1769,1795]" fontSize="11" level="3" pageId="42" pageNumber="43" reason="3">
<emphasis id="B917EAF6FFD4FFEBFF1544E2BE243D39" bold="true" box="[189,342,1769,1795]" pageId="42" pageNumber="43">Distribution</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF154513BB6A3D6F" blockId="42.[189,1399,1816,1878]" pageId="42" pageNumber="43">
<collectingCountry id="F3747674FFD4FFEBFF154513BE443D08" box="[189,310,1816,1842]" name="Nicaragua" pageId="42" pageNumber="43">Nicaragua</collectingCountry>
and north-Caribbean 
<collectingCountry id="F3747674FFD4FFEBFD924513BDC93D08" box="[570,699,1816,1842]" name="Costa Rica" pageId="42" pageNumber="43">Costa Rica</collectingCountry>
(
<figureCitation id="13582A61FFD4FFEBFD644513BC573D09" box="[716,805,1816,1843]" captionStart="Fig" captionStartId="51.[189,232,1233,1259]" captionTargetBox="[193,1395,270,1197]" captionTargetId="figure-241@51.[189,1398,265,1199]" captionTargetPageId="51" captionText="Fig. 16. Distribution of the Phanaeus endymion species group in Central America (in part)." figureDoi="http://doi.org/10.5281/zenodo.4836798" httpUri="https://zenodo.org/record/4836798/files/figure.png" pageId="42" pageNumber="43">Fig. 16</figureCitation>
). The distributions of 
<taxonomicName id="4C634D67FFD4FFEBFB864512BBE03D08" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[1070,1170,1817,1842]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD4FFEBFB864512BBE03D08" box="[1070,1170,1817,1842]" italics="true" pageId="42" pageNumber="43">P. pyrois</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C634D67FFD4FFEBFB674512BA053D08" authority="Moctezuma &amp; Halffter, 2021" authorityName="Moctezuma &amp; Halffter" authorityYear="2021" box="[1231,1399,1817,1842]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="panamensis" status="sp. nov.">
<emphasis id="B917EAF6FFD4FFEBFB674512BA053D08" box="[1231,1399,1817,1842]" italics="true" pageId="42" pageNumber="43">P. panamensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A224578DFFD4FFEBFF154537BE6A3D6C" box="[189,280,1852,1878]" pageId="42" pageNumber="43" rank="species">sp. nov.</taxonomicNameLabel>
show an important sympatry area in north Caribbean 
<collectingCountry id="F3747674FFD4FFEBFC384530BB663D6F" box="[912,1044,1851,1877]" name="Costa Rica" pageId="42" pageNumber="43">Costa Rica</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="C379656FFFD4FFEDFF154576BE123858" lastPageId="44" lastPageNumber="45" pageId="42" pageNumber="43" type="discussion">
<paragraph id="8BDC36E4FFD4FFEBFF154576BE5D3DAD" blockId="42.[189,303,1917,1943]" box="[189,303,1917,1943]" pageId="42" pageNumber="43">
<heading id="D0948188FFD4FFEBFF154576BE5D3DAD" bold="true" box="[189,303,1917,1943]" fontSize="11" level="3" pageId="42" pageNumber="43" reason="3">
<emphasis id="B917EAF6FFD4FFEBFF154576BE5D3DAD" bold="true" box="[189,303,1917,1943]" pageId="42" pageNumber="43">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="8BDC36E4FFD4FFEBFF1545A0BA063DD2" blockId="42.[187,1398,1963,2025]" pageId="42" pageNumber="43">
<taxonomicName id="4C634D67FFD4FFEBFF1545A0BE0C3DFF" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[189,382,1963,1989]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="42" pageNumber="43" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD4FFEBFF1545A0BE0C3DFF" box="[189,382,1963,1989]" italics="true" pageId="42" pageNumber="43">Phanaeus pyrois</emphasis>
