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<document id="244C0FB07F05A50B365D42A0BE4CC9D0" ID-DOI="10.5281/zenodo.7887628" ID-ISSN="1942-1354" ID-Zenodo-Dep="7887628" ID-ZooBank="E701D60F-A455-4048-8279-DA450930ACB3" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatmentCitations_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1683049820006" checkinUser="felipe" docAuthor="Solís, Ángel &amp; Kohlmann, Bert" docDate="2023" docId="B31B87B01F7E101B4FFCB4CAFADDFB6B" docLanguage="en" docName="InsectaMundi.2023.0972.1–21.pdf" docOrigin="Insecta Mundi 2023 (972)" docStyle="DocumentStyle:F9976E9B7F8A094D890004515A7A0592.3:InsectaMundi.2020-.journal_article" docStyleId="F9976E9B7F8A094D890004515A7A0592" docStyleName="InsectaMundi.2020-.journal_article" docStyleVersion="3" docTitle="Phanaeus (Notiophanaeus) pyrois Bates 1887" docType="treatment" docVersion="3" lastPageNumber="6" masterDocId="4F22FFC81F7B101C4F3CB23FFFA6FFEB" masterDocTitle="New synonyms and records of Costa Rican and Panamanian dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae)" masterLastPageNumber="21" masterPageNumber="1" pageNumber="4" updateTime="1699495782299" updateUser="plazi" zenodo-license-document="CC-BY-NC-3.0">
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<mods:title id="5701A92439F1397DF1CFCC755AA9A328">New synonyms and records of Costa Rican and Panamanian dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae)</mods:title>
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<subSubSection id="73A8652D1F7E10194FFCB4CAFCA4F8F9" box="[192,770,1781,1810]" pageId="5" pageNumber="4" type="nomenclature">
<paragraph id="3B0D36A61F7E10194FFCB4CAFCA4F8F9" blockId="5.[192,770,1781,1810]" box="[192,770,1781,1810]" pageId="5" pageNumber="4">
<heading id="604581CA1F7E10194FFCB4CAFCA4F8F9" box="[192,770,1781,1810]" fontSize="12" level="2" pageId="5" pageNumber="4" reason="2">
<taxonomicName id="FCB24D251F7E10194FFCB4CAFCA4F8F9" ID-CoL="4FSJX" ID-ENA="506469" authority="Bates, 1887" authorityName="Bates" authorityYear="1887" box="[192,770,1781,1810]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="4" phylum="Arthropoda" rank="species" species="pyrois" subGenus="Notiophanaeus">
<emphasis id="09C6EAB41F7E10194FFCB4CAFCA4F8F9" bold="true" box="[192,770,1781,1810]" pageId="5" pageNumber="4">
<emphasis id="09C6EAB41F7E10194FFCB4CAFDCDF8FA" bold="true" box="[192,619,1781,1810]" italics="true" pageId="5" pageNumber="4">Phanaeus (Notiophanaeus) pyrois</emphasis>
<bibRefCitation id="5F234B571F7E10194D4EB4CAFCA4F8F9" author="Bates HW" box="[626,770,1781,1810]" pageId="5" pageNumber="4" refId="ref10136" refString="Bates HW. 1887. Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis; London. 432 p. + 24 Tables." type="book" year="1887">Bates, 1887</bibRefCitation>
</emphasis>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="73A8652D1F7E101B4FDCB51FFC67FEF8" lastPageId="7" lastPageNumber="6" pageId="5" pageNumber="4" type="reference_group">
<paragraph id="3B0D36A61F7E10194FDCB51FFDB8F8D3" blockId="5.[224,542,1824,1849]" box="[224,542,1824,1849]" pageId="5" pageNumber="4">
<treatmentCitationGroup id="1BA211881F7E10194FDCB51FFDB8F8D3" box="[224,542,1824,1849]" pageId="5" pageNumber="4">
<treatmentCitation id="BA1310B71F7E10194FDCB51FFDBCF8D3" author="Bates HW" box="[224,538,1824,1849]" page="58" pageId="5" pageNumber="4" year="1887">
<taxonomicName id="FCB24D251F7E10194FDCB51FFDBCF8D3" ID-CoL="4FSJX" authority="Bates 1887: 58" authorityName="Bates" authorityPageNumber="58" authorityYear="1887" box="[224,538,1824,1849]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="4" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7E10194FDCB51FFE24F8D3" box="[224,386,1824,1848]" italics="true" pageId="5" pageNumber="4">Phanaeus pyrois</emphasis>
