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<document id="B724FCAB1A68276FAE005D590C3183B1" ID-CLB-Dataset="20763" ID-DOI="10.37520/aemnp.2021.025" ID-GBIF-Dataset="0839d72f-9521-4f11-916a-4b4c8ea24e8f" ID-ISSN="1804-6487" ID-Zenodo-Dep="5821143" ID-ZooBank="85434EFF-F859-4BBF-8AB5-F50B9BA08771" IM.bibliography_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1641390032322" checkinUser="plazi" docAuthor="Moctezuma, Victor, Halffter, Gonzalo &amp; Lizardo, Viridiana" docDate="2021" docId="1B2B878A8A2EFFBBFC7BFF45EFE4F7A7" docLanguage="en" docName="ActaEntMusNatPra.61.2.447-482.pdf" docOrigin="Acta Entomologica Musei Nationalis Pragae 61 (2)" docStyle="DocumentStyle:BA2C5C8540539042E893A65A182CEA85.8:ActaEntMusNatPra.2018-.journal_article" docStyleId="BA2C5C8540539042E893A65A182CEA85" docStyleName="ActaEntMusNatPra.2018-.journal_article" docStyleVersion="8" docTitle="Phanaeus tridens Castelnau 1840" docType="treatment" docVersion="8" lastPageNumber="453" masterDocId="E712FFF28A2AFFBDFFD5FFAAEF26FFB2" masterDocTitle="The Phanaeus tridens species group (Coleoptera: Scarabaeoidea): a dung beetle group with genital morphological stasis but a changing ecological niche" masterLastPageNumber="482" masterPageNumber="447" pageNumber="451" updateTime="1699295799786" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:title id="C08B0EBB9FA90B678D7C88F72E6D8213">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="142C882D1672C23D6AD256576AF8EFE8">Moctezuma, Victor</mods:namePart>
<mods:affiliation id="6D2B6B486BA24E6350C5ECC26696B167">Centro Tlaxcala de Biología de la Conducta, Universidad Autónoma de Tlaxcala, Tlaxcala de Xicohténcatl, Tlaxcala 90070, Mexico, Km 1.5 Carretera Tlaxcala-Puebla S / N, La Loma Xicohtencatl.</mods:affiliation>
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<mods:namePart id="7C3A048FF92CE7DFF70CE5E8FEB88217">Halffter, Gonzalo</mods:namePart>
<mods:affiliation id="8FD428630D54D7AA759BCFCAE9B012E2">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="179214A49CD1E5B748C51AB0FB41E57A">Lizardo, Viridiana</mods:namePart>
<mods:affiliation id="7AC1242892FCFC545B8DCDBFA9186A9D">Facultad de Ciencias, Universidad Nacional Autónoma de México, Mexico City 04510, Mexico, Apartado Postal 70 - 399.</mods:affiliation>
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<mods:title id="A123708E169D9165FBA2AC6DBE630BEA">Acta Entomologica Musei Nationalis Pragae</mods:title>
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<mods:date id="395BB168F9453ED41363565739E275D9">2021</mods:date>
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<mods:number id="6E47D56608895A145A68080A376552C1">2021-12-18</mods:number>
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<mods:number id="93CC45EE8470DD3CDDA23D40DCBE6238">61</mods:number>
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<treatment id="1B2B878A8A2EFFBBFC7BFF45EFE4F7A7" ID-DOI="http://doi.org/10.5281/zenodo.6303352" ID-GBIF-Taxon="193278662" ID-Zenodo-Dep="6303352" LSID="urn:lsid:plazi:treatment:1B2B878A8A2EFFBBFC7BFF45EFE4F7A7" httpUri="http://treatment.plazi.org/id/1B2B878A8A2EFFBBFC7BFF45EFE4F7A7" lastPageId="6" lastPageNumber="453" pageId="4" pageNumber="451">
<subSubSection id="DB9865178A2EFFB9FC7BFF45EA0FFEB4" box="[942,1321,239,262]" pageId="4" pageNumber="451" type="nomenclature">
<paragraph id="933D369C8A2EFFB9FC7BFF45EA0FFEB4" blockId="4.[942,1321,239,262]" box="[942,1321,239,262]" pageId="4" pageNumber="451">
<heading id="C87581F08A2EFFB9FC7BFF45EA0FFEB4" box="[942,1321,239,262]" centered="true" fontSize="10" level="2" pageId="4" pageNumber="451" reason="3">
<taxonomicName id="54824D1F8A2EFFB9FC7BFF45EA0FFEB4" ID-CoL="4FSL8" authority="Castelnau, 1840" authorityName="Castelnau" authorityYear="1840" box="[942,1321,239,262]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2EFFB9FC7BFF45EA0FFEB4" bold="true" box="[942,1321,239,262]" pageId="4" pageNumber="451">
<emphasis id="A1F6EA8E8A2EFFB9FC7BFF45EB49FEB4" bold="true" box="[942,1135,239,262]" italics="true" pageId="4" pageNumber="451">Phanaeus tridens</emphasis>
<bibRefCitation id="F7134B6D8A2EFFB9FBA0FF45EA0FFEB4" author="Castelnau" box="[1141,1321,239,262]" firstAuthor="Castelnau" pageId="4" pageNumber="451" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">Castelnau, 1840</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="DB9865178A2EFFB9FBD5FEA7EBFEFE92" box="[1024,1240,269,288]" pageId="4" pageNumber="451" type="description">
<paragraph id="933D369C8A2EFFB9FBD5FEA7EBFEFE92" blockId="4.[828,1438,269,757]" box="[1024,1240,269,288]" pageId="4" pageNumber="451">
(
<figureCitation id="0BB92A198A2EFFB9FBD3FEA7EB74FE92" box="[1030,1106,269,288]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 1–5</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FB89FEA7EB55FE92" box="[1116,1139,269,288]" captionStart="Figs 32–42" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="4" pageNumber="451">32</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FBA9FEA7EBB5FE92" box="[1148,1171,269,288]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">43</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FB49FEA7EB95FE92" box="[1180,1203,269,288]" captionStart="Figs 55–62" captionStartId="23.[151,188,2010,2029]" captionTargetBox="[201,1359,189,1996]" captionTargetId="figure-13@23.[200,1387,189,1996]" captionTargetPageId="23" captionText="Figs 55–62. 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="4" pageNumber="451">55</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FB69FEA7EBF4FE92" box="[1212,1234,269,288]" captionStart="Fig" captionStartId="24.[828,858,545,564]" captionTargetBox="[841,1417,205,525]" captionTargetId="figure-740@24.[825,1429,186,526]" captionTargetPageId="24" captionText="Fig.63. Phallobase, parameres and endophallite copulatrix of the P.tridens species group. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821192" httpUri="https://zenodo.org/record/5821192/files/figure.png" pageId="4" pageNumber="451">63</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A2EFFB9FCE9FE85ECC2FD88" pageId="4" pageNumber="451" type="reference_group">
<paragraph id="933D369C8A2EFFB9FCE9FE85ECE7FEEF" blockId="4.[828,1438,269,757]" pageId="4" pageNumber="451">
<treatmentCitationGroup id="B39211B28A2EFFB9FCE9FE85ECE7FEEF" pageId="4" pageNumber="451">
<treatmentCitation id="1223108D8A2EFFB9FCE9FE85EBA6FEF0" author="CASTELNAU C. D." box="[828,1152,303,322]" page="81" pageId="4" pageNumber="451" year="1840">
<taxonomicName id="54824D1F8A2EFFB9FCE9FE85EBA6FEF0" ID-CoL="4FSL8" authority="Castelnau, 1840: 81" authorityName="Castelnau" authorityPageNumber="81" authorityYear="1840" box="[828,1152,303,322]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2EFFB9FCE9FE85ECE9FEF0" box="[828,975,303,322]" italics="true" pageId="4" pageNumber="451">Phanaeus tridens</emphasis>
<bibRefCitation id="F7134B6D8A2EFFB9FC01FE85EBA6FEF0" author="Castelnau" box="[980,1152,303,322]" firstAuthor="Castelnau" page="81" pageId="4" pageNumber="451" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">Castelnau, 1840: 81</bibRefCitation>
</taxonomicName>
</treatmentCitation>
.