</taxonomicName>
and several closely related species were incorrectly lumped together by previous authors (
<bibRefCitation id="EFF24B15FFD4FFEBFF6A45C5BEAF3DD2" author="Howden H. F. &amp; Young O. P." box="[194,477,1998,2025]" pageId="42" pageNumber="43" pagination="1 - 204" refId="ref39249" refString="Howden H. F. &amp; Young O. P. 1981. Panamanian Scarabaeinae: Taxonomy, distribution, and habits (Coleoptera, Scarabaeidae). Contributions of the American Entomological Institute 18: 1 - 204." type="journal article" year="1981">Howden &amp; Young 1981</bibRefCitation>
; 
<bibRefCitation id="EFF24B15FFD4FFEBFE4145C5BDEF3DD2" author="Edmonds W. D." box="[489,669,1998,2024]" pageId="42" pageNumber="43" 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="EFF24B15FFD4FFEBFD0145C5BCB73DD2" author="Edmonds W. D. &amp; Zidek J." box="[681,965,1998,2024]" pageId="42" pageNumber="43" 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>
; 
<bibRefCitation id="EFF24B15FFD4FFEBFC7945C5BB833DD2" author="Solis A. &amp; Kohlmann B." box="[977,1265,1998,2024]" pageId="42" pageNumber="43" pagination="1 - 32" refId="ref42322" refString="Solis A. &amp; Kohlmann B. 2012. Checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica. Zootaxa 3482: 1 - 32." type="journal article" year="2012">Solís &amp; Kohlmann 2012</bibRefCitation>
; Chamorro
</paragraph>
<caption id="DF1C666CFFD5FFEAFF1545A1BA053DD2" ID-DOI="http://doi.org/10.5281/zenodo.4836790" ID-Zenodo-Dep="4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="43" pageNumber="44" startId="43.[189,232,1962,1988]" targetBox="[319,1279,265,1926]" targetPageId="43">
<paragraph id="8BDC36E4FFD5FFEAFF1545A1BA053DD2" blockId="43.[189,1399,1962,2025]" pageId="43" pageNumber="44">
<emphasis id="B917EAF6FFD5FFEAFF1545A1BE6B3DFE" bold="true" box="[189,281,1962,1988]" pageId="43" pageNumber="44">Fig. 12.</emphasis>
<taxonomicName id="4C634D67FFD5FFEAFE8845A0BD0B3DFF" authority="Bates, 1887" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[288,633,1963,1989]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="43" pageNumber="44" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD5FFEAFE8845A0BE963DFE" box="[288,484,1963,1989]" italics="true" pageId="43" pageNumber="44">Phanaeus pyrois</emphasis>
<bibRefCitation id="EFF24B15FFD5FFEAFE4345A0BD0B3DFF" author="Bates H. W." box="[491,633,1963,1989]" pageId="43" pageNumber="44" refId="ref36135" refString="Bates H. W. 1887. Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor &amp; Francis, London." type="book" year="1887">Bates, 1887</bibRefCitation>
</taxonomicName>
. 
<emphasis id="B917EAF6FFD5FFEAFD2C45A1BDEB3DFE" bold="true" box="[644,665,1962,1988]" pageId="43" pageNumber="44">A</emphasis>
. ♂, red morph (VCM). 
<emphasis id="B917EAF6FFD5FFEAFC1B45A0BCB53DFF" bold="true" box="[947,967,1963,1989]" pageId="43" pageNumber="44">B</emphasis>
. ♂, blue morph (VMC). 
<emphasis id="B917EAF6FFD5FFEAFB4545A1BA703DFE" bold="true" box="[1261,1282,1962,1988]" pageId="43" pageNumber="44">C</emphasis>
. ♀, green morph (TAMU). 
<emphasis id="B917EAF6FFD5FFEAFE2B45C5BEEA3DD2" bold="true" box="[387,408,1998,2024]" pageId="43" pageNumber="44">D</emphasis>
. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH).
