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<mods:title id="4B8FEF654007E5EE6AFA313D4040D6E5">The Phanaeus tridens species group (Coleoptera: Scarabaeoidea): a dung beetle group with genital morphological stasis but a changing ecological niche</mods:title>
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<mods:namePart id="04771F6AF99DDAEA19D4413F5939ECE5">Halffter, Gonzalo</mods:namePart>
<mods:affiliation id="BB6425F023411C5E50080C1D78F43555">Emeritus Researcher, Instituto de Ecología A. C., Xalapa, Veracruz 91073, Mexico, Carretera antigua a Coatepec 351, El Haya.</mods:affiliation>
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<mods:namePart id="5E1177EBD691CBF650D96AA90F1FB687">Lizardo, Viridiana</mods:namePart>
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<treatment id="1B2B878A8A20FFB1FC57FDFFEEF9FEE7" ID-DOI="http://doi.org/10.5281/zenodo.5821249" ID-GBIF-Taxon="193278669" ID-Zenodo-Dep="5821249" LSID="urn:lsid:plazi:treatment:1B2B878A8A20FFB1FC57FDFFEEF9FEE7" httpUri="http://treatment.plazi.org/id/1B2B878A8A20FFB1FC57FDFFEEF9FEE7" lastPageId="12" lastPageNumber="459" pageId="10" pageNumber="457">
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<paragraph id="933D369C8A20FFB7FC57FDFFEA70FDDE" blockId="10.[898,1366,597,620]" box="[898,1366,597,620]" pageId="10" pageNumber="457">
<heading id="C87581F08A20FFB7FC57FDFFEA70FDDE" box="[898,1366,597,620]" centered="true" fontSize="10" level="2" pageId="10" pageNumber="457" reason="3">
<emphasis id="A1F6EA8E8A20FFB7FC57FDFFEA70FDDE" bold="true" box="[898,1366,597,620]" pageId="10" pageNumber="457">
<taxonomicName id="54824D1F8A20FFB7FC57FDFFEBCCFDDE" ID-CoL="8TJQP" authority="Bates, 1887" authorityName="Bates" authorityYear="1887" box="[898,1258,597,620]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="457" phylum="Arthropoda" rank="species" species="coeruleus" status="stat. nov.">
<emphasis id="A1F6EA8E8A20FFB7FC57FDFFEB47FDDE" bold="true" box="[898,1121,597,620]" italics="true" pageId="10" pageNumber="457">Phanaeus coeruleus</emphasis>
<bibRefCitation id="F7134B6D8A20FFB7FBB2FDFFEBCCFDDE" author="Bates" box="[1127,1258,597,620]" firstAuthor="Bates" pageId="10" pageNumber="457" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">Bates, 1887</bibRefCitation>
</taxonomicName>
,
<taxonomicNameLabel id="BAC557F58A20FFB7FB23FDFFEA70FDDE" box="[1270,1366,597,620]" pageId="10" pageNumber="457">stat. rev.</taxonomicNameLabel>
</emphasis>
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</paragraph>
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<subSubSection id="DB9865178A20FFB7FBE0FDD9EB85FD34" box="[1077,1187,627,646]" pageId="10" pageNumber="457" type="description">
<paragraph id="933D369C8A20FFB7FBE0FDD9EB85FD34" blockId="10.[828,1437,627,707]" box="[1077,1187,627,646]" pageId="10" pageNumber="457">
(
<figureCitation id="0BB92A198A20FFB7FBEEFDD9EB5BFD34" box="[1083,1149,627,646]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A20FFB7FB53FDD9EBBAFD34" box="[1158,1180,627,646]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">47</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A20FFB7FCE9FD3FECBDFD71" pageId="10" pageNumber="457" type="reference_group">
<paragraph id="933D369C8A20FFB7FCE9FD3FECBDFD71" blockId="10.[828,1437,627,707]" pageId="10" pageNumber="457">
<treatmentCitationGroup id="B39211B28A20FFB7FCE9FD3FECBDFD71" pageId="10" pageNumber="457">
<treatmentCitation id="1223108D8A20FFB7FCE9FD3FEBFFFD1A" author="BATES H. W." box="[828,1241,661,680]" page="61" pageId="10" pageNumber="457" year="1887">
<taxonomicName id="54824D1F8A20FFB7FCE9FD3FEBFFFD1A" ID-CoL="8TJQP" authority="Bates, 1887: 61" authorityName="Bates" authorityPageNumber="61" authorityYear="1887" box="[828,1241,661,680]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="457" phylum="Arthropoda" rank="variety" species="daphnis" variety="coeruleus">
<emphasis id="A1F6EA8E8A20FFB7FCE9FD3FECF0FD1A" box="[828,982,661,680]" italics="true" pageId="10" pageNumber="457">Phanaeus daphnis</emphasis>
var.
<emphasis id="A1F6EA8E8A20FFB7FC2BFD3FEB77FD1A" box="[1022,1105,661,680]" italics="true" pageId="10" pageNumber="457">coeruleus</emphasis>
<bibRefCitation id="F7134B6D8A20FFB7FB81FD3FEBFFFD1A" author="Bates" box="[1108,1241,661,680]" firstAuthor="Bates" page="61" pageId="10" pageNumber="457" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">Bates, 1887: 61</bibRefCitation>
</taxonomicName>
</treatmentCitation>
.