<typeStatus id="4C39883E8A2EFFB9FB52FE85EB97FEF0" box="[1159,1201,303,322]" pageId="4" pageNumber="451">Type</typeStatus>
locality:
<collectingCountry id="EB95760C8A2EFFB9FAD5FE85EA65FEF0" box="[1280,1347,303,322]" name="Mexico" pageId="4" pageNumber="451">Mexico</collectingCountry>
,
<collectingRegion id="5146F87E8A2EFFB9FA9EFE85EABFFEF0" box="[1355,1433,303,322]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="4" pageNumber="451">Veracruz</collectingRegion>
, Palma Sola.
</treatmentCitationGroup>
</paragraph>
<treatmentCitationGroup id="B39211B28A2EFFB9FCE9FECEEB79FE58" pageId="4" pageNumber="451">
<paragraph id="933D369C8A2EFFB9FCE9FECEEAB8FE20" blockId="4.[828,1438,269,757]" pageId="4" pageNumber="451">
<treatmentCitation id="1223108D8A2EFFB9FCE9FECEEBF7FEC5" author="BALTHASAR V." box="[828,1233,356,375]" page="245" pageId="4" pageNumber="451" year="1939">
<taxonomicName id="54824D1F8A2EFFB9FCE9FECEEBF7FEC5" ID-CoL="4FSMG" authority="Balthasar, 1939: 245" authorityName="Balthasar" authorityPageNumber="245" authorityYear="1939" box="[828,1233,356,375]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="frankenbergeri">
<emphasis id="A1F6EA8E8A2EFFB9FCE9FECEEB33FEC5" box="[828,1045,356,375]" italics="true" pageId="4" pageNumber="451">Phanaeus frankenbergeri</emphasis>
<bibRefCitation id="F7134B6D8A2EFFB9FBCEFECEEBF7FEC5" author="Balthasar" box="[1051,1233,356,375]" firstAuthor="Balthasar" page="245" pageId="4" pageNumber="451" 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: 245</bibRefCitation>
</taxonomicName>
</treatmentCitation>
.
<typeStatus id="4C39883E8A2EFFB9FB0CFECEEA25FEC5" box="[1241,1283,356,375]" pageId="4" pageNumber="451">Type</typeStatus>
locality:
<collectingRegion id="5146F87E8A2EFFB9FA83FECEEABCFEC5" box="[1366,1434,356,375]" country="Mexico" name="Mexico" pageId="4" pageNumber="451">Mexico</collectingRegion>
. Synonymy established by
<bibRefCitation id="F7134B6D8A2EFFB9FBE0FED5EBD7FE20" author="ARNAUD" box="[1077,1265,383,402]" firstAuthor="ARNAUD" page="125" pageId="4" pageNumber="451" pagination="125 - 126" refId="ref24144" refString="ARNAUD P. 1982 b: Synonymies dans le genre Phanaeus (Col. Scarabaeidae). Miscellanea Entomologica 49: 125 - 126." type="journal article" year="1982" yearSuffix="b">ARNAUD (1982b: 125)</bibRefCitation>
and confirmed here.
</paragraph>
<paragraph id="933D369C8A2EFFB9FCE9FE0BEB79FE58" blockId="4.[828,1438,269,757]" pageId="4" pageNumber="451">
<emphasis id="A1F6EA8E8A2EFFB9FCE9FE0BEB0DFE06" bold="true" box="[828,1067,417,436]" pageId="4" pageNumber="451">
<typeStatus id="4C39883E8A2EFFB9FCE9FE0BEC4CFE06" box="[828,874,417,436]" pageId="4" pageNumber="451">Type</typeStatus>
material examined.
</emphasis>
<taxonomicName id="54824D1F8A2EFFB9FBE6FE0BEA09FE06" ID-CoL="4FSL8" authority=": N EOTYPE" authorityName="N EOTYPE" box="[1075,1327,417,436]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2EFFB9FBE6FE0BEBF7FE06" box="[1075,1233,417,436]" italics="true" pageId="4" pageNumber="451">Phanaeus tridens</emphasis>
: N EOTYPE
</taxonomicName>
(suggested, examined from photographs;
<figureCitation id="0BB92A198A2EFFB9FBE2FE16EB4CFE7D" box="[1079,1130,444,463]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Fig. 3</figureCitation>
):
<emphasis id="A1F6EA8E8A2EFFB9FBAEFE16EA61FE7D" bold="true" box="[1147,1351,444,463]" pageId="4" pageNumber="451">
<collectingCountry id="EB95760C8A2EFFB9FBAEFE16EBFEFE7D" box="[1147,1240,444,463]" name="Mexico" pageId="4" pageNumber="451">MEXICO</collectingCountry>
: V ERACRUZ:
</emphasis>
♂, Palma Sola (MNHN: MNHN EC10552).
</paragraph>
</treatmentCitationGroup>
<paragraph id="933D369C8A2EFFB9FC8FFE5BECC2FD88" blockId="4.[828,1438,269,757]" pageId="4" pageNumber="451">
<taxonomicName id="54824D1F8A2EFFB9FC8FFE5BEB15FDB6" ID-CoL="4FSMG" authorityName="Balthasar" authorityYear="1939" box="[858,1075,497,516]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="frankenbergeri">
<emphasis id="A1F6EA8E8A2EFFB9FC8FFE5BEB15FDB6" box="[858,1075,497,516]" italics="true" pageId="4" pageNumber="451">Phanaeus frankenbergeri</emphasis>
</taxonomicName>
: HOLOTYPE (examined from photographs;
<figureCitation id="0BB92A198A2EFFB9FCE9FDA6EC54FDAD" box="[828,882,524,543]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Fig. 4</figureCitation>
): ♂,
<collectingCountry id="EB95760C8A2EFFB9FC74FDA6ECC4FDAD" box="[929,994,524,543]" name="Mexico" pageId="4" pageNumber="451">Mexico</collectingCountry>
(without further locality data) (NMPC: Mus. Nat. Pragae Inv. 26351).
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A2EFFB9FCE9FDEBEA0AFD46" pageId="4" pageNumber="451" type="materials_examined">
<paragraph id="933D369C8A2EFFB9FCE9FDEBEA0AFD46" blockId="4.[828,1438,269,757]" pageId="4" pageNumber="451">
<emphasis id="A1F6EA8E8A2EFFB9FCE9FDEBEA08FDE6" bold="true" box="[828,1326,577,596]" pageId="4" pageNumber="451">
Non-type material examined.
