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<document id="DBCD269894AF0C3F65038144AF2BBC99" ID-DOI="10.11646/zootaxa.5512.3.4" ID-ISSN="1175-5326" ID-Zenodo-Dep="13848909" ID-ZooBank="39FED4E6-2067-4D95-AA9F-FF3D8449019A" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1727439787925" checkinUser="plazi" docAuthor="Celante, Gabriel Luiz, Carvalho, Adhan Gabriel, Jahyny, Benoit Jean Bernard, Araújo, Kaylla Brisley Silva &amp; Santos, Paula Batista Dos" docDate="2024" docId="03EAE8783518FFA5C6FCFF60FDCF1C5E" docLanguage="en" docName="zootaxa.5512.3.4.pdf" docOrigin="Zootaxa 5512 (3)" docSource="http://dx.doi.org/10.11646/zootaxa.5512.3.4" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Athyreus arretado Celante &amp; Carvalho &amp; Jahyny &amp; Araújo &amp; Santos 2024, new species" docType="treatment" docUuid="1CA6E1C3-F6DE-452F-A177-AA2730DEC2CF" docUuidSource="ZooBank" docVersion="2" lastPageNumber="394" masterDocId="FFD39000351BFFA0C66BFFF7FFA01B72" masterDocTitle="A new species of Athyreus MacLeay, 1819 (Coleoptera: Geotrupidae: Bolboceratinae: Athyreini) from northeastern Brazil" masterLastPageNumber="401" masterPageNumber="389" pageNumber="392" updateTime="1727440894639" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="2991966CDF0F4B79B30C7E2D748C24A4">A new species of Athyreus MacLeay, 1819 (Coleoptera: Geotrupidae: Bolboceratinae: Athyreini) from northeastern Brazil</mods:title>
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<mods:namePart id="08C18F9BB5E73610B18B1AA19B813DE5">Celante, Gabriel Luiz</mods:namePart>
<mods:affiliation id="E07FA6DA19676035A592D761130A631F">Laboratório de Mirmecologia do Sertão, Universidade Federal do Vale do São Francisco, Petrolina, Pernambuco, Brazil &amp; Programa de pós-graduação Ciências da Saúde e Biológicas, Universidade Federal do Vale do São Francisco, Petrolina, Pernambuco, Brazil</mods:affiliation>
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<mods:namePart id="6C48155480D46B6E4A04A9D46440826C">Carvalho, Adhan Gabriel</mods:namePart>
<mods:affiliation id="DCC2DBD9C3683FC63245E882018D1EC0">Laboratório de Mirmecologia do Sertão, Universidade Federal do Vale do São Francisco, Petrolina, Pernambuco, Brazil</mods:affiliation>
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<mods:namePart id="1D2ABF1BC93285B5E471A383D7A938E0">Jahyny, Benoit Jean Bernard</mods:namePart>
<mods:affiliation id="B1613B43461792CB6E5D1B0C58BD014E">Laboratório de Mirmecologia do Sertão, Universidade Federal do Vale do São Francisco, Petrolina, Pernambuco, Brazil</mods:affiliation>
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<mods:namePart id="FCEEAB03C783C18BE93DB4ECB62F532F">Araújo, Kaylla Brisley Silva</mods:namePart>
<mods:affiliation id="C5B77186E6AC80164519B0D30BE3916D">Museu de Fauna da Caatinga, Centro de Conservação e Manejo de Fauna, Universidade Federal do Vale do São Francisco, Petrolina, Pernambuco, Brazil</mods:affiliation>
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<mods:namePart id="BE48ADC9554C199AFED2B9E671E92814">Santos, Paula Batista Dos</mods:namePart>
<mods:affiliation id="AD0E52E641163168AB41079E46ECE599">Museu de Fauna da Caatinga, Centro de Conservação e Manejo de Fauna, Universidade Federal do Vale do São Francisco, Petrolina, Pernambuco, Brazil</mods:affiliation>
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<mods:date id="D1CD368D4BA43A0FE434BD1616ECB4EE">2024</mods:date>
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<taxonomicName id="4C4322ED3518FFA3C6FCFF60FDD81BC3" authority="Celante &amp; Carvalho" authorityName="Celante &amp; Carvalho &amp; Jahyny &amp; Araújo &amp; Santos" authorityYear="2024" box="[151,632,151,178]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="392" phylum="Arthropoda" rank="species" species="arretado" status="sp. nov.">
<emphasis id="B937857C3518FFA3C6FCFF60FED31BC0" bold="true" box="[151,371,151,178]" italics="true" pageId="3" pageNumber="392">Athyreus arretado</emphasis>
Celante &amp; Carvalho
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,
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<paragraph id="8BFC596E3518FFA3C6FCFF4CFB101B88" blockId="3.[151,1200,151,250]" pageId="3" pageNumber="392">
