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<document id="2EF88FB5AA461ED423CB02D0EE5C4208" ID-CLB-Dataset="21282" ID-DOI="10.1080/00222933.2015.1091099" ID-GBIF-Dataset="a61c3d84-bf86-420e-b8f3-18fc805c901a" ID-ISSN="1464-5262" ID-Zenodo-Dep="3990441" ID-ZooBank="FDF059DC-5D4C-4B85-80F5-841D020C307D" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1597772103698" checkinUser="carolina" docAuthor="Cupello, Mario &amp; Vaz-de-Mello, Fernando Z." docDate="2015" docId="039B9101BC582A5A16D9FBF735D0FBC3" docLanguage="en" docName="JNATHIST.50.15-16.943-969.pdf.imf" docOrigin="Journal of Natural History 50" docStyle="DocumentStyle:3C762F9783B768A90048B82ACD135A62.3:JNatHist.2017-.journal_article.0cover" docStyleId="3C762F9783B768A90048B82ACD135A62" docStyleName="JNatHist.2017-.journal_article.0cover" docStyleVersion="3" docTitle="Gromphas inermis Harold 1869" docType="treatment" docVersion="6" lastPageNumber="966" masterDocId="FFA2E979BC542A4D1709FFE4360EFFA2" masterDocTitle="A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)" masterLastPageNumber="969" masterPageNumber="943" pageNumber="955" updateTime="1698844722418" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
<mods:mods id="DBF2E26536EEC4BBBF0A240D90E0623A" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="F98F492974F52773E766C8517265DDEC">
<mods:title id="C475700F96CC4881F6CC161E159FA28A">A new species and the phylogeny of the South American genus Gromphas Brullé, 1837 (Coleoptera: Scarabaeidae: Scarabaeinae: Phanaeini)</mods:title>
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<mods:name id="DFF7982280567AA81D2F92AA2C8C803C" type="personal">
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<mods:roleTerm id="2C3109E7F1495DD35C9FEAF3376FD0B7">Author</mods:roleTerm>
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<mods:namePart id="39DE01031BA8ACF56FD4500174F336C6">Cupello, Mario</mods:namePart>
<mods:affiliation id="4ECD68A92E7465403D42F7D5976E9668">Departamento de Entomologia, Museu Nacional, Universidade Federal do Rio de Janeiro, UFRJ, Rio de Janeiro, RJ, Brazil;</mods:affiliation>
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<mods:namePart id="CA894E40FB3CE5086E9CA3A2827541EE">Vaz-de-Mello, Fernando Z.</mods:namePart>
<mods:affiliation id="FD507576A686E5A3CE4DD1D8BBCEE0CC">Departamento de Biologia e Zoologia, Instituto de Biociências, Universidade Federal de Mato Grosso, Cuiabá, MT, Brazil; &amp; Fellow of the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), Brazil</mods:affiliation>
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<mods:title id="8D5BBDD5B49AE12533A927CE5B35A6F3">Journal of Natural History</mods:title>
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<mods:part id="2F159B2C709DEB94F40CDC06E64D785C">
<mods:date id="5FE7B80EABE39EF1C093F5D4A4675137">2015</mods:date>
<mods:detail id="C408208ACE7A9D267FF7749899A22242" type="pubDate">
<mods:number id="4E29EE4D1452957720DF8F488B5CD88F">2015-09-30</mods:number>
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<mods:number id="9AD067FB5576756FCA7DF9B02B3F2634">50</mods:number>
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<mods:classification id="DEE0503730EF9D1D89E986713F561B2A">journal article</mods:classification>
<mods:identifier id="F6E35B94AA445F876A0ED30F4280AABF" type="CLB-Dataset">21282</mods:identifier>
<mods:identifier id="AC7C684AAA9E960AE571A4014019F631" type="DOI">10.1080/00222933.2015.1091099</mods:identifier>
<mods:identifier id="31EBA2C4FB21969A9BAC07E7F605319A" type="GBIF-Dataset">a61c3d84-bf86-420e-b8f3-18fc805c901a</mods:identifier>
<mods:identifier id="04E30D25189602F4D4E52024FBFFF7C3" type="ISSN">1464-5262</mods:identifier>
<mods:identifier id="B4CE619BF9F8F394B364189E4BB6CA06" type="Zenodo-Dep">3990441</mods:identifier>
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<treatment id="039B9101BC582A5A16D9FBF735D0FBC3" ID-DOI="http://doi.org/10.5281/zenodo.4328804" ID-GBIF-Taxon="166254660" ID-Zenodo-Dep="4328804" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3" httpUri="http://treatment.plazi.org/id/039B9101BC582A5A16D9FBF735D0FBC3" lastPageId="23" lastPageNumber="966" pageId="12" pageNumber="955">
<subSubSection id="C328739CBC582A4116D9FBF7355CFB8E" box="[464,850,1043,1068]" pageId="12" pageNumber="955" type="nomenclature">
<paragraph id="8B8D2017BC582A4116D9FBF7355CFB8E" blockId="12.[464,850,1043,1103]" box="[464,850,1043,1068]" pageId="12" pageNumber="955">
<heading id="D0C5977BBC582A4116D9FBF7355CFB8E" box="[464,850,1043,1068]" centered="true" fontSize="11" level="2" pageId="12" pageNumber="955" reason="2">
<taxonomicName id="4C325B94BC582A4116D9FBF7355CFB8E" ID-CoL="6L4GZ" authority="Harold, 1869" authorityName="Harold" authorityYear="1869" box="[464,850,1043,1068]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="955" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC582A4116D9FBF734BCFB8E" bold="true" box="[464,690,1043,1068]" inLineHeading="true" italics="true" pageId="12" pageNumber="955" reason="3">Gromphas inermis</emphasis>
<bibRefCitation id="EFA35DE6BC582A4115B5FBF7355CFB8E" author="Harold E" box="[700,850,1043,1068]" pageId="12" pageNumber="955" pagination="46 - 70" refId="ref15433" refString="Harold E. 1869. Ueber coprophage Lamellicornien mit besonderer Berucksichtigung der Parises Sammlungen. Coleopterol Hefte. 5: 46 - 70." type="journal article" year="1869">Harold, 1869</bibRefCitation>
</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="C328739CBC582A411524FBD237D6FB04" pageId="12" pageNumber="955" type="description">
<paragraph id="8B8D2017BC582A411524FBD234F8FBED" blockId="12.[464,850,1043,1103]" box="[557,758,1078,1103]" pageId="12" pageNumber="955">
(
<figureCitation id="13093C92BC582A41153CFBD234ACFBED" box="[565,674,1078,1103]" captionStart="Figure 5" captionStartId="10.[160,225,1281,1304]" captionTargetBox="[163,1153,142,1250]" captionTargetId="figure@10.[163,1153,142,1250]" captionTargetPageId="10" captionText="Figure 5. Updated distribution of the six species of Gromphas." figureDoi="http://doi.org/10.5281/zenodo.3990451" httpUri="https://zenodo.org/record/3990451/files/figure.png" pageId="12" pageNumber="955">Figures 5</figureCitation>
,
<figureCitation id="13093C92BC582A4115B8FBD234B1FBED" box="[689,703,1078,1103]" captionStart="Figure 6" captionStartId="16.[160,225,923,946]" captionTargetBox="[163,1153,142,892]" captionTargetId="figure@16.[163,1153,142,892]" captionTargetPageId="16" captionText="Figure 6. Phylogeny of Gromphas. Above: Cladogram showing the phylogenetic relationship of the species of Gromphas and the evolution of their characters. Black circles indicate uncontroverted synapormorphies; white circles with black margin, controverted synapomorphies; red circles, ambiguous uncontroverted synapomorphies; and white circles with red margin, ambiguous controverted synapomorphies. Below: Support values given for each clade are decay index (above branch) and bootstrap (below branch)." figureDoi="http://doi.org/10.5281/zenodo.3990455" httpUri="https://zenodo.org/record/3990455/files/figure.png" pageId="12" pageNumber="955">6</figureCitation>
,
<figureCitation id="13093C92BC582A4115C6FBD234E0FBED" box="[719,750,1078,1103]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="12" pageNumber="955">8A</figureCitation>
)
</paragraph>
<paragraph id="8B8D2017BC582A4117A9FB6937D6FB04" blockId="12.[160,1156,1165,1363]" box="[160,472,1165,1190]" pageId="12" pageNumber="955">
<heading id="D0C5977BBC582A4117A9FB6937D6FB04" box="[160,472,1165,1190]" fontSize="11" level="3" pageId="12" pageNumber="955" reason="5">
<emphasis id="B946FC05BC582A4117A9FB6937D6FB04" bold="true" box="[160,472,1165,1190]" inLineHeading="true" italics="true" pageId="12" pageNumber="955" reason="3">New geographical records</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C328739CBC582A4117A9FB5434F8FAF1" pageId="12" pageNumber="955" type="materials_examined">
<paragraph id="8B8D2017BC582A4117A9FB5434F8FAF1" blockId="12.[160,1156,1165,1363]" pageId="12" pageNumber="955">
<materialsCitation id="3B5A2A4ABC582A4117A9FB54346FFB6A" ID-GBIF-Occurrence="2831865350" box="[160,609,1200,1224]" collectorName="Gran Chaco &amp; Yacuiba" country="Bolivia" location="Tarija" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Tarija">
<emphasis id="B946FC05BC582A4117A9FB54370DFB6A" bold="true" box="[160,259,1200,1224]" pageId="12" pageNumber="955">
<collectingCountry id="F3256087BC582A4117A9FB54370DFB6A" box="[160,259,1200,1224]" name="Bolivia" pageId="12" pageNumber="955">BOLIVIA</collectingCountry>
</emphasis>
:
<collectingRegion id="49F6EEF5BC582A41161CFB54375AFB6A" box="[277,340,1200,1224]" country="Bolivia" name="Tarija" pageId="12" pageNumber="955">Tarija</collectingRegion>
:
<collectorName id="26C745C1BC582A41166AFB5437E5FB6A" box="[355,491,1200,1224]" pageId="12" pageNumber="955">Gran Chaco</collectorName>
(
<collectorName id="26C745C1BC582A4116F4FB543455FB6A" box="[509,603,1200,1224]" pageId="12" pageNumber="955">Yacuiba</collectorName>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A411579FB5435EFFB6A" ID-GBIF-Occurrence="2831865393" box="[624,993,1200,1224]" collectorName="Sapucai" country="Paraguay" location="Paraguari" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Paraguari">
<emphasis id="B946FC05BC582A411579FB5434F6FB6A" bold="true" box="[624,760,1200,1224]" pageId="12" pageNumber="955">
<collectingCountry id="F3256087BC582A411579FB5434F6FB6A" box="[624,760,1200,1224]" name="Paraguay" pageId="12" pageNumber="955">PARAGUAY</collectingCountry>
</emphasis>
:
<collectingRegion id="49F6EEF5BC582A411402FB543576FB6A" box="[779,888,1200,1224]" country="Paraguay" name="Paraguari" pageId="12" pageNumber="955">Paraguarí</collectingRegion>
:
<collectorName id="26C745C1BC582A41148EFB5435EFFB6A" box="[903,993,1200,1224]" pageId="12" pageNumber="955">Sapucaí</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A4114E6FB543410FB49" ID-GBIF-Occurrence="2831865463" collectorName="Cordoba &amp; Leones" country="Argentina" location="Jujuy" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Jujuy">
<emphasis id="B946FC05BC582A4114E6FB543273FB6A" bold="true" box="[1007,1149,1200,1224]" pageId="12" pageNumber="955">
<collectingCountry id="F3256087BC582A4114E6FB543273FB6A" box="[1007,1149,1200,1224]" name="Argentina" pageId="12" pageNumber="955">ARGENTINA</collectingCountry>
</emphasis>
:
<collectingRegion id="49F6EEF5BC582A4117A9FB3736D3FB49" box="[160,221,1235,1259]" country="Argentina" name="Jujuy" pageId="12" pageNumber="955">Jujuy</collectingRegion>
.
<collectingRegion id="49F6EEF5BC582A4117E1FB37375DFB49" box="[232,339,1235,1259]" country="Argentina" name="Tucuman" pageId="12" pageNumber="955">Tucumán</collectingRegion>
.
<collectorName id="26C745C1BC582A411657FB3737CFFB49" box="[350,449,1235,1259]" pageId="12" pageNumber="955">Córdoba</collectorName>
:
<collectorName id="26C745C1BC582A4116C5FB373410FB49" box="[460,542,1235,1259]" pageId="12" pageNumber="955">Leones</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A411520FB3736B3FAAF" ID-GBIF-Occurrence="2831865304" collectorName="Rosario" country="Argentina" location="General Obligado" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Santa Fe">
<collectingRegion id="49F6EEF5BC582A411520FB373483FB49" box="[553,653,1235,1259]" country="Argentina" name="Santa Fe" pageId="12" pageNumber="955">Santa Fe</collectingRegion>
:
<location id="8EED76CCBC582A411591FB373550FB49" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC582A411591FB373550FB49" box="[664,862,1235,1259]" country="Argentina" name="General Obligado" pageId="12" pageNumber="955" stateProvince="Santa Fe">General Obligado</location>
(Villa Ana),
<collectorName id="26C745C1BC582A4114ECFB373234FB49" box="[997,1082,1235,1259]" pageId="12" pageNumber="955">Rosario</collectorName>
,
<collectingRegion id="49F6EEF5BC582A41134CFB3736B3FAAF" country="Argentina" name="Santa Fe" pageId="12" pageNumber="955">Santa Fe</collectingRegion>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A4117D9FB11349CFAAF" ID-GBIF-Occurrence="2831865399" box="[208,658,1269,1293]" collectorName="Puerto Iguazu" country="Argentina" location="San Ignacio" municipality="Misiones" pageId="12" pageNumber="955" specimenCount="1">
<collectingMunicipality id="6BE9BA6DBC582A4117D9FB11373BFAAF" box="[208,309,1269,1293]" pageId="12" pageNumber="955">Misiones</collectingMunicipality>
:
<location id="8EED76CCBC582A411641FB1137D8FAAF" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC582A411641FB1137D8FAAF" box="[328,470,1269,1293]" country="Argentina" municipality="Misiones" name="San Ignacio" pageId="12" pageNumber="955">San Ignacio</location>
,
<collectorName id="26C745C1BC582A4116E0FB11349CFAAF" box="[489,658,1269,1293]" pageId="12" pageNumber="955">Puerto Iguazú</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A4115ACFB1135BCFAAF" ID-GBIF-Occurrence="2831865473" box="[677,946,1269,1293]" collectorName="San Roque" country="Argentina" location="Corrientes" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Corrientes">
<collectingRegion id="49F6EEF5BC582A4115ACFB113514FAAF" box="[677,794,1269,1293]" country="Argentina" name="Corrientes" pageId="12" pageNumber="955">Corrientes</collectingRegion>
:
<collectorName id="26C745C1BC582A411427FB1135BCFAAF" box="[814,946,1269,1293]" pageId="12" pageNumber="955">San Roque</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A4114CCFB113717FA92" ID-GBIF-Occurrence="2831865439" collectorName="Villa Paranacito" country="Argentina" location="Entre Rios" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Entre Rios">
<collectingRegion id="49F6EEF5BC582A4114CCFB11324CFAAF" box="[965,1090,1269,1293]" country="Argentina" name="Entre Rios" pageId="12" pageNumber="955">Entre Ríos</collectingRegion>
:
<collectorName id="26C745C1BC582A41135CFB113717FA92" pageId="12" pageNumber="955">Villa Paranacito</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A411638FAFC35F0FA92" ID-GBIF-Occurrence="2831865371" box="[305,1022,1304,1328]" collectorName="Pergamino &amp; San Pedro" country="Argentina" location="Isla Martin Garcia" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Ciudad Autonoma de Buenos Aires">
<collectingRegion id="49F6EEF5BC582A411638FAFC37DDFA92" box="[305,467,1304,1328]" country="Argentina" name="Ciudad Autonoma de Buenos Aires" pageId="12" pageNumber="955">Buenos Aires</collectingRegion>
:
<location id="8EED76CCBC582A4116E5FAFC34C3FA92" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC582A4116E5FAFC34C3FA92" box="[492,717,1304,1328]" country="Argentina" name="Isla Martin Garcia" pageId="12" pageNumber="955" stateProvince="Ciudad Autonoma de Buenos Aires">Isla Martín García</location>
,
<collectorName id="26C745C1BC582A4115ECFAFC356CFA92" box="[741,866,1304,1328]" pageId="12" pageNumber="955">Pergamino</collectorName>
,
<collectorName id="26C745C1BC582A411472FAFC35F0FA92" box="[891,1022,1304,1328]" pageId="12" pageNumber="955">San Pedro</collectorName>
</materialsCitation>
.
<collectingRegion id="49F6EEF5BC582A41131FFAFC328EFA92" box="[1046,1152,1304,1328]" country="Argentina" name="Mendoza" pageId="12" pageNumber="955">Mendoza</collectingRegion>
.
<materialsCitation id="3B5A2A4ABC582A4117A9FADF37F2FAF1" ID-GBIF-Occurrence="2831865395" box="[160,508,1339,1363]" collectorName="Riachuelo" country="Uruguay" location="Colonia" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Colonia">
<emphasis id="B946FC05BC582A4117A9FADF3714FAF1" bold="true" box="[160,282,1339,1363]" pageId="12" pageNumber="955">
<collectingCountry id="F3256087BC582A4117A9FADF3714FAF1" box="[160,282,1339,1363]" name="Uruguay" pageId="12" pageNumber="955">URUGUAY</collectingCountry>
</emphasis>
:
<collectingRegion id="49F6EEF5BC582A411621FADF378EFAF1" box="[296,384,1339,1363]" country="Uruguay" name="Colonia" pageId="12" pageNumber="955">Colonia</collectingRegion>
:
<collectorName id="26C745C1BC582A411685FADF37F2FAF1" box="[396,508,1339,1363]" pageId="12" pageNumber="955">Riachuelo</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC582A41150EFADF34FDFAF1" ID-GBIF-Occurrence="2831865450" box="[519,755,1339,1363]" collectorName="Atlantida" country="Uruguay" location="Canelones" pageId="12" pageNumber="955" specimenCount="1" stateProvince="Canelones">
<collectingRegion id="49F6EEF5BC582A41150EFADF3470FAF1" box="[519,638,1339,1363]" country="Uruguay" name="Canelones" pageId="12" pageNumber="955">Canelones</collectingRegion>
:
<collectorName id="26C745C1BC582A411583FADF34FDFAF1" box="[650,755,1339,1363]" pageId="12" pageNumber="955">Atlántida</collectorName>
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C328739CBC582A5A17A9FA7535D0FBC3" lastPageId="23" lastPageNumber="966" pageId="12" pageNumber="955" type="discussion">
<paragraph id="8B8D2017BC582A4117A9FA753710FA08" blockId="12.[160,1156,1425,1727]" box="[160,286,1425,1450]" pageId="12" pageNumber="955">
<heading id="D0C5977BBC582A4117A9FA753710FA08" box="[160,286,1425,1450]" fontSize="11" level="3" pageId="12" pageNumber="955" reason="5">
<emphasis id="B946FC05BC582A4117A9FA753710FA08" bold="true" box="[160,286,1425,1450]" inLineHeading="true" italics="true" pageId="12" pageNumber="955" reason="3">Comments</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC582A4017A9FA513459FF6B" blockId="12.[160,1156,1425,1727]" lastBlockId="13.[160,1156,142,651]" lastPageId="13" lastPageNumber="956" pageId="12" pageNumber="955">
In
<bibRefCitation id="EFA35DE6BC582A4117B4FA513423FA6F" author="Cupello M &amp; Vaz-de-Mello FZ" box="[189,557,1461,1485]" pageId="12" pageNumber="955" pagination="439 - 482" refId="ref14473" 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>
, we cited the occurrence of
<taxonomicName id="4C325B94BC582A411467FA5135D5FA6F" authorityName="Harold" authorityYear="1869" box="[878,987,1461,1485]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="955" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC582A411467FA5135D5FA6F" box="[878,987,1461,1485]" italics="true" pageId="12" pageNumber="955">G. inermis</emphasis>
</taxonomicName>
in the Bolivian department of Beni based on the report of this species (cited as
<emphasis id="B946FC05BC582A41147FFA333572FA4D" box="[886,892,1495,1519]" italics="true" pageId="12" pageNumber="955">‘</emphasis>
<taxonomicName id="4C325B94BC582A411475FA33325DFA4D" authority="Brulle" authorityName="Brulle" authorityYear="1834" box="[892,1107,1495,1519]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="955" phylum="Arthropoda" rank="species" species="lacordairei">
<emphasis id="B946FC05BC582A411475FA333205FA4D" box="[892,1035,1495,1519]" italics="true" pageId="12" pageNumber="955">G. lacordairei</emphasis>
Brullé
</taxonomicName>
<emphasis id="B946FC05BC582A41135AFA333257FA4D" box="[1107,1113,1495,1519]" italics="true" pageId="12" pageNumber="955">’</emphasis>
) by
<bibRefCitation id="EFA35DE6BC582A4117A9FA1E37C6F9B0" author="Hamel-Leigue AC &amp; Herzog SK &amp; Mann DJ &amp; Larsen TH &amp; Gill BD &amp; Edmonds WD &amp; Spector S" box="[160,456,1530,1554]" pageId="12" pageNumber="955" pagination="43 - 95" refId="ref15324" refString="Hamel-Leigue AC, Herzog SK, Mann DJ, Larsen TH, Gill BD, Edmonds WD, Spector S. 2009. Distribucion e historia natural de escarabajos coprofagos de la tribu Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae) en Bolivia. Kempffiana. 5: 43 - 95." type="journal article" year="2009">Hamel-Leigue et al. (2009)</bibRefCitation>
. Nonetheless, we know now that the specimens examined by them from Beni are, in fact,
<taxonomicName id="4C325B94BC582A4116E0F9F93441F997" box="[489,591,1565,1589]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="955" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC582A4116E0F9F93441F997" box="[489,591,1565,1589]" italics="true" pageId="12" pageNumber="955">G. jardim</emphasis>
</taxonomicName>
, and so Beni and the biogeographic province of Rondônia (former Pantanal province, as cited in
<bibRefCitation id="EFA35DE6BC582A4115CCF9DB3226F9F5" author="Cupello M &amp; Vaz-de-Mello FZ" box="[709,1064,1599,1623]" pageId="12" pageNumber="955" pagination="439 - 482" refId="ref14473" 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>
) should be disregarded as an area of occurrence of
<taxonomicName id="4C325B94BC582A41158DF98634FEF9D8" authorityName="Harold" authorityYear="1869" box="[644,752,1634,1658]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="955" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC582A41158DF98634FEF9D8" box="[644,752,1634,1658]" italics="true" pageId="12" pageNumber="955">G. inermis</emphasis>
</taxonomicName>
. On the other hand, localities of the Chacoan province in the departments of Tarija and Santa Cruz, also cited by Hamel- Leigue et al. (2009), are indeed places where
<taxonomicName id="4C325B94BC582A411595F9433507F91D" authorityName="Harold" authorityYear="1869" box="[668,777,1703,1727]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="12" pageNumber="955" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC582A411595F9433507F91D" box="[668,777,1703,1727]" italics="true" pageId="12" pageNumber="955">G. inermis</emphasis>
</taxonomicName>
is present. The occurrence of this species in
<collectingCountry id="F3256087BC592A40161DFF6A37A1FF04" box="[276,431,142,166]" name="Argentina" pageId="13" pageNumber="956">the Argentine</collectingCountry>
province of
<collectingRegion id="49F6EEF5BC592A401535FF6A34A8FF04" box="[572,678,142,166]" country="Argentina" name="Mendoza" pageId="13" pageNumber="956">Mendoza</collectingRegion>
, as first reported here, is the most western record known for
<taxonomicName id="4C325B94BC592A401667FF5537D2FF6B" authorityName="Harold" authorityYear="1869" box="[366,476,177,201]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC592A401667FF5537D2FF6B" box="[366,476,177,201]" italics="true" pageId="13" pageNumber="956">G. inermis</emphasis>
</taxonomicName>
(
<figureCitation id="13093C92BC592A4016E5FF553444FF6B" box="[492,586,177,201]" captionStart="Figure 5" captionStartId="10.[160,225,1281,1304]" captionTargetBox="[163,1153,142,1250]" captionTargetId="figure@10.[163,1153,142,1250]" captionTargetPageId="10" captionText="Figure 5. Updated distribution of the six species of Gromphas." figureDoi="http://doi.org/10.5281/zenodo.3990451" httpUri="https://zenodo.org/record/3990451/files/figure.png" pageId="13" pageNumber="956">Figure 5</figureCitation>
).