<typeStatus id="4C39883E8A20FFB7FB0AFD3FEA2FFD1A" box="[1247,1289,661,680]" pageId="10" pageNumber="457">Type</typeStatus>
locality:
<collectingCountry id="EB95760C8A20FFB7FA83FD3FEABFFD1A" box="[1366,1433,661,680]" name="Mexico" pageId="10" pageNumber="457">Mexico</collectingCountry>
,
<collectingRegion id="5146F87E8A20FFB7FC8FFD1AECB1FD71" box="[858,919,688,707]" country="Mexico" name="Puebla" pageId="10" pageNumber="457">Puebla</collectingRegion>
.
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A20FFB7FCE9FD70EB74FC90" pageId="10" pageNumber="457" type="materials_examined">
<paragraph id="933D369C8A20FFB7FCE9FD70EB74FC90" blockId="10.[828,1437,730,802]" pageId="10" pageNumber="457">
<emphasis id="A1F6EA8E8A20FFB7FCE9FD70EBAFFD5F" bold="true" box="[828,1161,730,749]" pageId="10" pageNumber="457">
<typeStatus id="4C39883E8A20FFB7FCE9FD70EC4FFD5F" box="[828,873,730,749]" pageId="10" pageNumber="457">Type</typeStatus>
material examined.
<collectingCountry id="EB95760C8A20FFB7FBF2FD70EBA2FD5F" box="[1063,1156,730,749]" name="Mexico" pageId="10" pageNumber="457">MEXICO</collectingCountry>
:
</emphasis>
HOLOTYPE (fixed by monotypy by
<bibRefCitation id="F7134B6D8A20FFB7FC8CFD5EECE6FCB5" author="BATES" box="[857,960,756,775]" firstAuthor="BATES" pageId="10" pageNumber="457" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">BATES 1887</bibRefCitation>
, examined from photographs;
<figureCitation id="0BB92A198A20FFB7FB18FD5EEA36FCB5" box="[1229,1296,756,775]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A20FFB7FAC9FD5EEA13FCB5" box="[1308,1333,756,775]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">47</figureCitation>
): ♂,
<collectingRegion id="5146F87E8A20FFB7FAB7FD5EEABAFCB5" box="[1378,1436,756,775]" country="Mexico" name="Puebla" pageId="10" pageNumber="457">Puebla</collectingRegion>
(
<collectionCode id="F593AE598A20FFB7FC97FCA5ECA0FC90" box="[834,902,783,802]" pageId="10" pageNumber="457">BMNH</collectionCode>
: NHMUK 013667754).
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A20FFB7FCE9FC91EA20FB60" pageId="10" pageNumber="457" type="diagnosis">
<paragraph id="933D369C8A20FFB7FCE9FC91EA20FB60" blockId="10.[828,1437,827,1234]" pageId="10" pageNumber="457">
<emphasis id="A1F6EA8E8A20FFB7FCE9FC91EC96FCE0" bold="true" box="[828,944,827,850]" pageId="10" pageNumber="457">Diagnosis.</emphasis>
Metallic dark blue with turquoise sheen (
<figureCitation id="0BB92A198A20FFB7FAB8FC91EC7FFCC0" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A20FFB7FCB1FCF1ECA4FCC0" box="[868,898,859,882]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">47</figureCitation>
). Sides of pronotal disc granulate anteriorly, becoming punctate with effaced granules posteriorly. Pronotal disc coarsely rugose (
<figureCitation id="0BB92A198A20FFB7FBF5FC31EB55FC00" box="[1056,1139,923,946]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A20FFB7FBA8FC31EBBDFC00" box="[1149,1179,923,946]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">47</figureCitation>
). Posteromedial process of pronotum produced into short denticle, not bifurcated but truncate apically (
<figureCitation id="0BB92A198A20FFB7FBFEFC71EB58FC40" box="[1067,1150,987,1010]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">Fig. 47</figureCitation>
). Anterolateral margins of pronotal disc without ridge or line of tubercles (
<figureCitation id="0BB92A198A20FFB7FAEAFC51EAB6FBA0" box="[1343,1424,1019,1042]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">Fig. 47</figureCitation>
). Posterolateral angles of pronotum much longer than posteromedial process (
<figureCitation id="0BB92A198A20FFB7FBC2FB91EB4EFBE0" box="[1047,1128,1083,1106]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="10" pageNumber="457">Fig. 47</figureCitation>
). Elytral striae scabriculous, distinctly impressed, superficially punctate (
<figureCitation id="0BB92A198A20FFB7FAECFBF1EAA8FBC0" box="[1337,1422,1115,1138]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457">Fig. 13</figureCitation>
). Elytral interstriae scabriculous, distinctly smooth (except for interstriae VI and VII with lightly roughened integument), superficially punctate, flat (
<figureCitation id="0BB92A198A20FFB7FB7FFB11EBDFFB60" box="[1194,1273,1211,1234]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457">Fig. 13</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="C7FD66148A20FFB7FF42F7A7EB2AF792" ID-DOI="http://doi.org/10.5281/zenodo.5976087" ID-Zenodo-Dep="5976087" box="[151,1036,2061,2080]" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="10" pageNumber="457" startId="10.[151,182,2061,2080]" targetBox="[329,1263,1275,2048]" targetPageId="10">
<paragraph id="933D369C8A20FFB7FF42F7A7EB2AF792" blockId="10.[151,1036,2061,2080]" box="[151,1036,2061,2080]" pageId="10" pageNumber="457">
Fig. 13.