<bibRefCitation id="5F234B571F7E10194EB4B51EFDBCF8D3" author="Bates HW" box="[392,538,1824,1849]" pageId="5" pageNumber="4" refId="ref10136" refString="Bates HW. 1887. Biologia Centrali-Americana. Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. Taylor and Francis; London. 432 p. + 24 Tables." type="book" year="1887">Bates 1887: 58</bibRefCitation>
</taxonomicName>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph id="3B0D36A61F7E101B4FFCB575FC67FEF8" blockId="5.[191,1439,1865,1927]" lastBlockId="7.[191,1442,214,969]" lastPageId="7" lastPageNumber="6" pageId="5" pageNumber="4">
<bibRefCitation id="5F234B571F7E10194FFCB575FDE1F888" author="Moctezuma V &amp; Halffter G." box="[192,583,1865,1892]" pageId="5" pageNumber="4" pagination="1 - 71" refId="ref11792" refString="Moctezuma V, Halffter G. 2021. Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species. European Journal of Taxonomy 747: 1 - 71." type="journal article" year="2021">Moctezuma and Halffter (2021: 42)</bibRefCitation>
stated that the elytral striae of
<taxonomicName id="FCB24D251F7E10194CA5B575FC55F888" ID-CoL="4FSJX" authorityName="Bates" authorityYear="1887" box="[921,1011,1866,1891]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="4" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7E10194CA5B575FC55F888" box="[921,1011,1866,1891]" italics="true" pageId="5" pageNumber="4">P. pyrois</emphasis>
</taxonomicName>
are not strongly impressed basally and according to their key’s couplet 10 (2021: 54), the “elytral interstriae are distinctly flattened”. We do not agree with this statement; the elytral striae are indeed strongly impressed basally and the interstriae are convex (
<figureCitation id="A3892A231F7C101B4A1CB2E9FAC4FF1B" box="[1312,1378,214,240]" captionStart="Figures 1–5" captionStartId="6.[192,279,1799,1825]" captionTargetBox="[231,1401,213,1770]" captionTargetId="figure-14@6.[231,1401,213,1770]" captionTargetPageId="6" captionText="Figures 1–5. Phanaeus spp. 1) Elytra of P. olsoufieffi. 2) Elytra of Phanaeus pyrois. 3) Lamella copulatrix of P. olsoufieffi from Valle de Antón, Panama. 4) Lamella copulatrix of P. olsoufieffi from Barro Colorado Island, Panama. 5) Known distribution of Phanaeus malyi, P. pyrois, and P. olsoufieffi in lower Central America. Right arrow, Barro Colorado Island; left arrow, Valle de Antón." figureDoi="http://doi.org/10.5281/zenodo.7887630" httpUri="https://zenodo.org/record/7887630/files/figure.png" pageId="7" pageNumber="6">Fig. 2</figureCitation>
), and these characters easily separate this species from
<taxonomicName id="FCB24D251F7C101B4DDDB2C6FCC5FEF9" ID-CoL="874GF" authorityName="Balthasar" authorityYear="1939" box="[737,867,248,274]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="olsoufieffi">
<emphasis id="09C6EAB41F7C101B4DDDB2C6FCC5FEF9" box="[737,867,248,274]" italics="true" pageId="7" pageNumber="6">P. olsoufieffi</emphasis>
</taxonomicName>
(
<figureCitation id="A3892A231F7C101B4C4EB2C7FC16FEF8" box="[882,944,248,275]" captionStart="Figures 1–5" captionStartId="6.[192,279,1799,1825]" captionTargetBox="[231,1401,213,1770]" captionTargetId="figure-14@6.[231,1401,213,1770]" captionTargetPageId="6" captionText="Figures 1–5. Phanaeus spp. 1) Elytra of P. olsoufieffi. 2) Elytra of Phanaeus pyrois. 3) Lamella copulatrix of P. olsoufieffi from Valle de Antón, Panama. 4) Lamella copulatrix of P. olsoufieffi from Barro Colorado Island, Panama. 5) Known distribution of Phanaeus malyi, P. pyrois, and P. olsoufieffi in lower Central America. Right arrow, Barro Colorado Island; left arrow, Valle de Antón." figureDoi="http://doi.org/10.5281/zenodo.7887630" httpUri="https://zenodo.org/record/7887630/files/figure.png" pageId="7" pageNumber="6">Fig. 1</figureCitation>
).