<collectingCountry id="EB95760C8A2EFFB9FB8FFDEBEB9FFDE6" box="[1114,1209,577,596]" name="Mexico" pageId="4" pageNumber="451">MEXICO</collectingCountry>
: V ERACRUZ:
</emphasis>
<specimenCount id="8584FD158A2EFFB9FAE3FDEBEA4FFDE6" box="[1334,1385,571,596]" pageId="4" pageNumber="451" type="female">2 ♀♀</specimenCount>
, Alto Lucero (
<collectionCode id="F593AE598A2EFFB9FC51FDF6ECECFDDD" box="[900,970,604,623]" pageId="4" pageNumber="451">GHVM</collectionCode>
); 2 ♂♂,
<quantity id="547A9B798A2EFFB9FBC0FDF6EB6AFDDD" box="[1045,1100,604,623]" metricMagnitude="2" metricUnit="m" metricValue="7.0" pageId="4" pageNumber="451" unit="m" value="700.0">700 m</quantity>
NE Apazapan (
<collectionCode id="F593AE598A2EFFB9FB01FDF6EA2AFDDD" box="[1236,1292,604,623]" pageId="4" pageNumber="451">IEXA</collectionCode>
);
<specimenCount id="8584FD158A2EFFB9FACEFDF6EA6CFDDC" box="[1307,1354,597,623]" pageId="4" pageNumber="451" type="female">2 ♀♀</specimenCount>
, Conejos (
<collectionCode id="F593AE598A2EFFB9FC97FDDDECAEFD38" box="[834,904,631,650]" pageId="4" pageNumber="451">GHVM</collectionCode>
); 3 ♂♂
<specimenCount id="8584FD158A2EFFB9FC1EFDDDECCCFD3B" box="[971,1002,624,650]" pageId="4" pageNumber="451" type="female">1 ♀</specimenCount>
, Cotaxtla (
<collectionCode id="F593AE598A2EFFB9FB9FFDDDEBB6FD38" box="[1098,1168,631,650]" pageId="4" pageNumber="451">GHVM</collectionCode>
); 1 ♂, Laguna Verde (
<collectionCode id="F593AE598A2EFFB9FA9EFDDDEAA5FD38" box="[1355,1411,631,650]" pageId="4" pageNumber="451">IEXA</collectionCode>
);
<specimenCount id="8584FD158A2EFFB9FA44FDDDEC6DFD16" pageId="4" pageNumber="451" type="female">1 ♀</specimenCount>
, Motzorongo (
<collectionCode id="F593AE598A2EFFB9FC19FD3BEB34FD16" box="[972,1042,657,676]" pageId="4" pageNumber="451">GHVM</collectionCode>
); 24 ♂♂
<specimenCount id="8584FD158A2EFFB9FBB4FD3BEBBDFD16" box="[1121,1179,651,676]" pageId="4" pageNumber="451" type="female">30 ♀♀</specimenCount>
, Palma Sola (
<collectionCode id="F593AE598A2EFFB9FAC6FD3BEA6CFD16" box="[1299,1354,657,676]" pageId="4" pageNumber="451">IEXA</collectionCode>
: 11 ♂♂
<specimenCount id="8584FD158A2EFFB9FA44FD3BEC7DFD0C" pageId="4" pageNumber="451" type="female">2 ♀♀</specimenCount>
; GHVM: 10 ♂♂
<specimenCount id="8584FD158A2EFFB9FC24FD06EB0DFD0C" box="[1009,1067,677,703]" pageId="4" pageNumber="451" type="female">24 ♀♀</specimenCount>
:
<collectionCode id="F593AE598A2EFFB9FBE3FD06EB5DFD0D" box="[1078,1147,684,703]" pageId="4" pageNumber="451">VMPM</collectionCode>
: 3 ♂♂
<specimenCount id="8584FD158A2EFFB9FB62FD06EBC0FD0C" box="[1207,1254,677,703]" pageId="4" pageNumber="451" type="female">4 ♀♀</specimenCount>
);
<specimenCount id="8584FD158A2EFFB9FB2DFD06EA3EFD0C" box="[1272,1304,677,703]" pageId="4" pageNumber="451" type="female">1 ♀</specimenCount>
, Plan de Hayas (
<collectionCode id="F593AE598A2EFFB9FC97FD6DECAEFD68" box="[834,904,711,730]" pageId="4" pageNumber="451">GHVM</collectionCode>
);
<specimenCount id="8584FD158A2EFFB9FC4DFD6DECEEFD6B" box="[920,968,704,730]" pageId="4" pageNumber="451" type="female">2 ♀♀</specimenCount>
, Presidio (
<collectionCode id="F593AE598A2EFFB9FBFDFD6DEB48FD68" box="[1064,1134,711,730]" pageId="4" pageNumber="451">GHVM</collectionCode>
); 1 ♂
<specimenCount id="8584FD158A2EFFB9FB73FD6DEBECFD6B" box="[1190,1226,704,730]" pageId="4" pageNumber="451" type="female">1 ♀</specimenCount>
<collectingRegion id="5146F87E8A2EFFB9FB1BFD6DEA3CFD68" box="[1230,1306,711,730]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="4" pageNumber="451">Veracruz</collectingRegion>
(
<collectionCode id="F593AE598A2EFFB9FAF2FD6DEA4BFD68" box="[1319,1389,711,730]" pageId="4" pageNumber="451">GHVM</collectionCode>
:
<specimenCount id="8584FD158A2EFFB9FAA3FD6DEAB1FD6B" box="[1398,1431,704,730]" pageId="4" pageNumber="451" type="female">1 ♀</specimenCount>
,
<collectionCode id="F593AE598A2EFFB9FCE9FD4BECA7FD46" box="[828,897,737,756]" pageId="4" pageNumber="451">VMPM</collectionCode>
: 1♂); 1 ♂
<specimenCount id="8584FD158A2EFFB9FC0FFD4BECDCFD46" box="[986,1018,731,756]" pageId="4" pageNumber="451" type="female">1 ♀</specimenCount>
, without specific locality (
<collectionCode id="F593AE598A2EFFB9FB09FD4BEA04FD46" box="[1244,1314,737,756]" pageId="4" pageNumber="451">GHVM</collectionCode>
).
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A2EFFB9FCE9FCA7EA7DF976" pageId="4" pageNumber="451" type="diagnosis">
<paragraph id="933D369C8A2EFFB9FCE9FCA7EB4AFA96" blockId="4.[828,1437,781,2084]" pageId="4" pageNumber="451">
<emphasis id="A1F6EA8E8A2EFFB9FCE9FCA7EC96FC96" bold="true" box="[828,944,781,804]" pageId="4" pageNumber="451">Diagnosis.</emphasis>
Metallic green, deep green to blue green (
<figureCitation id="0BB92A198A2EFFB9FAB8FCA7EC43FCF6" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 3–5</figureCitation>
), occasionally with red sheen (
<figureCitation id="0BB92A198A2EFFB9FB4BFC87EBFBFCF6" box="[1182,1245,813,836]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Fig. 4</figureCitation>
). Sides of pronotal disc finely granulate, becoming granulorugose on raised outer margin of disc (
<figureCitation id="0BB92A198A2EFFB9FBF4FCC7EBA6FC36" box="[1057,1152,877,900]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 3–5</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FB59FCC7EB8CFC36" box="[1164,1194,877,900]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">43</figureCitation>
). Pronotal disc weakly but coarsely rugose (
<figureCitation id="0BB92A198A2EFFB9FBFDFC27EBA8FC16" box="[1064,1166,909,932]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 3–5</figureCitation>
). Posteromedial process of pronotum produced into denticle, distinctly widened laterally (not reaching anteromedial carina), elongate, and apically bifurcated (
<figureCitation id="0BB92A198A2EFFB9FBDAFC47EB44FBB6" box="[1039,1122,1005,1028]" captionStart="Figs 32–42" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="4" pageNumber="451">Figs 32</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FBB9FC47EBACFBB6" box="[1132,1162,1005,1028]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">43</figureCitation>
). Anteromedial portion of pronotal disc with two denticles, usually joint by medial carina (
<figureCitation id="0BB92A198A2EFFB9FC5DFB87ECF3FBF6" box="[904,981,1069,1092]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">Fig. 43</figureCitation>
).Anterolateral margins of disc with distinctly developed ridge of tubercles (
<figureCitation id="0BB92A198A2EFFB9FB77FBE7EBD2FBD6" box="[1186,1268,1101,1124]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">Fig. 43</figureCitation>
). Posterolateral angles of pronotum shorter than posteromedial process of pronotum (
<figureCitation id="0BB92A198A2EFFB9FC67FB27EB20FB16" box="[946,1030,1165,1188]" captionStart="Figs 32–42" captionStartId="21.[828,865,1898,1917]" captionTargetBox="[211,1440,197,2079]" captionTargetId="figure-13@21.[211,1376,189,2079]" captionTargetPageId="21" captionText="Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821182" httpUri="https://zenodo.org/record/5821182/files/figure.png" pageId="4" pageNumber="451">Figs 32</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FBC4FB27EB09FB16" box="[1041,1071,1165,1188]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">43</figureCitation>
). Deep green to green blue elytra, occasionally with red sheen (
<figureCitation id="0BB92A198A2EFFB9FBBCFB07EBE1FB76" box="[1129,1223,1197,1220]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 1–5</figureCitation>
). Elytral striae scabriculous, distinctly impressed, superficially punctate (
<figureCitation id="0BB92A198A2EFFB9FAB8FB67EC43FAB6" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 1–5</figureCitation>
). Elytral interstriae scabriculous, smooth, superficially punctate, convex (
<figureCitation id="0BB92A198A2EFFB9FC2BFAA7EB79FA96" box="[1022,1119,1293,1316]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 1–5</figureCitation>
).