<taxonomicName id="54824D1F8A20FFB7FF0BF7A7EED1F792" authority="Bates, 1887" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[222,503,2061,2080]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="457" phylum="Arthropoda" rank="species" species="coeruleus" status="stat. nov.">
<emphasis id="A1F6EA8E8A20FFB7FF0BF7A7EEADF792" box="[222,395,2061,2080]" italics="true" pageId="10" pageNumber="457">Phanaeus coeruleus</emphasis>
<bibRefCitation id="F7134B6D8A20FFB7FE45F7A7EED1F792" author="Bates" box="[400,503,2061,2080]" firstAuthor="Bates" pageId="10" pageNumber="457" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">Bates, 1887</bibRefCitation>
</taxonomicName>
,
<taxonomicNameLabel id="BAC557F58A20FFB7FE2AF7A7ED6CF792" box="[511,586,2061,2080]" pageId="10" pageNumber="457">stat. rev.</taxonomicNameLabel>
, holotype and labels (by Keita Matsumoto, BMNH).
</paragraph>
</caption>
<subSubSection id="DB9865178A21FFB1FF42FF14EEF9FEE7" lastPageId="12" lastPageNumber="459" pageId="11" pageNumber="458" type="description">
<paragraph id="933D369C8A21FFB6FF42FF14EDC0FF67" blockId="11.[151,742,190,213]" box="[151,742,190,213]" pageId="11" pageNumber="458">
<heading id="C87581F08A21FFB6FF42FF14EDC0FF67" box="[151,742,190,213]" centered="true" fontSize="36" level="2" pageId="11" pageNumber="458" reason="4">
<emphasis id="A1F6EA8E8A21FFB6FF42FF14EEBEFF67" bold="true" box="[151,408,190,213]" pageId="11" pageNumber="458">
Variation.
<emphasis id="A1F6EA8E8A21FFB6FEC5FF14EEBEFF67" bold="true" box="[272,408,190,213]" italics="true" pageId="11" pageNumber="458">
<taxonomicName id="54824D1F8A21FFB6FEC5FF14EE70FF67" authorityName="Greenslade P &amp; Greenslade Pjm" authorityYear="1987" box="[272,342,190,213]" class="Insecta" family="Scarabaeidae" genus="Minor" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="genus">Minor</taxonomicName>
male.
</emphasis>
</emphasis>
Unknown.
<emphasis id="A1F6EA8E8A21FFB6FDC0FF14ED49FF67" bold="true" box="[533,623,190,213]" italics="true" pageId="11" pageNumber="458">Female.</emphasis>
Unknown.
</heading>
</paragraph>
<paragraph id="933D369C8A21FFB6FF42FF74EDA3FD67" blockId="11.[151,760,222,1013]" pageId="11" pageNumber="458">
<emphasis id="A1F6EA8E8A21FFB6FF42FF74EE3EFF47" bold="true" box="[151,280,222,245]" pageId="11" pageNumber="458">Comments.</emphasis>
<bibRefCitation id="F7134B6D8A21FFB6FECEFF74ED24FF47" author="ARNAUD" box="[283,514,222,245]" firstAuthor="ARNAUD" page="113" pageId="11" pageNumber="458" pagination="113 - 118" refId="ref24103" refString="ARNAUD P. 1982 a: Liste des types de Phanaeini du Museum National d'Histoire Naturelle de Paris [Coleoptera, Scarabaeidae]. Revue Francaise d'Entomologie (Nouvelle S erie) 4 (3): 113 - 118." type="journal article" year="1982" yearSuffix="a">ARNAUD (1982a: 113)</bibRefCitation>
collectively designated
<typeStatus id="4C39883E8A21FFB6FF42FF54EE23FEA7" box="[151,261,254,277]" pageId="11" pageNumber="458" type="lectotype">lectotypes</typeStatus>
for some species-group names of
<taxonomicName id="54824D1F8A21FFB6FD58FF54EDD1FEA7" authorityName="MacLeay" authorityYear="1819" box="[653,759,254,277]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="A1F6EA8E8A21FFB6FD58FF54EDD1FEA7" box="[653,759,254,277]" italics="true" pageId="11" pageNumber="458">Phanaeus</emphasis>
</taxonomicName>
housed at BMNH. More specifically, instead of pointing out which individual
<typeStatus id="4C39883E8A21FFB6FEACFE94EEEAFEE7" box="[377,460,318,341]" pageId="11" pageNumber="458" type="syntype">syntype</typeStatus>
was being designated as the
<typeStatus id="4C39883E8A21FFB6FF42FEF4EFDFFEC7" box="[151,249,350,373]" pageId="11" pageNumber="458" type="lectotype">lectotype</typeStatus>
of each name,
<bibRefCitation id="F7134B6D8A21FFB6FE4AFEF4ED71FEC7" author="ARNAUD" box="[415,599,350,373]" firstAuthor="ARNAUD" pageId="11" pageNumber="458" pagination="113 - 118" refId="ref24103" refString="ARNAUD P. 1982 a: Liste des types de Phanaeini du Museum National d'Histoire Naturelle de Paris [Coleoptera, Scarabaeidae]. Revue Francaise d'Entomologie (Nouvelle S erie) 4 (3): 113 - 118." type="journal article" year="1982" yearSuffix="a">ARNAUD (1982a)</bibRefCitation>
simply said in a general statement that he considered
<typeStatus id="4C39883E8A21FFB6FDEBFED4EDBDFE27" box="[574,667,382,405]" pageId="11" pageNumber="458" type="syntype">syntypes</typeStatus>
labelled as “sp. figured” in the BMNH to be
<typeStatus id="4C39883E8A21FFB6FDF4FE34EDABFE07" box="[545,653,414,437]" pageId="11" pageNumber="458" type="lectotype">lectotypes</typeStatus>
of names established by
<bibRefCitation id="F7134B6D8A21FFB6FEEFFE14EEEAFE67" author="BATES" box="[314,460,446,469]" firstAuthor="BATES" pageId="11" pageNumber="458" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">BATES (1887)</bibRefCitation>
. Nevertheless, as explained by