</paragraph>
</caption>
<paragraph id="8BDC36E4FFD2FFEDFF154301BE123858" blockId="44.[188,1399,265,610]" pageId="44" pageNumber="45">
<emphasis id="B917EAF6FFD2FFEDFF154301BF813B19" box="[189,243,265,291]" italics="true" pageId="44" pageNumber="45">et al</emphasis>
. 2018, 2019; 
<bibRefCitation id="EFF24B15FFD2FFEDFE354302BDCD3B1E" author="GBIF Secretariat" box="[413,703,265,292]" pageId="44" pageNumber="45" refId="ref38117" refString="GBIF Secretariat. 2019 b. Phanaeus pyrois Bates, 1887. GBIF Backbone Taxonomy. Checklist dataset. Available from https: // doi. org / 10.15468 / 39 omei [accessed Jul. 2020]." type="book" year="2019">GBIF Secretariat 2019b</bibRefCitation>
). Differences in body colour and the pronotal, elytral and genital morphology were found to confidently diagnose 
<taxonomicName id="4C634D67FFD2FFEDFCE44326BCC23B7C" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[844,944,301,326]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD2FFEDFCE44326BCC23B7C" box="[844,944,301,326]" italics="true" pageId="44" pageNumber="45">P. pyrois</emphasis>
</taxonomicName>
and all the closely related species. The blue chromatic morph of 
<taxonomicName id="4C634D67FFD2FFEDFE5A435ABD2A3B50" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[498,600,337,362]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD2FFEDFE5A435ABD2A3B50" box="[498,600,337,362]" italics="true" pageId="44" pageNumber="45">P. pyrois</emphasis>
</taxonomicName>
(
<figureCitation id="13582A61FFD2FFEDFDC1435BBDA63B50" box="[617,724,336,362]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="44" pageNumber="45">Fig. 12B</figureCitation>
) was suggested by 
<bibRefCitation id="EFF24B15FFD2FFEDFC69435BBBF53B50" author="Edmonds W. D." box="[961,1159,336,362]" pageId="44" pageNumber="45" 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>
to be a hybrid with 
<taxonomicName id="4C634D67FFD2FFEDFF17437FBE393BB7" authorityName="Harold. Ex-Mus" authorityYear="1863" box="[191,331,371,397]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="endymion">
<emphasis id="B917EAF6FFD2FFEDFF17437FBE393BB7" box="[191,331,371,397]" italics="true" pageId="44" pageNumber="45">P. endymion</emphasis>
</taxonomicName>
. Nevertheless, blue specimens of 
<taxonomicName id="4C634D67FFD2FFEDFD7E437FBC483BB7" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[726,826,372,397]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD2FFEDFD7E437FBC483BB7" box="[726,826,372,397]" italics="true" pageId="44" pageNumber="45">P. pyrois</emphasis>
</taxonomicName>
(
<figureCitation id="13582A61FFD2FFEDFCE34378BCC03BB7" box="[843,946,371,397]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="44" pageNumber="45">Fig. 12B</figureCitation>
) do not share the diagnostic characters with 
<taxonomicName id="4C634D67FFD2FFEDFF51439CBEFB3B8B" authorityName="Harold. Ex-Mus" authorityYear="1863" box="[249,393,407,433]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="endymion">
<emphasis id="B917EAF6FFD2FFEDFF51439CBEFB3B8B" box="[249,393,407,433]" italics="true" pageId="44" pageNumber="45">P. endymion</emphasis>