</paragraph>
<paragraph id="933D369C8A2EFFB9FCE9FA87EA7DF976" blockId="4.[828,1437,781,2084]" pageId="4" pageNumber="451">
<emphasis id="A1F6EA8E8A2EFFB9FCE9FA87EB6DFAF6" bold="true" box="[828,1099,1325,1348]" pageId="4" pageNumber="451">
Variability.
<emphasis id="A1F6EA8E8A2EFFB9FC17FA87EB6DFAF6" bold="true" box="[962,1099,1325,1348]" italics="true" pageId="4" pageNumber="451">
<taxonomicName id="54824D1F8A2EFFB9FC17FA87EB2EFAF6" authorityName="Greenslade P &amp; Greenslade Pjm" authorityYear="1987" box="[962,1032,1325,1348]" class="Insecta" family="Scarabaeidae" genus="Minor" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="genus">Minor</taxonomicName>
male.
</emphasis>
</emphasis>
Similar to major males, except for reduction of secondary sexual characters (i.e., cephalic horn, pronotal processes and posterolateral angles).
<emphasis id="A1F6EA8E8A2EFFB9FA81FAC7EC70FA16" bold="true" italics="true" pageId="4" pageNumber="451">Female.</emphasis>
Similar to male, except for head showing trituberculate carina; pronotal sculpture granulate; pronotum with anteromedial black macula, and anteromedial trapezoidal carina, followed by posterior concavity (
<figureCitation id="0BB92A198A2EFFB9FB4BFA47EBC8F9B6" box="[1182,1262,1517,1540]" captionStart="Figs 55–62" captionStartId="23.[151,188,2010,2029]" captionTargetBox="[201,1359,189,1996]" captionTargetId="figure-13@23.[200,1387,189,1996]" captionTargetPageId="23" captionText="Figs 55–62. 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="4" pageNumber="451">Fig. 55</figureCitation>
). This species is variable in colour: typical individuals are green dorsally with blue sheen (
<figureCitation id="0BB92A198A2EFFB9FC24F987EB4DF9F6" box="[1009,1131,1581,1604]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Figs 1, 3–5</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FBA3F987EBB5F9F6" box="[1142,1171,1581,1604]" captionStart="Figs 43–54" captionStartId="22.[151,188,1979,1998]" captionTargetBox="[195,1407,189,1941]" captionTargetId="figure-13@22.[179,1408,189,1947]" captionTargetPageId="22" captionText="Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821186" httpUri="https://zenodo.org/record/5821186/files/figure.png" pageId="4" pageNumber="451">43</figureCitation>
,
<figureCitation id="0BB92A198A2EFFB9FB4BF987EB9DF9F6" box="[1182,1211,1581,1604]" captionStart="Figs 55–62" captionStartId="23.[151,188,2010,2029]" captionTargetBox="[201,1359,189,1996]" captionTargetId="figure-13@23.[200,1387,189,1996]" captionTargetPageId="23" captionText="Figs 55–62. 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="4" pageNumber="451">55</figureCitation>
), but specimens with metallic red sheen on head and pronotum are occasionally observed (
<figureCitation id="0BB92A198A2EFFB9FC72F9C7ECC0F936" box="[935,998,1645,1668]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="4" pageNumber="451">Fig. 2</figureCitation>
).
<bibRefCitation id="F7134B6D8A2EFFB9FC2DF9C7EA2EF936" author="EDMONDS &amp; ZIDEK" box="[1016,1288,1645,1668]" firstAuthor="EDMONDS" pageId="4" pageNumber="451" pagination="1 - 108" refId="ref25590" refString="EDMONDS W. D. &amp; ZIDEK J. 2012: Taxonomy of Phanaeus revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Insecta Mundi 274: 1 - 108." type="journal article" year="2012">EDMONDS &amp; ZÍDEK (2012)</bibRefCitation>
figured a dark blue female specimen of
<taxonomicName id="54824D1F8A2EFFB9FB90F927EB8FF916" authorityName="Castelnau" authorityYear="1840" box="[1093,1193,1677,1700]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2EFFB9FB90F927EB8FF916" box="[1093,1193,1677,1700]" italics="true" pageId="4" pageNumber="451">P. tridens</emphasis>
</taxonomicName>
. Nevertheless, none of the specimens revised for this study was dark blue.
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A2EFFBBFCE9F967ED35F867" lastPageId="6" lastPageNumber="453" pageId="4" pageNumber="451" type="discussion">
<paragraph id="933D369C8A2EFFB8FCE9F967ED61F8ED" blockId="4.[828,1437,781,2084]" lastBlockId="5.[151,763,1832,2079]" lastPageId="5" lastPageNumber="452" pageId="4" pageNumber="451">
<emphasis id="A1F6EA8E8A2EFFB9FCE9F967EC98F956" bold="true" box="[828,958,1741,1764]" pageId="4" pageNumber="451">Comments.</emphasis>
<bibRefCitation id="F7134B6D8A2EFFB9FC12F967EBA7F956" author="EDMONDS" box="[967,1153,1741,1764]" firstAuthor="EDMONDS" pageId="4" pageNumber="451" 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>
commented that the type material for many phanaeine species described by Castelnau was probably lost. As discussed by different authors (
<bibRefCitation id="F7134B6D8A2EFFB9FC96F887ECCBF8F6" author="EVENHUIS" box="[835,1005,1837,1860]" firstAuthor="EVENHUIS" pageId="4" pageNumber="451" pagination="53 - 63" refId="ref25820" refString="EVENHUIS N. L. 2012: Francois-Louis Comte de Castelnau (1802 - 1880) and the mysterious disappearance of his original insect collection. Zootaxa 3168: 53 - 63." type="journal article" year="2012">EVENHUIS 2012</bibRefCitation>
,
<bibRefCitation id="F7134B6D8A2EFFB9FC2DF887EBCAF8F6" author="MALDANER" box="[1016,1260,1837,1860]" etAl="et al." firstAuthor="MALDANER" pageId="4" pageNumber="451" pagination="83 - 102" refId="ref26840" refString="MALDANER M. E., CUPELLO M., FERREIRA D. C. &amp; VAZ-DE- MELLO F. Z. 2017: Type specimens and names assigned to Coprophanaeus (Megaphanaeus) d'Olsoufieff, 1924, the largest New World dung beetles (Coleoptera: Scarabaeidae: Phanaeini). Zootaxa 4272 (1): 83 - 102." type="journal article" year="2017">MALDANER et al. 2017</bibRefCitation>
,
<bibRefCitation id="F7134B6D8A2EFFB9FB22F887ECD9F8D6" author="VAZ-DE-MELLO &amp; CUPELLO" firstAuthor="VAZ-DE-MELLO" pageId="4" pageNumber="451" pagination="41 - 58" refId="ref28657" refString="VAZ-DE-MELLO F. Z. &amp; CUPELLO M. 2018: The type specimens of South American dung beetles, Part II: The species described by the Austrian coleopterist Ludwig Redtenbacher (1814 - 1876) in the 1868 part of the &quot; Reise der osterreichischen Fregatte Novara um die Erde &quot; (Insecta: Coleoptera, Scarabaeidae, Scarabaeinae). Annalen des Naturhistorischen Museums in Wien 120: 41 - 58." type="journal article" year="2018">VAZ-DE-MELLO &amp; CUPELLO 2018</bibRefCitation>
), Castelnau’s first personal collection was donated to the today’s Smithsonian Institution (
<collectingCountry id="EB95760C8A2EFFB9FABFF8C7EC83F816" name="United States of America" pageId="4" pageNumber="451">United States</collectingCountry>
) in 1841, probably including the type series of species names established by
<bibRefCitation id="F7134B6D8A2EFFB9FBA8F807EA6CF876" author="CASTELNAU" box="[1149,1354,1965,1988]" firstAuthor="CASTELNAU" pageId="4" pageNumber="451" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">CASTELNAU (1840)</bibRefCitation>
, that of
<taxonomicName id="54824D1F8A2EFFB9FCE9F867EC86F856" authorityName="Castelnau" authorityYear="1840" box="[828,928,1997,2020]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="451" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2EFFB9FCE9F867EC86F856" box="[828,928,1997,2020]" italics="true" pageId="4" pageNumber="451">P. tridens</emphasis>
</taxonomicName>