<figureCitation id="137845EB3518FFA3C6FCFF4CFF4F1BA7" box="[151,239,187,213]" captionStart="FIGURE 2" captionStartId="2.[152,255,1833,1857]" captionTargetBox="[151,1435,525,1809]" captionTargetId="figure-165@2.[151,1435,525,1809]" captionTargetPageId="2" captionText="FIGURE 2AD. Collecting localities.A, São Francisco River Sand Dunes in Casa Nova, Bahia, Brazil; B, Caatinga in Petrolina, Pernambuco, Brazil; C, flight intercept trap in Refúgio de Vida Silvestre Mata do Buraquinho, João Pessoa, Paraíba, Brazil; D, Reserva Biológica Guaribas, Mamanguape, Paraíba, Brazil." figureDoi="http://doi.org/10.5281/zenodo.13848915" httpUri="https://zenodo.org/record/13848915/files/figure.png" pageId="3" pageNumber="392">Fig. 2A</figureCitation>
,
<figureCitation id="137845EB3518FFA3C690FF4CFEE21BA7" box="[251,322,187,213]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">3AG</figureCitation>
,
<figureCitation id="137845EB3518FFA3C726FF4CFE2E1BA7" box="[333,398,187,213]" captionStart="FIGURE 4" captionStartId="5.[152,255,1343,1367]" captionTargetBox="[161,1424,192,1320]" captionTargetId="figure-20@5.[151,1436,181,1320]" captionTargetPageId="5" captionText="FIGURE 4AF. Athyreus arretado new species aedeagus. AB, ventral view; CD, dorsal view; EF, lateral view. Scale bar: A, C, E: 1.0 mm; B, D, F: 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848921" httpUri="https://zenodo.org/record/13848921/files/figure.png" pageId="3" pageNumber="392">4AF</figureCitation>
,
<figureCitation id="137845EB3518FFA3C7F2FF4CFE1A1BA4" box="[409,442,187,214]" captionStart="FIGURE 5" captionStartId="6.[152,255,1716,1740]" captionTargetBox="[151,1435,181,1692]" captionTargetId="figure-24@6.[151,1435,181,1692]" captionTargetPageId="6" captionText="FIGURE 5AD. Distribution records of some Bolboceratinae species. A, Athyreus arretado new species, Athyreus vavini Boucomont, 1902 and Athyreus tuberifer Felsche, 1909 distribution records; B, Neoathyreus anthracinus (Klug, 1843) and Neoathyreus braziliensis Howden, 1985 distribution records; C, Neoathyreus centromaculatus (Felsche, 1909), Neoathyreus lanuginosus (Klug, 1843), and Parathyreus trituberculatus (Klug, 1843) distribution records; D, Bolbapium caesum (Klug, 1843), Bolbapium parcepunctatum (Arrow, 1913), and Bolbapium sulcifrons Ide &amp; Martínez, 1993 distribution records. The black symbols represent new records, the unfilled symbols are literature records." figureDoi="http://doi.org/10.5281/zenodo.13848925" httpUri="https://zenodo.org/record/13848925/files/figure.png" pageId="3" pageNumber="392">5A</figureCitation>
http://zoobank.org/
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<emphasis id="B937857C3518FFA3C6FCFED0FE9E1A33" bold="true" box="[151,318,295,321]" pageId="3" pageNumber="392">Type material</emphasis>
.
<materialsCitation id="3B2B53333518FFA3C720FED0FBB81A17" collectingDate="2023-01-27" collectionCode="MFCE" collectorName="Celante, G." country="Brazil" location="Areia Grande" municipality="Casa Nova" pageId="3" pageNumber="392" specimenCode="MFCE014265" specimenCount="1" stateProvince="Bahia" typeStatus="holotype">
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male,
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<collectingRegion id="4987978C3518FFA3C5ECFED0FC431A33" box="[903,995,295,321]" country="Brazil" name="Bahia" pageId="3" pageNumber="392">BAHIA</collectingRegion>
,
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,
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, LMS16AB,
<date id="FFFD7FAE3518FFA3C6FCFEBCFEA41A17" box="[151,260,331,357]" pageId="3" pageNumber="392" value="2023-01-27">
<collectingDate id="EFB986463518FFA3C6FCFEBCFEA41A17" box="[151,260,331,357]" pageId="3" pageNumber="392" value="2023-01-27">27.i.2023</collectingDate>
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,
<collectorName id="26B63CB83518FFA3C779FEBCFE331A17" box="[274,403,331,357]" pageId="3" pageNumber="392">Celante, G.</collectorName>
leg.” (
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, deposited at
<collectionCode id="ED52C1AB3518FFA3C542FEBCFCD71A17" box="[809,887,331,357]" pageId="3" pageNumber="392">MFCE</collectionCode>
) (
<figureCitation id="137845EB3518FFA3C5FAFEBCFBB01A17" box="[913,1040,331,357]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3AH</figureCitation>
)
</materialsCitation>
.