</paragraph>
<paragraph id="8B8D2017BC592A4017C9FF3037D7FD29" blockId="13.[160,1156,142,651]" pageId="13" pageNumber="956">
During our examination of specimens for the revision of
<taxonomicName id="4C325B94BC592A40147DFF3035EBFF4E" authorityName="Brulle" authorityYear="1837" box="[884,997,212,236]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC592A40147DFF3035EBFF4E" box="[884,997,212,236]" italics="true" pageId="13" pageNumber="956">Gromphas</emphasis>
</taxonomicName>
, we found a female from
<collectingRegion id="49F6EEF5BC592A40163AFF12378AFEAC" box="[307,388,246,270]" country="Uruguay" name="Artigas" pageId="13" pageNumber="956">Artigas</collectingRegion>
,
<collectingCountry id="F3256087BC592A401699FF1237FCFEAC" box="[400,498,246,270]" name="Uruguay" pageId="13" pageNumber="956">Uruguay</collectingCountry>
, that is very interesting by being matte black and having the pronotal surface completely irregular; nonetheless, taking into account all the other characteristics, it fits in
<taxonomicName id="4C325B94BC592A4016A3FED83418FEF6" authorityName="Harold" authorityYear="1869" box="[426,534,316,340]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC592A4016A3FED83418FEF6" box="[426,534,316,340]" italics="true" pageId="13" pageNumber="956">G. inermis</emphasis>
</taxonomicName>
, and so we identified that specimen with this name in
<bibRefCitation id="EFA35DE6BC592A4017A9FEBA3411FED4" author="Cupello M &amp; Vaz-de-Mello FZ" box="[160,543,350,374]" pageId="13" pageNumber="956" pagination="439 - 482" refId="ref14473" 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>
. Since then, we examined the Hermann Burmeister collection, housed in MACN, and there we found two other Uruguayan females (from
<emphasis id="B946FC05BC592A4017A9FE4736A8FE19" box="[160,166,419,443]" italics="true" pageId="13" pageNumber="956">‘</emphasis>
Banda Orient.
<emphasis id="B946FC05BC592A401637FE47374DFE19" box="[318,323,419,443]" italics="true" pageId="13" pageNumber="956">’</emphasis>
) with a very similar colour and irregular pronotum that were identified by Burmeister with the unavailable name
<emphasis id="B946FC05BC592A401555FE22346CFE7C" box="[604,610,454,478]" italics="true" pageId="13" pageNumber="956">‘</emphasis>
<taxonomicName id="4C325B94BC592A40156BFE223574FE7C" authority="Nob." authorityName="Nob." box="[610,890,454,478]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="species" species="rugicollis">
<emphasis id="B946FC05BC592A40156BFE223535FE7C" box="[610,827,454,478]" italics="true" pageId="13" pageNumber="956">Gromphas rugicollis</emphasis>
Nob.
</taxonomicName>
<emphasis id="B946FC05BC592A401470FE223570FE7C" box="[889,894,454,478]" italics="true" pageId="13" pageNumber="956">’</emphasis>
. For now, we consider these differences as intraspecific variations of
<taxonomicName id="4C325B94BC592A4015B6FE0D353EFDA3" authorityName="Harold" authorityYear="1869" box="[703,816,489,513]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC592A4015B6FE0D353EFDA3" box="[703,816,489,513]" italics="true" pageId="13" pageNumber="956">G. inermis</emphasis>
</taxonomicName>
(possibly teratological specimens); we observed black specimens of
<taxonomicName id="4C325B94BC592A401563FDE834D6FD86" authorityName="Harold" authorityYear="1869" box="[618,728,524,548]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC592A401563FDE834D6FD86" box="[618,728,524,548]" italics="true" pageId="13" pageNumber="956">G. inermis</emphasis>
</taxonomicName>
from other regions and, in fact, black and brown colorations seems to be related to teneral specimens, which would also explain the irregular surface of pronotum. All the other Uruguayan specimens examined by us are typical
<taxonomicName id="4C325B94BC592A40166CFD9737DCFD29" authorityName="Harold" authorityYear="1869" box="[357,466,627,651]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="13" pageNumber="956" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC592A40166CFD9737DCFD29" box="[357,466,627,651]" italics="true" pageId="13" pageNumber="956">G. inermis</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B8D2017BC592A4017A9FD5D340BFD70" blockId="13.[160,1156,697,1727]" box="[160,517,697,722]" pageId="13" pageNumber="956">
<heading id="D0C5977BBC592A4017A9FD5D340BFD70" box="[160,517,697,722]" fontSize="11" level="3" pageId="13" pageNumber="956" reason="5">
<emphasis id="B946FC05BC592A4017A9FD5D340BFD70" bold="true" box="[160,517,697,722]" inLineHeading="true" italics="true" pageId="13" pageNumber="956" reason="3">Additional material examined</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC592A4017A9FD38328AFAE0" blockId="13.[160,1156,697,1727]" pageId="13" pageNumber="956">
<emphasis id="B946FC05BC592A4017A9FD38373FFD56" bold="true" box="[160,305,732,756]" pageId="13" pageNumber="956">
<collectingCountry id="F3256087BC592A4017A9FD38373FFD56" box="[160,305,732,756]" name="Argentina" pageId="13" pageNumber="956">ARGENTINA</collectingCountry>
</emphasis>
: 1904, O
<materialsCitation id="3B5A2A4ABC592A401696FD3837C8FD56" ID-GBIF-Occurrence="2831865343" box="[415,454,732,756]" country="Argentina" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">. W</materialsCitation>
. Thomas col
<materialsCitation id="3B5A2A4ABC592A401554FD383751FCB5" ID-GBIF-Occurrence="2831865468" collectionCode="BMNH" country="Argentina" location="Rio Bermejo" pageId="13" pageNumber="956" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A401565FD383477FD56" box="[620,633,732,756]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40158DFD3834DBFD56" box="[644,725,732,756]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A40141FFD383572FD56" box="[790,892,732,756]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401484FD3835D5FD56" box="[909,987,732,756]" pageId="13" pageNumber="956">BMNH</collectionCode>
);
<location id="8EED76CCBC592A4014F9FD38328EFD56" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A4014F9FD38328EFD56" box="[1008,1152,732,756]" country="Argentina" name="Rio Bermejo" pageId="13" pageNumber="956">Río Bermejo</location>
, without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401656FD1B37FFFCB5" ID-GBIF-Occurrence="2831865432" box="[351,497,767,791]" collectionCode="BMNH" collectorName="Richter" country="Argentina" isTailEnumeration="true" location="Rio Bermejo" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A401679FD1B37CEFCB5" box="[368,448,767,791]" pageId="13" pageNumber="956">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4016F8FD1B3471FC98" ID-GBIF-Occurrence="2831865308" collectionCode="MLPA, BMNH" country="Argentina" location="Rio Salado" pageId="13" pageNumber="956" specimenCount="6" specimenCount-female="3" specimenCount-male="3">
.
<emphasis id="B946FC05BC592A40150BFD1B3401FCB5" box="[514,527,767,791]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401515FD1B3461FCB5" box="[540,623,767,791]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A4015BDFD1B3512FCB5" box="[692,796,767,791]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401426FD1B3579FCB5" box="[815,887,767,791]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<location id="8EED76CCBC592A401484FD1B3202FCB5" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A401484FD1B3202FCB5" box="[909,1036,767,791]" country="Argentina" name="Rio Salado" pageId="13" pageNumber="956">Río Salado</location>
, no more data
<emphasis id="B946FC05BC592A4017E8FCC636E0FC98" box="[225,238,802,826]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4017F5FCC63754FC98" box="[252,346,802,826]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A4016AAFCC63419FC98" box="[419,535,802,826]" pageId="13" pageNumber="956" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401522FCC63477FC98" box="[555,633,802,826]" pageId="13" pageNumber="956">BMNH</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC592A401599FCC634A3FCFE" ID-GBIF-Occurrence="2831865348" collectionCode="BMNH, MACN" country="Argentina" location="BUENOS AIRES" pageId="13" pageNumber="956" specimenCount="3" specimenCount-female="1" specimenCount-male="2" stateProvince="Ciudad Autonoma de Buenos Aires">
<collectingRegion id="49F6EEF5BC592A401599FCC6354CFC98" box="[656,834,802,826]" country="Argentina" name="Ciudad Autonoma de Buenos Aires" pageId="13" pageNumber="956">BUENOS AIRES</collectingRegion>
: no more data
<emphasis id="B946FC05BC592A401300FCC63218FC98" box="[1033,1046,802,826]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40132CFCC6328DFC98" box="[1061,1155,802,826]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017AFFCA036FAFCFE" box="[166,244,836,860]" pageId="13" pageNumber="956">BMNH</collectionCode>
),
<specimenCount id="9D34EB9EBC592A401602FCA0377DFCFE" box="[267,371,836,860]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A40168EFCA037DBFCFE" box="[391,469,836,860]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
); without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4015A4FCA03531FCFE" ID-GBIF-Occurrence="2831865417" box="[685,831,836,860]" collectionCode="BMNH, MACN" collectorName="Richter" country="Argentina" isTailEnumeration="true" location="BUENOS AIRES" pageId="13" pageNumber="956" specimenCount="1" stateProvince="Ciudad Autonoma de Buenos Aires">
.
<collectorName id="26C745C1BC592A4015B7FCA03500FCFE" box="[702,782,836,860]" pageId="13" pageNumber="956">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401449FCA037B9FCDD" ID-GBIF-Occurrence="2831865317" collectionCode="MLPA" country="Argentina" pageId="13" pageNumber="956" specimenCount="5" specimenCount-female="3" specimenCount-male="2">
.
<emphasis id="B946FC05BC592A401459FCA03553FCFE" box="[848,861,836,860]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401462FCA035C7FCFE" box="[875,969,836,860]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401319FCA0328DFCFE" box="[1040,1155,836,860]" pageId="13" pageNumber="956" type="female">3 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017AFFC8336E0FCDD" box="[166,238,871,895]" pageId="13" pageNumber="956">MLPA</collectionCode>
); without date, J
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4016B1FC833420FCDD" ID-GBIF-Occurrence="2831865362" box="[440,558,871,895]" collectionCode="MLPA" collectorName="Bosq" country="Argentina" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A4016CEFC8337F0FCDD" box="[455,510,871,895]" pageId="13" pageNumber="956">Bosq</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401527FC83328EFCDD" ID-GBIF-Occurrence="2831865303" box="[558,1152,871,895]" collectingDate="1898-10" collectionCode="MLPA" country="Argentina" pageId="13" pageNumber="956" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A401535FC833447FCDD" box="[572,585,871,895]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40155CFC8334A9FCDD" box="[597,679,871,895]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A4015E1FC833541FCDD" box="[744,847,871,895]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401469FC8335A6FCDD" box="[864,936,871,895]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<date id="FF8C06D7BC592A4014B4FC833251FCDD" box="[957,1119,871,895]" pageId="13" pageNumber="956" value="1898-10">
<collectingDate id="EFC8FF3FBC592A4014B4FC833251FCDD" box="[957,1119,871,895]" pageId="13" pageNumber="956" value="1898-10">October 1898</collectingDate>
</date>
, C
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401389FC833718FC00" ID-GBIF-Occurrence="2831865349" collectingDate="1898-10" collectionCode="MLPA" collectorName="Bruch" country="Argentina" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A4017A9FC6E36ECFC00" box="[160,226,906,930]" pageId="13" pageNumber="956">Bruch</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A40161FFC6E35F2FC00" ID-GBIF-Occurrence="2831865311" box="[278,1020,906,930]" collectingDate="1903-09-10" collectionCode="MLPA" collectorName="Carlos Bruch" country="Argentina" pageId="13" pageNumber="956" specimenCount="1" specimenCount-female="1">
.
<emphasis id="B946FC05BC592A401620FC6E3738FC00" box="[297,310,906,930]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40164EFC6E37BCFC00" box="[327,434,906,930]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4016C1FC6E341EFC00" box="[456,528,906,930]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<date id="FF8C06D7BC592A401520FC6E3513FC00" box="[553,797,906,930]" pageId="13" pageNumber="956" value="1903-09-10">
<collectingDate id="EFC8FF3FBC592A401520FC6E3513FC00" box="[553,797,906,930]" pageId="13" pageNumber="956" value="1903-09-10">10 September 1903</collectingDate>
</date>
,
<collectorName id="26C745C1BC592A401439FC6E35C6FC00" box="[816,968,906,930]" pageId="13" pageNumber="956">Carlos Bruch</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4014F4FC6E34CCFC66" ID-GBIF-Occurrence="2831865474" collectingDate="1904-03-02" collectionCode="MACN" collectorName="Carlos Bruch" country="Argentina" pageId="13" pageNumber="956" specimenCount="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A401319FC6E3213FC00" box="[1040,1053,906,930]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401327FC6E328AFC00" box="[1070,1156,906,930]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017AFFC4836FEFC66" box="[166,240,940,964]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC592A4017F2FC483706FC66" box="[251,264,940,964]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<collectorName id="26C745C1BC592A40161DFC4837A9FC66" box="[276,423,940,964]" pageId="13" pageNumber="956">Carlos Bruch</collectorName>
coll.);
<date id="FF8C06D7BC592A4016F3FC4834AEFC66" box="[506,672,940,964]" pageId="13" pageNumber="956" value="1904-03-02">
<collectingDate id="EFC8FF3FBC592A4016F3FC4834AEFC66" box="[506,672,940,964]" pageId="13" pageNumber="956" value="1904-03-02">2 March 1904</collectingDate>
</date>
, A
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4015CBFC483533FC66" ID-GBIF-Occurrence="2831865352" box="[706,829,940,964]" collectingDate="1904-03-02" collectionCode="MACN" collectorName="Zotta" country="Argentina" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A4015D9FC483503FC66" box="[720,781,940,964]" pageId="13" pageNumber="956">Zotta</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401434FC483770FBA8" ID-GBIF-Occurrence="2831865305" collectionCode="MACN" country="Argentina" location="Deceber" pageId="13" pageNumber="956" specimenCount="3" specimenCount-female="1" specimenCount-male="2">
.
<emphasis id="B946FC05BC592A401445FC483557FC66" box="[844,857,940,964]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40146CFC4835B8FC66" box="[869,950,940,964]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4014CEFC48321BFC66" box="[967,1045,940,964]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
); without locality,
<location id="8EED76CCBC592A40160CFC2B376AFC45" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A40160CFC2B376AFC45" box="[261,356,975,999]" country="Argentina" name="Deceber" pageId="13" pageNumber="956">Deceber</location>
1921, without collector
<emphasis id="B946FC05BC592A40159DFC2B34AFFC45" box="[660,673,975,999]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4015A6FC2B350DFC45" box="[687,771,975,999]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401440FC2B35BFFC45" box="[841,945,975,999]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4014CCFC2B321DFC45" box="[965,1043,975,999]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
); without locality and date, J
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401677FC1637E1FBA8" ID-GBIF-Occurrence="2831865380" box="[382,495,1010,1034]" collectionCode="MACN" collectorName="Boso" country="Argentina" isTailEnumeration="true" location="Deceber" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A401682FC1637CCFBA8" box="[395,450,1010,1034]" pageId="13" pageNumber="956">Boso</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4016E6FC1637CCFB8E" ID-GBIF-Occurrence="2831865375" collectingDate="1980-01-08" collectionCode="MACN" country="Argentina" location="Buenos Aires" pageId="13" pageNumber="956" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A4016F5FC163407FBA8" box="[508,521,1010,1034]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40151AFC16346CFBA8" box="[531,610,1010,1034]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401596FC16350DFBA8" box="[671,771,1010,1034]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A40141AFC16356FFBA8" box="[787,865,1010,1034]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<quantity id="4CCA8DF2BC592A40147DFC1635B2FBA8" box="[884,956,1010,1034]" metricMagnitude="4" metricUnit="m" metricValue="1.7" pageId="13" pageNumber="956" unit="km" value="17.0">17 km</quantity>
south of
<location id="8EED76CCBC592A401326FC1636D5FB8E" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A401326FC1636D5FB8E" country="Argentina" name="Buenos Aires" pageId="13" pageNumber="956">Buenos Aires</location>
,
<date id="FF8C06D7BC592A4017E3FBF037AFFB8E" box="[234,417,1044,1068]" pageId="13" pageNumber="956" value="1980-01-08">
<collectingDate id="EFC8FF3FBC592A4017E3FBF037AFFB8E" box="[234,417,1044,1068]" pageId="13" pageNumber="956" value="1980-01-08">8 January 1980</collectingDate>
</date>
, C
</materialsCitation>
. and M.
<materialsCitation id="3B5A2A4ABC592A401526FBF034A2FB8E" ID-GBIF-Occurrence="2831865451" box="[559,684,1044,1068]" country="Argentina" location="Vardy" pageId="13" pageNumber="956" specimenCount="1">
<location id="8EED76CCBC592A401526FBF0347EFB8E" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A401526FBF0347EFB8E" box="[559,624,1044,1068]" country="Argentina" name="Vardy" pageId="13" pageNumber="956">Vardy</location>
cols
</materialsCitation>
.
<emphasis id="B946FC05BC592A4015B2FBF034C6FB8E" box="[699,712,1044,1068]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<materialsCitation id="3B5A2A4ABC592A4015DDFBF03749FBED" ID-GBIF-Occurrence="2831865374" collectingDate="1937-04" collectionCode="BMNH" collectorName="Isla Martin Garcia" country="Argentina" pageId="13" pageNumber="956" specimenCount="1" specimenCount-female="1">
<specimenCount id="9D34EB9EBC592A4015DDFBF03534FB8E" box="[724,826,1044,1068]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401444FBF03595FB8E" box="[845,923,1044,1068]" pageId="13" pageNumber="956">BMNH</collectionCode>
);
<collectorName id="26C745C1BC592A4014A6FBF0328EFB8E" box="[943,1152,1044,1068]" pageId="13" pageNumber="956">Isla Martín Garcia</collectorName>
,
<date id="FF8C06D7BC592A4017A9FBD33710FBED" box="[160,286,1079,1103]" pageId="13" pageNumber="956" value="1937-04">
<collectingDate id="EFC8FF3FBC592A4017A9FBD33710FBED" box="[160,286,1079,1103]" pageId="13" pageNumber="956" value="1937-04">April 1937</collectingDate>
</date>
, M
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A40164EFBD3376BFBED" ID-GBIF-Occurrence="2831865321" box="[327,357,1079,1103]" collectingDate="1937-04" collectionCode="BMNH" collectorName="Isla Martin Garcia" country="Argentina" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">. J</materialsCitation>
. Viana col
<materialsCitation id="3B5A2A4ABC592A4016EEFBD33528FBD0" ID-GBIF-Occurrence="2831865387" collectionCode="MACN, BMNH" collectorName="La Plata" country="Argentina" location="La Plata" pageId="13" pageNumber="956" specimenCount="4" specimenCount-male="4">
.
<emphasis id="B946FC05BC592A4016F1FBD3340BFBED" box="[504,517,1079,1103]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40151DFBD33469FBED" box="[532,615,1079,1103]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401572FBD334C7FBED" box="[635,713,1079,1103]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<location id="8EED76CCBC592A4015E9FBD3354DFBED" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A4015E9FBD3354DFBED" box="[736,835,1079,1103]" country="Argentina" name="La Plata" pageId="13" pageNumber="956">La Plata</location>
, no more data
<emphasis id="B946FC05BC592A401303FBD33219FBED" box="[1034,1047,1079,1103]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40132CFBD3328DFBED" box="[1061,1155,1079,1103]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017AFFBBE36FCFBD0" box="[166,242,1114,1138]" pageId="13" pageNumber="956">BMNH</collectionCode>
) and
<specimenCount id="9D34EB9EBC592A401630FBBE3784FBD0" box="[313,394,1114,1138]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401692FBBE37E7FBD0" box="[411,489,1114,1138]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC592A4016F4FBBE3453FBD0" box="[509,605,1114,1138]" pageId="13" pageNumber="956">La Plata</collectorName>
, without date, A
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A40142FFBBE354BFBD0" ID-GBIF-Occurrence="2831865408" box="[806,837,1114,1138]" collectionCode="MACN, BMNH" collectorName="La Plata" country="Argentina" isTailEnumeration="true" location="La Plata" pageId="13" pageNumber="956" specimenCount="1">. R</materialsCitation>
. Bezzi col
<materialsCitation id="3B5A2A4ABC592A4014B4FBBE347FFB15" ID-GBIF-Occurrence="2831865410" collectingDate="1924-02-16" collectingDateMax="1949-02" collectingDateMin="1924-02-16" collectionCode="MLPA" collectorName="San Isidro &amp; Martinez" country="Argentina" location="Pergamino" pageId="13" pageNumber="956" specimenCount="3" specimenCount-female="1" specimenCount-male="2">
.
<emphasis id="B946FC05BC592A4014C2FBBE35D6FBD0" box="[971,984,1114,1138]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4014EDFBBE323AFBD0" box="[996,1076,1114,1138]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A40137CFBBE36E3FB36" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401609FB983746FB36" box="[256,328,1148,1172]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<location id="8EED76CCBC592A401657FB9837D3FB36" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A401657FB9837D3FB36" box="[350,477,1148,1172]" country="Argentina" name="Pergamino" pageId="13" pageNumber="956">Pergamino</location>
,
<date id="FF8C06D7BC592A4016E5FB983496FB36" box="[492,664,1148,1172]" pageId="13" pageNumber="956" value="1949-02">
<collectingDate id="EFC8FF3FBC592A4016E5FB983496FB36" box="[492,664,1148,1172]" pageId="13" pageNumber="956" value="1949-02">February 1949</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC592A401474FB983584FB36" box="[893,906,1148,1172]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40149EFB9835E4FB36" box="[919,1002,1148,1172]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4014F5FB98324AFB36" box="[1020,1092,1148,1172]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<collectorName id="26C745C1BC592A401353FB9836ECFB15" pageId="13" pageNumber="956">San Isidro</collectorName>
,
<collectorName id="26C745C1BC592A4017FDFB7B3754FB15" box="[244,346,1183,1207]" pageId="13" pageNumber="956">Martínez</collectorName>
,
<date id="FF8C06D7BC592A401664FB7B3448FB15" box="[365,582,1183,1207]" pageId="13" pageNumber="956" value="1924-02-16">
<collectingDate id="EFC8FF3FBC592A401664FB7B3448FB15" box="[365,582,1183,1207]" pageId="13" pageNumber="956" value="1924-02-16">16 February 1924</collectingDate>
</date>
, M
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401579FB7B34E2FB15" ID-GBIF-Occurrence="2831865313" box="[624,748,1183,1207]" collectingDate="1924-02-16" collectingDateMax="1949-02" collectingDateMin="1924-02-16" collectionCode="MLPA" collectorName="Sires" country="Argentina" isTailEnumeration="true" location="Pergamino" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A40158DFB7B34B7FB15" box="[644,697,1183,1207]" pageId="13" pageNumber="956">Sires</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4015E5FB7B3753FB78" ID-GBIF-Occurrence="2831865458" collectionCode="MACN" collectorName="San Pedro" country="Argentina" pageId="13" pageNumber="956" specimenCount="1" specimenCount-female="1">
.
<emphasis id="B946FC05BC592A4015F6FB7B3502FB15" box="[767,780,1183,1207]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401412FB7B3588FB15" box="[795,902,1183,1207]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401492FB7B35E7FB15" box="[923,1001,1183,1207]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC592A40130BFB7B328EFB15" box="[1026,1152,1183,1207]" pageId="13" pageNumber="956">San Pedro</collectorName>
, without date, A
</materialsCitation>
. G.
<materialsCitation id="3B5A2A4ABC592A401686FB2634BFFB78" ID-GBIF-Occurrence="2831865341" box="[399,689,1218,1242]" collectionCode="MACN" country="Argentina" location="Frears" pageId="13" pageNumber="956" specimenCount="1" specimenCount-male="1">
<location id="8EED76CCBC592A401686FB2637DAFB78" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A401686FB2637DAFB78" box="[399,468,1218,1242]" country="Argentina" name="Frears" pageId="13" pageNumber="956">Frears</location>
<emphasis id="B946FC05BC592A4016E9FB2637E3FB78" box="[480,493,1218,1242]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4016F3FB263445FB78" box="[506,587,1218,1242]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401554FB2634A5FB78" box="[605,683,1218,1242]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC592A4015C9FB26350EFB5E" ID-GBIF-Occurrence="2831865365" collectingDate="1895-12" collectionCode="MACN" collectorName="Carlos Bruch" country="Argentina" location="CHACO" pageId="13" pageNumber="956" specimenCount="3" specimenCount-female="1" specimenCount-male="2" stateProvince="Chaco">
<collectingRegion id="49F6EEF5BC592A4015C9FB263519FB78" box="[704,791,1218,1242]" country="Argentina" name="Chaco" pageId="13" pageNumber="956">CHACO</collectingRegion>
: no more data
<emphasis id="B946FC05BC592A4014DFFB2635EDFB78" box="[982,995,1218,1242]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4014F9FB263245FB78" box="[1008,1099,1218,1242]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A4017A9FB00370BFB5E" box="[160,261,1252,1276]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A40161FFB00376AFB5E" box="[278,356,1252,1276]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<date id="FF8C06D7BC592A40167EFB00343CFB5E" box="[375,562,1252,1276]" pageId="13" pageNumber="956" value="1895-12">
<collectingDate id="EFC8FF3FBC592A40167EFB00343CFB5E" box="[375,562,1252,1276]" pageId="13" pageNumber="956" value="1895-12">December 1895</collectingDate>
</date>
,
<collectorName id="26C745C1BC592A401536FB0034DCFB5E" box="[575,722,1252,1276]" pageId="13" pageNumber="956">Carlos Bruch</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401409FB0036D6FABD" ID-GBIF-Occurrence="2831865442" collectionCode="MACN" collectorName="Carlos Bruch" country="Argentina" pageId="13" pageNumber="956" specimenCount="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A401404FB003514FB5E" box="[781,794,1252,1276]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40142CFB00357BFB5E" box="[805,885,1252,1276]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A40148CFB0035C1FB5E" box="[901,975,1252,1276]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC592A4014D0FB0035E8FB5E" box="[985,998,1252,1276]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<collectorName id="26C745C1BC592A4014F8FB00328DFB5E" box="[1009,1155,1252,1276]" pageId="13" pageNumber="956">Carlos Bruch</collectorName>
coll.)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC592A4017E1FAE335B3FABD" ID-GBIF-Occurrence="2831865366" box="[232,957,1287,1311]" collectionCode="BMNH" country="Argentina" location="CORDOBA" pageId="13" pageNumber="956" specimenCount="1" specimenCount-male="1" stateProvince="CORDOBA">
<collectingRegion id="49F6EEF5BC592A4017E1FAE3376EFABD" box="[232,352,1287,1311]" country="Argentina" name="CORDOBA" pageId="13" pageNumber="956">CÓRDOBA</collectingRegion>
: no more data
<emphasis id="B946FC05BC592A401516FAE33422FABD" box="[543,556,1287,1311]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401531FAE33484FABD" box="[568,650,1287,1311]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401595FAE334E4FABD" box="[668,746,1287,1311]" pageId="13" pageNumber="956">BMNH</collectionCode>
); without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4014B4FAE33243FABD" ID-GBIF-Occurrence="2831865430" box="[957,1101,1287,1311]" collectionCode="BMNH" collectorName="Richter" country="Argentina" isTailEnumeration="true" location="CORDOBA" pageId="13" pageNumber="956" specimenCount="1" stateProvince="CORDOBA">
.