<bibRefCitation id="F7134B6D8A21FFB6FF62FE74EE4BFE47" author="EDMONDS" box="[183,365,478,501]" firstAuthor="EDMONDS" pageId="11" pageNumber="458" pagination="1 - 105" refId="ref25450" refString="EDMONDS W. D. 1994: Revision of Phanaeus MacLeay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science of the Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">EDMONDS (1994)</bibRefCitation>
, the International Code on Zoological Nomenclature prohibits collective designation of
<typeStatus id="4C39883E8A21FFB6FD73FE54EF9DFD87" pageId="11" pageNumber="458" type="lectotype">lectotypes</typeStatus>
like
<bibRefCitation id="F7134B6D8A21FFB6FF25FDB4EE8EFD87" author="ARNAUD" box="[240,424,542,565]" firstAuthor="ARNAUD" pageId="11" pageNumber="458" pagination="113 - 118" refId="ref24103" refString="ARNAUD P. 1982 a: Liste des types de Phanaeini du Museum National d'Histoire Naturelle de Paris [Coleoptera, Scarabaeidae]. Revue Francaise d'Entomologie (Nouvelle S erie) 4 (3): 113 - 118." type="journal article" year="1982" yearSuffix="a">ARNAUD (1982a)</bibRefCitation>
did. Indeed, the designation of
<typeStatus id="4C39883E8A21FFB6FF42FD94EE25FDE7" box="[151,259,574,597]" pageId="11" pageNumber="458" type="lectotype">lectotypes</typeStatus>
must be individual under Article 74.3 (
<bibRefCitation id="F7134B6D8A21FFB6FD6DFD94EFF7FDC7" author="ICZN" firstAuthor="ICZN" pageId="11" pageNumber="458" refId="ref26459" refString="ICZN [= INTERNATIONAL COMMISION ON ZOOLOGICAL NOMENCLATURE] 1999: International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature, London, 306 pp." type="book" year="1999">ICZN 1999</bibRefCitation>
). Consequently, all the
<typeStatus id="4C39883E8A21FFB6FE0FFDF4ED1BFDC7" box="[474,573,606,629]" pageId="11" pageNumber="458" type="lectotype">lectotype</typeStatus>
designations for the BMNH specimens by
<bibRefCitation id="F7134B6D8A21FFB6FE78FDD4ED43FD27" author="ARNAUD" box="[429,613,638,661]" firstAuthor="ARNAUD" pageId="11" pageNumber="458" pagination="113 - 118" refId="ref24103" refString="ARNAUD P. 1982 a: Liste des types de Phanaeini du Museum National d'Histoire Naturelle de Paris [Coleoptera, Scarabaeidae]. Revue Francaise d'Entomologie (Nouvelle S erie) 4 (3): 113 - 118." type="journal article" year="1982" yearSuffix="a">ARNAUD (1982a)</bibRefCitation>
were invalid. Among the invalid
<typeStatus id="4C39883E8A21FFB6FEADFD34EECEFD07" box="[376,488,670,693]" pageId="11" pageNumber="458" type="lectotype">lectotypes</typeStatus>
designated by
<bibRefCitation id="F7134B6D8A21FFB6FD4CFD34EFCCFD67" author="ARNAUD" firstAuthor="ARNAUD" pageId="11" pageNumber="458" pagination="113 - 118" refId="ref24103" refString="ARNAUD P. 1982 a: Liste des types de Phanaeini du Museum National d'Histoire Naturelle de Paris [Coleoptera, Scarabaeidae]. Revue Francaise d'Entomologie (Nouvelle S erie) 4 (3): 113 - 118." type="journal article" year="1982" yearSuffix="a">ARNAUD (1982a)</bibRefCitation>
was that of
<taxonomicName id="54824D1F8A21FFB6FEB9FD14EEC9FD67" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[364,495,702,725]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FEB9FD14EEC9FD67" box="[364,495,702,725]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FE29FD14ED69FD67" box="[508,591,702,725]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FD8FFD14ED5EFD67" box="[602,632,702,725]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">47</figureCitation>
).
</paragraph>
<paragraph id="933D369C8A21FFB6FF60FD74EEBCFC47" blockId="11.[151,760,222,1013]" pageId="11" pageNumber="458">
<bibRefCitation id="F7134B6D8A21FFB6FF60FD74EE62FD47" author="BATES" box="[181,324,734,757]" firstAuthor="BATES" pageId="11" pageNumber="458" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">BATES (1887)</bibRefCitation>
did not provide information on the size of the type series of
<taxonomicName id="54824D1F8A21FFB6FEA6FD54EED0FCA7" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[371,502,766,789]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FEA6FD54EED0FCA7" box="[371,502,766,789]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
. However, he explicitly cited
<bibRefCitation id="F7134B6D8A21FFB6FF01FCB4EE59FC87" author="HAROLD" box="[212,383,798,821]" firstAuthor="HAROLD" pageId="11" pageNumber="458" pagination="161 - 176" refId="ref26135" refString="HAROLD E. 1863: Note sur les especes mexicaines du genre Phanaeus et descriptions de quelques especes nouvelles de coleopteres mexicains. Annales de la Societe Entomologique de France 4 (3): 161 - 176." type="journal article" year="1863">HAROLD (1863)</bibRefCitation>
to describe
<taxonomicName id="54824D1F8A21FFB6FDDCFCB4EDA8FC87" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[521,654,798,821]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FDDCFCB4EDA8FC87" box="[521,654,798,821]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
.