<materialsCitation id="3B2B53333518FFA3C24DFEBCFC861ADF" collectingDate="2024-02-17" collectionCode="INPA" country="Brazil" location="BAHIA" pageId="3" pageNumber="392" specimenCode="MFCE018115" specimenCount="1" stateProvince="Bahia" typeStatus="paratype">
<emphasis id="B937857C3518FFA3C24DFEBCFB331A17" bold="true" box="[1062,1171,331,357]" pageId="3" pageNumber="392">
<typeStatus id="54F8E7CC3518FFA3C24DFEBCFB331A17" box="[1062,1171,331,357]" pageId="3" pageNumber="392" type="paratype">Paratype</typeStatus>
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male,
<collectingCountry id="F35419FE3518FFA3C28FFEBCFAEE1A17" box="[1252,1358,331,357]" name="Brazil" pageId="3" pageNumber="392">BRAZIL</collectingCountry>
: “24 L 241461 8960156,
<collectingRegion id="4987978C3518FFA3C70CFE98FE631AFB" box="[359,451,367,393]" country="Brazil" name="Bahia" pageId="3" pageNumber="392">BAHIA</collectingRegion>
, Casa Nova, Areia Grande, LMS98AB,
<date id="FFFD7FAE3518FFA3C5FFFE98FBA81AFB" box="[916,1032,367,393]" pageId="3" pageNumber="392" value="2024-02-17">
<collectingDate id="EFB986463518FFA3C5FFFE98FBA81AFB" box="[916,1032,367,393]" pageId="3" pageNumber="392" value="2024-02-17">17.ii.2024</collectingDate>
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, Celante, G., Gurgel, M., Melo. A, Silva-Junior, A. legs.” (
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, deposited at
<collectionCode id="ED52C1AB3518FFA3C4B2FE64FCBD1ADF" box="[729,797,403,429]" country="Brazil" httpUri="http://grbio.org/cool/rm88-499z" name="Instituto Nacional de Pesquisas da Amazonia" pageId="3" pageNumber="392">INPA</collectionCode>
)
</materialsCitation>
.
</paragraph>
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<subSubSection id="C3590AE53518FFA3C6ACFE40FEDF19F7" pageId="3" pageNumber="392" type="diagnosis">
<paragraph id="8BFC596E3518FFA3C6ACFE40FEDF19F7" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACFE40FEE21AA3" bold="true" box="[199,322,439,465]" pageId="3" pageNumber="392">Diagnosis.</emphasis>
This species can be distinguished from other
<taxonomicName id="4C4322ED3518FFA3C536FE40FBBB1AA3" box="[861,1051,439,465]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C536FE40FC601AA3" box="[861,960,439,465]" italics="true" pageId="3" pageNumber="392">Athyreus</emphasis>
species
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by the following combination of characters: clypeal horn absent (
<figureCitation id="137845EB3518FFA3C46EFE2CFD651A87" box="[517,709,475,501]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, CE, G</figureCitation>
); pronotum with only one horn in the middle of anterior margin (
<figureCitation id="137845EB3518FFA3C6F4FE08FEFA196B" box="[159,346,511,537]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, CE, G</figureCitation>
); presence of a pair of central carinae that begin in the anterior region of the pronotum that extend divergently to the medial region, where they change direction, extending convergently to the posterior region where they curve towards the center of the pronotum, giving a “J” pattern (
<figureCitation id="137845EB3518FFA3C5CCFDB0FB851913" box="[935,1061,583,609]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, D</figureCitation>
); protibia with six teeth on outer margin (
<figureCitation id="137845EB3518FFA3C69EFD9CFED219F7" box="[245,370,619,645]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3D, G</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C3590AE53518FFA3C6ACFD78FE3C1D23" pageId="3" pageNumber="392" type="description">
<paragraph id="8BFC596E3518FFA3C6ACFD78FB9D19D8" blockId="3.[151,1437,295,1870]" box="[199,1085,655,682]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACFD78FE7419DB" bold="true" box="[199,468,655,681]" pageId="3" pageNumber="392">
Description.
<typeStatus id="54F8E7CC3518FFA3C70BFD78FE6F19DB" box="[352,463,655,681]" pageId="3" pageNumber="392" type="holotype">Holotype</typeStatus>
.
</emphasis>
Male (
<figureCitation id="137845EB3518FFA3C448FD78FD0119DB" box="[547,673,655,681]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3AH</figureCitation>
). Length:
<quantity id="4CBBF48B3518FFA3C579FD78FCD819D8" box="[786,888,655,682]" metricMagnitude="-2" metricUnit="m" metricValue="2.36" pageId="3" pageNumber="392" unit="mm" value="23.6">23.6 mm</quantity>
, width:
<quantity id="4CBBF48B3518FFA3C5B9FD78FB9919D8" box="[978,1081,655,682]" metricMagnitude="-2" metricUnit="m" metricValue="1.45" pageId="3" pageNumber="392" unit="mm" value="14.5">14.5 mm</quantity>
.
</paragraph>
<paragraph id="8BFC596E3518FFA3C6ACFD44FE9C1983" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACFD44FEB019BF" bold="true" box="[199,272,691,717]" pageId="3" pageNumber="392">Color.</emphasis>
Dorsally brown, head and pronotum reddish brown, elytra dark brown; laterally slightly lighter; ventrally orange-brown.