<collectorName id="26C745C1BC592A4014C5FAE33212FABD" box="[972,1052,1287,1311]" pageId="13" pageNumber="956">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401344FAE33538FAE0" ID-GBIF-Occurrence="2831865309" collectingDate="1946-01-30" collectionCode="MLPA" collectorName="Leones" country="Argentina" pageId="13" pageNumber="956" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A401355FAE33267FABD" box="[1116,1129,1287,1311]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40137CFAE336D6FAE0" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401611FACE3773FAE0" box="[280,381,1322,1346]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401684FACE37DBFAE0" box="[397,469,1322,1346]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<collectorName id="26C745C1BC592A4016E0FACE3433FAE0" box="[489,573,1322,1346]" pageId="13" pageNumber="956">Leones</collectorName>
,
<date id="FF8C06D7BC592A401543FACE3502FAE0" box="[586,780,1322,1346]" pageId="13" pageNumber="956" value="1946-01-30">
<collectingDate id="EFC8FF3FBC592A401543FACE3502FAE0" box="[586,780,1322,1346]" pageId="13" pageNumber="956" value="1946-01-30">30 January 1946</collectingDate>
</date>
, W
</materialsCitation>
.N.P. col.
<emphasis id="B946FC05BC592A401497FACE35A5FAE0" box="[926,939,1322,1346]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<materialsCitation id="3B5A2A4ABC592A4014BFFACE328EFAE0" ID-GBIF-Occurrence="2831865447" box="[950,1152,1322,1346]" collectionCode="MACN" country="Argentina" pageId="13" pageNumber="956" specimenCount="1" specimenCount-female="1">
<specimenCount id="9D34EB9EBC592A4014BFFACE3215FAE0" box="[950,1051,1322,1346]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401325FACE3274FAE0" box="[1068,1146,1322,1346]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8D2017BC592A4317A9FAA8351AF9B3" blockId="13.[160,1156,697,1727]" lastBlockId="14.[160,1157,142,1553]" lastPageId="14" lastPageNumber="957" pageId="13" pageNumber="956">
<materialsCitation id="3B5A2A4ABC592A4017A9FAA836CAFA08" ID-GBIF-Occurrence="2831865360" collectingDate="1920-02" collectionCode="MACN, MLPA" collectorName="Carlos Bruch &amp; San Roque &amp; Bosq" pageId="13" pageNumber="956" specimenCount="11" specimenCount-female="7" specimenCount-male="4">
CORRIENTES: no more data
<emphasis id="B946FC05BC592A4016E0FAA837F8FAC6" box="[489,502,1356,1380]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401509FAA8345EFAC6" box="[512,592,1356,1380]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401599FAA8350EFAC6" box="[656,768,1356,1380]" pageId="13" pageNumber="956" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401419FAA83550FAC6" box="[784,862,1356,1380]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
),
<specimenCount id="9D34EB9EBC592A40147BFAA835CCFAC6" box="[882,962,1356,1380]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4014D8FAA83215FAC6" box="[977,1051,1356,1380]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC592A40132FFAA8323DFAC6" box="[1062,1075,1356,1380]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<collectorName id="26C745C1BC592A401334FAA836ECFA25" pageId="13" pageNumber="956">Carlos Bruch</collectorName>
coll.), and
<specimenCount id="9D34EB9EBC592A401678FA8B37C0FA25" box="[369,462,1391,1415]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A40151CFA8B3486FA25" box="[533,648,1391,1415]" pageId="13" pageNumber="956" type="female">5 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401595FA8B34EAFA25" box="[668,740,1391,1415]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<collectorName id="26C745C1BC592A4015F3FA8B3573FA25" box="[762,893,1391,1415]" pageId="13" pageNumber="956">San Roque</collectorName>
,
<date id="FF8C06D7BC592A401487FA8B3235FA25" box="[910,1083,1391,1415]" pageId="13" pageNumber="956" value="1920-02">
<collectingDate id="EFC8FF3FBC592A401487FA8B3235FA25" box="[910,1083,1391,1415]" pageId="13" pageNumber="956" value="1920-02">February 1920</collectingDate>
</date>
,
<collectorName id="26C745C1BC592A401345FA8B328AFA25" box="[1100,1156,1391,1415]" pageId="13" pageNumber="956">Bosq</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4017CDFA76357EFA4D" ID-GBIF-Occurrence="2831865331" collectingDate="1925-10" collectingDateMax="1926-09" collectingDateMin="1925-10" collectionCode="MLPA, MACN" collectorName="Santo Tome" location="Santo Tome" pageId="13" pageNumber="956" specimenCount="17" specimenCount-female="7" specimenCount-male="10">
.
<emphasis id="B946FC05BC592A4017DBFA7636D1FA08" box="[210,223,1426,1450]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4017E2FA763732FA08" box="[235,316,1426,1450]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401644FA76379BFA08" box="[333,405,1426,1450]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<location id="8EED76CCBC592A4016A0FA763436FA08" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC592A4016A0FA763436FA08" box="[425,568,1426,1450]" name="Santo Tome" pageId="13" pageNumber="956">Santo Tomé</location>
, no more data
<emphasis id="B946FC05BC592A4015FDFA76350FFA08" box="[756,769,1426,1450]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A401404FA763566FA08" box="[781,872,1426,1450]" pageId="13" pageNumber="956" type="male">4 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A4014A3FA763215FA08" box="[938,1051,1426,1450]" pageId="13" pageNumber="956" type="female">6 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401325FA763274FA08" box="[1068,1146,1426,1450]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC592A4017A9FA503720FA6E" box="[160,302,1460,1484]" pageId="13" pageNumber="956">Santo Tomé</collectorName>
,
<date id="FF8C06D7BC592A401635FA5037D2FA6E" box="[316,476,1460,1484]" pageId="13" pageNumber="956" value="1925-10">
<collectingDate id="EFC8FF3FBC592A401635FA5037D2FA6E" box="[316,476,1460,1484]" pageId="13" pageNumber="956" value="1925-10">October 1925</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC592A4015B3FA5034C9FA6E" box="[698,711,1460,1484]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4015DBFA503522FA6E" box="[722,812,1460,1484]" pageId="13" pageNumber="956" type="male">4 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401462FA5035DEFA6E" box="[875,976,1460,1484]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4014E8FA503221FA6E" box="[993,1071,1460,1484]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC592A40134BFA5036EDFA4D" pageId="13" pageNumber="956">Santo Tomé</collectorName>
,
<date id="FF8C06D7BC592A4017FAFA3337B9FA4D" box="[243,439,1495,1519]" pageId="13" pageNumber="956" value="1926-09">
<collectingDate id="EFC8FF3FBC592A4017FAFA3337B9FA4D" box="[243,439,1495,1519]" pageId="13" pageNumber="956" value="1926-09">September 1926</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC592A401594FA3334A4FA4D" box="[669,682,1495,1519]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4015BEFA333504FA4D" box="[695,778,1495,1519]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401415FA333564FA4D" box="[796,874,1495,1519]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
)
</materialsCitation>
. ENTRE RÍOS: no more data
<emphasis id="B946FC05BC592A4017D3FA1D36E9F9B3" box="[218,231,1529,1553]" italics="true" pageId="13" pageNumber="956">–</emphasis>
H
<materialsCitation id="3B5A2A4ABC592A40160AFA1E3785F9B0" ID-GBIF-Occurrence="2831865346" box="[259,395,1530,1554]" collectingDate="1925-10" collectingDateMax="1926-09" collectingDateMin="1925-10" collectionCode="MLPA, MACN" collectorName="Richter" isTailEnumeration="true" location="Santo Tome" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A401606FA1E3751F9B0" box="[271,351,1530,1554]" pageId="13" pageNumber="956">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401682FA1E3734F996" ID-GBIF-Occurrence="2831865368" collectionCode="MLPA, MACN" collectorName="Villa Paranacito" pageId="13" pageNumber="956" specimenCount="3" specimenCount-female="1" specimenCount-male="2">
.
<emphasis id="B946FC05BC592A40169FFA1D37ADF9B3" box="[406,419,1529,1553]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4016A5FA1E37F7F9B0" box="[428,505,1530,1554]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A40153DFA1E3499F9B0" box="[564,663,1530,1554]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4015ACFA1E34E3F9B0" box="[677,749,1530,1554]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<collectorName id="26C745C1BC592A4015F7FA1E35BFF9B0" box="[766,945,1530,1554]" pageId="13" pageNumber="956">Villa Paranacito</collectorName>
, no more data
<emphasis id="B946FC05BC592A401369FA1D3263F9B3" box="[1120,1133,1529,1553]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40137CFA1E36D6F996" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017EFF9F8373AF996" box="[230,308,1564,1588]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC592A40164FF9F83745F9F5" ID-GBIF-Occurrence="2831865329" collectionCode="MACN, MLPA" pageId="13" pageNumber="956" specimenCount="5" specimenCount-female="2" specimenCount-male="3">
JUJUY: no more data
<emphasis id="B946FC05BC592A401549F9F83443F996" box="[576,589,1564,1588]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40155FF9F834AAF996" box="[598,676,1564,1588]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A4015E9F9F8354AF996" box="[736,836,1564,1588]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A40145BF9F83593F996" box="[850,925,1564,1588]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
) and
<specimenCount id="9D34EB9EBC592A4014E9F9F83237F996" box="[992,1081,1564,1588]" pageId="13" pageNumber="956" type="male">2 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A40137CF9F836E3F9F5" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017F4F9DB374BF9F5" box="[253,325,1599,1623]" pageId="13" pageNumber="956">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC592A40165EF9DB320FF9F5" ID-GBIF-Occurrence="2831865461" box="[343,1025,1599,1623]" collectionCode="BMNH" pageId="13" pageNumber="956" specimenCount="3" specimenCount-female="2" specimenCount-male="1">
MENDOZA: no more data
<emphasis id="B946FC05BC592A401580F9DB3498F9F5" box="[649,662,1599,1623]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4015A9F9DB34E1F9F5" box="[672,751,1599,1623]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401427F9DB3593F9F5" box="[814,925,1599,1623]" pageId="13" pageNumber="956" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4014A4F9DB35F5F9F5" box="[941,1019,1599,1623]" pageId="13" pageNumber="956">BMNH</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC592A401307F9DB3404F93F" ID-GBIF-Occurrence="2831865456" collectionCode="MACN, MLPA" pageId="13" pageNumber="956" specimenCount="12" specimenCount-female="4" specimenCount-male="8">
MISIONES: no more data
<emphasis id="B946FC05BC592A401645F9853757F9DB" box="[332,345,1633,1657]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40166DF98637B1F9D8" box="[356,447,1634,1658]" pageId="13" pageNumber="956" type="male">8 males</specimenCount>
and
<specimenCount id="9D34EB9EBC592A401509F986347EF9D8" box="[512,624,1634,1658]" pageId="13" pageNumber="956" type="female">3 females</specimenCount>
(
<collectionCode id="ED23B8D2BC592A401588F98634C1F9D8" box="[641,719,1634,1658]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="13" pageNumber="956" type="Museum">MACN</collectionCode>
); without date, illegible collector
<emphasis id="B946FC05BC592A401354F9853264F9DB" box="[1117,1130,1633,1657]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A40137CF98636E3F93E" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4017F5F961374AF93F" box="[252,324,1669,1693]" pageId="13" pageNumber="956">MLPA</collectionCode>
); without date, C
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A401503F961348AF93F" ID-GBIF-Occurrence="2831865322" box="[522,644,1669,1693]" collectionCode="MACN, MLPA" collectorName="Bruch" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A40151FF9613456F93F" box="[534,600,1669,1693]" pageId="13" pageNumber="956">Bruch</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A40158DF9613226F93F" ID-GBIF-Occurrence="2831865377" box="[644,1064,1668,1693]" collectionCode="MLPA" pageId="13" pageNumber="956" specimenCount="1" specimenCount-female="1">
.
<emphasis id="B946FC05BC592A401599F9603493F93E" box="[656,669,1668,1692]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4015ACF9613507F93F" box="[677,777,1669,1693]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A40141EF9613551F93F" box="[791,863,1669,1693]" pageId="13" pageNumber="956">MLPA</collectionCode>
); without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A40132EF96136CAF91D" ID-GBIF-Occurrence="2831865364" collectionCode="MLPA" collectorName="Richter" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">
.
<collectorName id="26C745C1BC592A40133AF961328DF93F" box="[1075,1155,1669,1693]" pageId="13" pageNumber="956">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4017CDF94335F9F91D" ID-GBIF-Occurrence="2831865376" box="[196,1015,1703,1727]" collectingDate="1933-12-01" collectingDateMax="1933-12-18" collectingDateMin="1933-12-01" collectionCode="MLPA" collectorName="Alto Parana" pageId="13" pageNumber="956" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC592A4017D8F94336D0F91D" box="[209,222,1703,1727]" italics="true" pageId="13" pageNumber="956">–</emphasis>
<specimenCount id="9D34EB9EBC592A4017E0F9433736F91D" box="[233,312,1703,1727]" pageId="13" pageNumber="956" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC592A40167EF94337D2F91D" box="[375,476,1703,1727]" pageId="13" pageNumber="956" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC592A4016E5F943343AF91D" box="[492,564,1703,1727]" pageId="13" pageNumber="956">MLPA</collectionCode>
);
<collectorName id="26C745C1BC592A40154EF94334DFF91D" box="[583,721,1703,1727]" pageId="13" pageNumber="956">Alto Paraná</collectorName>
,
<date id="FF8C06D7BC592A4015D7F94335D7F91D" box="[734,985,1703,1727]" pageId="13" pageNumber="956" value="1933-12-01" valueMax="1933-12-18" valueMin="1933-12-01">
<collectingDate id="EFC8FF3FBC592A4015D7F94335D7F91D" box="[734,985,1703,1727]" pageId="13" pageNumber="956" value="1933-12-01" valueMax="1933-12-18" valueMin="1933-12-01">
1
<emphasis id="B946FC05BC592A4015E5F94334F7F91D" box="[748,761,1703,1727]" italics="true" pageId="13" pageNumber="956">–</emphasis>
18 December 1933
</collectingDate>
</date>
, K
</materialsCitation>
<materialsCitation id="3B5A2A4ABC592A4014F1F943321FF91D" ID-GBIF-Occurrence="2831865302" box="[1016,1041,1703,1727]" collectingDate="1933-12-01" collectingDateMax="1933-12-18" collectingDateMin="1933-12-01" collectionCode="MLPA" collectorName="Alto Parana" isTailEnumeration="true" pageId="13" pageNumber="956" specimenCount="1">. J</materialsCitation>
. Hayward col
<materialsCitation id="3B5A2A4ABC5A2A4317CDFF6A3565FF04" ID-GBIF-Occurrence="2831865372" box="[196,875,142,166]" collectingDate="1928" collectingDateMax="1929" collectingDateMin="1928" collectionCode="BMNH" collectorName="San Ignacio &amp; Quiroga" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1">
.
<emphasis id="B946FC05BC5A2A4317D8FF6A36D0FF04" box="[209,222,142,166]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4317E1FF6A3743FF04" box="[232,333,142,166]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431654FF6A37A5FF04" box="[349,427,142,166]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<collectorName id="26C745C1BC5A2A4316B7FF6A3449FF04" box="[446,583,142,166]" pageId="14" pageNumber="957">San Ignacio</collectorName>
,
<date id="FF8C06D7BC5A2A43155DFF6A34DAFF04" box="[596,724,142,166]" pageId="14" pageNumber="957" value="1928" valueMax="1929">
<collectingDate id="EFC8FF3FBC5A2A43155DFF6A34DAFF04" box="[596,724,142,166]" pageId="14" pageNumber="957" value="1928" valueMax="1929">
1928
<emphasis id="B946FC05BC5A2A431585FF6A3497FF04" box="[652,665,142,166]" italics="true" pageId="14" pageNumber="957">–</emphasis>
1929
</collectingDate>
</date>
,
<collectorName id="26C745C1BC5A2A4315E8FF6A3533FF04" box="[737,829,142,166]" pageId="14" pageNumber="957">Quiroga</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431462FF6A3490FF4E" ID-GBIF-Occurrence="2831865478" collectingDate="1927-10" collectingDateMax="1929-10-21" collectingDateMin="1927-10" collectionCode="MACN, MLPA" collectorName="Puerto Iguazu" location="San Ignacio" pageId="14" pageNumber="957" specimenCount="4" specimenCount-female="1" specimenCount-male="3">
.
<emphasis id="B946FC05BC5A2A431471FF6A358BFF04" box="[888,901,142,166]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431486FF6A35E7FF04" box="[911,1001,142,166]" pageId="14" pageNumber="957" type="male">2 males</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4314F0FF6A3249FF04" box="[1017,1095,142,166]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
);
<location id="8EED76CCBC5A2A431353FF6A36F8FF6B" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A431353FF6A36F8FF6B" name="San Ignacio" pageId="14" pageNumber="957">San Ignacio</location>
,
<date id="FF8C06D7BC5A2A43160AFF5537C7FF6B" box="[259,457,177,201]" pageId="14" pageNumber="957" value="1929-10-21">
<collectingDate id="EFC8FF3FBC5A2A43160AFF5537C7FF6B" box="[259,457,177,201]" pageId="14" pageNumber="957" value="1929-10-21">21 October 1929</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC5A2A4315ACFF5534BCFF6B" box="[677,690,177,201]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4315B5FF553502FF6B" box="[700,780,177,201]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431412FF55356DFF6B" box="[795,867,177,201]" pageId="14" pageNumber="957">MLPA</collectionCode>
);
<collectorName id="26C745C1BC5A2A43147CFF553214FF6B" box="[885,1050,177,201]" pageId="14" pageNumber="957">Puerto Iguazu</collectorName>
,
<date id="FF8C06D7BC5A2A43132EFF5536D4FF4E" pageId="14" pageNumber="957" value="1927-10">
<collectingDate id="EFC8FF3FBC5A2A43132EFF5536D4FF4E" pageId="14" pageNumber="957" value="1927-10">October 1927</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC5A2A4316B3FF3737C9FF49" box="[442,455,211,235]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4316DAFF303437FF4E" box="[467,569,212,236]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431543FF303496FF4E" box="[586,664,212,236]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
)
</materialsCitation>
. SALTA: without date, H
<materialsCitation id="3B5A2A4ABC5A2A4314C8FF303241FF4E" ID-GBIF-Occurrence="2831865421" box="[961,1103,212,236]" collectingDate="1927-10" collectingDateMax="1929-10-21" collectingDateMin="1927-10" collectionCode="MACN, MLPA" collectorName="Richter" isTailEnumeration="true" location="San Ignacio" pageId="14" pageNumber="957" specimenCount="1">
.
<collectorName id="26C745C1BC5A2A4314D9FF30322EFF4E" box="[976,1056,212,236]" pageId="14" pageNumber="957">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431346FF30373BFEAC" ID-GBIF-Occurrence="2831865390" collectionCode="MLPA" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC5A2A431354FF373264FF49" box="[1117,1130,211,235]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43137CFF3036D6FEAC" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317EEFF123721FEAC" box="[231,303,246,270]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431648FF1234DEFEAC" ID-GBIF-Occurrence="2831865323" box="[321,720,246,270]" location="Chaco" pageId="14" pageNumber="957" specimenCount="1">
SANTA FÉ:
<location id="8EED76CCBC5A2A4316C8FF123405FEAC" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4316C8FF123405FEAC" box="[449,523,246,270]" name="Chaco" pageId="14" pageNumber="957">Chaco</location>
, without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4315D9FF123554FEAC" ID-GBIF-Occurrence="2831865423" box="[720,858,246,270]" collectorName="Richter" isTailEnumeration="true" location="Chaco" pageId="14" pageNumber="957" specimenCount="1">
.
<collectorName id="26C745C1BC5A2A4315D5FF123522FEAC" box="[732,812,246,270]" pageId="14" pageNumber="957">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431453FF1235A3FE93" ID-GBIF-Occurrence="2831865394" collectingDate="1911-12-08" collectingDateMax="1911-12-20" collectingDateMin="1911-12-08" collectionCode="MLPA" location="Rio San Javier" municipality="Estancia La Noria" pageId="14" pageNumber="957" specimenCount="11" specimenCount-female="4" specimenCount-male="7">
.
<emphasis id="B946FC05BC5A2A43146FFF12357DFEAC" box="[870,883,246,270]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431474FF1235D8FEAC" box="[893,982,246,270]" pageId="14" pageNumber="957" type="male">7 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A43131DFF12328DFEAC" box="[1044,1155,246,270]" pageId="14" pageNumber="957" type="female">4 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFEFD36E0FE93" box="[166,238,281,305]" pageId="14" pageNumber="957">MLPA</collectionCode>
);
<collectingMunicipality id="6BE9BA6DBC5A2A431608FEFD37C3FE93" box="[257,461,281,305]" pageId="14" pageNumber="957">Estancia La Noria</collectingMunicipality>
,
<location id="8EED76CCBC5A2A4316D2FEFD348FFE93" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4316D2FEFD348FFE93" box="[475,641,281,305]" municipality="Estancia La Noria" name="Rio San Javier" pageId="14" pageNumber="957">Rio San Javier</location>
,
<date id="FF8C06D7BC5A2A431586FEFD3585FE93" box="[655,907,281,305]" pageId="14" pageNumber="957" value="1911-12-08" valueMax="1911-12-20" valueMin="1911-12-08">
<collectingDate id="EFC8FF3FBC5A2A431586FEFD3585FE93" box="[655,907,281,305]" pageId="14" pageNumber="957" value="1911-12-08" valueMax="1911-12-20" valueMin="1911-12-08">
8
<emphasis id="B946FC05BC5A2A431594FEFD34A4FE93" box="[669,682,281,305]" italics="true" pageId="14" pageNumber="957">–</emphasis>
20 December 1911
</collectingDate>
</date>
, G
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4314A5FEFD35C4FE93" ID-GBIF-Occurrence="2831865354" box="[940,970,281,305]" collectingDate="1911-12-08" collectingDateMax="1911-12-20" collectingDateMin="1911-12-08" collectionCode="MLPA" isTailEnumeration="true" location="Rio San Javier" municipality="Estancia La Noria" pageId="14" pageNumber="957" specimenCount="1">. E</materialsCitation>
. Bryant col
<materialsCitation id="3B5A2A4ABC5A2A431359FEFD3269FEF6" ID-GBIF-Occurrence="2831865326" collectingDate="1924-11" collectionCode="BMNH" collectorName="General Obligado &amp; Villa Ana" pageId="14" pageNumber="957" specimenCount="17" specimenCount-female="9" specimenCount-male="8">
.
<emphasis id="B946FC05BC5A2A431357FEFD3265FE93" box="[1118,1131,281,305]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43137CFEFD36ECFEF6" pageId="14" pageNumber="957" type="male">8 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A431616FED83780FEF6" box="[287,398,316,340]" pageId="14" pageNumber="957" type="female">9 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431694FED837E5FEF6" box="[413,491,316,340]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<collectorName id="26C745C1BC5A2A4316F4FED834C4FEF6" box="[509,714,316,340]" pageId="14" pageNumber="957">General Obligado</collectorName>
,
<collectorName id="26C745C1BC5A2A4315DFFED83533FEF6" box="[726,829,316,340]" pageId="14" pageNumber="957">Villa Ana</collectorName>
, FCSF,
<date id="FF8C06D7BC5A2A431486FED83244FEF6" box="[911,1098,316,340]" pageId="14" pageNumber="957" value="1924-11">
<collectingDate id="EFC8FF3FBC5A2A431486FED83244FEF6" box="[911,1098,316,340]" pageId="14" pageNumber="957" value="1924-11">November 1924</collectingDate>
</date>
, K
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43136EFED8328EFEF6" ID-GBIF-Occurrence="2831865396" box="[1127,1152,316,340]" collectingDate="1924-11" collectionCode="BMNH" collectorName="General Obligado &amp; Villa Ana" isTailEnumeration="true" pageId="14" pageNumber="957" specimenCount="1">. J</materialsCitation>
. Hayward col
<materialsCitation id="3B5A2A4ABC5A2A43163EFEBA3555FED4" ID-GBIF-Occurrence="2831865438" box="[311,859,350,374]" collectionCode="BMNH" collectorName="Rosario" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC5A2A431641FEBA375BFED4" box="[328,341,350,374]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43166DFEBA37B6FED4" box="[356,440,350,374]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4316C5FEBA3414FED4" box="[460,538,350,374]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<collectorName id="26C745C1BC5A2A43153BFEBA3487FED4" box="[562,649,350,374]" pageId="14" pageNumber="957">Rosario</collectorName>
, without date, A
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431452FEBA320CFED4" ID-GBIF-Occurrence="2831865345" box="[859,1026,350,374]" collectionCode="BMNH" collectorName="Stevenin" isTailEnumeration="true" pageId="14" pageNumber="957" specimenCount="1">
.