</paragraph>
</subSubSection>
<caption id="6FCD662E1F7D101A4FFCB538FD4DF86D" ID-DOI="http://doi.org/10.5281/zenodo.7887630" ID-Zenodo-Dep="7887630" httpUri="https://zenodo.org/record/7887630/files/figure.png" pageId="6" pageNumber="5" startId="6.[192,279,1799,1825]" targetBox="[231,1401,213,1770]" targetPageId="6" targetType="figure">
<paragraph id="3B0D36A61F7D101A4FFCB538FD4DF86D" blockId="6.[192,1441,1799,1927]" pageId="6" pageNumber="5">
<emphasis id="09C6EAB41F7D101A4FFCB538FEE9F8CA" bold="true" box="[192,335,1799,1825]" pageId="6" pageNumber="5">Figures 1–5.</emphasis>
<taxonomicName id="FCB24D251F7D101A4E69B538FDAFF8CA" box="[341,521,1799,1826]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="undefined-1">
<emphasis id="09C6EAB41F7D101A4E69B538FE67F8CA" box="[341,449,1799,1825]" italics="true" pageId="6" pageNumber="5">Phanaeus</emphasis>
spp.
<emphasis id="09C6EAB41F7D101A4EC6B538FDAFF8CA" bold="true" box="[506,521,1799,1825]" pageId="6" pageNumber="5">1</emphasis>
</taxonomicName>
<emphasis id="09C6EAB41F7D101A4D35B538FDB5F8CA" bold="true" box="[521,531,1799,1825]" pageId="6" pageNumber="5">)</emphasis>
Elytra of
<taxonomicName id="FCB24D251F7D101A4DBCB537FCA4F8CA" authorityName="Balthasar" authorityYear="1939" box="[640,770,1799,1825]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="olsoufieffi">
<emphasis id="09C6EAB41F7D101A4DBCB537FCA4F8CA" box="[640,770,1799,1825]" italics="true" pageId="6" pageNumber="5">P. olsoufieffi</emphasis>
</taxonomicName>
.
<emphasis id="09C6EAB41F7D101A4C33B538FC81F8CA" bold="true" box="[783,807,1799,1825]" pageId="6" pageNumber="5">2)</emphasis>
Elytra of
<taxonomicName id="FCB24D251F7D101A4CA9B538FBEFF8CA" authorityName="Bates" authorityYear="1887" box="[917,1097,1799,1825]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7D101A4CA9B538FBEFF8CA" box="[917,1097,1799,1825]" italics="true" pageId="6" pageNumber="5">Phanaeus pyrois</emphasis>
</taxonomicName>
.
<emphasis id="09C6EAB41F7D101A4B6AB538FBC8F8CA" bold="true" box="[1110,1134,1799,1825]" pageId="6" pageNumber="5">3)</emphasis>
<emphasis id="09C6EAB41F7D101A4B49B538FAE1F8CA" box="[1141,1351,1799,1825]" italics="true" pageId="6" pageNumber="5">Lamella copulatrix</emphasis>
of
<taxonomicName id="FCB24D251F7D101A4A56B537FEB3F8A8" authorityName="Balthasar" authorityYear="1939" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="olsoufieffi">
<emphasis id="09C6EAB41F7D101A4A56B537FEB3F8A8" italics="true" pageId="6" pageNumber="5">P. olsoufieffi</emphasis>
</taxonomicName>
from Valle de Antón, Panama.