</paragraph>
<paragraph id="8BFC596E3518FFA3C6ACFD0CFDB51F47" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACFD0CFEAE1867" bold="true" box="[199,270,763,789]" pageId="3" pageNumber="392">Head.</emphasis>
Labrum 2.3x wider than long, in dorsal view, lateral margins rounded, anterior margin concave, surface irregularly punctate (
<figureCitation id="137845EB3518FFA3C7EBFCE8FE56184B" box="[384,502,799,825]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, E</figureCitation>
). Clypeus without a central horn (
<figureCitation id="137845EB3518FFA3C502FCE8FB81184B" box="[873,1057,799,825]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, CE, G</figureCitation>
); in dorsal view, presence of a pair of carinae that originate, posteriorly, close to the height of the insertion of the antennae, extending obliquely towards the center of the anterior margin of the clypeus, without being connected (
<figureCitation id="137845EB3518FFA3C588FC90FB9B18F3" box="[995,1083,871,897]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3E</figureCitation>
); in lateral view, central region elevated, with a triangular appearance, approximately the height of the apex of the mandibles (
<figureCitation id="137845EB3518FFA3C2A6FC7CFA8718D7" box="[1229,1319,907,933]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3C</figureCitation>
). Gena, in dorsal view, anteriorly and posteriorly concave; ocular canthus with rounded apex; carinate margins, starting at the clypeal carina and surrounding the laterodorsal margin of the head to the anterior region (
<figureCitation id="137845EB3518FFA3C2EFFC24FB7C189F" box="[1156,1244,979,1005]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3E</figureCitation>
). Front, in dorsal view, with a pair of carinae that begin at the proximal anterior angle of the gena and extend obliquely towards the vertex, without meeting (
<figureCitation id="137845EB3518FFA3C7DAFBECFDA81F47" box="[433,520,1051,1077]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3E</figureCitation>
).
</paragraph>
<paragraph id="8BFC596E3518FFA3C6ACFBC8FC5F1D7B" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACFBC8FEE71F2B" bold="true" box="[199,327,1087,1113]" pageId="3" pageNumber="392">Pronotum.</emphasis>
Middle of anterior margin with a single horn, directed upwards (
<figureCitation id="137845EB3518FFA3C270FBC8FB771F2B" box="[1051,1239,1087,1113]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, CE, G</figureCitation>
); posterior region with a groove extending from behind the central horn to a concavity located in the anterior region (
<figureCitation id="137845EB3518FFA3C297FB94FADA1F0F" box="[1276,1402,1123,1149]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, G</figureCitation>
); a carina extends around the outer margins, thicker in the posterior region (
<figureCitation id="137845EB3518FFA3C5A2FB70FB831FD3" box="[969,1059,1159,1185]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A</figureCitation>
); a pair of central carinae on disc extend from the base of the anterior pronotal horn divergently to the medial region, where they change direction, extending convergently to the region posterior where they curve towards the center of the pronotum giving a “J” appearance (
<figureCitation id="137845EB3518FFA3C74DFB04FE6C1E7F" box="[294,460,1267,1294]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A, D, G</figureCitation>
); median region of the central carinae with a tooth in lateral view (where the carinae change direction, from divergent to convergent) (
<figureCitation id="137845EB3518FFA3C4A0FAE0FC861E43" box="[715,806,1303,1329]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3C</figureCitation>
); posterior region with two trapezoidal lobes in frontal view, slightly exceeding the level of the elytra (
<figureCitation id="137845EB3518FFA3C4ABFACCFCBC1E27" box="[704,796,1339,1365]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3D</figureCitation>
); cavity of the procoxae connected to the most anterior margin of the pronotum by a carina (
<figureCitation id="137845EB3518FFA3C429FAA8FD3A1E0B" box="[578,666,1375,1401]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3F</figureCitation>
).
<emphasis id="B937857C3518FFA3C4DBFAA8FCA31E0B" bold="true" box="[688,771,1375,1401]" pageId="3" pageNumber="392">Elytra.</emphasis>
Six striae between the base of the elytra and the humeral callus (
<figureCitation id="137845EB3518FFA3C68CFA74FEE01EEF" box="[231,320,1411,1437]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3A</figureCitation>
).
<emphasis id="B937857C3518FFA3C738FA74FE321EEF" bold="true" box="[339,402,1411,1437]" pageId="3" pageNumber="392">Legs.</emphasis>
Protibiae with five well-defined teeth on the outer lateral margin, elongate apical tooth, teeth 25 triangular, gradually decreasing as they move away from the distal region, a reduced sixth tooth is present in the proximal region (
<figureCitation id="137845EB3518FFA3C7E9FA3CFE5E1E97" box="[386,510,1483,1509]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="3" pageNumber="392">Fig. 3D, G</figureCitation>
).
<emphasis id="B937857C3518FFA3C47AFA3CFD2C1E97" bold="true" box="[529,652,1483,1509]" pageId="3" pageNumber="392">Aedeagus.</emphasis>
Parallel and separate parameres, regularly curved from the base to the apex, divergent; in lateral view, parameres flattened and hooked. (
<figureCitation id="137845EB3518FFA3C511FA18FC531D7B" box="[890,1011,1519,1545]" captionStart="FIGURE 4" captionStartId="5.[152,255,1343,1367]" captionTargetBox="[161,1424,192,1320]" captionTargetId="figure-20@5.[151,1436,181,1320]" captionTargetPageId="5" captionText="FIGURE 4AF. Athyreus arretado new species aedeagus. AB, ventral view; CD, dorsal view; EF, lateral view. Scale bar: A, C, E: 1.0 mm; B, D, F: 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848921" httpUri="https://zenodo.org/record/13848921/files/figure.png" pageId="3" pageNumber="392">Fig. 4AF</figureCitation>
).