<collectorName id="26C745C1BC5A2A431465FEBA35C1FED4" box="[876,975,350,374]" pageId="14" pageNumber="957">Stévenin</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43130BFEBA34D0FE3B" ID-GBIF-Occurrence="2831865312" collectionCode="MACN, MLPA" collectorName="Santa Fe" pageId="14" pageNumber="957" specimenCount="2" specimenCount-male="2">
.
<emphasis id="B946FC05BC5A2A43131DFEBA322FFED4" box="[1044,1057,350,374]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431339FEBA328AFED4" box="[1072,1156,350,374]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFE6536FAFE3B" box="[166,244,385,409]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC5A2A43160FFE653765FE3B" box="[262,363,385,409]" pageId="14" pageNumber="957">Santa Fé</collectorName>
, no more data
<emphasis id="B946FC05BC5A2A431514FE653424FE3B" box="[541,554,385,409]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43153AFE65348CFE3B" box="[563,642,385,409]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431599FE6534D6FE3B" box="[656,728,385,409]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4315E3FE6537C6FE1E" ID-GBIF-Occurrence="2831865386" collectorName="Wagner" location="Rio Salado" pageId="14" pageNumber="957" specimenCount="1">
SANTIAGO DEL ESTERO:
<location id="8EED76CCBC5A2A43130CFE65328EFE3B" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A43130CFE65328EFE3B" box="[1029,1152,385,409]" name="Rio Salado" pageId="14" pageNumber="957">Río Salado</location>
, without date,
<collectorName id="26C745C1BC5A2A43164AFE403795FE1E" box="[323,411,420,444]" pageId="14" pageNumber="957">Wagner</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4316C1FE4034BAFE1E" ID-GBIF-Occurrence="2831865306" box="[456,692,419,444]" collectionCode="MLPA" pageId="14" pageNumber="957" specimenCount="2" specimenCount-female="2">
.
<emphasis id="B946FC05BC5A2A4316DDFE4737EFFE19" box="[468,481,419,443]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4316E3FE403456FE1E" box="[490,600,420,444]" pageId="14" pageNumber="957" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43156FFE4034A0FE1E" box="[614,686,420,444]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4315C9FE40344BFE7C" ID-GBIF-Occurrence="2831865391" collectionCode="BMNH" pageId="14" pageNumber="957" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
TUCUMÁN: no more data
<emphasis id="B946FC05BC5A2A4314E5FE4735F7FE19" box="[1004,1017,419,443]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43130BFE40325EFE1E" box="[1026,1104,420,444]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A4317A9FE223707FE7C" box="[160,265,454,478]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431614FE223765FE7C" box="[285,363,454,478]" pageId="14" pageNumber="957">BMNH</collectionCode>
); without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43154DFE2234D6FE7C" ID-GBIF-Occurrence="2831865415" box="[580,728,454,478]" collectionCode="BMNH" collectorName="Richter" isTailEnumeration="true" pageId="14" pageNumber="957" specimenCount="1">
.
<collectorName id="26C745C1BC5A2A43155FFE2234A8FE7C" box="[598,678,454,478]" pageId="14" pageNumber="957">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4315D1FE22328EFE7C" ID-GBIF-Occurrence="2831865342" box="[728,1152,454,478]" collectionCode="MLPA" pageId="14" pageNumber="957" specimenCount="6" specimenCount-female="3" specimenCount-male="3">
.
<emphasis id="B946FC05BC5A2A4315E0FE2234F8FE7C" box="[745,758,454,478]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43140CFE22356CFE7C" box="[773,866,454,478]" pageId="14" pageNumber="957" type="male">3 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A4314A3FE223210FE7C" box="[938,1054,454,478]" pageId="14" pageNumber="957" type="female">3 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43133BFE223274FE7C" box="[1074,1146,454,478]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4317A9FE0D352DFD86" ID-GBIF-Occurrence="2831865384" collectingDate="1959-11" collectionCode="CMNC" country="Bolivia" latitude="-18.338057" location="Santiago" longLatPrecision="21" longitude="-59.59361" municipality="Chiquitos" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Santa Cruz">
<emphasis id="B946FC05BC5A2A4317A9FE0D370AFDA3" bold="true" box="[160,260,489,513]" pageId="14" pageNumber="957">
<collectingCountry id="F3256087BC5A2A4317A9FE0D370AFDA3" box="[160,260,489,513]" name="Bolivia" pageId="14" pageNumber="957">BOLÍVIA</collectingCountry>
</emphasis>
:
<collectingRegion id="49F6EEF5BC5A2A43161DFE0D37A3FDA3" box="[276,429,489,513]" country="Bolivia" name="Santa Cruz" pageId="14" pageNumber="957">SANTA CRUZ</collectingRegion>
:
<collectingMunicipality id="6BE9BA6DBC5A2A4316B0FE0D3426FDA3" box="[441,552,489,513]" pageId="14" pageNumber="957">Chiquitos</collectingMunicipality>
,
<location id="8EED76CCBC5A2A43153FFE0D3493FDA3" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A43153FFE0D3493FDA3" box="[566,669,489,513]" country="Bolivia" latitude="-18.338057" longLatPrecision="21" longitude="-59.59361" municipality="Chiquitos" name="Santiago" pageId="14" pageNumber="957" stateProvince="Santa Cruz">Santiago</location>
,
<geoCoordinate id="EE0646D0BC5A2A4315A0FE0D352BFDA3" box="[681,805,489,513]" degrees="18" direction="south" minutes="20" orientation="latitude" pageId="14" pageNumber="957" precision="15" seconds="17" value="-18.338057">
18°20
<emphasis id="B946FC05BC5A2A4315E3FE0D34FEFDA3" bold="true" box="[746,752,489,513]" italics="true" pageId="14" pageNumber="957">ʹ</emphasis>
17
<emphasis id="B946FC05BC5A2A431405FE0D351BFDA3" box="[780,789,489,513]" italics="true" pageId="14" pageNumber="957">”</emphasis>
S
</geoCoordinate>
,
<geoCoordinate id="EE0646D0BC5A2A43143BFE0D35B9FDA3" box="[818,951,489,513]" degrees="59" direction="west" minutes="35" orientation="longitude" pageId="14" pageNumber="957" precision="15" seconds="37" value="-59.59361">
59°35
<emphasis id="B946FC05BC5A2A43147AFE0D3577FDA3" bold="true" box="[883,889,489,513]" italics="true" pageId="14" pageNumber="957">ʹ</emphasis>
37
<emphasis id="B946FC05BC5A2A43149CFE0D3590FDA3" box="[917,926,489,513]" italics="true" pageId="14" pageNumber="957">”</emphasis>
W
</geoCoordinate>
,
<date id="FF8C06D7BC5A2A4314CDFE0D328FFDA3" box="[964,1153,489,513]" pageId="14" pageNumber="957" value="1959-11">
<collectingDate id="EFC8FF3FBC5A2A4314CDFE0D328FFDA3" box="[964,1153,489,513]" pageId="14" pageNumber="957" value="1959-11">November 1959</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC5A2A431664FDEF3774FD81" box="[365,378,523,547]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43168AFDE837DFFD86" box="[387,465,524,548]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4316D6FDE83425FD86" box="[479,555,524,548]" pageId="14" pageNumber="957">CMNC</collectionCode>
; examined by photo)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431426FDE83724FDCB" ID-GBIF-Occurrence="2831865406" collectingDate="2005-01-03" collectorName="Mann &amp; Hamel &amp; Herzog" country="Bolivia" elevation="620" latitude="-21.847778" location="Yaguacua-Caiza" longLatPrecision="21" longitude="-63.60722" municipality="Gran Chaco" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Tarija">
<collectingRegion id="49F6EEF5BC5A2A431426FDE8358EFD86" box="[815,896,524,548]" country="Bolivia" name="Tarija" pageId="14" pageNumber="957">TARIJA</collectingRegion>
:
<collectingMunicipality id="6BE9BA6DBC5A2A431482FDE8321AFD86" box="[907,1044,524,548]" pageId="14" pageNumber="957">Gran Chaco</collectingMunicipality>
, between
<location id="8EED76CCBC5A2A4317A9FDCA3759FDE4" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4317A9FDCA3759FDE4" box="[160,343,558,582]" country="Bolivia" latitude="-21.847778" longLatPrecision="21" longitude="-63.60722" municipality="Gran Chaco" name="Yaguacua-Caiza" pageId="14" pageNumber="957" stateProvince="Tarija">Yaguacua-Caiza</location>
,
<geoCoordinate id="EE0646D0BC5A2A43166FFDCA37ECFDE4" box="[358,482,558,582]" degrees="21" direction="south" minutes="50" orientation="latitude" pageId="14" pageNumber="957" precision="15" seconds="52" value="-21.847778">
21°50
<emphasis id="B946FC05BC5A2A4316AEFDCA37A3FDE4" bold="true" box="[423,429,558,582]" italics="true" pageId="14" pageNumber="957">ʹ</emphasis>
52
<emphasis id="B946FC05BC5A2A4316C0FDCA37DCFDE4" box="[457,466,558,582]" italics="true" pageId="14" pageNumber="957">”</emphasis>
S
</geoCoordinate>
,
<geoCoordinate id="EE0646D0BC5A2A4316FBFDCA3476FDE4" box="[498,632,558,582]" degrees="63" direction="west" minutes="36" orientation="longitude" pageId="14" pageNumber="957" precision="15" seconds="26" value="-63.60722">
63º36
<emphasis id="B946FC05BC5A2A43153AFDCA3437FDE4" bold="true" box="[563,569,558,582]" italics="true" pageId="14" pageNumber="957">ʹ</emphasis>
26
<emphasis id="B946FC05BC5A2A43155CFDCA3450FDE4" box="[597,606,558,582]" italics="true" pageId="14" pageNumber="957">”</emphasis>
W
</geoCoordinate>
,
<quantity id="4CCA8DF2BC5A2A431581FDCA34D8FDE4" box="[648,726,558,582]" metricMagnitude="2" metricUnit="m" metricValue="6.2" pageId="14" pageNumber="957" unit="m" value="620.0">
<elevation id="001FC724BC5A2A431581FDCA34D8FDE4" box="[648,726,558,582]" metricMagnitude="2" metricUnit="m" metricValue="6.2" pageId="14" pageNumber="957" unit="m" value="620.0">620 m</elevation>
</quantity>
,
<date id="FF8C06D7BC5A2A4315EEFDCA3591FDE4" box="[743,927,558,582]" pageId="14" pageNumber="957" value="2005-01-03">
<collectingDate id="EFC8FF3FBC5A2A4315EEFDCA3591FDE4" box="[743,927,558,582]" pageId="14" pageNumber="957" value="2005-01-03">3 January 2005</collectingDate>
</date>
,
<collectorName id="26C745C1BC5A2A4314A6FDCA35FCFDE4" box="[943,1010,558,582]" pageId="14" pageNumber="957">Mann</collectorName>
,
<collectorName id="26C745C1BC5A2A43130BFDCA3245FDE4" box="[1026,1099,558,582]" pageId="14" pageNumber="957">Hamel</collectorName>
and
<collectorName id="26C745C1BC5A2A4317A9FDB536FCFDCB" box="[160,242,593,617]" pageId="14" pageNumber="957">Herzog</collectorName>
cols
</materialsCitation>
.
<emphasis id="B946FC05BC5A2A43163FFDB5374DFDCB" box="[310,323,593,617]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<materialsCitation id="3B5A2A4ABC5A2A431644FDB53776FD0C" ID-GBIF-Occurrence="2831865429" collectingDate="2005-01-03" collectionCode="BMNH, OUMNH" collectorName="Gran Chaco &amp; Yacuiba &amp; Mann &amp; Hamel &amp; Herzog" country="Bolivia" elevation="622" latitude="-21.900833" longLatPrecision="21" longitude="-63.631664" pageId="14" pageNumber="957" specimenCount="82" specimenCount-female="44" specimenCount-male="38" stateProvince="Tarija">
<specimenCount id="9D34EB9EBC5A2A431644FDB537A8FDCB" box="[333,422,593,617]" pageId="14" pageNumber="957" type="male">3 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A4316EDFDB53446FDCB" box="[484,584,593,617]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43155EFDB534ADFDCB" box="[599,675,593,617]" pageId="14" pageNumber="957">BMNH</collectionCode>
) and
<specimenCount id="9D34EB9EBC5A2A4315EFFDB53543FDCB" box="[742,845,593,617]" pageId="14" pageNumber="957" type="male">35 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A431482FDB53206FDCB" box="[907,1032,593,617]" pageId="14" pageNumber="957" type="female">43 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43131EFDB53275FDCB" box="[1047,1147,593,617]" pageId="14" pageNumber="957">OUMNH</collectionCode>
);
<collectorName id="26C745C1BC5A2A4317A9FD973722FD29" box="[160,300,627,651]" pageId="14" pageNumber="957">Gran Chaco</collectorName>
,
<collectorName id="26C745C1BC5A2A431633FD973799FD29" box="[314,407,627,651]" pageId="14" pageNumber="957">Yacuiba</collectorName>
,
<quantity id="4CCA8DF2BC5A2A4316AFFD9737FDFD29" box="[422,499,627,651]" metricMagnitude="2" metricUnit="m" metricValue="6.22" pageId="14" pageNumber="957" unit="m" value="622.0">
<elevation id="001FC724BC5A2A4316AFFD9737FDFD29" box="[422,499,627,651]" metricMagnitude="2" metricUnit="m" metricValue="6.22" pageId="14" pageNumber="957" unit="m" value="622.0">622 m</elevation>
</quantity>
,
<geoCoordinate id="EE0646D0BC5A2A43150AFD973471FD29" box="[515,639,627,651]" degrees="21" direction="south" minutes="54" orientation="latitude" pageId="14" pageNumber="957" precision="15" seconds="03" value="-21.900833">
21°54
<emphasis id="B946FC05BC5A2A43154AFD973447FD29" bold="true" box="[579,585,627,651]" italics="true" pageId="14" pageNumber="957">ʹ</emphasis>
03
<emphasis id="B946FC05BC5A2A43156CFD973460FD29" box="[613,622,627,651]" italics="true" pageId="14" pageNumber="957">”</emphasis>
S
</geoCoordinate>
,
<geoCoordinate id="EE0646D0BC5A2A431587FD97351DFD29" box="[654,787,627,651]" degrees="63" direction="west" minutes="37" orientation="longitude" pageId="14" pageNumber="957" precision="15" seconds="54" value="-63.631664">
63°37
<emphasis id="B946FC05BC5A2A4315C7FD9734DAFD29" bold="true" box="[718,724,627,651]" italics="true" pageId="14" pageNumber="957">ʹ</emphasis>
54
<emphasis id="B946FC05BC5A2A4315F9FD9734F7FD29" box="[752,761,627,651]" italics="true" pageId="14" pageNumber="957">”</emphasis>
W
</geoCoordinate>
,
<date id="FF8C06D7BC5A2A43142BFD9735D4FD29" box="[802,986,627,651]" pageId="14" pageNumber="957" value="2005-01-03">
<collectingDate id="EFC8FF3FBC5A2A43142BFD9735D4FD29" box="[802,986,627,651]" pageId="14" pageNumber="957" value="2005-01-03">3 January 2005</collectingDate>
</date>
,
<collectorName id="26C745C1BC5A2A4314E1FD973225FD29" box="[1000,1067,627,651]" pageId="14" pageNumber="957">Mann</collectorName>
,
<collectorName id="26C745C1BC5A2A431333FD97328DFD29" box="[1082,1155,627,651]" pageId="14" pageNumber="957">Hamel</collectorName>
and
<collectorName id="26C745C1BC5A2A4317E8FD72373DFD0C" box="[225,307,662,686]" pageId="14" pageNumber="957">Herzog</collectorName>
cols
</materialsCitation>
.
<emphasis id="B946FC05BC5A2A431698FD723790FD0C" box="[401,414,662,686]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<materialsCitation id="3B5A2A4ABC5A2A4316BDFD72349BFD0C" ID-GBIF-Occurrence="2831865401" box="[436,661,662,686]" collectionCode="BMNH" country="Bolivia" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1" stateProvince="Tarija">
<specimenCount id="9D34EB9EBC5A2A4316BDFD72342BFD0C" box="[436,549,662,686]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431548FD723481FD0C" box="[577,655,662,686]" pageId="14" pageNumber="957">BMNH</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4315A7FD723435FD73" ID-GBIF-Occurrence="2831865339" collectionCode="OUMNH" collectorName="Hope-Westwood" country="Brazil" location="BRAZIL" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1">
<emphasis id="B946FC05BC5A2A4315A7FD723508FD0C" bold="true" box="[686,774,662,686]" pageId="14" pageNumber="957">
<collectingCountry id="F3256087BC5A2A4315A7FD723508FD0C" box="[686,774,662,686]" name="Brazil" pageId="14" pageNumber="957">BRAZIL</collectingCountry>
</emphasis>
: no more data
<emphasis id="B946FC05BC5A2A4314E6FD7235F2FD0C" box="[1007,1020,662,686]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43131BFD72328DFD0C" box="[1042,1155,662,686]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFD5D3708FD73" box="[166,262,697,721]" pageId="14" pageNumber="957">OUMNH</collectionCode>
<emphasis id="B946FC05BC5A2A43161FFD5D372DFD73" box="[278,291,697,721]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A43163DFD5D37FDFD73" box="[308,499,697,721]" pageId="14" pageNumber="957">Hope-Westwood</collectorName>
coll.)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431559FD5D36CAFD51" ID-GBIF-Occurrence="2831865381" collectorName="Koslomosky" country="Brazil" location="MATO GROSSO" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Mato Grosso">
<collectingRegion id="49F6EEF5BC5A2A431559FD5D3504FD73" box="[592,778,697,721]" country="Brazil" name="Mato Grosso" pageId="14" pageNumber="957">MATO GROSSO</collectingRegion>
: without date,
<collectorName id="26C745C1BC5A2A4314D8FD5D3256FD73" box="[977,1112,697,721]" pageId="14" pageNumber="957">Koslomosky</collectorName>
(?) col
</materialsCitation>
.
<emphasis id="B946FC05BC5A2A4317DFFD3F36EDFD51" box="[214,227,731,755]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<materialsCitation id="3B5A2A4ABC5A2A4317FBFD3F3725FCB4" ID-GBIF-Occurrence="2831865383" collectingDate="2015-02-08" collectionCode="MLPA" collectorName="Pocone &amp; Rodovia Transpantaneira &amp; Mario Cupello" country="Brazil" pageId="14" pageNumber="957" specimenCount="3" specimenCount-male="3" stateProvince="Mato Grosso">
<specimenCount id="9D34EB9EBC5A2A4317FBFD3F375EFD51" box="[242,336,731,755]" pageId="14" pageNumber="957" type="male">3 males</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43166FFD3F37A0FD51" box="[358,430,731,755]" pageId="14" pageNumber="957">MLPA</collectionCode>
);
<collectorName id="26C745C1BC5A2A4316CCFD3F3413FD51" box="[453,541,731,755]" pageId="14" pageNumber="957">Poconé</collectorName>
,
<collectorName id="26C745C1BC5A2A431526FD3F355AFD51" box="[559,852,731,755]" pageId="14" pageNumber="957">Rodovia Transpantaneira</collectorName>
,
<date id="FF8C06D7BC5A2A43146FFD3F323FFD51" box="[870,1073,731,755]" pageId="14" pageNumber="957" value="2015-02-08">
<collectingDate id="EFC8FF3FBC5A2A43146FFD3F323FFD51" box="[870,1073,731,755]" pageId="14" pageNumber="957" value="2015-02-08">8 February 2015</collectingDate>
</date>
,
<collectorName id="26C745C1BC5A2A43134BFD3F36F9FCB4" pageId="14" pageNumber="957">Mario Cupello</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431622FD1A341CFCB4" ID-GBIF-Occurrence="2831865327" box="[299,530,766,790]" collectionCode="MNRJ" country="Brazil" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Mato Grosso">
.
<emphasis id="B946FC05BC5A2A431637FD1A3745FCB4" box="[318,331,766,790]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431652FD1A37BEFCB4" box="[347,432,766,790]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4316CCFD1A3402FCB4" box="[453,524,766,790]" country="Brazil" httpUri="http://grbio.org/cool/zi1i-a0b5" name="Museu Nacional/Universidade Federal de Rio de Janeiro" pageId="14" pageNumber="957">MNRJ</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A43152DFD1A3758FC9B" ID-GBIF-Occurrence="2831865465" collectorName="Alto Paraguai" country="Brazil" location="Corumba" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Mato Grosso do Sul">
<collectingRegion id="49F6EEF5BC5A2A43152DFD1A3542FCB4" box="[548,844,766,790]" country="Brazil" name="Mato Grosso do Sul" pageId="14" pageNumber="957">MATO GROSSO DO SUL</collectingRegion>
:
<location id="8EED76CCBC5A2A431454FD1A35C7FCB4" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A431454FD1A35C7FCB4" box="[861,969,766,790]" country="Brazil" name="Corumba" pageId="14" pageNumber="957" stateProvince="Mato Grosso do Sul">Corumbá</location>
,
<collectorName id="26C745C1BC5A2A4314D5FD1A328EFCB4" box="[988,1152,766,790]" pageId="14" pageNumber="957">Alto Paraguai</collectorName>
, without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43165EFCC537EEFC9B" ID-GBIF-Occurrence="2831865355" box="[343,480,801,825]" collectorName="Richter" country="Brazil" isTailEnumeration="true" location="Corumba" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Mato Grosso do Sul">
.
<collectorName id="26C745C1BC5A2A43166AFCC537BDFC9B" box="[355,435,801,825]" pageId="14" pageNumber="957">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4316E9FCC534A2FC9B" ID-GBIF-Occurrence="2831865448" box="[480,684,801,825]" collectionCode="MLPA" country="Brazil" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Mato Grosso do Sul">
.
<emphasis id="B946FC05BC5A2A4316E5FCC537F7FC9B" box="[492,505,801,825]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43150BFCC53441FC9B" box="[514,591,801,825]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431557FCC534A8FC9B" box="[606,678,801,825]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4315B1FCC534E0FCF9" ID-GBIF-Occurrence="2831865419" collectingDate="1959-11-07" collectionCode="BMNH" country="Brazil" location="RIO GRANDE DO SUL" pageId="14" pageNumber="957" specimenCount="5" specimenCount-female="1" specimenCount-male="4" stateProvince="Rio Grande do Sul">
<collectingRegion id="49F6EEF5BC5A2A4315B1FCC535A0FC9B" box="[696,942,801,825]" country="Brazil" name="Rio Grande do Sul" pageId="14" pageNumber="957">RIO GRANDE DO SUL</collectingRegion>
: no more data
<emphasis id="B946FC05BC5A2A431369FCC53263FC9B" box="[1120,1133,801,825]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43137CFCC536ECFCF9" pageId="14" pageNumber="957" type="male">4 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A43162AFCA73786FCF9" box="[291,392,835,859]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431691FCA737E8FCF9" box="[408,486,835,859]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<date id="FF8C06D7BC5A2A4316F3FCA734C1FCF9" box="[506,719,835,859]" pageId="14" pageNumber="957" value="1959-11-07">
<collectingDate id="EFC8FF3FBC5A2A4316F3FCA734C1FCF9" box="[506,719,835,859]" pageId="14" pageNumber="957" value="1959-11-07">7 November 1959</collectingDate>
</date>
, C
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4315E6FCA7359FFCF9" ID-GBIF-Occurrence="2831865363" box="[751,913,835,859]" collectingDate="1959-11-07" collectionCode="BMNH" collectorName="Biezanko" country="Brazil" isTailEnumeration="true" location="RIO GRANDE DO SUL" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Rio Grande do Sul">
.
<collectorName id="26C745C1BC5A2A4315F5FCA7356CFCF9" box="[764,866,835,859]" pageId="14" pageNumber="957">Biezanko</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431499FCA7378AFCDC" ID-GBIF-Occurrence="2831865426" collectingDate="1961-10-20" collectionCode="BMNH" country="Brazil" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1" stateProvince="Rio Grande do Sul">
.
<emphasis id="B946FC05BC5A2A431497FCA735A5FCF9" box="[926,939,835,859]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4314BFFCA73215FCF9" box="[950,1051,835,859]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431325FCA73274FCF9" box="[1068,1146,835,859]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<date id="FF8C06D7BC5A2A4317A9FC82376BFCDC" box="[160,357,870,894]" pageId="14" pageNumber="957" value="1961-10-20">
<collectingDate id="EFC8FF3FBC5A2A4317A9FC82376BFCDC" box="[160,357,870,894]" pageId="14" pageNumber="957" value="1961-10-20">20 October 1961</collectingDate>
</date>
, C
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43168DFC82342AFCDC" ID-GBIF-Occurrence="2831865443" box="[388,548,870,894]" collectingDate="1961-10-20" collectionCode="BMNH" collectorName="Biezanko" country="Brazil" isTailEnumeration="true" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Rio Grande do Sul">
.
<collectorName id="26C745C1BC5A2A431698FC8237F9FCDC" box="[401,503,870,894]" pageId="14" pageNumber="957">Biezanko</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43152DFC8235C8FCDC" ID-GBIF-Occurrence="2831865449" box="[548,966,870,894]" collectingDate="1995-01" collectionCode="BMNH" country="Brazil" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Rio Grande do Sul">
.