<bibRefCitation id="F7134B6D8A21FFB6FD4BFCB4EFF8FCE7" author="HAROLD" firstAuthor="HAROLD" pageId="11" pageNumber="458" pagination="161 - 176" refId="ref26135" refString="HAROLD E. 1863: Note sur les especes mexicaines du genre Phanaeus et descriptions de quelques especes nouvelles de coleopteres mexicains. Annales de la Societe Entomologique de France 4 (3): 161 - 176." type="journal article" year="1863">HAROLD (1863)</bibRefCitation>
commented that he examined only
<specimenCount id="8584FD158A21FFB6FDB1FC94EDDEFCE7" box="[612,760,830,853]" pageId="11" pageNumber="458" type="generic">one specimen</specimenCount>
from Sallés Collection. Since
<bibRefCitation id="F7134B6D8A21FFB6FE09FCF4ED4FFCC7" author="BATES" box="[476,617,862,885]" firstAuthor="BATES" pageId="11" pageNumber="458" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">BATES (1887)</bibRefCitation>
also relied in part on Sallés material, it is very likely that both referred to the same specimen. Consequently, that specimen should be considered the
<typeStatus id="4C39883E8A21FFB6FEB0FC14EEE5FC67" box="[357,451,958,981]" pageId="11" pageNumber="458" type="holotype">holotype</typeStatus>
of
<taxonomicName id="54824D1F8A21FFB6FE3BFC14ED53FC67" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[494,629,958,981]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FE3BFC14ED53FC67" box="[494,629,958,981]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
by original monotypy (
<figureCitation id="0BB92A198A21FFB6FEC4FC74EE42FC47" box="[273,356,990,1013]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FEBAFC74EEABFC47" box="[367,397,990,1013]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">47</figureCitation>
).
</paragraph>
<paragraph id="933D369C8A21FFB6FC8FFF14EB50FC87" blockId="11.[828,1440,190,1013]" pageId="11" pageNumber="458">
<bibRefCitation id="F7134B6D8A21FFB6FC8FFF14EB3EFF67" author="EDMONDS" box="[858,1048,190,213]" firstAuthor="EDMONDS" pageId="11" pageNumber="458" pagination="1 - 105" refId="ref25450" refString="EDMONDS W. D. 1994: Revision of Phanaeus MacLeay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science of the Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">EDMONDS (1994)</bibRefCitation>
considered
<taxonomicName id="54824D1F8A21FFB6FB71FF14EA0AFF67" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[1188,1324,190,213]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FB71FF14EA0AFF67" box="[1188,1324,190,213]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FAEAFF14EAB1FF67" box="[1343,1431,190,213]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FCE9FF74EC7EFF47" box="[828,856,222,245]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">47</figureCitation>
) to be a junior subjective synonym of
<taxonomicName id="54824D1F8A21FFB6FB25FF74EA78FF47" authorityName="Harold" authorityYear="1863" box="[1264,1374,222,245]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="daphnis">
<emphasis id="A1F6EA8E8A21FFB6FB25FF74EA78FF47" box="[1264,1374,222,245]" italics="true" pageId="11" pageNumber="458">P. daphnis</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FAB8FF74EC59FEA7" captionStart-0="Figs 1012" captionStart-1="Fig" captionStartId-0="9.[760,797,2010,2029]" captionStartId-1="10.[151,182,2061,2080]" captionTargetBox-0="[166,1438,335,2082]" captionTargetBox-1="[329,1263,1275,2048]" captionTargetId-0="figure-13@9.[151,1436,335,2082]" captionTargetId-1="figure-548@10.[325,1263,1272,2051]" captionTargetPageId-0="9" captionTargetPageId-1="10" captionText-0="Figs 1012. Phanaeus daphnis Harold, 1863. 10 male green phase; 11 male deep blue-green phase; 12 lectotype and labels (by Christophe Rivier, MNHN). Scale bar = 1.0 mm." captionText-1="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi-0="http://doi.org/10.5281/zenodo.5821158" figureDoi-1="http://doi.org/10.5281/zenodo.5976087" httpUri-0="https://zenodo.org/record/5821158/files/figure.png" httpUri-1="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 1113</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FC5CFF54EC80FEA7" box="[905,934,254,277]" captionStart="Figs 3242" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 3242. Posterior view of pronotum of major male. 32 P. tridens Castelnau, 1840; 33 P. moroni Arnaud, 2001, stat. rev.; 34 P. bal- thasari Arnaud, 2001, stat. rev.; 35 P. daphnis Harold, 1863; 36 P. substriolatus Balthasar, 1939, stat. rev.; 37 P.herbeus Bates, 1887, stat. rev.; 38 P.furiosus Bates, 1887; 39 P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 P. nimrod Harold, 1863; 41 P. victoriae Moctezuma sp. nov.; 42 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="11" pageNumber="458">35</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FC7AFF54ECEAFEA7" box="[943,972,254,277]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">46</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FC00FF54ECD5FEA7" box="[981,1011,254,277]" captionStart="Figs 5562" captionStartId="23.[151,188,2010,2029]" captionTargetBox="[201,1359,189,1996]" captionTargetId="figure-13@23.[200,1387,189,1996]" captionTargetPageId="23" captionText="Figs 5562. Pronotum of female. 55 P. tridens Castelnau, 1840; 56 P. balthasari Arnaud, 2001, stat. rev.; 57 P. daphnis Harold, 1863; 58 P. substriolatus Balthasar, 1939, stat. rev. (dark blue-black phase); 59 P.herbeus Bates, 1887, stat. rev. (green phase); 60 P. nimrod Harold, 1863 (blue-green phase); 61 P. furiosus Bates, 1887 (dark blue phase); 62 P. pseudofurcosus Balthasar, 1939, stat. rev. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821190" httpUri="https://zenodo.org/record/5821190/files/figure.png" pageId="11" pageNumber="458">57</figureCitation>