among them. Subsequently, this collection was destroyed by a fire in 1865 (
<bibRefCitation id="F7134B6D8A2EFFB9FBBBF847EA3CF7B6" author="EVENHUIS" box="[1134,1306,2029,2052]" firstAuthor="EVENHUIS" pageId="4" pageNumber="451" pagination="53 - 63" refId="ref25820" refString="EVENHUIS N. L. 2012: Francois-Louis Comte de Castelnau (1802 - 1880) and the mysterious disappearance of his original insect collection. Zootaxa 3168: 53 - 63." type="journal article" year="2012">EVENHUIS 2012</bibRefCitation>
). Therefore, the original type series was most probably destroyed in that year. Consequently, a
<typeStatus id="4C39883E8A2FFFB8FEA9F882EEF6F88D" box="[380,464,1832,1855]" pageId="5" pageNumber="452" type="neotype">neotype</typeStatus>
deposited at NMVA (
<figureCitation id="0BB92A198A2FFFB8FD64F882EDC9F88D" box="[689,751,1832,1855]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="5" pageNumber="452">Fig. 5</figureCitation>
) was designated by
<bibRefCitation id="F7134B6D8A2FFFB8FEB4F8E2ED62F8ED" author="EDMONDS" box="[353,580,1864,1887]" firstAuthor="EDMONDS" page="53" pageId="5" pageNumber="452" 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: 53)</bibRefCitation>
.
</paragraph>
<caption id="C7FD66148A2FFFB8FF42F909EDFBF959" ID-DOI="http://doi.org/10.5281/zenodo.5821151" ID-Zenodo-Dep="5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="5" pageNumber="452" startId="5.[151,188,1699,1718]" targetBox="[153,1436,206,1678]" targetPageId="5">
<paragraph id="933D369C8A2FFFB8FF42F909EDFBF959" blockId="5.[151,1438,1699,1771]" pageId="5" pageNumber="452">
Figs 1–5.
<taxonomicName id="54824D1F8A2FFFB8FF3FF909ED28F904" authority="Castelnau, 1840" authorityName="Castelnau" authorityYear="1840" box="[234,526,1699,1718]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="452" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2FFFB8FF3FF909EE58F904" box="[234,382,1699,1718]" italics="true" pageId="5" pageNumber="452">Phanaeus tridens</emphasis>
<bibRefCitation id="F7134B6D8A2FFFB8FE56F909ED28F904" author="Castelnau" box="[387,526,1699,1718]" firstAuthor="Castelnau" pageId="5" pageNumber="452" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">Castelnau, 1840</bibRefCitation>
</taxonomicName>
. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 –
<taxonomicName id="54824D1F8A2FFFB8FE9AF917EEC2F962" authorityName="Balthasar" authorityYear="1939" box="[335,484,1725,1744]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="452" phylum="Arthropoda" rank="species" species="frankenbergeri">
<emphasis id="A1F6EA8E8A2FFFB8FE9AF917EEC2F962" box="[335,484,1725,1744]" italics="true" pageId="5" pageNumber="452">P. frankenbergeri</emphasis>
</taxonomicName>
junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – EDMONDS’ (1994) neotype of
<taxonomicName id="54824D1F8A2FFFB8FA58F917EFF4F959" authorityName="Castelnau" authorityYear="1840" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="452" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2FFFB8FA58F917EFF4F959" italics="true" pageId="5" pageNumber="452">P. tridens</emphasis>
</taxonomicName>
and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm.
</paragraph>
</caption>
<paragraph id="933D369C8A2FFFB8FF60F8C2EABBF7AD" blockId="5.[151,763,1832,2079]" lastBlockId="5.[828,1437,1832,2079]" pageId="5" pageNumber="452">
<bibRefCitation id="F7134B6D8A2FFFB8FF60F8C2EE78F8CD" author="ARNAUD" box="[181,350,1896,1919]" firstAuthor="ARNAUD" pageId="5" pageNumber="452" pagination="2 - 8" refId="ref24169" refString="ARNAUD P. 2001: Description de nouvelles especes de Phanaeides (Col. Scarabaeidae). Besoiro 6: 2 - 8." type="journal article" year="2001">ARNAUD (2001)</bibRefCitation>
suggested that the
<typeStatus id="4C39883E8A2FFFB8FDF1F8C2ED5EF8CD" box="[548,632,1896,1919]" pageId="5" pageNumber="452" type="neotype">neotype</typeStatus>
designation by
<bibRefCitation id="F7134B6D8A2FFFB8FF63F822EE4EF82D" author="EDMONDS" box="[182,360,1928,1951]" firstAuthor="EDMONDS" pageId="5" pageNumber="452" 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>
was inadequate, since the
<typeStatus id="4C39883E8A2FFFB8FDA0F822EDEEF82D" box="[629,712,1928,1951]" pageId="5" pageNumber="452" type="neotype">neotype</typeStatus>
specimen was labeled as coming from
<collectingCountry id="EB95760C8A2FFFB8FDDCF802ED65F80D" box="[521,579,1960,1983]" name="Chile" pageId="5" pageNumber="452">Chile</collectingCountry>
(
<figureCitation id="0BB92A198A2FFFB8FD84F802EDB6F80D" box="[593,656,1960,1983]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="5" pageNumber="452">Fig. 5</figureCitation>
), while
<taxonomicName id="54824D1F8A2FFFB8FD31F802EFF9F86D" authorityName="Castelnau" authorityYear="1840" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="452" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2FFFB8FD31F802EFF9F86D" italics="true" pageId="5" pageNumber="452">P. tridens</emphasis>
</taxonomicName>
is endemic to
<collectingCountry id="EB95760C8A2FFFB8FEBAF862EEE4F86D" box="[367,450,1992,2015]" name="Mexico" pageId="5" pageNumber="452">Mexico</collectingCountry>
.Accordingly,
<bibRefCitation id="F7134B6D8A2FFFB8FD87F862EDDEF86D" author="ARNAUD" box="[594,760,1992,2015]" firstAuthor="ARNAUD" pageId="5" pageNumber="452" pagination="2 - 8" refId="ref24169" refString="ARNAUD P. 2001: Description de nouvelles especes de Phanaeides (Col. Scarabaeidae). Besoiro 6: 2 - 8." type="journal article" year="2001">ARNAUD (2001)</bibRefCitation>
designated a new
<typeStatus id="4C39883E8A2FFFB8FE8FF842EE88F84D" box="[346,430,2024,2047]" pageId="5" pageNumber="452" type="neotype">neotype</typeStatus>
for
<taxonomicName id="54824D1F8A2FFFB8FE0BF842ED62F84D" authorityName="Castelnau" authorityYear="1840" box="[478,580,2024,2047]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="452" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2FFFB8FE0BF842ED62F84D" box="[478,580,2024,2047]" italics="true" pageId="5" pageNumber="452">P. tridens</emphasis>
</taxonomicName>
and commented that his designation would be accompanied by a request to the International Commission on Zoological Nomenclature (ICZN) to set aside EDMONDS’ S (1994)
<typeStatus id="4C39883E8A2FFFB8FB18F8E2EA02F8ED" box="[1229,1316,1864,1887]" pageId="5" pageNumber="452" type="neotype">neotype</typeStatus>
. Our search in the
<emphasis id="A1F6EA8E8A2FFFB8FC54F8C2EA2AF8CD" box="[897,1292,1896,1919]" italics="true" pageId="5" pageNumber="452">Bulletin of Zoological Nomenclature</emphasis>
for Arnaud’s request has been unsuccessful. Indeed, Patrick Arnaud personally confirmed to the first author of this work that he had probably never sent the request to the ICZN. Consequently, the
<typeStatus id="4C39883E8A2FFFB8FC05F842EB02F84D" box="[976,1060,2024,2047]" pageId="5" pageNumber="452" type="neotype">neotype</typeStatus>
designation by
<bibRefCitation id="F7134B6D8A2FFFB8FB13F842EA48F84D" author="ARNAUD" box="[1222,1390,2024,2047]" firstAuthor="ARNAUD" pageId="5" pageNumber="452" pagination="2 - 8" refId="ref24169" refString="ARNAUD P. 2001: Description de nouvelles especes de Phanaeides (Col. Scarabaeidae). Besoiro 6: 2 - 8." type="journal article" year="2001">ARNAUD (2001)</bibRefCitation>
was not made available and Edmonds’s one was not set aside.