<emphasis id="B946FC05BC5A2A431538FC823430FCDC" box="[561,574,870,894]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431541FC823499FCDC" box="[584,663,870,894]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4315AFFC8234FAFCDC" box="[678,756,870,894]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<date id="FF8C06D7BC5A2A43140FFC8235AFFCDC" box="[774,929,870,894]" pageId="14" pageNumber="957" value="1995-01">
<collectingDate id="EFC8FF3FBC5A2A43140FFC8235AFFCDC" box="[774,929,870,894]" pageId="14" pageNumber="957" value="1995-01">January 1995</collectingDate>
</date>
, M
</materialsCitation>
. A.
<materialsCitation id="3B5A2A4ABC5A2A4314FBFC8236CAFC03" ID-GBIF-Occurrence="2831865460" country="Brazil" location="Fernando" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Rio Grande do Sul">
<location id="8EED76CCBC5A2A4314FBFC823251FCDC" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4314FBFC823251FCDC" box="[1010,1119,870,894]" country="Brazil" name="Fernando" pageId="14" pageNumber="957" stateProvince="Rio Grande do Sul">Fernando</location>
(?) col
</materialsCitation>
.
<emphasis id="B946FC05BC5A2A4317DDFC6D36EFFC03" box="[212,225,905,929]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<materialsCitation id="3B5A2A4ABC5A2A4317E7FC6D352DFC03" ID-GBIF-Occurrence="2831865338" box="[238,803,905,929]" collectingDate="1953-11-10" collectionCode="MACN" collectorName="Pelotas" country="Brazil" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Rio Grande do Sul">
<specimenCount id="9D34EB9EBC5A2A4317E7FC6D374EFC03" box="[238,320,905,929]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43165BFC6D37AEFC03" box="[338,416,905,929]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC5A2A4316BFFC6D3405FC03" box="[438,523,905,929]" pageId="14" pageNumber="957">Pelotas</collectorName>
,
<date id="FF8C06D7BC5A2A431512FC6D350FFC03" box="[539,769,905,929]" pageId="14" pageNumber="957" value="1953-11-10">
<collectingDate id="EFC8FF3FBC5A2A431512FC6D350FFC03" box="[539,769,905,929]" pageId="14" pageNumber="957" value="1953-11-10">10 November 1953</collectingDate>
</date>
, C
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43142DFC6D3545FC03" ID-GBIF-Occurrence="2831865425" box="[804,843,905,929]" collectingDate="1953-11-10" collectionCode="MACN" collectorName="Pelotas" country="Brazil" isTailEnumeration="true" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Rio Grande do Sul">. M</materialsCitation>
. Biezanko col
<materialsCitation id="3B5A2A4ABC5A2A4314FBFC6D36F4FC61" ID-GBIF-Occurrence="2831865416" collectionCode="BMNH" country="Brazil" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1" stateProvince="Rio Grande do Sul">
.
<emphasis id="B946FC05BC5A2A43130BFC6D3201FC03" box="[1026,1039,905,929]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431315FC6D328DFC03" box="[1052,1155,905,929]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFC4F36FAFC61" box="[166,244,939,963]" pageId="14" pageNumber="957">BMNH</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A43160FFC4F342BFC44" ID-GBIF-Occurrence="2831865409" collectingDate="1820-03" collectionCode="BMNH, OUMNH" collectorName="Hope-Westwood" country="Brazil" location="SANTA CATARINA" pageId="14" pageNumber="957" specimenCount="4" specimenCount-female="4" stateProvince="Santa Catarina">
<collectingRegion id="49F6EEF5BC5A2A43160FFC4F37DBFC61" box="[262,469,939,963]" country="Brazil" name="Santa Catarina" pageId="14" pageNumber="957">SANTA CATARINA</collectingRegion>
: no more data
<emphasis id="B946FC05BC5A2A43158EFC4F349AFC61" box="[647,660,939,963]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431594FC4F3505FC61" box="[669,779,939,963]" pageId="14" pageNumber="957" type="female">3 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431413FC4F3566FC61" box="[794,872,939,963]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<date id="FF8C06D7BC5A2A431473FC4F320FFC61" box="[890,1025,939,963]" pageId="14" pageNumber="957" value="1820-03">
<collectingDate id="EFC8FF3FBC5A2A431473FC4F320FFC61" box="[890,1025,939,963]" pageId="14" pageNumber="957" value="1820-03">March 1820</collectingDate>
</date>
<emphasis id="B946FC05BC5A2A431300FC4F3218FC61" box="[1033,1046,939,963]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431329FC4F328DFC61" box="[1056,1155,939,963]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFC2A3708FC44" box="[166,262,974,998]" pageId="14" pageNumber="957">OUMNH</collectionCode>
<emphasis id="B946FC05BC5A2A431607FC2A3715FC44" box="[270,283,974,998]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A43162CFC2A37EAFC44" box="[293,484,974,998]" pageId="14" pageNumber="957">Hope-Westwood</collectorName>
coll.)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A43153BFC2A37F1FBAB" ID-GBIF-Occurrence="2831865459" collectionCode="BMNH, MACN" country="Paraguay" location="PARAGUAY" pageId="14" pageNumber="957" specimenCount="2" specimenCount-male="2">
<emphasis id="B946FC05BC5A2A43153BFC2A34B2FC44" bold="true" box="[562,700,974,998]" pageId="14" pageNumber="957">
<collectingCountry id="F3256087BC5A2A43153BFC2A34B2FC44" box="[562,700,974,998]" name="Paraguay" pageId="14" pageNumber="957">PARAGUAY</collectingCountry>
</emphasis>
: no more data
<emphasis id="B946FC05BC5A2A43147BFC2A3571FC44" box="[882,895,974,998]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431480FC2A35D6FC44" box="[905,984,974,998]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4314EFFC2A323FFC44" box="[998,1073,974,998]" pageId="14" pageNumber="957">BMNH</collectionCode>
) and
<specimenCount id="9D34EB9EBC5A2A43137CFC2A36D6FBAB" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317EEFC15373BFBAB" box="[231,309,1009,1033]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
); without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4316F7FC153486FBAB" ID-GBIF-Occurrence="2831865358" box="[510,648,1009,1033]" collectionCode="BMNH, MACN" collectorName="Richter" country="Paraguay" isTailEnumeration="true" location="PARAGUAY" pageId="14" pageNumber="957" specimenCount="1">
.
<collectorName id="26C745C1BC5A2A431502FC153455FBAB" box="[523,603,1009,1033]" pageId="14" pageNumber="957">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431581FC1535B6FBAB" ID-GBIF-Occurrence="2831865453" box="[648,952,1009,1033]" collectionCode="MLPA" country="Paraguay" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC5A2A43159DFC1534AFFBAB" box="[660,673,1009,1033]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4315A2FC1534F4FBAB" box="[683,762,1009,1033]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431401FC15355EFBAB" box="[776,848,1009,1033]" pageId="14" pageNumber="957">MLPA</collectionCode>
); 1908, F
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4314B1FC1535D4FBAB" ID-GBIF-Occurrence="2831865324" box="[952,986,1009,1033]" collectionCode="MLPA" country="Paraguay" isTailEnumeration="true" pageId="14" pageNumber="957" specimenCount="1">. O</materialsCitation>
. Lucas col
<materialsCitation id="3B5A2A4ABC5A2A43135AFC1537EBFB89" ID-GBIF-Occurrence="2831865433" collectionCode="MLPA" country="Paraguay" pageId="14" pageNumber="957" specimenCount="2" specimenCount-female="1" specimenCount-male="1">
.
<emphasis id="B946FC05BC5A2A431356FC153262FBAB" box="[1119,1132,1009,1033]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43137CFC1536D6FB89" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A431614FBF7378AFB89" box="[285,388,1043,1067]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43169EFBF737D1FB89" box="[407,479,1043,1067]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4316FCFBF736F7FBEC" ID-GBIF-Occurrence="2831865477" collectingDate="1922-09" collectingDateMax="1923-04" collectingDateMin="1922-09" country="Paraguay" location="Asuncion" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Asuncion">
DISTRITO CAPITAL:
<collectingRegion id="49F6EEF5BC5A2A4315D4FBF73544FB89" box="[733,842,1043,1067]" country="Paraguay" name="Asuncion" pageId="14" pageNumber="957">Asunción</collectingRegion>
,
<date id="FF8C06D7BC5A2A431453FBF73215FB89" box="[858,1051,1043,1067]" pageId="14" pageNumber="957" value="1922-09">
<collectingDate id="EFC8FF3FBC5A2A431453FBF73215FB89" box="[858,1051,1043,1067]" pageId="14" pageNumber="957" value="1922-09">September 1922</collectingDate>
</date>
to
<date id="FF8C06D7BC5A2A431344FBF736D4FBEC" pageId="14" pageNumber="957" value="1923-04">
<collectingDate id="EFC8FF3FBC5A2A431344FBF736D4FBEC" pageId="14" pageNumber="957" value="1923-04">April 1923</collectingDate>
</date>
, E
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4317F1FBD23715FBEC" ID-GBIF-Occurrence="2831865335" box="[248,283,1078,1102]" collectingDate="1922-09" collectingDateMax="1923-04" collectingDateMin="1922-09" country="Paraguay" isTailEnumeration="true" location="Asuncion" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Asuncion">. G</materialsCitation>
. Kent col
<materialsCitation id="3B5A2A4ABC5A2A431685FBD23470FBEC" ID-GBIF-Occurrence="2831865420" box="[396,638,1078,1102]" collectionCode="BMNH" country="Paraguay" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1" stateProvince="Asuncion">
.
<emphasis id="B946FC05BC5A2A431693FBD237A9FBEC" box="[410,423,1078,1102]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4316BAFBD23417FBEC" box="[435,537,1078,1102]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431523FBD23476FBEC" box="[554,632,1078,1102]" pageId="14" pageNumber="957">BMNH</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431585FBD2322BFBEC" ID-GBIF-Occurrence="2831865351" box="[652,1061,1078,1102]" country="Paraguay" location="Villarrica" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Guaira">
<collectingRegion id="49F6EEF5BC5A2A431585FBD234E6FBEC" box="[652,744,1078,1102]" country="Paraguay" name="Guaira" pageId="14" pageNumber="957">GUAIRÁ</collectingRegion>
:
<location id="8EED76CCBC5A2A4315FEFBD23555FBEC" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4315FEFBD23555FBEC" box="[759,859,1078,1102]" country="Paraguay" name="Villarrica" pageId="14" pageNumber="957" stateProvince="Guaira">Villarrica</location>
, without date, H
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43132CFBD236CAFBD3" ID-GBIF-Occurrence="2831865424" collectorName="Richter" country="Paraguay" isTailEnumeration="true" location="Villarrica" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Guaira">
.
<collectorName id="26C745C1BC5A2A43133AFBD2328DFBEC" box="[1075,1155,1078,1102]" pageId="14" pageNumber="957">Richter</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4317CDFBBD345DFBD3" ID-GBIF-Occurrence="2831865400" box="[196,595,1113,1137]" collectionCode="MLPA" country="Paraguay" pageId="14" pageNumber="957" specimenCount="3" specimenCount-female="2" specimenCount-male="1" stateProvince="Guaira">
.
<emphasis id="B946FC05BC5A2A4317DAFBBD36EEFBD3" box="[211,224,1113,1137]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4317E4FBBD3730FBD3" box="[237,318,1113,1137]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A431688FBBD37FDFBD3" box="[385,499,1113,1137]" pageId="14" pageNumber="957" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43150CFBBD3443FBD3" box="[517,589,1113,1137]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A43156BFBBD35C9FBD3" ID-GBIF-Occurrence="2831865332" box="[610,967,1113,1137]" country="Paraguay" location="Sapucai" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Paraguari">
<collectingRegion id="49F6EEF5BC5A2A43156BFBBD34E2FBD3" box="[610,748,1113,1137]" country="Paraguay" name="Paraguari" pageId="14" pageNumber="957">PARAGUARÍ</collectingRegion>
:
<location id="8EED76CCBC5A2A4315F2FBBD3558FBD3" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4315F2FBBD3558FBD3" box="[763,854,1113,1137]" country="Paraguay" name="Sapucai" pageId="14" pageNumber="957" stateProvince="Paraguari">Sapucai</location>
, 1903, W
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A4314C1FBBD3243FBD3" ID-GBIF-Occurrence="2831865472" box="[968,1101,1113,1137]" collectorName="Foster" country="Paraguay" isTailEnumeration="true" location="Sapucai" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Paraguari">
.
<collectorName id="26C745C1BC5A2A4314DEFBBD3213FBD3" box="[983,1053,1113,1137]" pageId="14" pageNumber="957">Foster</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431344FBBD376CFB31" ID-GBIF-Occurrence="2831865471" collectionCode="BMNH" country="Paraguay" pageId="14" pageNumber="957" specimenCount="2" specimenCount-female="2" stateProvince="Paraguari">
.
<emphasis id="B946FC05BC5A2A431355FBBD3267FBD3" box="[1116,1129,1113,1137]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43137CFBBD36F6FB31" pageId="14" pageNumber="957" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431604FB9F3755FB31" box="[269,347,1147,1171]" pageId="14" pageNumber="957">BMNH</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A43167DFB9F3493FB14" ID-GBIF-Occurrence="2831865446" collectionCode="BMNH, MACN" collectorName="Hermann Burmeister" country="Uruguay" location="URUGUAY" pageId="14" pageNumber="957" specimenCount="6" specimenCount-female="4" specimenCount-male="2">
<emphasis id="B946FC05BC5A2A43167DFB9F37FEFB31" bold="true" box="[372,496,1147,1171]" pageId="14" pageNumber="957">
<collectingCountry id="F3256087BC5A2A43167DFB9F37FEFB31" box="[372,496,1147,1171]" name="Uruguay" pageId="14" pageNumber="957">URUGUAY</collectingCountry>
</emphasis>
: no more data
<emphasis id="B946FC05BC5A2A4315B6FB9F34C2FB31" box="[703,716,1147,1171]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4315D2FB9F355EFB31" box="[731,848,1147,1171]" pageId="14" pageNumber="957" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43146FFB9F35BAFB31" box="[870,948,1147,1171]" pageId="14" pageNumber="957">BMNH</collectionCode>
),
<specimenCount id="9D34EB9EBC5A2A4314C5FB9F3225FB31" box="[972,1067,1147,1171]" pageId="14" pageNumber="957" type="male">2 males</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A43137CFB9F36F6FB14" pageId="14" pageNumber="957" type="female">2 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43160FFB7A375EFB14" box="[262,336,1182,1206]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC5A2A431650FB7A3768FB14" box="[345,358,1182,1206]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A431666FB7A3453FB14" box="[367,605,1182,1206]" pageId="14" pageNumber="957">Hermann Burmeister</collectorName>
coll.)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4315A3FB7A3732FB7B" ID-GBIF-Occurrence="2831865427" collectionCode="MACN" country="Uruguay" location="Atlantida" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Canelones">
<collectingRegion id="49F6EEF5BC5A2A4315A3FB7A3536FB14" box="[682,824,1182,1206]" country="Uruguay" name="Canelones" pageId="14" pageNumber="957">CANELONES</collectingRegion>
:
<location id="8EED76CCBC5A2A43144AFB7A35A0FB14" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A43144AFB7A35A0FB14" box="[835,942,1182,1206]" country="Uruguay" name="Atlantida" pageId="14" pageNumber="957" stateProvince="Canelones">Atlántida</location>
, no more data
<emphasis id="B946FC05BC5A2A431369FB7A3263FB14" box="[1120,1133,1182,1206]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43137CFB7A36D6FB7B" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317E1FB253738FB7B" box="[232,310,1217,1241]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431640FB25328EFB7B" ID-GBIF-Occurrence="2831865392" box="[329,1152,1217,1241]" collectionCode="MACN" collectorName="Riachuelo" country="Uruguay" location="COLONIA" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Colonia">
<collectingRegion id="49F6EEF5BC5A2A431640FB2537B9FB7B" box="[329,439,1217,1241]" country="Uruguay" name="Colonia" pageId="14" pageNumber="957">COLONIA</collectingRegion>
: no more data
<emphasis id="B946FC05BC5A2A431566FB253472FB7B" box="[623,636,1217,1241]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43158FFB2534D8FB7B" box="[646,726,1217,1241]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4315EFFB25353AFB7B" box="[742,820,1217,1241]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
);
<collectorName id="26C745C1BC5A2A43144EFB2535B7FB7B" box="[839,953,1217,1241]" pageId="14" pageNumber="957">Riachuelo</collectorName>
, without date, A
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A431389FB253739FB59" ID-GBIF-Occurrence="2831865445" collectionCode="MACN" collectorName="Stevenin" country="Uruguay" isTailEnumeration="true" location="COLONIA" pageId="14" pageNumber="957" specimenCount="1" stateProvince="Colonia">
.
<collectorName id="26C745C1BC5A2A4317A9FB07370DFB59" box="[160,259,1251,1275]" pageId="14" pageNumber="957">Stévenin</collectorName>
col
</materialsCitation>
<materialsCitation id="3B5A2A4ABC5A2A43163EFB07342BFB59" ID-GBIF-Occurrence="2831865325" box="[311,549,1251,1275]" collectionCode="MACN" country="Uruguay" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Colonia">
.
<emphasis id="B946FC05BC5A2A431643FB073759FB59" box="[330,343,1251,1275]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43166EFB0737B2FB59" box="[359,444,1251,1275]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4316D8FB073411FB59" box="[465,543,1251,1275]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431531FB0736FAFAC1" ID-GBIF-Occurrence="2831865457" collectingDate="1929-12-20" collectingDateMax="1933-10-10" collectingDateMin="1929-12-20" collectionCode="BMNH, MACN, MLPA" collectorName="Penarol" country="Uruguay" location="Penarol" municipality="Carrasco" pageId="14" pageNumber="957" specimenCount="4" specimenCount-female="3" specimenCount-male="1" stateProvince="Montevideo">
<collectingRegion id="49F6EEF5BC5A2A431531FB0734D6FB59" box="[568,728,1251,1275]" country="Uruguay" name="Montevideo" pageId="14" pageNumber="957">MONTEVIDEO</collectingRegion>
: no more data
<emphasis id="B946FC05BC5A2A4314ACFB0735BCFB59" box="[933,946,1251,1275]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4314CBFB073219FB59" box="[962,1047,1251,1275]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431325FB073274FB59" box="[1068,1146,1251,1275]" pageId="14" pageNumber="957">BMNH</collectionCode>
);
<collectingMunicipality id="6BE9BA6DBC5A2A4317A9FAE2370BFABC" box="[160,261,1286,1310]" pageId="14" pageNumber="957">Carrasco</collectingMunicipality>
, no more data
<emphasis id="B946FC05BC5A2A4316C2FAE237D6FABC" box="[459,472,1286,1310]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A4316EFFAE23440FABC" box="[486,590,1286,1310]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43156AFAE234BFFABC" box="[611,689,1286,1310]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
);
<location id="8EED76CCBC5A2A4315C1FAE2352FFABC" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4315C1FAE2352FFABC" box="[712,801,1286,1310]" country="Uruguay" municipality="Carrasco" name="Penarol" pageId="14" pageNumber="957" stateProvince="Montevideo">Peñarol</location>
,
<date id="FF8C06D7BC5A2A43143BFAE23217FABC" box="[818,1049,1286,1310]" pageId="14" pageNumber="957" value="1929-12-20">
<collectingDate id="EFC8FF3FBC5A2A43143BFAE23217FABC" box="[818,1049,1286,1310]" pageId="14" pageNumber="957" value="1929-12-20">20 December 1929</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC5A2A431607FACD3715FAE3" box="[270,283,1321,1345]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43162FFACD3785FAE3" box="[294,395,1321,1345]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431695FACD37EAFAE3" box="[412,484,1321,1345]" pageId="14" pageNumber="957">MLPA</collectionCode>
);
<collectorName id="26C745C1BC5A2A4316FEFACD345EFAE3" box="[503,592,1321,1345]" pageId="14" pageNumber="957">Peñarol</collectorName>
,
<date id="FF8C06D7BC5A2A431557FACD3528FAE3" box="[606,806,1321,1345]" pageId="14" pageNumber="957" value="1933-10-10">
<collectingDate id="EFC8FF3FBC5A2A431557FACD3528FAE3" box="[606,806,1321,1345]" pageId="14" pageNumber="957" value="1933-10-10">10 October 1933</collectingDate>
</date>
, without collector
<emphasis id="B946FC05BC5A2A43130FFACD321DFAE3" box="[1030,1043,1321,1345]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A431317FACD328DFAE3" box="[1054,1155,1321,1345]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFAAF36E0FAC1" box="[166,238,1355,1379]" pageId="14" pageNumber="957">MLPA</collectionCode>
)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431608FAAF374EFA24" ID-GBIF-Occurrence="2831865385" collectionCode="MACN" collectorName="Carlos Bruch &amp; Parana" country="Uruguay" county="Ambiguous" location="Bruch-Waiser" municipality="Fives Lille" pageId="14" pageNumber="957" specimenCount="2" specimenCount-male="2" stateProvince="Montevideo">
<emphasis id="B946FC05BC5A2A431608FAAF37C3FAC1" bold="true" box="[257,461,1355,1379]" pageId="14" pageNumber="957">
<collectingCounty id="62EC589BBC5A2A431608FAAF3783FAC1" box="[257,397,1355,1379]" pageId="14" pageNumber="957">Ambiguous</collectingCounty>
data
</emphasis>
:
<emphasis id="B946FC05BC5A2A4316D5FAAF37ECFAC1" box="[476,482,1355,1379]" italics="true" pageId="14" pageNumber="957">‘</emphasis>
<collectingMunicipality id="6BE9BA6DBC5A2A4316EBFAAF345BFAC1" box="[482,597,1355,1379]" pageId="14" pageNumber="957">Fives Lille</collectingMunicipality>
,
<location id="8EED76CCBC5A2A43156BFAAF34F6FAC1" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A43156BFAAF34F6FAC1" box="[610,760,1355,1379]" country="Uruguay" county="Ambiguous" municipality="Fives Lille" name="Bruch-Waiser" pageId="14" pageNumber="957" stateProvince="Montevideo">Bruch-Waiser</location>
<emphasis id="B946FC05BC5A2A4315F1FAAF351AFAC1" box="[760,788,1355,1379]" italics="true" pageId="14" pageNumber="957">’ –</emphasis>
<specimenCount id="9D34EB9EBC5A2A431417FAAF3576FAC1" box="[798,888,1355,1379]" pageId="14" pageNumber="957" type="male">2 males</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43148EFAAF35DFFAC1" box="[903,977,1355,1379]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC5A2A4314D2FAAF35E6FAC1" box="[987,1000,1355,1379]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A4314FBFAAF328DFAC1" box="[1010,1155,1355,1379]" pageId="14" pageNumber="957">Carlos Bruch</collectorName>
coll.);
<emphasis id="B946FC05BC5A2A4317E0FA8A36E1FA24" box="[233,239,1390,1414]" italics="true" pageId="14" pageNumber="957">‘</emphasis>
<collectorName id="26C745C1BC5A2A4317E6FA8A374EFA24" box="[239,320,1390,1414]" pageId="14" pageNumber="957">Parana</collectorName>
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431646FA8A349FFA24" ID-GBIF-Occurrence="2831865315" box="[335,657,1390,1414]" collectorName="Brazil" country="Uruguay" location="Nov." pageId="14" pageNumber="957" specimenCount="1" stateProvince="Montevideo">
<location id="8EED76CCBC5A2A431646FA8A378AFA24" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A431646FA8A378AFA24" box="[335,388,1390,1414]" country="Uruguay" name="Nov." pageId="14" pageNumber="957" stateProvince="Montevideo">Nov.</location>
<emphasis id="B946FC05BC5A2A43168AFA8A3787FA24" box="[387,393,1390,1414]" italics="true" pageId="14" pageNumber="957">’</emphasis>
(Argentina or
<collectorName id="26C745C1BC5A2A431548FA8A348EFA24" box="[577,640,1390,1414]" pageId="14" pageNumber="957">Brazil</collectorName>
?)
</materialsCitation>
<emphasis id="B946FC05BC5A2A431597FA8A34A5FA24" box="[670,683,1390,1414]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<materialsCitation id="3B5A2A4ABC5A2A4315B0FA8A36D6FA0B" ID-GBIF-Occurrence="2831865388" collectionCode="MACN" collectorName="Hermann Burmeister" country="Uruguay" pageId="14" pageNumber="957" specimenCount="1" specimenCount-male="1" stateProvince="Montevideo">
<specimenCount id="9D34EB9EBC5A2A4315B0FA8A3502FA24" box="[697,780,1390,1414]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431416FA8A3567FA24" box="[799,873,1390,1414]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC5A2A43147FFA8A358DFA24" box="[886,899,1390,1414]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A431498FA8A328AFA24" box="[913,1156,1390,1414]" pageId="14" pageNumber="957">Hermann Burmeister</collectorName>
coll.)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A4317F9FA753780FA69" ID-GBIF-Occurrence="2831865336" collectionCode="BMNH" collectorName="Frey" country="Colombia" location="Specimens" pageId="14" pageNumber="957" specimenCount="1" specimenCount-female="1">
<emphasis id="B946FC05BC5A2A4317F9FA753503FA0B" bold="true" box="[240,781,1425,1449]" pageId="14" pageNumber="957">
<location id="8EED76CCBC5A2A4317F9FA75377CFA0B" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A4317F9FA75377CFA0B" box="[240,370,1425,1449]" country="Colombia" name="Specimens" pageId="14" pageNumber="957">Specimens</location>
surely mislabeled:
<collectingCountry id="F3256087BC5A2A43158EFA753503FA0B" box="[647,781,1425,1449]" name="Colombia" pageId="14" pageNumber="957">COLOMBIA</collectingCountry>
</emphasis>
: no more data
<emphasis id="B946FC05BC5A2A4314F8FA7535F0FA0B" box="[1009,1022,1425,1449]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<specimenCount id="9D34EB9EBC5A2A43131AFA75328DFA0B" box="[1043,1155,1425,1449]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317AFFA5736FEFA69" box="[166,240,1459,1483]" pageId="14" pageNumber="957">BMNH</collectionCode>
<emphasis id="B946FC05BC5A2A4317F4FA573704FA69" box="[253,266,1459,1483]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A431611FA573746FA69" box="[280,328,1459,1483]" pageId="14" pageNumber="957">Frey</collectorName>
coll.)