</taxonomicName>
(
<figureCitation id="13582A61FFD2FFEDFE32439CBE893B8B" box="[410,507,407,433]" captionStart="Fig" captionStartId="8.[189,232,1681,1707]" captionTargetBox="[334,1251,265,1642]" captionTargetId="figure-153@8.[333,1254,265,1655]" captionTargetPageId="8" captionText="Fig. 1. Endophallite copulatrix of the Phanaeus endymion species group (except for P. arletteae Arnaud, 2018). A. Phanaeus bravoensis Moctezuma, Sánchez-Huerta &amp; Halffter, 2017 (IEXA). B. Phanaeus chiapanecus sp. nov. (TAMU). C. Phanaeus edmondsi Moctezuma, Deloya &amp; Halffter, 2019 (IEXA). D. Phanaeus endymion Harold, 1863 (VMC). E. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). F. Phanaeus halffterorum Edmonds, 1979 (TAMU). G. Phanaeus huichol Moctezuma, Sánchez- Huerta &amp; Halffter, 2017 (IEXA). H. Phanaeus jackenioi sp. nov. (UVGC). I. Phanaeus malyi Arnaud, 2002 (VMC). J. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). K. Phanaeus pacificus sp. nov. (UVGC). L. Phanaeus panamensis sp. nov. (TAMU). M. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). N. Phanaeus pyrois Bates, 1887 (VMC).O. Phanaeus rzedowskii sp. nov. (TAMU).P. Phanaeus zapotecus Edmonds, 2006 (VMC). Q. Phanaeus zoque Moctezuma &amp; Halffter, 2017 (IEXA)." figureDoi="http://doi.org/10.5281/zenodo.4836748" httpUri="https://zenodo.org/record/4836748/files/figure.png" pageId="44" pageNumber="45">Figs 1D</figureCitation>
, 
<figureCitation id="13582A61FFD2FFEDFDA1439CBD5C3B8B" box="[521,558,407,433]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="44" pageNumber="45">2B</figureCitation>
, 
<figureCitation id="13582A61FFD2FFEDFD93439CBD3F3B8B" box="[571,589,407,433]" captionStart="Fig" captionStartId="13.[189,232,1925,1951]" captionTargetBox="[218,1398,491,1881]" captionTargetId="figure-72@13.[189,1398,491,1889]" captionTargetPageId="13" captionText="Fig. 4. Phanaeus endymion Harold, 1863. A. ♂, blue morph (typical; VMC). B. ♂, green morph (VMC). C. ♀, blue morph (typical; VMC). D. Lectotype, ♂ (by Christophe Rivier, Muséum national d’histoire naturelle, Paris, France) (MNHN)." figureDoi="http://doi.org/10.5281/zenodo.4836760" httpUri="https://zenodo.org/record/4836760/files/figure.png" pageId="44" pageNumber="45">4</figureCitation>
). As a consequence, there is no evidence to consider a hybridization between 
<taxonomicName id="4C634D67FFD2FFEDFE8143B0BEC93BEE" authorityName="Harold. Ex-Mus" authorityYear="1863" box="[297,443,442,468]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="endymion">
<emphasis id="B917EAF6FFD2FFEDFE8143B0BEC93BEE" box="[297,443,442,468]" italics="true" pageId="44" pageNumber="45">P. endymion</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C634D67FFD2FFEDFE5343B0BD113BEE" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[507,611,443,468]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD2FFEDFE5343B0BD113BEE" box="[507,611,443,468]" italics="true" pageId="44" pageNumber="45">P. pyrois</emphasis>
</taxonomicName>
. 