). The results of the present study suggest that
<taxonomicName id="54824D1F8A21FFB6FCBCFEB4ECCFFE87" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[873,1001,286,309]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FCBCFEB4ECCFFE87" box="[873,1001,286,309]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
deserves full species status because it has a unique combination of characters: the posteromedial process of pronotum is produced into a denticle that is more truncate apically and more widened at the base in
<taxonomicName id="54824D1F8A21FFB6FCE9FE34EC99FE07" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[828,959,414,437]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FCE9FE34EC99FE07" box="[828,959,414,437]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FC1BFE34EB3BFE07" box="[974,1053,414,437]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">Fig. 47</figureCitation>
) than in
<taxonomicName id="54824D1F8A21FFB6FBA8FE34EBCAFE07" authorityName="Harold" authorityYear="1863" box="[1149,1260,414,437]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="daphnis">
<emphasis id="A1F6EA8E8A21FFB6FBA8FE34EBCAFE07" box="[1149,1260,414,437]" italics="true" pageId="11" pageNumber="458">P. daphnis</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FB29FE34EA6BFE07" box="[1276,1357,414,437]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">Fig. 46</figureCitation>
), while the elytral interstriae IV are smooth in
<taxonomicName id="54824D1F8A21FFB6FB37FE14EA45FE67" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[1250,1379,446,469]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FB37FE14EA45FE67" box="[1250,1379,446,469]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FAA4FE14EC7FFE47" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Fig. 13</figureCitation>
). It is curious that the same differences had already been noted by
<bibRefCitation id="F7134B6D8A21FFB6FC0BFE54EBACFDA7" author="HAROLD" box="[990,1162,510,533]" firstAuthor="HAROLD" pageId="11" pageNumber="458" pagination="161 - 176" refId="ref26135" refString="HAROLD E. 1863: Note sur les especes mexicaines du genre Phanaeus et descriptions de quelques especes nouvelles de coleopteres mexicains. Annales de la Societe Entomologique de France 4 (3): 161 - 176." type="journal article" year="1863">HAROLD (1863)</bibRefCitation>
, who regarded the single specimen that he examined as an odd individual of
<taxonomicName id="54824D1F8A21FFB6FA5DFDB4ECB6FDE7" authorityName="Harold" authorityYear="1863" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="daphnis">
<emphasis id="A1F6EA8E8A21FFB6FA5DFDB4ECB6FDE7" italics="true" pageId="11" pageNumber="458">P. daphnis</emphasis>
</taxonomicName>
, and by
<bibRefCitation id="F7134B6D8A21FFB6FC24FD94EBA4FDE7" author="BATES" box="[1009,1154,574,597]" firstAuthor="BATES" pageId="11" pageNumber="458" pagination="2 - 3" refId="ref24392" refString="BATES H. W. 1887: Tribe Lamellicornia. Pp. 25 - 64 + pl. 2 - 3. In: BATES H. W. 1886 - 1889: Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis, London, xii + 432 p. + 24 pl." type="book chapter" year="1887">BATES (1887)</bibRefCitation>
, who treated
<taxonomicName id="54824D1F8A21FFB6FAC3FD94EA58FDE7" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[1302,1406,574,597]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FAC3FD94EA58FDE7" box="[1302,1406,574,597]" italics="true" pageId="11" pageNumber="458">coeruleus</emphasis>
</taxonomicName>
as a variety of
<taxonomicName id="54824D1F8A21FFB6FC15FDF4EB09FDC7" authorityName="Harold" authorityYear="1863" box="[960,1071,606,629]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="daphnis">
<emphasis id="A1F6EA8E8A21FFB6FC15FDF4EB09FDC7" box="[960,1071,606,629]" italics="true" pageId="11" pageNumber="458">P. daphnis</emphasis>
</taxonomicName>
. In addition, the dark blue colour with turquoise sheen on the whole dorsal surface of
<taxonomicName id="54824D1F8A21FFB6FA5CFDD4EC82FD07" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FA5CFDD4EC82FD07" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FC65FD34EB25FD07" box="[944,1027,670,693]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FBD8FD34EB0FFD07" box="[1037,1065,670,693]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">47</figureCitation>
) differs from the dark blue to black colouration of
<taxonomicName id="54824D1F8A21FFB6FC00FD14EB5EFD67" authorityName="Balthasar" authorityYear="1939" box="[981,1144,702,725]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="substriolatus">