</materialsCitation>
.
<materialsCitation id="3B5A2A4ABC5A2A431696FA57351EF9B3" ID-GBIF-Occurrence="2831865397" collectionCode="MACN, OUMNH" collectorName="Carlos Bruch &amp; Hermann Burmeister &amp; Westwood &amp; Gory" country="Colombia" location="No" pageId="14" pageNumber="957" specimenCount="9" specimenCount-female="6" specimenCount-male="3">
<emphasis id="B946FC05BC5A2A431696FA57340BFA69" bold="true" box="[415,517,1459,1483]" pageId="14" pageNumber="957">
<location id="8EED76CCBC5A2A431696FA5737CFFA69" LSID="urn:lsid:plazi:treatment:039B9101BC582A5A16D9FBF735D0FBC3:8EED76CCBC5A2A431696FA5737CFFA69" box="[415,449,1459,1483]" country="Colombia" name="No" pageId="14" pageNumber="957">No</location>
data
</emphasis>
:
<specimenCount id="9D34EB9EBC5A2A431510FA573462FA69" box="[537,620,1459,1483]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A431576FA5734C7FA69" box="[639,713,1459,1483]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC5A2A4315DFFA5734EDFA69" box="[726,739,1459,1483]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A4315F8FA57358BFA69" box="[753,901,1459,1483]" pageId="14" pageNumber="957">Carlos Bruch</collectorName>
coll.),
<specimenCount id="9D34EB9EBC5A2A4314D5FA573221FA69" box="[988,1071,1459,1483]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A43137CFA5736E3FA4C" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4317F4FA323749FA4C" box="[253,327,1494,1518]" country="Argentina" httpUri="http://grbio.org/cool/pyng-6456" name="Museo Argentino de Ciencias Naturales Bernardino Rivadavia" pageId="14" pageNumber="957" type="Museum">MACN</collectionCode>
<emphasis id="B946FC05BC5A2A43165BFA323751FA4C" box="[338,351,1494,1518]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A431663FA323457FA4C" box="[362,601,1494,1518]" pageId="14" pageNumber="957">Hermann Burmeister</collectorName>
coll.),
<specimenCount id="9D34EB9EBC5A2A4315A3FA3234F5FA4C" box="[682,763,1494,1518]" pageId="14" pageNumber="957" type="male">1 male</specimenCount>
and
<specimenCount id="9D34EB9EBC5A2A431432FA3235A5FA4C" box="[827,939,1494,1518]" pageId="14" pageNumber="957" type="female">4 females</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A4314B2FA323215FA4C" box="[955,1051,1494,1518]" pageId="14" pageNumber="957">OUMNH</collectionCode>
<emphasis id="B946FC05BC5A2A43132CFA32323CFA4C" box="[1061,1074,1494,1518]" italics="true" pageId="14" pageNumber="957">–</emphasis>
Hope-
<collectorName id="26C745C1BC5A2A4317A9FA1D3717F9B3" box="[160,281,1529,1553]" pageId="14" pageNumber="957">Westwood</collectorName>
coll.), and
<specimenCount id="9D34EB9EBC5A2A4316A9FA1D340BF9B3" box="[416,517,1529,1553]" pageId="14" pageNumber="957" type="female">1 female</specimenCount>
(
<collectionCode id="ED23B8D2BC5A2A43151FFA1D3478F9B3" box="[534,630,1529,1553]" pageId="14" pageNumber="957">OUMNH</collectionCode>
<emphasis id="B946FC05BC5A2A431576FA1C3482F9B2" box="[639,652,1528,1552]" italics="true" pageId="14" pageNumber="957">–</emphasis>
<collectorName id="26C745C1BC5A2A43159EFA1D34C3F9B3" box="[663,717,1529,1553]" pageId="14" pageNumber="957">Gory</collectorName>
coll.)
</materialsCitation>
.
</paragraph>
<paragraph id="8B8D2017BC5A2A4317A9F9AB37B1F9CB" blockId="14.[160,447,1615,1641]" box="[160,447,1615,1641]" pageId="14" pageNumber="957">
<emphasis id="B946FC05BC5A2A4317A9F9AB37B1F9CB" bold="true" box="[160,447,1615,1641]" pageId="14" pageNumber="957">Phylogenetic analysis</emphasis>
</paragraph>
<paragraph id="8B8D2017BC5A2A4217A9F96137FDFD73" blockId="14.[160,1156,1669,1727]" lastBlockId="15.[160,1156,142,1102]" lastPageId="15" lastPageNumber="958" pageId="14" pageNumber="957">
The phylogenetic analysis performed here had three main goals: to test the monophyly of
<taxonomicName id="4C325B94BC5A2A431611F9433785F91D" authorityName="Brulle" authorityYear="1837" box="[280,395,1703,1727]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="14" pageNumber="957" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5A2A431611F9433785F91D" box="[280,395,1703,1727]" italics="true" pageId="14" pageNumber="957">Gromphas</emphasis>
</taxonomicName>
, to know the phylogenetic relationship of its species, and, therewith, to raise hypotheses about the transformation of their characters through the evolutionary process. In recent works (
<bibRefCitation id="EFA35DE6BC5B2A4215ADFF553571FF6B" author="Philips TK &amp; Edmonds WD &amp; Scholtz CH" box="[676,895,177,201]" pageId="15" pageNumber="958" pagination="43 - 63" refId="ref15926" refString="Philips TK, Edmonds WD, Scholtz CH. 2004. A phylogenetic analysis of the New World tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae): hypothesis on relationship and origins. Insect Syst Evol. 35: 43 - 63." type="journal article" year="2004">Philips et al. 2004</bibRefCitation>
; Cupello and Vaz-de- Mello 2013), it was well demonstrated that
<taxonomicName id="4C325B94BC5B2A4215CFFF303537FF4E" authorityName="Brulle" authorityYear="1837" box="[710,825,212,236]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5B2A4215CFFF303537FF4E" box="[710,825,212,236]" italics="true" pageId="15" pageNumber="958">Gromphas</emphasis>
</taxonomicName>
is a member of the tribe
<taxonomicName id="4C325B94BC5B2A4217A9FF12371EFEAC" box="[160,272,246,270]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
and closely related to
<taxonomicName id="4C325B94BC5B2A421533FF1234A6FEAC" authorityName="Bates" authorityYear="1870" box="[570,680,246,270]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5B2A421533FF1234A6FEAC" box="[570,680,246,270]" italics="true" pageId="15" pageNumber="958">Oruscatus</emphasis>
</taxonomicName>
; together, these two genera form the monophyletic subtribe
<taxonomicName id="4C325B94BC5B2A4216B3FEFD3454FE93" box="[442,602,281,305]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
. The monotypic genus
<taxonomicName id="4C325B94BC5B2A421488FEFD3270FE93" authority="Harold, 1868" authorityName="Harold" authorityYear="1868" box="[897,1150,281,305]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5B2A421488FEFD35D6FE93" box="[897,984,281,305]" italics="true" pageId="15" pageNumber="958">Bolbites</emphasis>
<bibRefCitation id="EFA35DE6BC5B2A4214ECFEFD3270FE93" author="Harold E" box="[997,1150,281,305]" pageId="15" pageNumber="958" pagination="79 - 86" refId="ref15415" refString="Harold E. 1868. Diagnosen neuer Coprophagen. Coleopterol Hefte. 4: 79 - 86." type="journal article" year="1868">Harold, 1868</bibRefCitation>
</taxonomicName>
, previously considered to be related to this clade, was transferred to Phanaeina by us (
<bibRefCitation id="EFA35DE6BC5B2A4217AFFEBA3417FED4" author="Cupello M &amp; Vaz-de-Mello FZ" box="[166,537,350,374]" pageId="15" pageNumber="958" pagination="439 - 482" refId="ref14473" 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>
) based primarily on the results of the phylogenetic analysis of
<bibRefCitation id="EFA35DE6BC5B2A42162BFE6537F9FE3B" author="Philips TK &amp; Edmonds WD &amp; Scholtz CH" box="[290,503,385,409]" pageId="15" pageNumber="958" pagination="43 - 63" refId="ref15926" refString="Philips TK, Edmonds WD, Scholtz CH. 2004. A phylogenetic analysis of the New World tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae): hypothesis on relationship and origins. Insect Syst Evol. 35: 43 - 63." type="journal article" year="2004">Philips et al. (2004</bibRefCitation>
; morphological data) and of
<bibRefCitation id="EFA35DE6BC5B2A421444FE65328EFE3B" author="Ocampo FC &amp; Hawks DC" box="[845,1152,385,409]" pageId="15" pageNumber="958" pagination="57 - 570" refId="ref15730" refString="Ocampo FC, Hawks DC. 2006. Molecular phylogenetics and evolution of the food relocation behaviour of the dung beetle tribe Eucraniini (Coleoptera: Scarabaeidae: Scarabaeinae). Invertebr Syst. 20: 57 - 570." type="journal article" year="2006">Ocampo and Hawks (2006</bibRefCitation>
; molecular data), besides other published information about the nesting behaviour of these three genera (e.g.
<bibRefCitation id="EFA35DE6BC5B2A4216D9FE223564FE7C" author="Cabrera-Walsh G &amp; Gandolfo D" box="[464,874,454,478]" pageId="15" pageNumber="958" pagination="581 - 588" refId="ref14349" refString="Cabrera-Walsh G, Gandolfo D. 1996. Nidification of thirteen common Argentine dung beetles (Scarabaeidae: Scarabaeinae). Ann Entomol Soc Am. 89: 581 - 588." type="journal article" year="1996">Cabrera-Walsh and Gandolfo 1996</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC5B2A421474FE2236D6FDA3" author="Halffter G &amp; Edmonds WD" pageId="15" pageNumber="958" refId="ref15238" refString="Halffter G, Edmonds WD. 1982. The nesting behavior of dung beetles (Scarabaeinae). An ecological and evolutive approach. Mexico (DF): Instituto de Ecologia." type="book" year="1982">Halffter and Edmonds 1982</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC5B2A4217E7FE0D3423FDA3" author="Sanchez MV &amp; Genise JF" box="[238,557,489,513]" pageId="15" pageNumber="958" pagination="49 - 61" refId="ref16130" refString="Sanchez MV, Genise JF. 2008. Nest and brood chamber structure of two South American dung beetles: Gromphas lacordairei Brulle and Ontherus sulcator (Fabricius) (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopt Bull. 62: 49 - 61." type="journal article" year="2008">Sánchez and Genise 2008</bibRefCitation>
). In Phanaeina,
<taxonomicName id="4C325B94BC5B2A4215F7FE0D355AFDA3" authorityName="Harold" authorityYear="1868" box="[766,852,489,513]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5B2A4215F7FE0D355AFDA3" box="[766,852,489,513]" italics="true" pageId="15" pageNumber="958">Bolbites</emphasis>
</taxonomicName>
is sister of the lineage comprising the remaining genera of the subtribe (
<bibRefCitation id="EFA35DE6BC5B2A421409FDE835D4FD86" author="Philips TK &amp; Edmonds WD &amp; Scholtz CH" box="[768,986,524,548]" pageId="15" pageNumber="958" pagination="43 - 63" refId="ref15926" refString="Philips TK, Edmonds WD, Scholtz CH. 2004. A phylogenetic analysis of the New World tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae): hypothesis on relationship and origins. Insect Syst Evol. 35: 43 - 63." type="journal article" year="2004">Philips et al. 2004</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC5B2A4214E2FDE8373FFDE4" author="Ocampo FC &amp; Hawks DC" pageId="15" pageNumber="958" pagination="57 - 570" refId="ref15730" refString="Ocampo FC, Hawks DC. 2006. Molecular phylogenetics and evolution of the food relocation behaviour of the dung beetle tribe Eucraniini (Coleoptera: Scarabaeidae: Scarabaeinae). Invertebr Syst. 20: 57 - 570." type="journal article" year="2006">Ocampo and Hawks 2006</bibRefCitation>
). Taking these information into account, we selected as outgroups one of the two species of the undoubtedly monophyletic
<emphasis id="B946FC05BC5B2A42144BFDB535B8FDCB" box="[834,950,593,617]" italics="true" pageId="15" pageNumber="958">
<taxonomicName id="4C325B94BC5B2A42144BFDB535BCFDCB" authorityName="Bates" authorityYear="1870" box="[834,946,593,617]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">Oruscatus</taxonomicName>
,
</emphasis>
<taxonomicName id="4C325B94BC5B2A4214CCFDB53754FD29" authority="(Erichson, 1847)" baseAuthorityName="Erichson" baseAuthorityYear="1847" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="species" species="davus">
<emphasis id="B946FC05BC5B2A4214CCFDB5328DFDCB" box="[965,1155,593,617]" italics="true" pageId="15" pageNumber="958">Oruscatus davus</emphasis>
(
<bibRefCitation id="EFA35DE6BC5B2A4217AFFD97375FFD29" author="Erichson WF" box="[166,337,627,651]" pageId="15" pageNumber="958" pagination="552 - 950" refId="ref14975" refString="Erichson WF, editor. 1847. Scarabaeides. In: Naturgeschichte der Insecten Deutschlands. Berlin: Nicolaische Buchhandlung; p. 552 - 950." type="book chapter" year="1847">Erichson, 1847</bibRefCitation>
)
</taxonomicName>
, and one Phanaeina,
<taxonomicName id="4C325B94BC5B2A421557FD9735CAFD29" authority="Harold, 1868" authorityName="Harold" authorityYear="1868" box="[606,964,627,651]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="species" species="onitoides">
<emphasis id="B946FC05BC5B2A421557FD97352BFD29" box="[606,805,627,651]" italics="true" pageId="15" pageNumber="958">Bolbites onitoides</emphasis>
<bibRefCitation id="EFA35DE6BC5B2A421426FD9735CAFD29" author="Harold E" box="[815,964,627,651]" pageId="15" pageNumber="958" pagination="79 - 86" refId="ref15415" refString="Harold E. 1868. Diagnosen neuer Coprophagen. Coleopterol Hefte. 4: 79 - 86." type="journal article" year="1868">Harold, 1868</bibRefCitation>
</taxonomicName>
, the latter used to root the tree and selected among the Phanaeina by its close morphological similarity with
<taxonomicName id="4C325B94BC5B2A421646FD5D37E1FD73" box="[335,495,697,721]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
.
</paragraph>
<paragraph id="8B8D2017BC5B2A4217C9FD3F320EFBEC" blockId="15.[160,1156,142,1102]" pageId="15" pageNumber="958">
Based on 31 informative morphological characters (see below and
<tableCitation id="C6B015ACBC5B2A4214FDFD3F3243FD51" box="[1012,1101,731,755]" captionStart="Table 1" captionStartId="15.[160,216,1371,1394]" captionText="Table 1. Matrix used for the phylogenetic analysis." pageId="15" pageNumber="958">Table 1</tableCitation>
), an exhaustive search (implicit enumeration) was conducted using the program TNT (
<bibRefCitation id="EFA35DE6BC5B2A4217AFFCC5379BFC9B" author="Goloboff PA &amp; Farris JS &amp; Nixon KC" box="[166,405,801,825]" pageId="15" pageNumber="958" pagination="774 - 786" refId="ref15210" refString="Goloboff PA, Farris JS, Nixon KC. 2008. TNT, a free program for phylogenetic analysis. Cladistics. 24: 774 - 786." type="journal article" year="2008">Goloboff et al. 2008</bibRefCitation>
) with Fitch parsimony and equally weighted characters, which resulted in a single tree (
<figureCitation id="13093C92BC5B2A4216C4FCA73420FCF9" box="[461,558,835,859]" captionStart="Figure 6" captionStartId="16.[160,225,923,946]" captionTargetBox="[163,1153,142,892]" captionTargetId="figure@16.[163,1153,142,892]" captionTargetPageId="16" captionText="Figure 6. Phylogeny of Gromphas. Above: Cladogram showing the phylogenetic relationship of the species of Gromphas and the evolution of their characters. Black circles indicate uncontroverted synapormorphies; white circles with black margin, controverted synapomorphies; red circles, ambiguous uncontroverted synapomorphies; and white circles with red margin, ambiguous controverted synapomorphies. Below: Support values given for each clade are decay index (above branch) and bootstrap (below branch)." figureDoi="http://doi.org/10.5281/zenodo.3990455" httpUri="https://zenodo.org/record/3990455/files/figure.png" pageId="15" pageNumber="958">Figure 6</figureCitation>
) of length 46, consistency index 78 and retention index 77, both indices calculated in WinClada (
<bibRefCitation id="EFA35DE6BC5B2A4215F6FC823580FCDC" author="Nixon K" box="[767,910,870,894]" pageId="15" pageNumber="958" refId="ref15694" refString="Nixon K. 1999 - 2002. Winclada, Version 1.00.08. Published by the author." type="book" year="1999">
Nixon
<date id="FF8C06D7BC5B2A42145CFC823580FCDC" box="[853,910,870,894]" pageId="15" pageNumber="958" value="1999" valueMax="2002">1999</date>
</bibRefCitation>
<emphasis id="B946FC05BC5B2A421487FC823595FCDC" box="[910,923,870,894]" italics="true" pageId="15" pageNumber="958">–</emphasis>
2002). Supports of clades were calculated in TNT using both Bootstrap and decay index (Bremmer support) values, this last index based on a search of the 88 suboptimal trees with up to 10 steps longer than the optimal tree. The most parsimonious tree was edited in WinClada, where unambiguous characters were plotted. The ambiguous characters, on the other hand, were individually optimized based on scenarios considered more likely by us and discussed throughout the text below.
</paragraph>
<paragraph id="8B8D2017BC5B2A4217A9FB4A37B2FB6A" blockId="15.[160,444,1198,1224]" box="[160,444,1198,1224]" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4217A9FB4A37B2FB6A" bold="true" box="[160,444,1198,1224]" pageId="15" pageNumber="958">Character statements</emphasis>
</paragraph>
<paragraph id="8B8D2017BC5B2A4217A9FB07328CFABC" blockId="15.[160,1154,1251,1310]" pageId="15" pageNumber="958">
The character statements, i.e. the characters (including locators, variables and variable qualifiers) and their states, were presented following Sereno
<emphasis id="B946FC05BC5B2A42140EFAE23502FABC" box="[775,780,1286,1310]" italics="true" pageId="15" pageNumber="958">’</emphasis>
s (2007) proposals. As a result, 31
</paragraph>
<caption id="DF4D709FBC5B2A4217A9FABF34AEFAD0" box="[160,672,1371,1394]" pageId="15" pageNumber="958">
<paragraph id="8B8D2017BC5B2A4217A9FABF34AEFAD0" blockId="15.[160,672,1371,1394]" box="[160,672,1371,1394]" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4217A9FABF36FDFAD0" bold="true" box="[160,243,1371,1394]" pageId="15" pageNumber="958">Table 1.</emphasis>
Matrix used for the phylogenetic analysis.