<emphasis id="09C6EAB41F7D101A4D54B517FD27F8A9" bold="true" box="[616,641,1832,1858]" pageId="6" pageNumber="5">4)</emphasis>
<emphasis id="09C6EAB41F7D101A4DBAB516FCF1F8A8" box="[646,855,1833,1859]" italics="true" pageId="6" pageNumber="5">Lamella copulatrix</emphasis>
of
<taxonomicName id="FCB24D251F7D101A4C44B515FC5EF8A8" authorityName="Balthasar" authorityYear="1939" box="[888,1016,1833,1859]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="olsoufieffi">
<emphasis id="09C6EAB41F7D101A4C44B515FC5EF8A8" box="[888,1016,1833,1859]" italics="true" pageId="6" pageNumber="5">P. olsoufieffi</emphasis>
</taxonomicName>
from Barro Colorado Island, Panama.
<emphasis id="09C6EAB41F7D101A4FFCB575FF7FF88F" bold="true" box="[192,217,1866,1892]" pageId="6" pageNumber="5">5)</emphasis>
Known distribution of
<taxonomicName id="FCB24D251F7D101A4ED0B575FD06F88F" authorityName="Arnaud" authorityYear="2002" box="[492,672,1866,1892]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="malyi">
<emphasis id="09C6EAB41F7D101A4ED0B575FD06F88F" box="[492,672,1866,1892]" italics="true" pageId="6" pageNumber="5">Phanaeus malyi</emphasis>
</taxonomicName>
,
<taxonomicName id="FCB24D251F7D101A4D8CB574FCABF88F" authorityName="Bates" authorityYear="1887" box="[688,781,1867,1892]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7D101A4D8CB574FCABF88F" box="[688,781,1867,1892]" italics="true" pageId="6" pageNumber="5">P. pyrois</emphasis>
</taxonomicName>
, and
<taxonomicName id="FCB24D251F7D101A4C6DB574FC70F88F" authorityName="Balthasar" authorityYear="1939" box="[849,982,1866,1892]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="5" phylum="Arthropoda" rank="species" species="olsoufieffi">
<emphasis id="09C6EAB41F7D101A4C6DB574FC70F88F" box="[849,982,1866,1892]" italics="true" pageId="6" pageNumber="5">P. olsoufieffi</emphasis>
</taxonomicName>
in lower Central America. Right arrow, Barro Colorado Island; left arrow, Valle de Antón.
</paragraph>
</caption>
<subSubSection id="73A8652D1F7C101B4E3CB324FADDFB6B" pageId="7" pageNumber="6" type="description">
<paragraph id="3B0D36A61F7C101B4E3CB324FCB7FDC3" blockId="7.[191,1442,214,969]" pageId="7" pageNumber="6">
<taxonomicName id="FCB24D251F7C101B4E3CB324FE10FEDE" authorityName="Bates" authorityYear="1887" box="[256,438,283,309]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7C101B4E3CB324FE10FEDE" box="[256,438,283,309]" italics="true" pageId="7" pageNumber="6">Phanaeus pyrois</emphasis>
</taxonomicName>
is the most color variable taxon of the endymion species group (
<bibRefCitation id="5F234B571F7C101B4BB6B324FF5CFEB3" author="Moctezuma V &amp; Halffter G." pageId="7" pageNumber="6" pagination="1 - 71" refId="ref11792" refString="Moctezuma V, Halffter G. 2021. Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species. European Journal of Taxonomy 747: 1 - 71." type="journal article" year="2021">Moctezuma and Halffter 2021</bibRefCitation>