</paragraph>
<paragraph id="8BFC596E3518FFA3C6ACF9E4FC761D5C" blockId="3.[151,1437,295,1870]" box="[199,982,1555,1582]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACF9E4FE9F1D5F" bold="true" box="[199,319,1555,1581]" pageId="3" pageNumber="392">Variation.</emphasis>
No variation noted between the
<typeStatus id="54F8E7CC3518FFA3C4C7F9E4FCAE1D5F" box="[684,782,1555,1581]" pageId="3" pageNumber="392" type="holotype">holotype</typeStatus>
and the
<typeStatus id="54F8E7CC3518FFA3C505F9E3FC721D5C" box="[878,978,1556,1582]" pageId="3" pageNumber="392" type="paratype">paratype</typeStatus>
.
</paragraph>
<paragraph id="8BFC596E3518FFA3C6ACF9C0FE3C1D23" blockId="3.[151,1437,295,1870]" box="[199,412,1591,1617]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACF9C0FE851D23" bold="true" box="[199,293,1591,1617]" pageId="3" pageNumber="392">Female.</emphasis>
unknown.
</paragraph>
</subSubSection>
<subSubSection id="C3590AE53518FFA3C6ACF9ACFBE71DCF" pageId="3" pageNumber="392" type="etymology">
<paragraph id="8BFC596E3518FFA3C6ACF9ACFBE71DCF" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACF9ACFEEE1D07" bold="true" box="[199,334,1627,1653]" pageId="3" pageNumber="392">Etymology.</emphasis>
The species is named in reference to a typical expression from the northeastern region of
<collectingCountry id="F35419FE3518FFA3C33AF9ACFA391D07" box="[1361,1433,1627,1653]" name="Brazil" pageId="3" pageNumber="392">Brazil</collectingCountry>
. The term
<taxonomicName id="4C4322ED3518FFA3C768F988FECE1DEB" authorityName="Celante &amp; Carvalho &amp; Jahyny &amp; Araújo &amp; Santos" authorityYear="2024" box="[259,366,1663,1689]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="392" phylum="Arthropoda" rank="species" species="arretado">“arretado</taxonomicName>
” can have several meanings, including positively highlighting characteristics of people, objects, and places, and can be used as a synonym for beautiful, impressive, and spectacular.
</paragraph>
</subSubSection>
<subSubSection id="C3590AE53518FFA3C6ACF930FD831C77" pageId="3" pageNumber="392" type="biology_ecology">
<paragraph id="8BFC596E3518FFA3C6ACF930FD831C77" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACF930FE8C1D93" bold="true" box="[199,300,1735,1761]" pageId="3" pageNumber="392">Ecology.</emphasis>
The only known specimens of this species were attracted by flashlights and flew relatively close to the ground, approximately
<quantity id="4CBBF48B3518FFA3C7F6F91CFE421C74" box="[413,482,1771,1798]" metricMagnitude="-1" metricUnit="m" metricValue="6.0" pageId="3" pageNumber="392" unit="cm" value="60.0">60 cm</quantity>
high.
</paragraph>
</subSubSection>
<subSubSection id="C3590AE53518FFA5C6ACF8F8FDCF1C5E" lastPageId="5" lastPageNumber="394" pageId="3" pageNumber="392" type="distribution">
<paragraph id="8BFC596E3518FFA3C6ACF8F8FB351C3C" blockId="3.[151,1437,295,1870]" pageId="3" pageNumber="392">
<emphasis id="B937857C3518FFA3C6ACF8F8FEC11C5B" bold="true" box="[199,353,1807,1833]" pageId="3" pageNumber="392">Distribution.</emphasis>
The new species is only known from the
<typeStatus id="54F8E7CC3518FFA3C52FF8E7FCD41C58" box="[836,884,1808,1834]" pageId="3" pageNumber="392">type</typeStatus>
locality in the São Francisco River Sand Dunes Ecoregion (
<geoCoordinate id="EE773FA93518FFA3C772F8C4FED41C3C" box="[281,372,1843,1870]" degrees="9.3952" direction="south" orientation="latitude" pageId="3" pageNumber="392" precision="5" value="-9.3952">-9.3952</geoCoordinate>
,
<geoCoordinate id="EE773FA93518FFA3C714F8C4FE461C3F" box="[383,486,1843,1869]" degrees="41.3610" direction="west" orientation="longitude" pageId="3" pageNumber="392" precision="5" value="-41.361">-41.3610</geoCoordinate>
, Casa Nova Dunes Complex,
<collectingRegion id="4987978C3518FFA3C55DF8C4FCDA1C3F" box="[822,890,1843,1869]" country="Brazil" name="Bahia" pageId="3" pageNumber="392">Bahia</collectingRegion>
,
<collectingCountry id="F35419FE3518FFA3C5EEF8C4FC6E1C3F" box="[901,974,1843,1869]" name="Brazil" pageId="3" pageNumber="392">Brazil</collectingCountry>