</paragraph>
</caption>
<paragraph id="8B8D2017BC5B2A42163AFA9B3274F912" pageId="15" pageNumber="958">
<table id="F932D2B7BC5BD5B217A2FA9B3274F912" box="[171,1146,1407,1712]" gridcols="32" gridrows="13" pageId="15" pageNumber="958">
<tr id="35022255BC5BD5B217A2FA9B3274FA33" box="[171,1146,1407,1425]" gridrow="0" pageId="15" pageNumber="958" rowspan-0="1" rowspan-24="1" rowspan-25="1" rowspan-26="1" rowspan-27="1" rowspan-28="1" rowspan-29="1" rowspan-30="1" rowspan-31="1">
<th id="76D34B29BC5BD5B2163AFA9B3776FA33" box="[307,376,1407,1425]" gridcol="1" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42163AFA9B3776FA33" bold="true" box="[307,376,1407,1425]" pageId="15" pageNumber="958">Characters</emphasis>
</th>
<th id="76D34B29BC5BD5B21680FA9B379CFA33" box="[393,402,1407,1425]" gridcol="2" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421680FA9B379FFA33" bold="true" box="[393,401,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B216ABFA9B37A5FA33" box="[418,427,1407,1425]" gridcol="3" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216AAFA9B37A5FA33" bold="true" box="[419,427,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B216B5FA9B37CBFA33" box="[444,453,1407,1425]" gridcol="4" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216B5FA9B37CAFA33" bold="true" box="[444,452,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B216DCFA9B37D1FA33" box="[469,479,1407,1425]" gridcol="5" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216DFFA9B37D0FA33" bold="true" box="[470,478,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B216E6FA9B37F7FA33" box="[495,505,1407,1425]" gridcol="6" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216F9FA9B37F6FA33" bold="true" box="[496,504,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B21500FA9B341DFA33" box="[521,531,1407,1425]" gridcol="7" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421503FA9B341CFA33" bold="true" box="[522,530,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B2152AFA9B3422FA33" box="[547,556,1407,1425]" gridcol="8" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42152AFA9B3425FA33" bold="true" box="[547,555,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B21535FA9B3448FA33" box="[572,582,1407,1425]" gridcol="9" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421534FA9B344BFA33" bold="true" box="[573,581,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B2155FFA9B3451FA33" box="[598,607,1407,1425]" gridcol="10" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42155EFA9B3451FA33" bold="true" box="[599,607,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B21579FA9B3477FA33" box="[624,633,1407,1425]" gridcol="11" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421579FA9B3476FA33" bold="true" box="[624,632,1407,1425]" pageId="15" pageNumber="958">1</emphasis>
</th>
<th id="76D34B29BC5BD5B21580FA9B349CFA33" box="[649,658,1407,1425]" gridcol="12" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421583FA9B349CFA33" bold="true" box="[650,658,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B215AAFA9B34A2FA33" box="[675,684,1407,1425]" gridcol="13" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215AAFA9B34A5FA33" bold="true" box="[675,683,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B215B5FA9B34C8FA33" box="[700,710,1407,1425]" gridcol="14" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215B4FA9B34CBFA33" bold="true" box="[701,709,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B215DFFA9B34D1FA33" box="[726,735,1407,1425]" gridcol="15" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215DFFA9B34D0FA33" bold="true" box="[726,734,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B215E6FA9B34F7FA33" box="[751,761,1407,1425]" gridcol="16" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215F9FA9B34F6FA33" bold="true" box="[752,760,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B21400FA9B351DFA33" box="[777,787,1407,1425]" gridcol="17" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421403FA9B351CFA33" bold="true" box="[778,786,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B2142AFA9B3522FA33" box="[803,812,1407,1425]" gridcol="18" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42142DFA9B3522FA33" bold="true" box="[804,812,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B21434FA9B3548FA33" box="[829,838,1407,1425]" gridcol="19" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421434FA9B354BFA33" bold="true" box="[829,837,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B2145FFA9B3551FA33" box="[854,863,1407,1425]" gridcol="20" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42145EFA9B3551FA33" bold="true" box="[855,863,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B21479FA9B3577FA33" box="[880,889,1407,1425]" gridcol="21" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421479FA9B3576FA33" bold="true" box="[880,888,1407,1425]" pageId="15" pageNumber="958">2</emphasis>
</th>
<th id="76D34B29BC5BD5B21480FA9B359DFA33" box="[905,915,1407,1425]" gridcol="22" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421483FA9B359CFA33" bold="true" box="[906,914,1407,1425]" pageId="15" pageNumber="958">3</emphasis>
</th>
<th id="76D34B29BC5BD5B214AAFA9B35A3FA33" box="[931,941,1407,1425]" gridcol="23" gridrow="0" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4214ADFA9B35A2FA33" bold="true" box="[932,940,1407,1425]" pageId="15" pageNumber="958">3</emphasis>
</th>
</tr>
<tr id="35022255BC5BD5B217A2FA7A3274FA12" box="[171,1146,1438,1456]" gridrow="1" pageId="15" pageNumber="958">
<th id="76D34B29BC5BD5B217A2FA7A372DFA12" box="[171,291,1438,1456]" gridcol="0" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4217A2FA7A36D5FA12" bold="true" box="[171,219,1438,1456]" pageId="15" pageNumber="958">Species</emphasis>
</th>
<td id="76D34B29BC5BD5B2163AFA7A3776FA12" box="[307,376,1438,1456]" gridcol="1" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42165AFA7A3755FA12" bold="true" box="[339,347,1438,1456]" pageId="15" pageNumber="958">1</emphasis>
</td>
<td id="76D34B29BC5BD5B21680FA7A379CFA12" box="[393,402,1438,1456]" gridcol="2" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421683FA7A379CFA12" bold="true" box="[394,402,1438,1456]" pageId="15" pageNumber="958">2</emphasis>
</td>
<td id="76D34B29BC5BD5B216ABFA7A37A5FA12" box="[418,427,1438,1456]" gridcol="3" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216AAFA7A37A5FA12" bold="true" box="[419,427,1438,1456]" pageId="15" pageNumber="958">3</emphasis>
</td>
<td id="76D34B29BC5BD5B216B5FA7A37CBFA12" box="[444,453,1438,1456]" gridcol="4" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216B4FA7A37CBFA12" bold="true" box="[445,453,1438,1456]" pageId="15" pageNumber="958">4</emphasis>
</td>
<td id="76D34B29BC5BD5B216DCFA7A37D1FA12" box="[469,479,1438,1456]" gridcol="5" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216DEFA7A37D1FA12" bold="true" box="[471,479,1438,1456]" pageId="15" pageNumber="958">5</emphasis>
</td>
<td id="76D34B29BC5BD5B216E6FA7A37F7FA12" box="[495,505,1438,1456]" gridcol="6" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216F9FA7A37F6FA12" bold="true" box="[496,504,1438,1456]" pageId="15" pageNumber="958">6</emphasis>
</td>
<td id="76D34B29BC5BD5B21500FA7A341DFA12" box="[521,531,1438,1456]" gridcol="7" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421503FA7A341CFA12" bold="true" box="[522,530,1438,1456]" pageId="15" pageNumber="958">7</emphasis>
</td>
<td id="76D34B29BC5BD5B2152AFA7A3422FA12" box="[547,556,1438,1456]" gridcol="8" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42152DFA7A3422FA12" bold="true" box="[548,556,1438,1456]" pageId="15" pageNumber="958">8</emphasis>
</td>
<td id="76D34B29BC5BD5B21535FA7A3448FA12" box="[572,582,1438,1456]" gridcol="9" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421537FA7A3448FA12" bold="true" box="[574,582,1438,1456]" pageId="15" pageNumber="958">9</emphasis>
</td>
<td id="76D34B29BC5BD5B2155FFA7A3451FA12" box="[598,607,1438,1456]" gridcol="10" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42155EFA7A3451FA12" bold="true" box="[599,607,1438,1456]" pageId="15" pageNumber="958">0</emphasis>
</td>
<td id="76D34B29BC5BD5B21579FA7A3477FA12" box="[624,633,1438,1456]" gridcol="11" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421578FA7A3477FA12" bold="true" box="[625,633,1438,1456]" pageId="15" pageNumber="958">1</emphasis>
</td>
<td id="76D34B29BC5BD5B21580FA7A349CFA12" box="[649,658,1438,1456]" gridcol="12" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421583FA7A349CFA12" bold="true" box="[650,658,1438,1456]" pageId="15" pageNumber="958">2</emphasis>
</td>
<td id="76D34B29BC5BD5B215AAFA7A34A2FA12" box="[675,684,1438,1456]" gridcol="13" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215ADFA7A34A2FA12" bold="true" box="[676,684,1438,1456]" pageId="15" pageNumber="958">3</emphasis>
</td>
<td id="76D34B29BC5BD5B215B5FA7A34C8FA12" box="[700,710,1438,1456]" gridcol="14" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215B7FA7A34C8FA12" bold="true" box="[702,710,1438,1456]" pageId="15" pageNumber="958">4</emphasis>
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<td id="76D34B29BC5BD5B215DFFA7A34D1FA12" box="[726,735,1438,1456]" gridcol="15" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215DEFA7A34D1FA12" bold="true" box="[727,735,1438,1456]" pageId="15" pageNumber="958">5</emphasis>
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<td id="76D34B29BC5BD5B215E6FA7A34F7FA12" box="[751,761,1438,1456]" gridcol="16" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4215F8FA7A34F7FA12" bold="true" box="[753,761,1438,1456]" pageId="15" pageNumber="958">6</emphasis>
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<td id="76D34B29BC5BD5B21400FA7A351DFA12" box="[777,787,1438,1456]" gridcol="17" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421403FA7A351CFA12" bold="true" box="[778,786,1438,1456]" pageId="15" pageNumber="958">7</emphasis>
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<td id="76D34B29BC5BD5B2142AFA7A3522FA12" box="[803,812,1438,1456]" gridcol="18" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42142DFA7A3522FA12" bold="true" box="[804,812,1438,1456]" pageId="15" pageNumber="958">8</emphasis>
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<td id="76D34B29BC5BD5B21434FA7A3548FA12" box="[829,838,1438,1456]" gridcol="19" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421437FA7A3548FA12" bold="true" box="[830,838,1438,1456]" pageId="15" pageNumber="958">9</emphasis>
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<td id="76D34B29BC5BD5B2145FFA7A3551FA12" box="[854,863,1438,1456]" gridcol="20" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42145EFA7A3551FA12" bold="true" box="[855,863,1438,1456]" pageId="15" pageNumber="958">0</emphasis>
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<td id="76D34B29BC5BD5B21479FA7A3577FA12" box="[880,889,1438,1456]" gridcol="21" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421478FA7A3577FA12" bold="true" box="[881,889,1438,1456]" pageId="15" pageNumber="958">1</emphasis>
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<td id="76D34B29BC5BD5B21480FA7A359DFA12" box="[905,915,1438,1456]" gridcol="22" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421482FA7A359DFA12" bold="true" box="[907,915,1438,1456]" pageId="15" pageNumber="958">2</emphasis>
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<emphasis id="B946FC05BC5B2A4214ACFA7A35A3FA12" bold="true" box="[933,941,1438,1456]" pageId="15" pageNumber="958">3</emphasis>
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<td id="76D34B29BC5BD5B214B4FA7A35C9FA12" box="[957,967,1438,1456]" gridcol="24" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4214B6FA7A35C9FA12" bold="true" box="[959,967,1438,1456]" pageId="15" pageNumber="958">4</emphasis>
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<td id="76D34B29BC5BD5B214DEFA7A35EEFA12" box="[983,992,1438,1456]" gridcol="25" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4214D1FA7A35EEFA12" bold="true" box="[984,992,1438,1456]" pageId="15" pageNumber="958">5</emphasis>
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<td id="76D34B29BC5BD5B214F9FA7A35F4FA12" box="[1008,1018,1438,1456]" gridcol="26" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4214FBFA7A35F4FA12" bold="true" box="[1010,1018,1438,1456]" pageId="15" pageNumber="958">6</emphasis>
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<td id="76D34B29BC5BD5B21303FA7A321DFA12" box="[1034,1043,1438,1456]" gridcol="27" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421302FA7A321DFA12" bold="true" box="[1035,1043,1438,1456]" pageId="15" pageNumber="958">7</emphasis>
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<emphasis id="B946FC05BC5B2A42132CFA7A3223FA12" bold="true" box="[1061,1069,1438,1456]" pageId="15" pageNumber="958">8</emphasis>
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<td id="76D34B29BC5BD5B21334FA7A3248FA12" box="[1085,1094,1438,1456]" gridcol="29" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421337FA7A3248FA12" bold="true" box="[1086,1094,1438,1456]" pageId="15" pageNumber="958">9</emphasis>
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<td id="76D34B29BC5BD5B2135EFA7A326EFA12" box="[1111,1120,1438,1456]" gridcol="30" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421351FA7A326EFA12" bold="true" box="[1112,1120,1438,1456]" pageId="15" pageNumber="958">0</emphasis>
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<td id="76D34B29BC5BD5B21379FA7A3274FA12" box="[1136,1146,1438,1456]" gridcol="31" gridrow="1" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A42137BFA7A3274FA12" bold="true" box="[1138,1146,1438,1456]" pageId="15" pageNumber="958">1</emphasis>
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</tr>
<tr id="35022255BC5BD5B217A2FA503274FA65" box="[171,1146,1460,1479]" gridrow="2" pageId="15" pageNumber="958">
<th id="76D34B29BC5BD5B217A2FA50372DFA65" box="[171,291,1460,1479]" gridcol="0" gridrow="2" pageId="15" pageNumber="958">
<taxonomicName id="4C325B94BC5B2A4217A2FA50372EFA64" authorityName="Harold" authorityYear="1868" box="[171,288,1460,1478]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="species" species="onitoides">
<emphasis id="B946FC05BC5B2A4217A2FA50372EFA64" bold="true" box="[171,288,1460,1478]" italics="true" pageId="15" pageNumber="958">Bolbites onitoides</emphasis>
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<td id="76D34B29BC5BD5B21680FA50379CFA65" box="[393,402,1460,1479]" gridcol="2" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216ABFA5037A5FA65" box="[418,427,1460,1479]" gridcol="3" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216B5FA5037CBFA65" box="[444,453,1460,1479]" gridcol="4" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216DCFA5037D1FA65" box="[469,479,1460,1479]" gridcol="5" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216E6FA5037F7FA65" box="[495,505,1460,1479]" gridcol="6" gridrow="2" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216E6FA5137F7FA65" box="[495,505,1461,1479]" italics="true" pageId="15" pageNumber="958">–</emphasis>
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<td id="76D34B29BC5BD5B21500FA50341DFA65" box="[521,531,1460,1479]" gridcol="7" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2152AFA503422FA65" box="[547,556,1460,1479]" gridcol="8" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21535FA503448FA65" box="[572,582,1460,1479]" gridcol="9" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2155FFA503451FA65" box="[598,607,1460,1479]" gridcol="10" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21579FA503477FA65" box="[624,633,1460,1479]" gridcol="11" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21580FA50349CFA65" box="[649,658,1460,1479]" gridcol="12" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215AAFA5034A2FA65" box="[675,684,1460,1479]" gridcol="13" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215B5FA5034C8FA65" box="[700,710,1460,1479]" gridcol="14" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215DFFA5034D1FA65" box="[726,735,1460,1479]" gridcol="15" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215E6FA5034F7FA65" box="[751,761,1460,1479]" gridcol="16" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21400FA50351DFA65" box="[777,787,1460,1479]" gridcol="17" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2142AFA503522FA65" box="[803,812,1460,1479]" gridcol="18" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21434FA503548FA65" box="[829,838,1460,1479]" gridcol="19" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2145FFA503551FA65" box="[854,863,1460,1479]" gridcol="20" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21479FA503577FA65" box="[880,889,1460,1479]" gridcol="21" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21480FA50359DFA65" box="[905,915,1460,1479]" gridcol="22" gridrow="2" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421480FA51359DFA65" box="[905,915,1461,1479]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
<td id="76D34B29BC5BD5B214AAFA5035A3FA65" box="[931,941,1460,1479]" gridcol="23" gridrow="2" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4214AAFA5135A3FA65" box="[931,941,1461,1479]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
<td id="76D34B29BC5BD5B214B4FA5035C9FA65" box="[957,967,1460,1479]" gridcol="24" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B214DEFA5035EEFA65" box="[983,992,1460,1479]" gridcol="25" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B214F9FA5035F4FA65" box="[1008,1018,1460,1479]" gridcol="26" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21303FA50321DFA65" box="[1034,1043,1460,1479]" gridcol="27" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2132DFA503223FA65" box="[1060,1069,1460,1479]" gridcol="28" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21334FA503248FA65" box="[1085,1094,1460,1479]" gridcol="29" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2135EFA50326EFA65" box="[1111,1120,1460,1479]" gridcol="30" gridrow="2" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21379FA503274FA65" box="[1136,1146,1460,1479]" gridcol="31" gridrow="2" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421379FA513274FA65" box="[1136,1146,1461,1479]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
</tr>
<tr id="35022255BC5BD5B217A2FA283274FA7C" box="[171,1146,1484,1502]" gridrow="3" pageId="15" pageNumber="958">
<th id="76D34B29BC5BD5B217A2FA28372DFA7C" box="[171,291,1484,1502]" gridcol="0" gridrow="3" pageId="15" pageNumber="958">
<taxonomicName id="4C325B94BC5B2A4217A2FA283718FA7C" baseAuthorityName="Erichson" baseAuthorityYear="1847" box="[171,278,1484,1502]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="species" species="davus">
<emphasis id="B946FC05BC5B2A4217A2FA283718FA7C" bold="true" box="[171,278,1484,1502]" italics="true" pageId="15" pageNumber="958">Oruscatus davus</emphasis>
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</th>
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<td id="76D34B29BC5BD5B21680FA28379CFA7C" box="[393,402,1484,1502]" gridcol="2" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B216ABFA2837A5FA7C" box="[418,427,1484,1502]" gridcol="3" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216B5FA2837CBFA7C" box="[444,453,1484,1502]" gridcol="4" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B216DCFA2837D1FA7C" box="[469,479,1484,1502]" gridcol="5" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216E6FA2837F7FA7C" box="[495,505,1484,1502]" gridcol="6" gridrow="3" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216E6FA2837F7FA7C" box="[495,505,1484,1502]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
<td id="76D34B29BC5BD5B21500FA28341DFA7C" box="[521,531,1484,1502]" gridcol="7" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2152AFA283422FA7C" box="[547,556,1484,1502]" gridcol="8" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21535FA283448FA7C" box="[572,582,1484,1502]" gridcol="9" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2155FFA283451FA7C" box="[598,607,1484,1502]" gridcol="10" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21579FA283477FA7C" box="[624,633,1484,1502]" gridcol="11" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21580FA28349CFA7C" box="[649,658,1484,1502]" gridcol="12" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215AAFA2834A2FA7C" box="[675,684,1484,1502]" gridcol="13" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215B5FA2834C8FA7C" box="[700,710,1484,1502]" gridcol="14" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B215DFFA2834D1FA7C" box="[726,735,1484,1502]" gridcol="15" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215E6FA2834F7FA7C" box="[751,761,1484,1502]" gridcol="16" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21400FA28351DFA7C" box="[777,787,1484,1502]" gridcol="17" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B2142AFA283522FA7C" box="[803,812,1484,1502]" gridcol="18" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21434FA283548FA7C" box="[829,838,1484,1502]" gridcol="19" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2145FFA283551FA7C" box="[854,863,1484,1502]" gridcol="20" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21479FA283577FA7C" box="[880,889,1484,1502]" gridcol="21" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21480FA28359DFA7C" box="[905,915,1484,1502]" gridcol="22" gridrow="3" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421480FA28359DFA7C" box="[905,915,1484,1502]" italics="true" pageId="15" pageNumber="958">–</emphasis>
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<td id="76D34B29BC5BD5B214AAFA2835A3FA7C" box="[931,941,1484,1502]" gridcol="23" gridrow="3" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4214AAFA2835A3FA7C" box="[931,941,1484,1502]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
<td id="76D34B29BC5BD5B214B4FA2835C9FA7C" box="[957,967,1484,1502]" gridcol="24" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B214DEFA2835EEFA7C" box="[983,992,1484,1502]" gridcol="25" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B214F9FA2835F4FA7C" box="[1008,1018,1484,1502]" gridcol="26" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21303FA28321DFA7C" box="[1034,1043,1484,1502]" gridcol="27" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2132DFA283223FA7C" box="[1060,1069,1484,1502]" gridcol="28" gridrow="3" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21334FA283248FA7C" box="[1085,1094,1484,1502]" gridcol="29" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B2135EFA28326EFA7C" box="[1111,1120,1484,1502]" gridcol="30" gridrow="3" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21379FA283274FA7C" box="[1136,1146,1484,1502]" gridcol="31" gridrow="3" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421379FA283274FA7C" box="[1136,1146,1484,1502]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
</tr>
<tr id="35022255BC5BD5B217A2FA073274FA57" box="[171,1146,1507,1525]" gridrow="4" pageId="15" pageNumber="958">
<th id="76D34B29BC5BD5B217A2FA07372DFA57" box="[171,291,1507,1525]" gridcol="0" gridrow="4" pageId="15" pageNumber="958">
<taxonomicName id="4C325B94BC5B2A4217A2FA0736E7FA57" authorityName="Brulle" authorityYear="1837" box="[171,233,1507,1525]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5B2A4217A2FA0736E7FA57" bold="true" box="[171,233,1507,1525]" italics="true" pageId="15" pageNumber="958">Gromphas</emphasis>
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</th>
<td id="76D34B29BC5BD5B2163AFA073776FA57" box="[307,376,1507,1525]" gridcol="1" gridrow="4" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21680FA07379CFA57" box="[393,402,1507,1525]" gridcol="2" gridrow="4" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216ABFA0737A5FA57" box="[418,427,1507,1525]" gridcol="3" gridrow="4" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216B5FA0737CBFA57" box="[444,453,1507,1525]" gridcol="4" gridrow="4" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B216DCFA0737D1FA57" box="[469,479,1507,1525]" gridcol="5" gridrow="4" pageId="15" pageNumber="958">2</td>
<td id="76D34B29BC5BD5B216E6FA0737F7FA57" box="[495,505,1507,1525]" gridcol="6" gridrow="4" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A4216E6FA0737F7FA57" box="[495,505,1507,1525]" italics="true" pageId="15" pageNumber="958">–</emphasis>
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<emphasis id="B946FC05BC5B2A4216E6F9F637F7F986" box="[495,505,1554,1572]" italics="true" pageId="15" pageNumber="958">–</emphasis>
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<emphasis id="B946FC05BC5B2A421400F9F6351DF986" box="[777,787,1554,1572]" italics="true" pageId="15" pageNumber="958">–</emphasis>
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<taxonomicName id="4C325B94BC5B2A4217A2F98B36E7F923" authorityName="Brulle" authorityYear="1837" box="[171,233,1647,1665]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC5B2A4217A2F98B36E7F923" bold="true" box="[171,233,1647,1665]" italics="true" pageId="15" pageNumber="958">Gromphas</emphasis>
</taxonomicName>
</th>
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<emphasis id="B946FC05BC5B2A421500F98B341DF923" box="[521,531,1647,1665]" italics="true" pageId="15" pageNumber="958">–</emphasis>
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<td id="76D34B29BC5BD5B2135EF98B326EF923" box="[1111,1120,1647,1665]" gridcol="30" gridrow="10" pageId="15" pageNumber="958">1</td>
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<tr id="35022255BC5BD5B217A2F9633274F93B" box="[171,1146,1671,1689]" gridrow="11" pageId="15" pageNumber="958" rowspan-1="1" rowspan-10="1" rowspan-11="1" rowspan-12="1" rowspan-13="1" rowspan-14="1" rowspan-15="1" rowspan-16="1" rowspan-17="1" rowspan-18="1" rowspan-19="1" rowspan-2="1" rowspan-20="1" rowspan-21="1" rowspan-22="1" rowspan-23="1" rowspan-24="1" rowspan-25="1" rowspan-26="1" rowspan-27="1" rowspan-28="1" rowspan-29="1" rowspan-3="1" rowspan-30="1" rowspan-31="1" rowspan-4="1" rowspan-5="1" rowspan-6="1" rowspan-7="1" rowspan-8="1" rowspan-9="1">
<th id="76D34B29BC5BD5B217A2F963372DF93B" box="[171,291,1671,1689]" gridcol="0" gridrow="11" pageId="15" pageNumber="958">
<taxonomicName id="4C325B94BC5B2A4217C9F963370CF93B" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[192,258,1671,1689]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="B946FC05BC5B2A4217C9F963370CF93B" bold="true" box="[192,258,1671,1689]" italics="true" pageId="15" pageNumber="958">amazonica</emphasis>
</taxonomicName>
</th>
</tr>
<tr id="35022255BC5BD5B217A2F97A3274F912" box="[171,1146,1694,1712]" gridrow="12" pageId="15" pageNumber="958">
<th id="76D34B29BC5BD5B217A2F97A372DF912" box="[171,291,1694,1712]" gridcol="0" gridrow="12" pageId="15" pageNumber="958">
<taxonomicName id="4C325B94BC5B2A4217A2F97A3713F912" box="[171,285,1694,1712]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="958" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC5B2A4217A2F97A3713F912" bold="true" box="[171,285,1694,1712]" italics="true" pageId="15" pageNumber="958">Gromphas jardim</emphasis>
</taxonomicName>
</th>
<td id="76D34B29BC5BD5B2163AF97A3776F912" box="[307,376,1694,1712]" gridcol="1" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21680F97A379CF912" box="[393,402,1694,1712]" gridcol="2" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216ABF97A37A5F912" box="[418,427,1694,1712]" gridcol="3" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B216B5F97A37CBF912" box="[444,453,1694,1712]" gridcol="4" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B216DCF97A37D1F912" box="[469,479,1694,1712]" gridcol="5" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B216E6F97A37F7F912" box="[495,505,1694,1712]" gridcol="6" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21500F97A341DF912" box="[521,531,1694,1712]" gridcol="7" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2152AF97A3422F912" box="[547,556,1694,1712]" gridcol="8" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21535F97A3448F912" box="[572,582,1694,1712]" gridcol="9" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2155FF97A3451F912" box="[598,607,1694,1712]" gridcol="10" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21579F97A3477F912" box="[624,633,1694,1712]" gridcol="11" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21580F97A349CF912" box="[649,658,1694,1712]" gridcol="12" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B215AAF97A34A2F912" box="[675,684,1694,1712]" gridcol="13" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B215B5F97A34C8F912" box="[700,710,1694,1712]" gridcol="14" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B215DFF97A34D1F912" box="[726,735,1694,1712]" gridcol="15" gridrow="12" pageId="15" pageNumber="958">2</td>
<td id="76D34B29BC5BD5B215E6F97A34F7F912" box="[751,761,1694,1712]" gridcol="16" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21400F97A351DF912" box="[777,787,1694,1712]" gridcol="17" gridrow="12" pageId="15" pageNumber="958">
<emphasis id="B946FC05BC5B2A421400F97A351DF912" box="[777,787,1694,1712]" italics="true" pageId="15" pageNumber="958">–</emphasis>
</td>
<td id="76D34B29BC5BD5B2142AF97A3522F912" box="[803,812,1694,1712]" gridcol="18" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21434F97A3548F912" box="[829,838,1694,1712]" gridcol="19" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B2145FF97A3551F912" box="[854,863,1694,1712]" gridcol="20" gridrow="12" pageId="15" pageNumber="958">0</td>
<td id="76D34B29BC5BD5B21479F97A3577F912" box="[880,889,1694,1712]" gridcol="21" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21480F97A359DF912" box="[905,915,1694,1712]" gridcol="22" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B214AAF97A35A3F912" box="[931,941,1694,1712]" gridcol="23" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B214B4F97A35C9F912" box="[957,967,1694,1712]" gridcol="24" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B214DEF97A35EEF912" box="[983,992,1694,1712]" gridcol="25" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B214F9F97A35F4F912" box="[1008,1018,1694,1712]" gridcol="26" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21303F97A321DF912" box="[1034,1043,1694,1712]" gridcol="27" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B2132DF97A3223F912" box="[1060,1069,1694,1712]" gridcol="28" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21334F97A3248F912" box="[1085,1094,1694,1712]" gridcol="29" gridrow="12" pageId="15" pageNumber="958">2</td>
<td id="76D34B29BC5BD5B2135EF97A326EF912" box="[1111,1120,1694,1712]" gridcol="30" gridrow="12" pageId="15" pageNumber="958">1</td>
<td id="76D34B29BC5BD5B21379F97A3274F912" box="[1136,1146,1694,1712]" gridcol="31" gridrow="12" pageId="15" pageNumber="958">2</td>
</tr>
</table>
</paragraph>
<caption id="DF4D709FBC442A5D17A9FC7F37ABFBE6" ID-DOI="http://doi.org/10.5281/zenodo.3990455" ID-Zenodo-Dep="3990455" httpUri="https://zenodo.org/record/3990455/files/figure.png" pageId="16" pageNumber="959" startId="16.[160,225,923,946]" targetBox="[163,1153,142,892]" targetPageId="16">
<paragraph id="8B8D2017BC442A5D17A9FC7F37ABFBE6" blockId="16.[160,1156,922,1092]" pageId="16" pageNumber="959">
<emphasis id="B946FC05BC442A5D17A9FC7F36F2FC10" bold="true" box="[160,252,923,946]" pageId="16" pageNumber="959">Figure 6.</emphasis>
Phylogeny of
<taxonomicName id="4C325B94BC442A5D1686FC7E37FFFC13" authorityName="Brulle" authorityYear="1837" box="[399,497,922,945]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="959" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC442A5D1686FC7E37FFFC13" box="[399,497,922,945]" italics="true" pageId="16" pageNumber="959">Gromphas</emphasis>
</taxonomicName>
. Above: Cladogram showing the phylogenetic relationship of the species of
<taxonomicName id="4C325B94BC442A5D1619FC53377CFC6C" authorityName="Brulle" authorityYear="1837" box="[272,370,951,974]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="959" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC442A5D1619FC53377CFC6C" box="[272,370,951,974]" italics="true" pageId="16" pageNumber="959">Gromphas</emphasis>
</taxonomicName>
and the evolution of their characters. Black circles indicate uncontroverted synapormorphies; white circles with black margin, controverted synapomorphies; red circles, ambiguous uncontroverted synapomorphies; and white circles with red margin, ambiguous controverted synapomorphies. Below: Support values given for each clade are decay index (above branch) and bootstrap (below branch).
</paragraph>
</caption>
<paragraph id="8B8D2017BC442A5D17A9FB6B371CFB6B" blockId="16.[160,1154,1167,1225]" pageId="16" pageNumber="959">informative morphological characters were found by us, of which 26 are binary and five multistate.</paragraph>
<paragraph id="8B8D2017BC442A5D17A9FACE36EEFAE1" blockId="16.[160,224,1322,1347]" box="[160,224,1322,1347]" pageId="16" pageNumber="959">
<heading id="D0C5977BBC442A5D17A9FACE36EEFAE1" box="[160,224,1322,1347]" fontSize="11" level="3" pageId="16" pageNumber="959" reason="5">
<emphasis id="B946FC05BC442A5D17A9FACE36EEFAE1" bold="true" box="[160,224,1322,1347]" inLineHeading="true" italics="true" pageId="16" pageNumber="959" reason="3">Head</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC442A5D17B4FABB3751FA3B" blockId="16.[189,1156,1375,1711]" pageId="16" pageNumber="959">1. Clypeus, apical margin, form: straight or only slightly upturned (0); clearly bent upward (1).</paragraph>
<paragraph id="8B8D2017BC442A5D17B4FA40328AFA1E" blockId="16.[189,1156,1375,1711]" box="[189,1156,1444,1468]" pageId="16" pageNumber="959">
2. Clypeus, apical margin, shape: with two lobes (0) (
<figureCitation id="13093C92BC442A5D141AFA403572FA1E" box="[787,892,1444,1468]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="16" pageNumber="959">Figure 1F</figureCitation>
); truncate (1) ({fig. 26}).