). However, no detailed analysis of the different color morphs has heretofore been undertaken. We present a study of several color morphs in
<collectingCountry id="43A576361F7C101B4D13B35EFD0FFE91" box="[559,681,352,379]" name="Costa Rica" pageId="7" pageNumber="6">Costa Rica</collectingCountry>
(
<figureCitation id="A3892A231F7C101B4D8BB35FFC8BFE91" box="[695,813,352,378]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Figs. 6–10</figureCitation>
) and their relative frequencies (
<figureCitation id="A3892A231F7C101B4BADB35FFB47FE90" box="[1169,1249,352,379]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Fig. 11</figureCitation>
). Our analysis of
<specimenCount id="2DB4FD2F1F7C101B4FFCB3BCFEA4FE76" box="[192,258,387,414]" pageId="7" pageNumber="6" type="generic">
665
<taxonomicName id="FCB24D251F7C101B4FD2B3BBFEA4FE76" authorityName="Bates" authorityYear="1887" box="[238,258,388,413]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7C101B4FD2B3BBFEA4FE76" box="[238,258,388,413]" italics="true" pageId="7" pageNumber="6">P.</emphasis>
</taxonomicName>
</specimenCount>
pyrois specimens shows metallic red (
<figureCitation id="A3892A231F7C101B4D9BB3BCFD4EFE76" box="[679,744,387,413]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Fig. 6</figureCitation>
) to be the most common color morph, followed by a dark blue (
<figureCitation id="A3892A231F7C101B4FF4B399FEAFFE2B" box="[200,265,422,448]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Fig. 7</figureCitation>
), a dark green (
<figureCitation id="A3892A231F7C101B4E8AB399FE51FE2B" box="[438,503,422,448]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Fig. 8</figureCitation>
), a red with green reflections (
<figureCitation id="A3892A231F7C101B4C75B399FC2DFE2B" box="[841,907,422,448]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Fig. 9</figureCitation>
), and finally a dark red (
<figureCitation id="A3892A231F7C101B4BA6B399FB4DFE2B" box="[1178,1259,422,448]" captionStart="Figures 6–11. 6" captionStartId="8.[192,279,1833,1859]" captionTargetBox="[239,1396,213,1803]" captionTargetId="figure-15@8.[235,1396,213,1803]" captionTargetPageId="8" captionText="Figures 6–11. 6) Red morph of P. pyrois.7). Dark blue morph of P. pyrois. 8). Dark green morph of P. pyrois. 9). Red with green reflections morph of P. pyrois. 10). Dark red morph of P. pyrois. 11). Color pie-chart of the Costa Rican P. pyrois populations, n = 665." figureDoi="http://doi.org/10.5281/zenodo.7887632" httpUri="https://zenodo.org/record/7887632/files/figure.png" pageId="7" pageNumber="6">Fig. 10</figureCitation>
). As can be seen from figures 12 and 13, these frequencies vary geographically. There is a greater percentage of red, red with green reflections, and dark red morphs in the northern part of its range, whereas there is a greater percentage of dark blue and dark green morphs in the southern portion.