). (
<figureCitation id="137845EB3518FFA3C583F8C4FB841C3F" box="[1000,1060,1843,1869]" captionStart="FIGURE 1" captionStartId="1.[152,255,1477,1501]" captionTargetBox="[161,1430,182,1451]" captionTargetId="figure-20@1.[158,1430,179,1451]" captionTargetPageId="1" captionText="FIGURE 1. Collection points of Bolboceratinae integrated into a map of a portion of South America, delineating the boundaries of the states within the northeastern region of Brazil, as well as the limits of the Seasonally Dry Tropical Forest and Woodland Biome (SDTFW) and the São Francisco River Sand Dunes Ecoregion (SFRSD). State acronyms: BA—Bahia, PB—Paraíba, PE—Pernambuco." figureDoi="http://doi.org/10.5281/zenodo.13848911" httpUri="https://zenodo.org/record/13848911/files/figure.png" pageId="3" pageNumber="392">Fig 1</figureCitation>
,
<figureCitation id="137845EB3518FFA3C259F8C4FBF71C3F" box="[1074,1111,1843,1869]" captionStart="FIGURE 2" captionStartId="2.[152,255,1833,1857]" captionTargetBox="[151,1435,525,1809]" captionTargetId="figure-165@2.[151,1435,525,1809]" captionTargetPageId="2" captionText="FIGURE 2AD. Collecting localities.A, São Francisco River Sand Dunes in Casa Nova, Bahia, Brazil; B, Caatinga in Petrolina, Pernambuco, Brazil; C, flight intercept trap in Refúgio de Vida Silvestre Mata do Buraquinho, João Pessoa, Paraíba, Brazil; D, Reserva Biológica Guaribas, Mamanguape, Paraíba, Brazil." figureDoi="http://doi.org/10.5281/zenodo.13848915" httpUri="https://zenodo.org/record/13848915/files/figure.png" pageId="3" pageNumber="392">2A</figureCitation>
,
<figureCitation id="137845EB3518FFA3C208F8C4FB281C3C" box="[1123,1160,1843,1870]" captionStart="FIGURE 5" captionStartId="6.[152,255,1716,1740]" captionTargetBox="[151,1435,181,1692]" captionTargetId="figure-24@6.[151,1435,181,1692]" captionTargetPageId="6" captionText="FIGURE 5AD. Distribution records of some Bolboceratinae species. A, Athyreus arretado new species, Athyreus vavini Boucomont, 1902 and Athyreus tuberifer Felsche, 1909 distribution records; B, Neoathyreus anthracinus (Klug, 1843) and Neoathyreus braziliensis Howden, 1985 distribution records; C, Neoathyreus centromaculatus (Felsche, 1909), Neoathyreus lanuginosus (Klug, 1843), and Parathyreus trituberculatus (Klug, 1843) distribution records; D, Bolbapium caesum (Klug, 1843), Bolbapium parcepunctatum (Arrow, 1913), and Bolbapium sulcifrons Ide &amp; Martínez, 1993 distribution records. The black symbols represent new records, the unfilled symbols are literature records." figureDoi="http://doi.org/10.5281/zenodo.13848925" httpUri="https://zenodo.org/record/13848925/files/figure.png" pageId="3" pageNumber="392">5A</figureCitation>
).
</paragraph>
<caption id="DF3C09E6351FFFA4C6F3F8D2FE201CF7" ID-DOI="http://doi.org/10.5281/zenodo.13848919" ID-Zenodo-Dep="13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="4" pageNumber="393" startId="4.[152,255,1829,1853]" targetBox="[151,1416,185,1806]" targetPageId="4" targetType="figure">
<paragraph id="8BFC596E351FFFA4C6F3F8D2FE201CF7" blockId="4.[151,1436,1829,1926]" pageId="4" pageNumber="393">
<emphasis id="B937857C351FFFA4C6F3F8D2FEEF1C4F" bold="true" box="[152,335,1829,1853]" pageId="4" pageNumber="393">FIGURE 3AG.</emphasis>
<taxonomicName id="4C4322ED351FFFA4C733F8D1FDB31C4C" authorityName="Celante &amp; Carvalho &amp; Jahyny &amp; Araújo &amp; Santos" authorityYear="2024" box="[344,531,1830,1854]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="393" phylum="Arthropoda" rank="species" species="arretado" status="sp. nov.">
<emphasis id="B937857C351FFFA4C733F8D1FDB31C4C" box="[344,531,1830,1854]" italics="true" pageId="4" pageNumber="393">Athyreus arretado</emphasis>
</taxonomicName>
<emphasis id="B937857C351FFFA4C477F8D1FD3C1C4F" bold="true" box="[540,668,1829,1854]" pageId="4" pageNumber="393">
<taxonomicNameLabel id="A2043807351FFFA4C477F8D1FD3C1C4F" box="[540,668,1829,1854]" pageId="4" pageNumber="393" rank="species">new species</taxonomicNameLabel>
</emphasis>
holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm.