</paragraph>
<paragraph id="933D369C8A2CFFBBFF60FF14EE5BFD47" blockId="6.[151,763,190,2069]" pageId="6" pageNumber="453">
According to the International Code of Zoological Nomenclature, a
<typeStatus id="4C39883E8A2CFFBBFE8DFF74EE88FF47" box="[344,430,222,245]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
designation is only available if it fulfils all the conditions under Article 75 (
<bibRefCitation id="F7134B6D8A2CFFBBFD63FF54EFF5FE87" author="ICZN" firstAuthor="ICZN" pageId="6" pageNumber="453" refId="ref26427" refString="ICZN [= INTERNATIONAL COMMISION ON ZOOLOGICAL NOMENCLATURE] 1985: International Code of Zoological Nomenclature. Third Edition. The International Trust for Zoological Nomenclature, London, 338 pp." type="book" year="1985">ICZN 1985</bibRefCitation>
). The
<typeStatus id="4C39883E8A2CFFBBFEF5FEB4EE5EFE87" box="[288,376,286,309]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
designation by
<bibRefCitation id="F7134B6D8A2CFFBBFDE3FEB4EDD1FE87" author="EDMONDS" box="[566,759,286,309]" firstAuthor="EDMONDS" pageId="6" pageNumber="453" 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>
is problematic because it does not comply with article 75.d.5 (evidence that the
<typeStatus id="4C39883E8A2CFFBBFE11FEF4ED3AFEC7" box="[452,540,350,373]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
came as nearly as practicable from the original type locality) of the Code. Indeed, the prevailing usage of the name
<taxonomicName id="54824D1F8A2CFFBBFDA4FE34EDFDFE07" authorityName="Castelnau" authorityYear="1840" box="[625,731,414,437]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFDA4FE34EDFDFE07" box="[625,731,414,437]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
is threatened by the identity of EDMONDS’ S (1994)
<typeStatus id="4C39883E8A2CFFBBFD77FE14EDDEFE67" box="[674,760,446,469]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
specimen which is correctly identified as
<taxonomicName id="54824D1F8A2CFFBBFDB1FE74EDEBFE47" authorityName="Castelnau" authorityYear="1840" box="[612,717,478,501]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFDB1FE74EDEBFE47" box="[612,717,478,501]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
but mislabelled, indicated to come from
<collectingCountry id="EB95760C8A2CFFBBFDF0FE54ED46FDA7" box="[549,608,510,533]" name="Chile" pageId="6" pageNumber="453">Chile</collectingCountry>
(
<figureCitation id="0BB92A198A2CFFBBFDBBFE54ED8BFDA7" box="[622,685,510,533]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Fig. 5</figureCitation>
) while this species is endemic to
<collectingCountry id="EB95760C8A2CFFBBFE7DFDB4EEDCFD87" box="[424,506,542,565]" name="Mexico" pageId="6" pageNumber="453">Mexico</collectingCountry>
(
<bibRefCitation id="F7134B6D8A2CFFBBFDDDFDB4EDE2FD87" author="CASTELNAU" box="[520,708,542,565]" firstAuthor="CASTELNAU" pageId="6" pageNumber="453" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">CASTELNAU 1840</bibRefCitation>
:
<figureCitation id="0BB92A198A2CFFBBFD18FDB4EF90FDE7" captionStart="Fig" captionStartId="28.[151,182,953,972]" captionTargetBox="[162,1425,196,930]" captionTargetId="figure-13@28.[162,1425,196,930]" captionTargetPageId="28" captionText="Fig. 64. Predicted distribution of P. tridens Castelnau, 1840, P. moroni Arnaud, 2001 and P. balthasari Arnaud, 2001. The distribution of P. coeruleus Bates, 1887 was not modelled because it is only known from a single locality that needs confirmation." figureDoi="http://doi.org/10.5281/zenodo.5821195" httpUri="https://zenodo.org/record/5821195/files/figure.png" pageId="6" pageNumber="453">Fig. 64</figureCitation>
). In order to clarify the application of the name
<taxonomicName id="54824D1F8A2CFFBBFD31FD94EFC5FDC7" authorityName="Castelnau" authorityYear="1840" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFD31FD94EFC5FDC7" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
and to conserve its current usage, we suggest to designate a
<typeStatus id="4C39883E8A2CFFBBFEC0FDD4EE4DFD27" box="[277,363,638,661]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
(
<figureCitation id="0BB92A198A2CFFBBFEACFDD4EE9CFD27" box="[377,442,638,661]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Fig. 3</figureCitation>
). In addition, this publication is followed by a request under Article 75.6 to the
<bibRefCitation id="F7134B6D8A2CFFBBFD6CFD34EFC6FD67" author="ICZN" firstAuthor="ICZN" pageId="6" pageNumber="453" 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>
to set aside EDMONDS’ S (1994)
<typeStatus id="4C39883E8A2CFFBBFD90FD14EDBDFD67" box="[581,667,702,725]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
(
<figureCitation id="0BB92A198A2CFFBBFD79FD14EDC9FD67" box="[684,751,702,725]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Fig. 5</figureCitation>
) by its plenary power.