</paragraph>
<paragraph id="3B0D36A61F7C101B4E3CB00EFEB9FD58" blockId="7.[191,1442,214,969]" pageId="7" pageNumber="6">
<bibRefCitation id="5F234B571F7C101B4E3CB00EFE4DFDA0" author="Cupello M &amp; Ribeiro-Costa CS &amp; Vaz-de-Mello FZ" box="[256,491,561,587]" pageId="7" pageNumber="6" pagination="973 - 1034" refId="ref10278" refString="Cupello M, Ribeiro-Costa CS, Vaz-de-Mello FZ. 2022. The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem. Zoological Journal of the Linnean Society 194: 973 - 1034." type="journal article" year="2022">Cupello et al. (2022)</bibRefCitation>
have undertaken an exquisitely detailed color analysis of
<taxonomicName id="FCB24D251F7C101B4BBDB00EFF50FD86" authority="Harold, 1868" authorityName="Harold" authorityYear="1868" class="Insecta" family="Scarabaeidae" genus="Bolbites" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="onitoides">
<emphasis id="09C6EAB41F7C101B4BBDB00EFAE5FDA0" box="[1153,1347,561,587]" italics="true" pageId="7" pageNumber="6">Bolbites onitoides</emphasis>
<bibRefCitation id="5F234B571F7C101B4A70B00EFF50FD86" author="Harold" pageId="7" pageNumber="6" pagination="80 - 86" refId="ref10850" refString="Harold E von. 1868. Diagnosen neuer Coprophagen. Coleopterologische Hefte 4: 80 - 86." type="journal article" year="1868">Harold, 1868</bibRefCitation>
</taxonomicName>
another
<taxonomicName id="FCB24D251F7C101B4E5DB06CFE73FD86" box="[353,469,595,621]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
, on the question of whether its two-color variants (blue and red) represent distinct subspecies. They concluded that
<taxonomicName id="FCB24D251F7C101B4D04B048FD1FFD7B" authorityName="Harold" authorityYear="1868" box="[568,697,630,656]" class="Insecta" family="Scarabaeidae" genus="Bolbites" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="onitoides">
<emphasis id="09C6EAB41F7C101B4D04B048FD1FFD7B" box="[568,697,630,656]" italics="true" pageId="7" pageNumber="6">B. onitoides</emphasis>
</taxonomicName>
should be treated as a single monotypic species for the following reasons:
</paragraph>
<paragraph id="3B0D36A61F7C101B4FD4B083FB4FFD3E" blockId="7.[191,1442,214,969]" box="[232,1257,699,726]" pageId="7" pageNumber="6">• Two color intermediates were found living among the main color variants (red and blue);</paragraph>
<paragraph id="3B0D36A61F7C101B4FD4B0E0FD66FD13" blockId="7.[191,1442,214,969]" box="[232,704,734,761]" pageId="7" pageNumber="6">• the distribution of the variants overlaps;</paragraph>
<paragraph id="3B0D36A61F7C101B4FD4B13EFCA6FCF0" blockId="7.[191,1442,214,969]" box="[232,768,769,795]" pageId="7" pageNumber="6">• no other characters paralleled color variation;</paragraph>
<paragraph id="3B0D36A61F7C101B4FD4B11BFAE6FC8B" blockId="7.[191,1442,214,969]" pageId="7" pageNumber="6">• the overall geographical pattern (blue western-half and a red eastern-half) can be explained by phenomena other than (incipient) speciation, such as phenotypic plasticity and distinct selective regimes; and</paragraph>
<paragraph id="3B0D36A61F7C101B4FD4B156FA06FC68" blockId="7.[191,1442,214,969]" box="[232,1440,873,899]" pageId="7" pageNumber="6">• color has been extensively shown not to be a reliable indicator of speciation processes among dung beetles,</paragraph>
<paragraph id="3B0D36A61F7C101B4E3CB1B4FA39FC4E" blockId="7.[191,1442,214,969]" box="[256,1439,907,934]" pageId="7" pageNumber="6">
especially among the
<taxonomicName id="FCB24D251F7C101B4ECDB1B4FDC4FC4E" box="[497,610,907,933]" class="Insecta" family="Scarabaeidae" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
(
<bibRefCitation id="5F234B571F7C101B4D4EB1B4FC8EFC4D" author="Edmonds WD" box="[626,808,907,934]" pageId="7" pageNumber="6" pagination="1 - 105" refId="ref10539" refString="Edmonds WD. 1994. Revision of Phanaeus Macleay, a New World genus of Scarabaeinae dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae). Contributions in Science, Natural History Museum of Los Angeles County 443: 1 - 105." type="journal article" year="1994">Edmonds, 1994</bibRefCitation>