</paragraph>
</caption>
<caption id="DF3C09E6351EFFA5C6F3FAC8FE581E09" ID-DOI="http://doi.org/10.5281/zenodo.13848921" ID-Zenodo-Dep="13848921" httpUri="https://zenodo.org/record/13848921/files/figure.png" pageId="5" pageNumber="394" startId="5.[152,255,1343,1367]" targetBox="[161,1424,192,1320]" targetPageId="5" targetType="figure">
<paragraph id="8BFC596E351EFFA5C6F3FAC8FE581E09" blockId="5.[151,1437,1343,1404]" pageId="5" pageNumber="394">
<emphasis id="B937857C351EFFA5C6F3FAC8FEE41E25" bold="true" box="[152,324,1343,1367]" pageId="5" pageNumber="394">FIGURE 4AF.</emphasis>
<taxonomicName id="4C4322ED351EFFA5C727FAB7FDA31E2A" authorityName="Celante &amp; Carvalho &amp; Jahyny &amp; Araújo &amp; Santos" authorityYear="2024" box="[332,515,1344,1368]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="arretado" status="sp. nov.">
<emphasis id="B937857C351EFFA5C727FAB7FDA31E2A" box="[332,515,1344,1368]" italics="true" pageId="5" pageNumber="394">Athyreus arretado</emphasis>
</taxonomicName>
<emphasis id="B937857C351EFFA5C461FAB7FD271E25" bold="true" box="[522,647,1343,1368]" pageId="5" pageNumber="394">
<taxonomicNameLabel id="A2043807351EFFA5C461FAB7FD271E25" box="[522,647,1343,1368]" pageId="5" pageNumber="394" rank="species">new species</taxonomicNameLabel>
</emphasis>
aedeagus. AB, ventral view; CD, dorsal view; EF, lateral view. Scale bar: A, C, E: 1.0 mm; B, D, F: 0.5 mm.
</paragraph>
</caption>
<paragraph id="8BFC596E351EFFA5C6ACFA3AFDCF1C5E" blockId="5.[151,1437,1485,1836]" pageId="5" pageNumber="394">
<emphasis id="B937857C351EFFA5C6ACFA3AFE1E1E95" bold="true" box="[199,446,1485,1511]" pageId="5" pageNumber="394">Taxonomic position.</emphasis>
<taxonomicName id="4C4322ED351EFFA5C7ADFA39FD361E9A" authorityName="Celante &amp; Carvalho &amp; Jahyny &amp; Araújo &amp; Santos" authorityYear="2024" box="[454,662,1486,1512]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="arretado">
<emphasis id="B937857C351EFFA5C7ADFA39FD361E9A" box="[454,662,1486,1512]" italics="true" pageId="5" pageNumber="394">Athyreus arretado</emphasis>
</taxonomicName>
is a representative of the
<taxonomicName id="4C4322ED351EFFA5C5A8FA39FB921E9A" authorityName="Klug" authorityYear="1843" box="[963,1074,1486,1512]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="aeneus">
<emphasis id="B937857C351EFFA5C5A8FA39FC741E9A" box="[963,980,1486,1512]" italics="true" pageId="5" pageNumber="394">A</emphasis>
.
<emphasis id="B937857C351EFFA5C588FA38FB921E9A" box="[995,1074,1487,1512]" italics="true" pageId="5" pageNumber="394">aeneus</emphasis>
</taxonomicName>
species group and differs from all species belonging to the
<taxonomicName id="4C4322ED351EFFA5C7B4FA05FDD41D7E" authorityName="MacLeay" authorityYear="1819" box="[479,628,1522,1548]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="bifurcatus">
<emphasis id="B937857C351EFFA5C7B4FA05FE501D7E" box="[479,496,1522,1548]" italics="true" pageId="5" pageNumber="394">A</emphasis>
.
<emphasis id="B937857C351EFFA5C46AFA05FDD41D7E" box="[513,628,1522,1548]" italics="true" pageId="5" pageNumber="394">bifurcatus</emphasis>
</taxonomicName>
species group and
<taxonomicName id="4C4322ED351EFFA5C53DFA05FBA11D7E" authorityName="Fairmaire" authorityYear="1892" box="[854,1025,1522,1548]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="chalybeatus">
<emphasis id="B937857C351EFFA5C53DFA05FCC71D7E" box="[854,871,1522,1548]" italics="true" pageId="5" pageNumber="394">A</emphasis>
.
<emphasis id="B937857C351EFFA5C513FA05FBA11D7E" box="[888,1025,1522,1548]" italics="true" pageId="5" pageNumber="394">chalybeatus</emphasis>
</taxonomicName>
species group by the presence of a complete longitudinal sulcus, which begins at posterior base of anterior pronotal horn, and extends to the posterior region of pronotum (
<figureCitation id="137845EB351EFFA5C7E7F9CDFDAF1D26" box="[396,527,1594,1620]" captionStart="FIGURE 3" captionStartId="4.[152,255,1829,1853]" captionTargetBox="[151,1416,185,1806]" captionTargetId="figure-24@4.[151,1436,181,1806]" captionTargetPageId="4" captionText="FIGURE 3AG. Athyreus arretado new species holotype, male, habitus. A, Dorsal habitus; B, ventral habitus; C, lateral habitus; D, head and pronotum (frontal view); E, head dorsal view; F, head ventral view; G, head and pronotum, laterofrontal view. Scale bar: 5 mm." figureDoi="http://doi.org/10.5281/zenodo.13848919" httpUri="https://zenodo.org/record/13848919/files/figure.png" pageId="5" pageNumber="394">Fig. 3A, G</figureCitation>
). The species groups established by
<bibRefCitation id="EFD2249F351EFFA5C5D7F9CDFAA21D26" author="Howden, H. F. &amp; Martinez, A." box="[956,1282,1594,1620]" pageId="5" pageNumber="394" pagination="1 - 70" refId="ref5829" refString="Howden, H. F. &amp; Martinez, A. (1978) A review of the New World genus Athyreus MacLeay (Scarabaeidae, Geotrupinae, Athyreini). Contributions of the American Entomological Institute, 15, 1 - 70." type="journal article" year="1978">Howden &amp; Martínez (1978)</bibRefCitation>