</paragraph>
<paragraph id="933D369C8A2CFFBBFF60FD54EE8FFB47" blockId="6.[151,763,190,2069]" pageId="6" pageNumber="453">
The characters for which
<taxonomicName id="54824D1F8A2CFFBBFE17FD54ED00FCA7" authorityName="Castelnau" authorityYear="1840" box="[450,550,766,789]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFE17FD54ED00FCA7" box="[450,550,766,789]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
is differentiated are provided in the diagnosis of this species. The designated
<typeStatus id="4C39883E8A2CFFBBFF14FC94EE33FCE7" box="[193,277,830,853]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
is a major male housed at MNHN (MNHN EC10552,
<figureCitation id="0BB92A198A2CFFBBFEDFFCF4EE6CFCC7" box="[266,330,862,885]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Fig. 3</figureCitation>
). The
<typeStatus id="4C39883E8A2CFFBBFE45FCF4EEC2FCC7" box="[400,484,862,885]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
specimen (
<figureCitation id="0BB92A198A2CFFBBFD8AFCF4EDB9FCC7" box="[607,671,862,885]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Fig. 3</figureCitation>
) fits the original description of
<taxonomicName id="54824D1F8A2CFFBBFE54FCD4ED88FC27" authority="(CASTELNAU 1840)" baseAuthorityName="CASTELNAU" baseAuthorityYear="1840" box="[385,686,894,917]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFE54FCD4EEC4FC27" box="[385,482,894,917]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
(
<bibRefCitation id="F7134B6D8A2CFFBBFE3BFCD4ED80FC27" author="CASTELNAU" box="[494,678,894,917]" firstAuthor="CASTELNAU" pageId="6" pageNumber="453" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">CASTELNAU 1840</bibRefCitation>
)
</taxonomicName>
: “D’un vert métallique éclatant; devant du chaperon et une longue corne arquée sur la tête noirs; corselet rugueux, profondément excavé dans son milieu, avec deux cornes latérales et une postérieure au milieu; élytres lisses, striées, avec des enfoncemens à la base des quatre premières stries; jambes et tarses noirs. —
<collectingCountry id="EB95760C8A2CFFBBFE87FB94EE95FBE7" box="[338,435,1086,1109]" name="Mexico" pageId="6" pageNumber="453">Mexique</collectingCountry>
” [translation to English: bright metallic green species; male with a large horn curved over the head; roughened pronotum, deeply excavated medially, with two lateral horns and a posteromedial horn; flattened elytra; impressed fossae at bases of the first four elytral striae; Mexican species)].
</paragraph>
<paragraph id="933D369C8A2CFFBBFF60FB54ED35F867" blockId="6.[151,763,190,2069]" pageId="6" pageNumber="453">
Nevertheless, at least four related species (
<taxonomicName id="54824D1F8A2CFFBBFD79FB54EFC2FA87" authorityName="Arnaud" authorityYear="2001" baseAuthorityName="Arnaud" baseAuthorityYear="2001" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="balthasari">
<emphasis id="A1F6EA8E8A2CFFBBFD79FB54EFC2FA87" italics="true" pageId="6" pageNumber="453">P. balthasari</emphasis>
</taxonomicName>
,
<figureCitation id="0BB92A198A2CFFBBFF20FAB4EE7FFA87" box="[245,345,1310,1333]" captionStart="Figs 8–9" captionStartId="8.[151,187,1301,1320]" captionTargetBox="[151,1395,200,1349]" captionTargetId="figure-13@8.[183,1405,189,1353]" captionTargetPageId="8" captionText="Figs 8–9.Phanaeus balthasari Arnaud,2001, stat.rev. 8 – male green phase;9 – holotype" figureDoi="http://doi.org/10.5281/zenodo.5821156" httpUri="https://zenodo.org/record/5821156/files/figure.png" pageId="6" pageNumber="453">Figs 8–9</figureCitation>
;
<taxonomicName id="54824D1F8A2CFFBBFEBDFAB4EEFDFA87" baseAuthorityName="EDMONDS" baseAuthorityYear="1994" box="[360,475,1310,1333]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="daphnis">
<emphasis id="A1F6EA8E8A2CFFBBFEBDFAB4EEFDFA87" box="[360,475,1310,1333]" italics="true" pageId="6" pageNumber="453">P. daphnis</emphasis>
</taxonomicName>
,
<figureCitation id="0BB92A198A2CFFBBFE39FAB4ED49FA87" box="[492,623,1310,1333]" captionStart="Figs 10–12" captionStartId="9.[760,797,2010,2029]" captionTargetBox="[166,1438,335,2082]" captionTargetId="figure-13@9.[151,1436,335,2082]" captionTargetPageId="9" captionText="Figs 10–12. 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." figureDoi="http://doi.org/10.5281/zenodo.5821158" httpUri="https://zenodo.org/record/5821158/files/figure.png" pageId="6" pageNumber="453">Figs 10, 12</figureCitation>
;
<taxonomicName id="54824D1F8A2CFFBBFDA8FAB4EDD7FA87" authorityName="Bates" authorityYear="1887" baseAuthorityName="Bates" baseAuthorityYear="1887" box="[637,753,1310,1333]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="herbeus">
<emphasis id="A1F6EA8E8A2CFFBBFDA8FAB4EDD7FA87" box="[637,753,1310,1333]" italics="true" pageId="6" pageNumber="453">P. herbeus</emphasis>
</taxonomicName>
,
<figureCitation id="0BB92A198A2CFFBBFF42FA94EE32FAE7" box="[151,276,1342,1365]" captionStart="Figs 16–18" captionStartId="13.[827,864,1721,1740]" captionTargetBox="[163,1438,199,1820]" captionTargetId="figure-13@13.[151,1436,199,1825]" captionTargetPageId="13" captionText="Figs 16–18. Phanaeus herbeus Bates, 1887, stat. rev. 16 – male green phase; 17 – holotype and labels (by Mario Cupello, UFPR); 18 – P. tricornis Olsoufieff, 1924, junior subjective synonymy, redrawn from Oඅඌඈඎൿංൾൿൿ (1924). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821162" httpUri="https://zenodo.org/record/5821162/files/figure.png" pageId="6" pageNumber="453">Figs 16–17</figureCitation>
; and the one to which the name
<taxonomicName id="54824D1F8A2CFFBBFDA2FA94EDF8FAE7" authorityName="Castelnau" authorityYear="1840" baseAuthorityName="CASTELNAU" baseAuthorityYear="1840" box="[631,734,1342,1365]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFDA2FA94EDF8FAE7" box="[631,734,1342,1365]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
is traditionally applied,
<figureCitation id="0BB92A198A2CFFBBFEABFAF4EED0FAC7" box="[382,502,1374,1397]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Figs 1, 3–5</figureCitation>
) fit the original description, being metallic green in colour and from
<collectingCountry id="EB95760C8A2CFFBBFD52FAD4EDFCFA27" box="[647,730,1406,1429]" name="Mexico" pageId="6" pageNumber="453">Mexico</collectingCountry>
as mentioned by
<bibRefCitation id="F7134B6D8A2CFFBBFEE7FA34ED24FA07" author="CASTELNAU" box="[306,514,1438,1461]" firstAuthor="CASTELNAU" pageId="6" pageNumber="453" refId="ref24732" refString="CASTELNAU C. D. 1840: Histoire naturelle des coleopteres. Histoire Naturelle des Animaux Articules, Annelides, Crustaces, Arachnides, Myriapodes et Insectes. Tome deuxieme. P. Dumenil, Paris, 583 pp." type="book" year="1840">CASTELNAU (1840)</bibRefCitation>
. This problem is made worse by the fact that details about the sizes, shapes and positions of the pronotal ornaments are not mentioned by him, which makes it impossible to determine what species Castelnau’s specimen actually belonged to.