,
<bibRefCitation id="5F234B571F7C101B4C0FB1B3FCCBFC4D" author="Edmonds WD" box="[819,877,908,934]" pageId="7" pageNumber="6" pagination="1 - 60" refId="ref10582" refString="Edmonds WD. 2000. Revision of the Neotropical dung beetle genus Sulcophanaeus (Coleoptera: Scarabaeidae: Scarabaeinae). Folia Heyrovskyana, Supplementum 6: 1 - 60." type="journal article" year="2000">2000</bibRefCitation>
;
<bibRefCitation id="5F234B571F7C101B4C44B1B4FB03FC4D" author="Edmonds WD &amp; Zidek J." box="[888,1189,907,934]" pageId="7" pageNumber="6" pagination="1 - 58" refId="ref10613" refString="Edmonds WD, Zidek J. 2004. Revision of the Neotropical dung beetle genus Oxysternon (Scarabaeidae: Scarabaeinae: Phanaeini). Folia Heyrovskyana 11: 1 - 58." type="journal article" year="2004">Edmonds and Zídek, 2004</bibRefCitation>
,
<bibRefCitation id="5F234B571F7C101B4B8CB1B3FB4EFC4D" author="Edmonds WD &amp; Zidek J." box="[1200,1256,908,934]" pageId="7" pageNumber="6" pagination="1 - 111" refId="ref10645" refString="Edmonds WD, Zidek J. 2010. A taxonomic review of the Neotropical genus Coprophanaeus Olsoufieff, 1924 (Coleoptera: Scarabaeidae, Scarabaeinae). Insecta Mundi 129: 1 - 111." type="journal article" year="2010">2010</bibRefCitation>
, 2012;
<bibRefCitation id="5F234B571F7C101B4A05B1B4FA39FC4E" author="Solis A &amp; Kohlmann B." box="[1337,1439,907,933]" pageId="7" pageNumber="6" pagination="1 - 32" refId="ref12379" refString="Solis A, Kohlmann B. 2012. A checklist and distribution atlas of the Scarabaeinae (Coleoptera: Scarabaeidae) of Costa Rica. Zootaxa 3482: 1 - 32." type="journal article" year="2012">Solís and</bibRefCitation>
</paragraph>
<paragraph id="3B0D36A61F7C101B4E3DB191FB3AFC23" blockId="7.[191,1442,214,969]" box="[257,1180,942,969]" pageId="7" pageNumber="6">
Kohlmann 2012;
<bibRefCitation id="5F234B571F7C101B4EF9B191FC8FFC23" author="Cupello M &amp; Vaz-de-Mello FZ" box="[453,809,942,968]" pageId="7" pageNumber="6" pagination="439 - 482" refId="ref10325" refString="Cupello M, Vaz-de-Mello FZ. 2013. Taxonomic revision of the South American dung beetle genus Gromphas Brulle, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini: Gromphadina). Zootaxa 3722: 439 - 482." type="journal article" year="2013">Cupello and Vaz-de-Mello 2013</bibRefCitation>
,
<bibRefCitation id="5F234B571F7C101B4C08B191FCCAFC23" author="Cupello M &amp; Vaz-de-Mello FZ" box="[820,876,942,968]" pageId="7" pageNumber="6" pagination="399 - 400" refId="ref10365" refString="Cupello M, Vaz-de-Mello FZ. 2014. Correction of the type species of the South American genus Gromphas Brulle, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Zootaxa 3790: 399 - 400." type="journal article" year="2014">2014</bibRefCitation>
,
<bibRefCitation id="5F234B571F7C101B4C4AB191FC16FC23" author="Cupello M &amp; Vaz-de-Mello FZ" box="[886,944,942,968]" pageId="7" pageNumber="6" pagination="943 - 969" refId="ref10404" refString="Cupello M, Vaz-de-Mello FZ. 2016. A new species and the phylogeny of the South American genus Gromphas Brulle, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini). Journal of Natural History 50 (15 - 16): 943 - 969." type="journal article" year="2016">2016</bibRefCitation>
;
<bibRefCitation id="5F234B571F7C101B4C87B191FB37FC23" author="Cupello M &amp; Ribeiro-Costa CS &amp; Vaz-de-Mello FZ" box="[955,1169,942,969]" pageId="7" pageNumber="6" pagination="973 - 1034" refId="ref10278" refString="Cupello M, Ribeiro-Costa CS, Vaz-de-Mello FZ. 2022. The evolution of Bolbites onitoides (Coleoptera: Scarabaeidae: Phanaeini): its phylogenetic significance, geographical polychromatism and the subspecies problem. Zoological Journal of the Linnean Society 194: 973 - 1034." type="journal article" year="2022">Cupello et al. 2022</bibRefCitation>
).