<emphasis id="A1F6EA8E8A21FFB6FC00FD14EB5EFD67" box="[981,1144,702,725]" italics="true" pageId="11" pageNumber="458">P. substriolatus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FB53FD14EA26FD67" box="[1158,1280,702,725]" captionStart="Figs 1415" captionStartId="11.[151,188,2063,2082]" captionTargetBox="[165,1436,1055,2025]" captionTargetId="figure-495@11.[151,1436,1044,2033]" captionTargetPageId="11" captionText="Figs 1415. Phanaeus substriolatus Balthasar, 1939, stat. rev. 14 male; 15 holotype and labels (by Jiří Hájek, NMPC). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821160" httpUri="https://zenodo.org/record/5821160/files/figure.png" pageId="11" pageNumber="458">Figs 1415</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FADFFD14EA01FD67" box="[1290,1319,702,725]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">48</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FAE4FD14EA68FD67" box="[1329,1358,702,725]" captionStart="Figs 5562" captionStartId="23.[151,188,2010,2029]" captionTargetBox="[201,1359,189,1996]" captionTargetId="figure-13@23.[200,1387,189,1996]" captionTargetPageId="23" captionText="Figs 5562. Pronotum of female. 55 P. tridens Castelnau, 1840; 56 P. balthasari Arnaud, 2001, stat. rev.; 57 P. daphnis Harold, 1863; 58 P. substriolatus Balthasar, 1939, stat. rev. (dark blue-black phase); 59 P.herbeus Bates, 1887, stat. rev. (green phase); 60 P. nimrod Harold, 1863 (blue-green phase); 61 P. furiosus Bates, 1887 (dark blue phase); 62 P. pseudofurcosus Balthasar, 1939, stat. rev. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821190" httpUri="https://zenodo.org/record/5821190/files/figure.png" pageId="11" pageNumber="458">58</figureCitation>
), while the elytral striae are distinctly impressed in
<taxonomicName id="54824D1F8A21FFB6FACCFD74EABAFD47" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[1305,1436,734,757]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FACCFD74EABAFD47" box="[1305,1436,734,757]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FC96FD54ECB5FCA7" box="[835,915,766,789]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
) but superficially impressed to partially erased in
<taxonomicName id="54824D1F8A21FFB6FCE9FCB4ECC6FC87" authorityName="Balthasar" authorityYear="1939" box="[828,992,798,821]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="substriolatus">
<emphasis id="A1F6EA8E8A21FFB6FCE9FCB4ECC6FC87" box="[828,992,798,821]" italics="true" pageId="11" pageNumber="458">P. substriolatus</emphasis>
</taxonomicName>
(
<figureCitation id="0BB92A198A21FFB6FC3BFCB4EB4DFC87" box="[1006,1131,798,821]" captionStart="Figs 1415" captionStartId="11.[151,188,2063,2082]" captionTargetBox="[165,1436,1055,2025]" captionTargetId="figure-495@11.[151,1436,1044,2033]" captionTargetPageId="11" captionText="Figs 1415. Phanaeus substriolatus Balthasar, 1939, stat. rev. 14 male; 15 holotype and labels (by Jiří Hájek, NMPC). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821160" httpUri="https://zenodo.org/record/5821160/files/figure.png" pageId="11" pageNumber="458">Figs 1415</figureCitation>
).
</paragraph>
<paragraph id="933D369C8A21FFB1FC8FFC94EEF5FE87" blockId="11.[828,1440,190,1013]" lastBlockId="12.[151,763,190,341]" lastPageId="12" lastPageNumber="459" pageId="11" pageNumber="458">
The only
<specimenCount id="8584FD158A21FFB6FC14FC94EB75FCE7" box="[961,1107,830,853]" pageId="11" pageNumber="458" type="generic">one specimen</specimenCount>
of
<taxonomicName id="54824D1F8A21FFB6FBA2FC94EBDCFCE7" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[1143,1274,830,853]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FBA2FC94EBDCFCE7" box="[1143,1274,830,853]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
known to us is the
<typeStatus id="4C39883E8A21FFB6FCB3FCF4ECE5FCC7" box="[870,963,862,885]" pageId="11" pageNumber="458" type="holotype">holotype</typeStatus>
(
<figureCitation id="0BB92A198A21FFB6FC00FCF4EB0DFCC7" box="[981,1067,862,885]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
,
<figureCitation id="0BB92A198A21FFB6FBECFCF4EB71FCC7" box="[1081,1111,862,885]" captionStart="Figs 4354" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 4354. Lateral view of pronotum of major male. 43 P. tridens Castelnau, 1840; 44 P. moroni Arnaud, 2001, stat. rev.; 45 P. balthasari Arnaud, 2001; 46 P. daphnis Harold, 1863; 47 P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 P. substriolatus Balthasar, 1939, stat. rev.; 49 P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 P. furiosus Bates, 1887 (green phase); 51 P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 P. nimrod Harold, 1863; 53 P. victoriae Moctezuma sp. nov. (holotype); 54 P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="11" pageNumber="458">47</figureCitation>
). The labels of this specimen indicate that it was collected from
<collectingRegion id="5146F87E8A21FFB6FB69FCD4EA21FC27" box="[1212,1287,894,917]" country="Mexico" name="Puebla" pageId="11" pageNumber="458">Puebla</collectingRegion>
, without specific locality (