<bibRefCitation id="F7134B6D8A2CFFBBFD49F9B4EFC6F9E7" author="HAROLD" firstAuthor="HAROLD" pageId="6" pageNumber="453" 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>
redescribed
<taxonomicName id="54824D1F8A2CFFBBFEBEF994EEF4F9E7" authorityName="Castelnau" authorityYear="1840" baseAuthorityName="CASTELNAU" baseAuthorityYear="1840" box="[363,466,1598,1621]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFEBEF994EEF4F9E7" box="[363,466,1598,1621]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
and suggested
<collectingRegion id="5146F87E8A2CFFBBFDACF994EDFCF9E7" box="[633,730,1598,1621]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="6" pageNumber="453">Veracruz</collectingRegion>
as the distribution area of this species, while
<bibRefCitation id="F7134B6D8A2CFFBBFDB2F9F4EDD1F9C7" author="BATES" box="[615,759,1630,1653]" firstAuthor="BATES" pageId="6" pageNumber="453" 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>
provided accurate localities from the north-central region of
<collectingRegion id="5146F87E8A2CFFBBFF67F934EE33F907" box="[178,277,1694,1717]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="6" pageNumber="453">Veracruz</collectingRegion>
. The only one of those four species to occur in north-central
<collectingRegion id="5146F87E8A2CFFBBFEFDF914EEAEF967" box="[296,392,1726,1749]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="6" pageNumber="453">Veracruz</collectingRegion>
(
<figureCitation id="0BB92A198A2CFFBBFE40F914EEC4F967" box="[405,482,1726,1749]" captionStart="Fig" captionStartId="28.[151,182,953,972]" captionTargetBox="[162,1425,196,930]" captionTargetId="figure-13@28.[162,1425,196,930]" captionTargetPageId="28" captionText="Fig. 64. Predicted distribution of P. tridens Castelnau, 1840, P. moroni Arnaud, 2001 and P. balthasari Arnaud, 2001. The distribution of P. coeruleus Bates, 1887 was not modelled because it is only known from a single locality that needs confirmation." figureDoi="http://doi.org/10.5281/zenodo.5821195" httpUri="https://zenodo.org/record/5821195/files/figure.png" pageId="6" pageNumber="453">Fig. 64</figureCitation>
) is indeed the one that has been traditionally called
<taxonomicName id="54824D1F8A2CFFBBFE71F974ED2CF947" authorityName="Castelnau" authorityYear="1840" baseAuthorityName="CASTELNAU" baseAuthorityYear="1840" box="[420,522,1758,1781]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFE71F974ED2CF947" box="[420,522,1758,1781]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
(or
<taxonomicName id="54824D1F8A2CFFBBFDE3F974EDC9F947" authority="tridens" authorityName="tridens (EDMONDS" authorityYear="1994" box="[566,751,1758,1781]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="subSpecies" species="tridens" subSpecies="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFDE3F974EDC9F947" box="[566,751,1758,1781]" italics="true" pageId="6" pageNumber="453">P. tridens tridens</emphasis>
</taxonomicName>
) for more than a century in collections and in literature (e.g.
<bibRefCitation id="F7134B6D8A2CFFBBFF1AF8B4EE4DF887" author="HAROLD" box="[207,363,1822,1845]" firstAuthor="HAROLD" pageId="6" pageNumber="453" 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>
,
<bibRefCitation id="F7134B6D8A2CFFBBFEA2F8B4EEDEF887" author="BATES" box="[375,504,1822,1845]" firstAuthor="BATES" pageId="6" pageNumber="453" 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>
,
<bibRefCitation id="F7134B6D8A2CFFBBFDD6F8B4EDE5F887" author="OLSOUFIEFF" box="[515,707,1822,1845]" firstAuthor="OLSOUFIEFF" pageId="6" pageNumber="453" pagination="4 - 172" refId="ref27631" 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</bibRefCitation>
,
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,
<bibRefCitation id="F7134B6D8A2CFFBBFEF4F894EEEAF8E7" author="EDMONDS" box="[289,460,1854,1877]" firstAuthor="EDMONDS" pageId="6" pageNumber="453" pagination="1 - 105" refId="ref25450" refString="EDMONDS W. D. 1994: Revision of Phanaeus MacLeay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science of the Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">EDMONDS 1994</bibRefCitation>
,
<bibRefCitation id="F7134B6D8A2CFFBBFE0DF894EDCAF8E7" author="EDMONDS &amp; ZIDEK" box="[472,748,1854,1877]" firstAuthor="EDMONDS" pageId="6" pageNumber="453" pagination="1 - 108" refId="ref25590" refString="EDMONDS W. D. &amp; ZIDEK J. 2012: Taxonomy of Phanaeus revisited: Revised keys to and comments on species of the New World dung beetle genus Phanaeus MacLeay, 1819 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Insecta Mundi 274: 1 - 108." type="journal article" year="2012">EDMONDS &amp; ZÍDEK 2012</bibRefCitation>
). Consequently, we believe that designating a specimen from a north-central
<collectingRegion id="5146F87E8A2CFFBBFE50F8D4EEC0F827" box="[389,486,1918,1941]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="6" pageNumber="453">Veracruz</collectingRegion>
population (such as the Palma Sola one,
<figureCitation id="0BB92A198A2CFFBBFE86F834EEB2F807" box="[339,404,1950,1973]" captionStart="Figs 1–5" captionStartId="5.[151,188,1699,1718]" captionTargetBox="[153,1436,206,1678]" captionTargetId="figure-13@5.[151,1436,205,1678]" captionTargetPageId="5" captionText="Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5821151" httpUri="https://zenodo.org/record/5821151/files/figure.png" pageId="6" pageNumber="453">Fig. 3</figureCitation>
) as the
<typeStatus id="4C39883E8A2CFFBBFE3BF834ED62F807" box="[494,580,1950,1973]" pageId="6" pageNumber="453" type="neotype">neotype</typeStatus>
for
<taxonomicName id="54824D1F8A2CFFBBFDA0F834EDFBF807" authorityName="Castelnau" authorityYear="1840" baseAuthorityName="CASTELNAU" baseAuthorityYear="1840" box="[629,733,1950,1973]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="453" phylum="Arthropoda" rank="species" species="tridens">
<emphasis id="A1F6EA8E8A2CFFBBFDA0F834EDFBF807" box="[629,733,1950,1973]" italics="true" pageId="6" pageNumber="453">P. tridens</emphasis>
</taxonomicName>
is appropriate and promotes stability.
</paragraph>
</subSubSection>
<subSubSection id="DB9865178A2CFFBBFF42F874EFE4F7A7" pageId="6" pageNumber="453" type="distribution">
<paragraph id="933D369C8A2CFFBBFF42F874EFE4F7A7" blockId="6.[151,763,190,2069]" pageId="6" pageNumber="453">
<emphasis id="A1F6EA8E8A2CFFBBFF42F874EE0CF847" bold="true" box="[151,298,2014,2037]" pageId="6" pageNumber="453">Distribution.</emphasis>
Northern to central
<collectingRegion id="5146F87E8A2CFFBBFDD5F874ED44F847" box="[512,610,2014,2037]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="6" pageNumber="453">Veracruz</collectingRegion>
,
<collectingCountry id="EB95760C8A2CFFBBFDB9F874ED9BF847" box="[620,701,2014,2037]" name="Mexico" pageId="6" pageNumber="453">Mexico</collectingCountry>
(
<figureCitation id="0BB92A198A2CFFBBFD19F874EF93F7A7" captionStart="Fig" captionStartId="28.[151,182,953,972]" captionTargetBox="[162,1425,196,930]" captionTargetId="figure-13@28.[162,1425,196,930]" captionTargetPageId="28" captionText="Fig. 64. Predicted distribution of P. tridens Castelnau, 1840, P. moroni Arnaud, 2001 and P. balthasari Arnaud, 2001. The distribution of P. coeruleus Bates, 1887 was not modelled because it is only known from a single locality that needs confirmation." figureDoi="http://doi.org/10.5281/zenodo.5821195" httpUri="https://zenodo.org/record/5821195/files/figure.png" pageId="6" pageNumber="453">Fig. 64</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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