</paragraph>
<paragraph id="3B0D36A61F7C101B4E3CB1E4FADDFB6B" blockId="7.[191,1440,987,1153]" pageId="7" pageNumber="6">
The same reasoning can be applied to
<taxonomicName id="FCB24D251F7C101B4D99B1E3FD59FC1E" authorityName="Bates" authorityYear="1887" box="[677,767,988,1013]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7C101B4D99B1E3FD59FC1E" box="[677,767,988,1013]" italics="true" pageId="7" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
, although the number of color variants and intermediaries is greater. Notably,
<taxonomicName id="FCB24D251F7C101B4E42B1C0FE7FFBF3" authorityName="Bates" authorityYear="1887" box="[382,473,1023,1048]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7C101B4E42B1C0FE7FFBF3" box="[382,473,1023,1048]" italics="true" pageId="7" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
represents the northernmost distribution of a taxon of this species group having a red color. The southernmost species with red color is
<taxonomicName id="FCB24D251F7C101B4DCDB61DFCCFFBD0" authorityName="Balthasar" authorityYear="1939" box="[753,873,1057,1083]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="funereus">
<emphasis id="09C6EAB41F7C101B4DCDB61DFCCFFBD0" box="[753,873,1057,1083]" italics="true" pageId="7" pageNumber="6">P. funereus</emphasis>
</taxonomicName>
in
<collectingCountry id="43A576361F7C101B4CB1B61EFC4AFBD0" box="[909,1004,1057,1083]" name="Ecuador" pageId="7" pageNumber="6">Ecuador</collectingCountry>
and is repeated in
<collectingCountry id="43A576361F7C101B4BFFB61EFA95FBD0" box="[1219,1331,1057,1083]" name="Colombia" pageId="7" pageNumber="6">Colombia</collectingCountry>
and Panama by
<taxonomicName id="FCB24D251F7C101B4E2BB67BFE3FFBB5" authorityName="Balthasar" authorityYear="1939" box="[279,409,1092,1118]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="olsoufieffi">
<emphasis id="09C6EAB41F7C101B4E2BB67BFE3FFBB5" box="[279,409,1092,1118]" italics="true" pageId="7" pageNumber="6">P. olsoufieffi</emphasis>
</taxonomicName>
. Interestingly, to the north of
<taxonomicName id="FCB24D251F7C101B4DD8B67BFC99FBB6" authorityName="Bates" authorityYear="1887" box="[740,831,1092,1117]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7C101B4DD8B67BFC99FBB6" box="[740,831,1092,1117]" italics="true" pageId="7" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
populations in
<collectingCountry id="43A576361F7C101B4CD2B67BFBCFFBB5" box="[1006,1129,1092,1118]" name="Costa Rica" pageId="7" pageNumber="6">Costa Rica</collectingCountry>
, only black, blue, and green species are present. Our taxonomic conclusion is that
<taxonomicName id="FCB24D251F7C101B4C27B658FCD3FB6B" authorityName="Bates" authorityYear="1887" box="[795,885,1127,1152]" class="Insecta" family="Scarabaeidae" genus="Phanaeus" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="6" phylum="Arthropoda" rank="species" species="pyrois">
<emphasis id="09C6EAB41F7C101B4C27B658FCD3FB6B" box="[795,885,1127,1152]" italics="true" pageId="7" pageNumber="6">P. pyrois</emphasis>
</taxonomicName>
is a single, monotypic, polychromatic species.
</paragraph>
</subSubSection>
</treatment>
</document>
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