<bibRefCitation id="EFF24B15FFD2FFEDFDDD43B1BCDE3BEE" author="Edmonds W. D. &amp; Zidek J." box="[629,940,442,468]" pageId="44" pageNumber="45" 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>
suggested that doubtful specimens of “ 
<taxonomicName id="4C634D67FFD2FFEDFF6243D6BE3B3BCD" authorityName="Olsoufieff" authorityYear="1924" box="[202,329,477,503]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="viridicollis">
<emphasis id="B917EAF6FFD2FFEDFF6243D6BE3B3BCD" box="[202,329,477,503]" italics="true" pageId="44" pageNumber="45">viridicollis</emphasis>
</taxonomicName>
” (
<figureCitation id="13582A61FFD2FFEDFECF43D6BEB23BC2" box="[359,448,477,504]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="44" pageNumber="45">Figs 2J</figureCitation>
, 
<figureCitation id="13582A61FFD2FFEDFE6543D5BE8D3BC2" box="[461,511,478,504]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="44" pageNumber="45">12C</figureCitation>
) were collected in 
<collectingCountry id="F3747674FFD2FFEDFD4A43D6BC293BCD" box="[738,859,477,503]" name="Nicaragua" pageId="44" pageNumber="45">Nicaragua</collectingCountry>
along with “normal” 
<taxonomicName id="4C634D67FFD2FFEDFBF643D5BBB63BCD" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[1118,1220,478,503]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD2FFEDFBF643D5BBB63BCD" box="[1118,1220,478,503]" italics="true" pageId="44" pageNumber="45">P. pyrois</emphasis>
</taxonomicName>
. After revising the doubtful specimens of “ 
<taxonomicName id="4C634D67FFD2FFEDFDAE400ABDF73821" authorityName="Olsoufieff" authorityYear="1924" box="[518,645,513,539]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="viridicollis">
<emphasis id="B917EAF6FFD2FFEDFDAE400ABDF73821" box="[518,645,513,539]" italics="true" pageId="44" pageNumber="45">viridicollis</emphasis>
</taxonomicName>
” from 
<collectingCountry id="F3747674FFD2FFEDFD74400ABC273821" box="[732,853,513,539]" name="Nicaragua" pageId="44" pageNumber="45">Nicaragua</collectingCountry>
(
<figureCitation id="13582A61FFD2FFEDFCCD400ABCCD3821" box="[869,959,513,539]" captionStart="Fig" captionStartId="9.[189,232,1786,1812]" captionTargetBox="[260,1330,265,1758]" captionTargetId="figure-104@9.[257,1330,265,1765]" captionTargetPageId="9" captionText="Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC)." figureDoi="http://doi.org/10.5281/zenodo.4836752" httpUri="https://zenodo.org/record/4836752/files/figure.png" pageId="44" pageNumber="45">Figs 2J</figureCitation>
, 
<figureCitation id="13582A61FFD2FFEDFC64400ABB723821" box="[972,1024,513,539]" captionStart="Fig" captionStartId="43.[189,232,1962,1988]" captionTargetBox="[319,1279,265,1926]" captionTargetId="figure-38@43.[306,1281,265,1926]" captionTargetPageId="43" captionText="Fig. 12. Phanaeus pyrois Bates, 1887. A. ♂, red morph (VCM). B. ♂, blue morph (VMC). C. ♀, green morph (TAMU). D. Lectotype, ♂ (by Mario Cupello, Universidade Federal do Paraná, Brazil) (BMNH)." figureDoi="http://doi.org/10.5281/zenodo.4836790" httpUri="https://zenodo.org/record/4836790/files/figure.png" pageId="44" pageNumber="45">12C</figureCitation>
), we disagree with 
<bibRefCitation id="EFF24B15FFD2FFEDFB40400ABE2A3805" author="Edmonds W. D. &amp; Zidek J." pageId="44" pageNumber="45" 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>
and conclude that they incorrectly referred to the green chromatic morph of 
<taxonomicName id="4C634D67FFD2FFEDFB58402EBA243804" authorityName="Bates, Det. W. D. Edmonds" authorityYear="1887" box="[1264,1366,549,574]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="B917EAF6FFD2FFEDFB58402EBA243804" box="[1264,1366,549,574]" italics="true" pageId="44" pageNumber="45">P. pyrois</emphasis>
</taxonomicName>
as 
<taxonomicName id="4C634D67FFD2FFEDFF164042BE2B3858" authorityName="Olsoufieff" authorityYear="1924" box="[190,345,584,610]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="44" pageNumber="45" phylum="Arthropoda" rank="species" species="viridicollis">
<emphasis id="B917EAF6FFD2FFEDFF164042BE2B3858" box="[190,345,584,610]" italics="true" pageId="44" pageNumber="45">P. viridicollis</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>