facilitate the identification key and the recognition of species. However, the phylogenetic validity of these species groups has never been tested and were questioned by
<bibRefCitation id="EFD2249F351EFFA5C414F975FC971DEE" author="Howden, H. F." box="[639,823,1666,1692]" pageId="5" pageNumber="394" pagination="339 - 354" refId="ref5703" refString="Howden, H. F. (1999) New species of Central and South American Athyreini (Coleoptera: Scarabaeidae: Geotrupinae). The Coleopterists Bulletin, 53, 339 - 354." type="journal article" year="1999">Howden (1999)</bibRefCitation>
, who noted that the males of
<taxonomicName id="4C4322ED351EFFA5C2E1F975FF701DB2" authority="Howden, 1999" authorityName="Howden" authorityYear="1999" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="fritzi">
<emphasis id="B937857C351EFFA5C2E1F975FA8C1DEE" box="[1162,1324,1666,1692]" italics="true" pageId="5" pageNumber="394">Athyreus fritzi</emphasis>
<bibRefCitation id="EFD2249F351EFFA5C35EF975FF701DB2" author="Howden, H. F." pageId="5" pageNumber="394" pagination="339 - 354" refId="ref5703" refString="Howden, H. F. (1999) New species of Central and South American Athyreini (Coleoptera: Scarabaeidae: Geotrupinae). The Coleopterists Bulletin, 53, 339 - 354." type="journal article" year="1999">Howden, 1999</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C4322ED351EFFA5C767F951FD931DB2" authority="Howden, 1999" authorityName="Howden" authorityYear="1999" box="[268,563,1702,1728]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="langeri">
<emphasis id="B937857C351EFFA5C767F951FE221DB2" box="[268,386,1702,1728]" italics="true" pageId="5" pageNumber="394">A. langeri</emphasis>
<bibRefCitation id="EFD2249F351EFFA5C7E2F951FD931DB2" author="Howden, H. F." box="[393,563,1702,1728]" pageId="5" pageNumber="394" pagination="339 - 354" refId="ref5703" refString="Howden, H. F. (1999) New species of Central and South American Athyreini (Coleoptera: Scarabaeidae: Geotrupinae). The Coleopterists Bulletin, 53, 339 - 354." type="journal article" year="1999">Howden, 1999</bibRefCitation>
</taxonomicName>
fit in the
<taxonomicName id="4C4322ED351EFFA5C4C7F951FCBD1DB2" authorityName="Klug" authorityYear="1843" box="[684,797,1702,1728]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="aeneus">
<emphasis id="B937857C351EFFA5C4C7F951FD1D1DB2" box="[684,701,1702,1728]" italics="true" pageId="5" pageNumber="394">A</emphasis>
.
<emphasis id="B937857C351EFFA5C4A5F950FCBD1DB2" box="[718,797,1703,1728]" italics="true" pageId="5" pageNumber="394">aeneus</emphasis>
</taxonomicName>
species group, while females fit in the
<taxonomicName id="4C4322ED351EFFA5C299F951FA3D1DB2" authorityName="Fairmaire" authorityYear="1892" box="[1266,1437,1702,1728]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="chalybeatus">
<emphasis id="B937857C351EFFA5C299F951FAA31DB2" box="[1266,1283,1702,1728]" italics="true" pageId="5" pageNumber="394">A</emphasis>
.
<emphasis id="B937857C351EFFA5C37FF951FA3D1DB2" box="[1300,1437,1702,1728]" italics="true" pageId="5" pageNumber="394">chalybeatus</emphasis>
</taxonomicName>
species group. Taking this discussion into account and considering the morphological similarity of
<taxonomicName id="4C4322ED351EFFA5C290F93DFACD1D96" authorityName="Howden" authorityYear="1999" box="[1275,1389,1738,1764]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="langeri">
<emphasis id="B937857C351EFFA5C290F93DFACD1D96" box="[1275,1389,1738,1764]" italics="true" pageId="5" pageNumber="394">A. langeri</emphasis>
</taxonomicName>
and the recently described species
<taxonomicName id="4C4322ED351EFFA5C798F919FD001C7A" box="[499,672,1774,1800]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="joseferreirai">
<emphasis id="B937857C351EFFA5C798F919FD001C7A" box="[499,672,1774,1800]" italics="true" pageId="5" pageNumber="394">A. joseferreirai</emphasis>
</taxonomicName>
, we consider the latter as a member of the
<taxonomicName id="4C4322ED351EFFA5C2E1F919FB591C7A" authorityName="Klug" authorityYear="1843" box="[1162,1273,1774,1800]" class="Insecta" family="Geotrupidae" genus="Athyreus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="394" phylum="Arthropoda" rank="species" species="aeneus">
<emphasis id="B937857C351EFFA5C2E1F919FB3B1C7A" box="[1162,1179,1774,1800]" italics="true" pageId="5" pageNumber="394">A</emphasis>
.
<emphasis id="B937857C351EFFA5C2C1F918FB591C7A" box="[1194,1273,1775,1800]" italics="true" pageId="5" pageNumber="394">aeneus</emphasis>
</taxonomicName>
species group and included it in the updated key below.
</paragraph>
</subSubSection>
</treatment>
</document>