<figureCitation id="0BB92A198A21FFB6FC04FC34EB03FC07" box="[977,1061,926,949]" captionStart="Fig" captionStartId="10.[151,182,2061,2080]" captionTargetBox="[329,1263,1275,2048]" captionTargetId="figure-548@10.[325,1263,1272,2051]" captionTargetPageId="10" captionText="Fig. 13. Phanaeus coeruleus Bates, 1887, stat. rev., holotype and labels (by Keita Matsumoto, BMNH)." figureDoi="http://doi.org/10.5281/zenodo.5976087" httpUri="https://zenodo.org/record/5976087/files/figure.png" pageId="11" pageNumber="458">Figs 13</figureCitation>
). Interestingly, none of the revised specimens from
<collectingRegion id="5146F87E8A21FFB6FC3EFC14EB12FC67" box="[1003,1076,958,981]" country="Mexico" name="Puebla" pageId="11" pageNumber="458">Puebla</collectingRegion>
where identical in morphology to
<taxonomicName id="54824D1F8A21FFB6FCE9FC74ECE6FC47" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[828,960,990,1013]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A21FFB6FCE9FC74ECE6FC47" box="[828,960,990,1013]" italics="true" pageId="11" pageNumber="458">P. coeruleus</emphasis>
</taxonomicName>
. Since
<taxonomicName id="54824D1F8A21FFB6FBC1FC74EB5BFC47" authorityName="MacLeay" authorityYear="1819" box="[1044,1149,990,1013]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="genus">
<emphasis id="A1F6EA8E8A21FFB6FBC1FC74EB5BFC47" box="[1044,1149,990,1013]" italics="true" pageId="11" pageNumber="458">Phanaeus</emphasis>
</taxonomicName>
dung beetles from
<collectingRegion id="5146F87E8A21FFB6FA81FC74EABBFC47" box="[1364,1437,990,1013]" country="Mexico" name="Puebla" pageId="11" pageNumber="458">Puebla</collectingRegion>
are scarcely studied (MORÓN et al. 2003), future intensive collecting is needed in this Mexican state to improve the knowledge of
<taxonomicName id="54824D1F8A26FFB1FEE2FF54EE9BFEA7" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[311,445,254,277]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="459" phylum="Arthropoda" rank="species" species="coeruleus">
<emphasis id="A1F6EA8E8A26FFB1FEE2FF54EE9BFEA7" box="[311,445,254,277]" italics="true" pageId="12" pageNumber="459">P. coeruleus</emphasis>
</taxonomicName>
and related species (e.g.,
<taxonomicName id="54824D1F8A26FFB1FD31FF54EFCDFE87" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="459" phylum="Arthropoda" rank="species" species="herbeus">
<emphasis id="A1F6EA8E8A26FFB1FD31FF54EFCDFE87" italics="true" pageId="12" pageNumber="459">P. herbeus</emphasis>
</taxonomicName>
and
<taxonomicName id="54824D1F8A26FFB1FECAFEB4EEE2FE87" authorityName="Balthasar" authorityYear="1939" box="[287,452,286,309]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="459" phylum="Arthropoda" rank="species" species="substriolatus">
<emphasis id="A1F6EA8E8A26FFB1FECAFEB4EEE2FE87" box="[287,452,286,309]" italics="true" pageId="12" pageNumber="459">P. substriolatus</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="C7FD66148A21FFB6FF42F7A5EA7AF790" ID-DOI="http://doi.org/10.5281/zenodo.5821160" ID-Zenodo-Dep="5821160" box="[151,1372,2063,2082]" httpUri="https://zenodo.org/record/5821160/files/figure.png" pageId="11" pageNumber="458" startId="11.[151,188,2063,2082]" targetBox="[165,1436,1055,2025]" targetPageId="11">
<paragraph id="933D369C8A21FFB6FF42F7A5EA7AF790" blockId="11.[151,1372,2063,2082]" box="[151,1372,2063,2082]" pageId="11" pageNumber="458">
Figs 1415.
<taxonomicName id="54824D1F8A21FFB6FED4F7A5ED70F790" authority="Balthasar, 1939" authorityName="Balthasar" authorityYear="1939" box="[257,598,2063,2082]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="458" phylum="Arthropoda" rank="species" species="substriolatus" status="stat. nov.">
<emphasis id="A1F6EA8E8A21FFB6FED4F7A5EEEFF790" box="[257,457,2063,2082]" italics="true" pageId="11" pageNumber="458">Phanaeus substriolatus</emphasis>
<bibRefCitation id="F7134B6D8A21FFB6FE1BF7A5ED70F790" author="Balthasar" box="[462,598,2063,2082]" firstAuthor="Balthasar" pageId="11" pageNumber="458" pagination="238 - 247" refId="ref24264" refString="BALTHASAR V. 1939: Neue Phanaeus - Arten. (7. Beitrag zur Kenntnis der Scarabaeiden der neotropischen Region). Folia Zoologica et Hydrobiologica 9 (2): 238 - 247." type="journal article" year="1939">Balthasar, 1939</bibRefCitation>
</taxonomicName>
,
<taxonomicNameLabel id="BAC557F58A21FFB6FD8BF7A5ED81F790" box="[606,679,2063,2082]" pageId="11" pageNumber="458">stat. rev.</taxonomicNameLabel>
14 male; 15 holotype and labels (by Jiří Hájek, NMPC). Scale bar = 1.0 mm.
</paragraph>
</caption>
<paragraph id="933D369C8A26FFB1FF42FE94EEF9FEE7" blockId="12.[151,763,190,341]" box="[151,479,318,341]" pageId="12" pageNumber="459">
<emphasis id="A1F6EA8E8A26FFB1FF42FE94EE0CFEE7" bold="true" box="[151,298,318,341]" pageId="12" pageNumber="459">Distribution.</emphasis>
<collectingCountry id="EB95760C8A26FFB1FEE5FE94EEA0FEE7" box="[304,390,318,341]" name="Mexico" pageId="12" pageNumber="459">Mexico</collectingCountry>
:
<collectingRegion id="5146F87E8A26FFB1FE5AFE94EEFDFEE7" box="[399,475,318,341]" country="Mexico" name="Puebla" pageId="12" pageNumber="459">Puebla</collectingRegion>
.
</paragraph>
</subSubSection>
</treatment>
</document>