</paragraph>
<paragraph id="8B8D2017BC442A5D17B4FA23326BFA7D" blockId="16.[189,1156,1375,1711]" box="[189,1125,1479,1503]" pageId="16" pageNumber="959">
3. Clypeus, lateral margin, shape: rounded (0) ({figs 24
<emphasis id="B946FC05BC442A5D1422FA233536FA7D" box="[811,824,1479,1503]" italics="true" pageId="16" pageNumber="959">–</emphasis>
26}); lobate (1) (
<figureCitation id="13093C92BC442A5D14E7FA233257FA7D" box="[1006,1113,1479,1503]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="16" pageNumber="959">Figure 1F</figureCitation>
).
</paragraph>
<paragraph id="8B8D2017BC442A5D17B4FA0D3776F986" blockId="16.[189,1156,1375,1711]" pageId="16" pageNumber="959">
4. Genae, tegument, sculpture adjacent to eyes: granulose (0) (
<figureCitation id="13093C92BC442A5D1483FA0D35FAF9A3" box="[906,1012,1513,1537]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="16" pageNumber="959">Figure 1F</figureCitation>
); smooth (1) ({figs 24, 25}).
</paragraph>
<paragraph id="8B8D2017BC442A5D17B4F9CB3598F9CB" blockId="16.[189,1156,1375,1711]" pageId="16" pageNumber="959">
5. Cephalic projection, shape: wide carina (0); carina with base wider than apex (1) (
<figureCitation id="13093C92BC442A5D17ECF9B5375AF9CB" box="[229,340,1617,1641]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="16" pageNumber="959">Figure 1G</figureCitation>
); horn flattened anteroposteriorly (2) ({figs 29,30}).
</paragraph>
<paragraph id="8B8D2017BC442A5D17B4F9903475F90D" blockId="16.[189,1156,1375,1711]" pageId="16" pageNumber="959">
6. Cephalic projection, carina with base wider than apex, apex, shape: truncate (0) ({fig. 33); emarginate (1) (
<figureCitation id="13093C92BC442A5D16F6F9733460F90D" box="[511,622,1687,1711]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="16" pageNumber="959">Figure 1G</figureCitation>
).
</paragraph>
<caption id="DF4D709FBC452A5C17A9FCFE3272FC43" ID-DOI="http://doi.org/10.5281/zenodo.3990459" ID-Zenodo-Dep="3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="17" pageNumber="960" startId="17.[160,225,794,817]" targetBox="[163,1153,142,763]" targetPageId="17">
<paragraph id="8B8D2017BC452A5C17A9FCFE3272FC43" blockId="17.[160,1156,793,993]" pageId="17" pageNumber="960">
<emphasis id="B946FC05BC452A5C17A9FCFE36F5FC93" bold="true" box="[160,251,794,817]" pageId="17" pageNumber="960">Figure 7.</emphasis>
Metepisterna. (A)
<taxonomicName id="4C325B94BC452A5C16A6FCFD345DFC92" box="[431,595,793,816]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="species" species="jardim" status="sp. nov.">
<emphasis id="B946FC05BC452A5C16A6FCFD345DFC92" box="[431,595,793,816]" italics="true" pageId="17" pageNumber="960">Gromphas jardim</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A275417EBC452A5C1551FCFE34ABFC93" box="[600,677,794,817]" pageId="17" pageNumber="960" rank="species">sp. nov.</taxonomicNameLabel>
:
<emphasis id="B946FC05BC452A5C15A4FCFD3505FC92" box="[685,779,793,816]" italics="true" pageId="17" pageNumber="960">gromphas</emphasis>
is unique among the
<taxonomicName id="4C325B94BC452A5C14E9FCFE3231FC93" box="[992,1087,794,817]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B)
<taxonomicName id="4C325B94BC452A5C160EFCB037D5FCCE" authority=": Both" authorityName="Both" box="[263,475,852,876]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="species" species="davus">
<emphasis id="B946FC05BC452A5C160EFCB037AFFCC9" box="[263,417,852,875]" italics="true" pageId="17" pageNumber="960">Oruscatus davus</emphasis>
: Both
</taxonomicName>
species of
<taxonomicName id="4C325B94BC452A5C154EFCB034ADFCC9" authorityName="Bates" authorityYear="1870" box="[583,675,852,875]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC452A5C154EFCB034ADFCC9" box="[583,675,852,875]" italics="true" pageId="17" pageNumber="960">Oruscatus</emphasis>
</taxonomicName>
have a small but evident MTab (indicated by red arrow). (C)
<taxonomicName id="4C325B94BC452A5C161BFC9537E5FC2B" authority=": The" authorityName="The" box="[274,491,881,905]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="species" species="onitoides">
<emphasis id="B946FC05BC452A5C161BFC9537B6FC2A" box="[274,440,881,904]" italics="true" pageId="17" pageNumber="960">Bolbites onitoides</emphasis>
: The
</taxonomicName>
MTab of
<taxonomicName id="4C325B94BC452A5C155BFC953494FC2A" authorityName="Harold" authorityYear="1868" box="[594,666,881,904]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC452A5C155BFC953494FC2A" box="[594,666,881,904]" italics="true" pageId="17" pageNumber="960">Bolbites</emphasis>
</taxonomicName>
is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D)
<taxonomicName id="4C325B94BC452A5C16FCFC483531FC66" authority="(Harold, 1867)" baseAuthorityName="Harold" baseAuthorityYear="1867" box="[501,831,940,964]" class="Insecta" family="Scarabaeidae" genus="Dichotomius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="species" species="fissus">
<emphasis id="B946FC05BC452A5C16FCFC4834A8FC61" box="[501,678,940,964]" italics="true" pageId="17" pageNumber="960">Dichotomius fissus</emphasis>
(
<bibRefCitation id="EFA35DE6BC452A5C15BFFC493538FC66" author="Harold E" box="[694,822,941,964]" pageId="17" pageNumber="960" pagination="94 - 100" refId="ref15397" refString="Harold E. 1867. Diagnosen neuer Coprophagen. Coleopterol Hefte. 2: 94 - 100." type="journal article" year="1867">Harold, 1867</bibRefCitation>
)
</taxonomicName>
: MTab in a non-Phanaeini. This structure is present also in a series of other
<taxonomicName id="4C325B94BC452A5C1541FC2E349EFC43" box="[584,656,970,993]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="960" phylum="Arthropoda" rank="tribe" tribe="Coprini">Coprini</taxonomicName>
, as well as in at least some Oniticellini and Onitini.
</paragraph>
</caption>
<paragraph id="8B8D2017BC452A5C17A9FBA436F8FBFB" blockId="17.[160,246,1088,1113]" box="[160,246,1088,1113]" pageId="17" pageNumber="960">
<heading id="D0C5977BBC452A5C17A9FBA436F8FBFB" box="[160,246,1088,1113]" fontSize="11" level="3" pageId="17" pageNumber="960" reason="5">
<emphasis id="B946FC05BC452A5C17A9FBA436F8FBFB" bold="true" box="[160,246,1088,1113]" inLineHeading="true" italics="true" pageId="17" pageNumber="960" reason="3">Thorax</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC452A5C17C2FB913798FB0D" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">7. Pronotum, centre, mid-longitudinal line of smooth and glossy tegument: absent (0); present (1).</paragraph>
<paragraph id="8B8D2017BC452A5C17C2FB5E3522FB70" blockId="17.[189,1156,1141,1685]" box="[203,812,1210,1234]" pageId="17" pageNumber="960">8. Pronotum, posterior fossae: present (0); absent (1).</paragraph>
<paragraph id="8B8D2017BC452A5C17C2FB39377BFAB5" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">
9. Pronotum, pronotal prominence: absent (0) (
<figureCitation id="13093C92BC452A5C1417FB39359FFB57" box="[798,913,1245,1269]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="17" pageNumber="960">Figures 1</figureCitation>
,
<figureCitation id="13093C92BC452A5C14A3FB3935B6FB57" box="[938,952,1245,1269]" captionStart="Figure 3" captionStartId="5.[160,225,1589,1612]" captionTargetBox="[163,1153,1108,1558]" captionTargetId="figure@5.[163,1153,1108,1558]" captionTargetPageId="5" captionText="Figure 3. Lateral view of pronotum. (A) Gromphas jardim sp. nov. (B) Gromphas amazonica. Note that the granulation extends much more posteriorly in G. jardim, reaching the pronotal posterior margin, than in G. amazonica, wherein the granulation is absent or rudimentary in posterolateral region after lateral fossa." figureDoi="http://doi.org/10.5281/zenodo.3990447" httpUri="https://zenodo.org/record/3990447/files/figure.png" pageId="17" pageNumber="960">3</figureCitation>
,
<figureCitation id="13093C92BC452A5C14D9FB3935D0FB57" box="[976,990,1245,1269]" captionStart="Figure 4" captionStartId="6.[160,225,837,860]" captionTargetBox="[164,1153,147,799]" captionTargetId="figure@6.[163,1153,142,806]" captionTargetPageId="6" captionText="Figure 4. Male paratype of Gromphas jardim sp. nov. and its labels (braces ‘{’ joining two sides of a same label). We believe that the French zoologist and explorer Alcide d’Orbigny was the collector of this specimen in 1832 (see the text)." figureDoi="http://doi.org/10.5281/zenodo.3990449" httpUri="https://zenodo.org/record/3990449/files/figure.png" pageId="17" pageNumber="960">4</figureCitation>
); present (1) ({figs 1
<emphasis id="B946FC05BC452A5C1631FB1B374BFAB5" box="[312,325,1279,1303]" italics="true" pageId="17" pageNumber="960">–</emphasis>
11}).
</paragraph>
<paragraph id="8B8D2017BC452A5C17B4FAC63499FADD" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">
10. Pronotum, form: irregular surface with single transverse depression or pair of depressions in its anterior region (0) ({figs 1
<emphasis id="B946FC05BC452A5C15DAFAA034EEFAFE" box="[723,736,1348,1372]" italics="true" pageId="17" pageNumber="960">–</emphasis>
11}); entirely convex and without any king of depression (1) (
<figureCitation id="13093C92BC452A5C16FDFA83345CFADD" box="[500,594,1383,1407]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="17" pageNumber="960">Figure 1</figureCitation>
,
<figureCitation id="13093C92BC452A5C1569FA833460FADD" box="[608,622,1383,1407]" captionStart="Figure 3" captionStartId="5.[160,225,1589,1612]" captionTargetBox="[163,1153,1108,1558]" captionTargetId="figure@5.[163,1153,1108,1558]" captionTargetPageId="5" captionText="Figure 3. Lateral view of pronotum. (A) Gromphas jardim sp. nov. (B) Gromphas amazonica. Note that the granulation extends much more posteriorly in G. jardim, reaching the pronotal posterior margin, than in G. amazonica, wherein the granulation is absent or rudimentary in posterolateral region after lateral fossa." figureDoi="http://doi.org/10.5281/zenodo.3990447" httpUri="https://zenodo.org/record/3990447/files/figure.png" pageId="17" pageNumber="960">3</figureCitation>
,
<figureCitation id="13093C92BC452A5C1575FA833484FADD" box="[636,650,1383,1407]" captionStart="Figure 4" captionStartId="6.[160,225,837,860]" captionTargetBox="[164,1153,147,799]" captionTargetId="figure@6.[163,1153,142,806]" captionTargetPageId="6" captionText="Figure 4. Male paratype of Gromphas jardim sp. nov. and its labels (braces ‘{’ joining two sides of a same label). We believe that the French zoologist and explorer Alcide d’Orbigny was the collector of this specimen in 1832 (see the text)." figureDoi="http://doi.org/10.5281/zenodo.3990449" httpUri="https://zenodo.org/record/3990449/files/figure.png" pageId="17" pageNumber="960">4</figureCitation>
).
</paragraph>
<paragraph id="8B8D2017BC452A5C17B4FA6E3238FA67" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">
11. Mesepimeron, metepisternum and outer sides of metasternum, pilosity: with long and abundant setae (0) (
<figureCitation id="13093C92BC452A5C150FFA493489FA67" box="[518,647,1453,1477]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="17" pageNumber="960">Figure 7B,C</figureCitation>
); entirely glabrous (1) (
<figureCitation id="13093C92BC452A5C1498FA4935F3FA67" box="[913,1021,1453,1477]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="17" pageNumber="960">Figure 1B</figureCitation>
,
<figureCitation id="13093C92BC452A5C1302FA493227FA67" box="[1035,1065,1453,1477]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="17" pageNumber="960">7A</figureCitation>
).
</paragraph>
<paragraph id="8B8D2017BC452A5C17B4FA2B3763F9A8" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">
12. Metepisternum, metepisternal tab: present (0) (
<figureCitation id="13093C92BC452A5C1459FA2B35D1FA45" box="[848,991,1487,1511]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="17" pageNumber="960">Figure 7B,C</figureCitation>
); absent (1) (
<figureCitation id="13093C92BC452A5C17FAFA16376EF9A8" box="[243,352,1522,1546]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="17" pageNumber="960">Figure 7A</figureCitation>
).
</paragraph>
<paragraph id="8B8D2017BC452A5C17B4F9F1342CF9ED" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">
13. Metasternum, sides of anteromedian angle, tegument, sculpture: smooth (0); granulose (1) ({figs 49
<emphasis id="B946FC05BC452A5C16EAF9D337FEF9ED" box="[483,496,1591,1615]" italics="true" pageId="17" pageNumber="960">–</emphasis>
52}).
</paragraph>
<paragraph id="8B8D2017BC452A5C17B4F9BE3555F937" blockId="17.[189,1156,1141,1685]" pageId="17" pageNumber="960">14. Metasternum, apex and region in front of anteromedian angle, pilosity: with evident pilosity, short or long (0); entirely glabrous (1).</paragraph>
<caption id="DF4D709FBC462A5F17A9FC253728FB92" ID-DOI="http://doi.org/10.5281/zenodo.3990461" ID-Zenodo-Dep="3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="18" pageNumber="961" startId="18.[160,225,961,984]" targetBox="[163,1153,142,930]" targetPageId="18">
<paragraph id="8B8D2017BC462A5F17A9FC253728FB92" blockId="18.[160,1155,960,1072]" pageId="18" pageNumber="961">
<emphasis id="B946FC05BC462A5F17A9FC2536F4FC7A" bold="true" box="[160,250,961,984]" pageId="18" pageNumber="961">Figure 8.</emphasis>
Lateral view of male protibia, ornamentation of ventral carina. (A)
<taxonomicName id="4C325B94BC462A5F14BCFC243265FC75" authorityName="Harold" authorityYear="1869" box="[949,1131,960,983]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="961" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC462A5F14BCFC243265FC75" box="[949,1131,960,983]" italics="true" pageId="18" pageNumber="961">Gromphas inermis</emphasis>
</taxonomicName>
: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of
<taxonomicName id="4C325B94BC462A5F14E7FC39325AFC56" authorityName="Brulle" authorityYear="1837" box="[1006,1108,989,1012]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="961" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC462A5F14E7FC39325AFC56" box="[1006,1108,989,1012]" italics="true" pageId="18" pageNumber="961">Gromphas</emphasis>
</taxonomicName>
. (B)
<taxonomicName id="4C325B94BC462A5F17A9FC1F3749FBB0" box="[160,327,1019,1042]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="961" phylum="Arthropoda" rank="species" species="davus">
<emphasis id="B946FC05BC462A5F17A9FC1F3749FBB0" box="[160,327,1019,1042]" italics="true" pageId="18" pageNumber="961">Oruscatus davus</emphasis>
</taxonomicName>
: a single large tooth. (C)
<taxonomicName id="4C325B94BC462A5F156AFC1F3518FBB0" box="[611,790,1019,1042]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="961" phylum="Arthropoda" rank="species" species="onitoides">
<emphasis id="B946FC05BC462A5F156AFC1F3518FBB0" box="[611,790,1019,1042]" italics="true" pageId="18" pageNumber="961">Bolbites onitoides</emphasis>
</taxonomicName>
: a large central tooth with minor lateral teeth.
</paragraph>
</caption>
<paragraph id="8B8D2017BC462A5F17B4FB673446FB1C" blockId="18.[189,1155,1155,1214]" pageId="18" pageNumber="961">15. Metasternum, centre, tegument, sculpture: irregular (0); densely punctate (1); with fine and sparse punctation (2).</paragraph>
<paragraph id="8B8D2017BC462A5F17A9FB11370FFAAC" blockId="18.[160,257,1269,1294]" box="[160,257,1269,1294]" pageId="18" pageNumber="961">
<heading id="D0C5977BBC462A5F17A9FB11370FFAAC" box="[160,257,1269,1294]" fontSize="11" level="3" pageId="18" pageNumber="961" reason="5">
<emphasis id="B946FC05BC462A5F17A9FB11370FFAAC" bold="true" box="[160,257,1269,1294]" inLineHeading="true" italics="true" pageId="18" pageNumber="961" reason="3">Protibia</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC462A5F17B4FACE3215FAC7" blockId="18.[189,1156,1322,1727]" pageId="18" pageNumber="961">16. Ventral longitudinal carina: ornamented with tubercles and/or teeth in males and simple in female (0) ({figs 20d, 23b}); simple in both sexes (1) ({fig. 22}).</paragraph>
<paragraph id="8B8D2017BC462A5F17B4FA8B3570FA6F" blockId="18.[189,1156,1322,1727]" pageId="18" pageNumber="961">
17. Ventral longitudinal carina, ornamentation, form: large tooth and small irregular teeth on the middle (0) (
<figureCitation id="13093C92BC462A5F1501FA76347BFA08" box="[520,629,1426,1450]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="18" pageNumber="961">Figure 8C</figureCitation>
); a single large tooth (1) (
<figureCitation id="13093C92BC462A5F14A8FA763203FA08" box="[929,1037,1426,1450]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="18" pageNumber="961">Figure 8B</figureCitation>
); a row of tubercles on the basal half (2) (
<figureCitation id="13093C92BC462A5F1558FA5134B1FA6F" box="[593,703,1461,1485]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="18" pageNumber="961">Figure 8A</figureCitation>
, {figs 20d, 23b}).
</paragraph>
<paragraph id="8B8D2017BC462A5F17B4FA333457F9B0" blockId="18.[189,1156,1322,1727]" pageId="18" pageNumber="961">18. Protibial spur (articulated spur), apex, shape: only slightly curved (0) ({fig. 22}); strongly expanded (1) ({fig. 23}).</paragraph>
<paragraph id="8B8D2017BC462A5F17B4F9F9327DF9F5" blockId="18.[189,1156,1322,1727]" pageId="18" pageNumber="961">19. Apical tuft of setae, sexual dimorphism: denser and longer in males than in females (0) ({fig. 23}); with same density and length in both sexes (1) ({fig. 22}).</paragraph>
<paragraph id="8B8D2017BC462A5F17B4F98634AFF93F" blockId="18.[189,1156,1322,1727]" pageId="18" pageNumber="961">20. Overall shape, sexual dimorphism: narrower in males than in females (0) ({fig. 23}); very broad in both sexes (1) ({fig. 22}).</paragraph>
<paragraph id="8B8D2017BC462A5F17B4F94335A3F91D" blockId="18.[189,1156,1322,1727]" box="[189,941,1703,1727]" pageId="18" pageNumber="961">21. Inner apical angle, tubercle: absent (0); present (1) ({fig. 23b}).</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FF6A3750FF4E" blockId="19.[189,1156,142,305]" pageId="19" pageNumber="962">
22. Inner apical angle, tubercle, form: tiny and almost imperceptible (0) ({fig. 23b}); developed into a strong spur clearly visible to the naked eye (1) (
<figureCitation id="13093C92BC472A5E14F0FF553273FF6B" box="[1017,1149,177,201]" captionStart="Figure 2" captionStartId="5.[160,225,933,956]" captionTargetBox="[178,1148,161,902]" captionTargetId="figure@5.[163,1153,142,902]" captionTargetPageId="5" captionText="Figure 2. Protibia of Gromphas jardim sp. nov. (A, B) Male protibia: (A) Ventral view, (B) Dorsal view. Red arrow indicates apical tubercle modified in a tapered spur. (C) Female protibia. Note that the apical tuft of setae is much denser and longer in male than in female." figureDoi="http://doi.org/10.5281/zenodo.3990445" httpUri="https://zenodo.org/record/3990445/files/figure.png" pageId="19" pageNumber="962">Figure 2A,B</figureCitation>
; {fig. 21b}).
</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FF123521FE93" blockId="19.[189,1156,142,305]" pageId="19" pageNumber="962">23. Apical protarsomere, apex: tapered and only slightly elongate at apex (0) ({figs 22, 46}); with a long spiniform projection (1) ({fig. 45}).</paragraph>
<paragraph id="8B8D2017BC472A5E17A9FE8B36E9FE2A" blockId="19.[160,231,367,392]" box="[160,231,367,392]" pageId="19" pageNumber="962">
<heading id="D0C5977BBC472A5E17A9FE8B36E9FE2A" box="[160,231,367,392]" fontSize="11" level="3" pageId="19" pageNumber="962" reason="5">
<emphasis id="B946FC05BC472A5E17A9FE8B36E9FE2A" bold="true" box="[160,231,367,392]" inLineHeading="true" italics="true" pageId="19" pageNumber="962" reason="3">Elytra</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FE403514FE1E" blockId="19.[189,1155,420,825]" box="[189,794,420,444]" pageId="19" pageNumber="962">24. Elytral striae, basal fossae: present (0); absent (1).</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FE223748FD86" blockId="19.[189,1155,420,825]" pageId="19" pageNumber="962">25. Elytral striae, sculpture: very fine and simple, not carinulate (0); especially those more internal, carinulate from base to at least mid-length of elytra (1) ({figs 37, 38}).</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FDCA379CFDCB" blockId="19.[189,1155,420,825]" pageId="19" pageNumber="962">26. Sutural margin, tegument, sculpture: densely punctate (0); with fine and sparse punctation (1).</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FD90322EFD73" blockId="19.[189,1155,420,825]" pageId="19" pageNumber="962">
27. Sutural margin, tegument: sheen and punctation limited to sutural margin (0) ({figs 1
<emphasis id="B946FC05BC472A5E163EFD72374AFD0C" box="[311,324,662,686]" italics="true" pageId="19" pageNumber="962">–</emphasis>
11}); in basal third or basal half, sheen and punctuation of sutural margin extend onto first or second interstriae (1) (
<figureCitation id="13093C92BC472A5E15DBFD5D3553FD73" box="[722,861,697,721]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="19" pageNumber="962">Figures 1A,E</figureCitation>
,
<figureCitation id="13093C92BC472A5E1465FD5D3574FD73" box="[876,890,697,721]" captionStart="Figure 4" captionStartId="6.[160,225,837,860]" captionTargetBox="[164,1153,147,799]" captionTargetId="figure@6.[163,1153,142,806]" captionTargetPageId="6" captionText="Figure 4. Male paratype of Gromphas jardim sp. nov. and its labels (braces ‘{’ joining two sides of a same label). We believe that the French zoologist and explorer Alcide d’Orbigny was the collector of this specimen in 1832 (see the text)." figureDoi="http://doi.org/10.5281/zenodo.3990449" httpUri="https://zenodo.org/record/3990449/files/figure.png" pageId="19" pageNumber="962">4</figureCitation>
); {figs 13
<emphasis id="B946FC05BC472A5E14EBFD5D35E1FD73" box="[994,1007,697,721]" italics="true" pageId="19" pageNumber="962">–</emphasis>
15}).
</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FD38343EFC9B" blockId="19.[189,1155,420,825]" pageId="19" pageNumber="962">28. Epipleura, shape: entirely horizontal and narrow or curved only at base (0); strongly curved and wide from base of elytron to metacoxa, remainder length horizontal and narrowed (1).</paragraph>
<paragraph id="8B8D2017BC472A5E17A9FC93371BFC32" blockId="19.[160,277,887,912]" box="[160,277,887,912]" pageId="19" pageNumber="962">
<heading id="D0C5977BBC472A5E17A9FC93371BFC32" box="[160,277,887,912]" fontSize="11" level="3" pageId="19" pageNumber="962" reason="5">
<emphasis id="B946FC05BC472A5E17A9FC93371BFC32" bold="true" box="[160,277,887,912]" inLineHeading="true" italics="true" pageId="19" pageNumber="962" reason="3">Abdomen</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FC4835EAFC45" blockId="19.[189,1155,940,999]" pageId="19" pageNumber="962">29. Pygidium, basal margin: complete (0) ({fig. 35}); present, but usually interrupted in the middle by the groove of propygidium (1); absent (2) ({fig. 36}).</paragraph>
<paragraph id="8B8D2017BC472A5E17A9FBC13716FB9C" blockId="19.[160,280,1061,1086]" box="[160,280,1061,1086]" pageId="19" pageNumber="962">
<heading id="D0C5977BBC472A5E17A9FBC13716FB9C" box="[160,280,1061,1086]" fontSize="11" level="3" pageId="19" pageNumber="962" reason="5">
<emphasis id="B946FC05BC472A5E17A9FBC13716FB9C" bold="true" box="[160,280,1061,1086]" inLineHeading="true" italics="true" pageId="19" pageNumber="962" reason="3">Aedeagus</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FBBE3406FB36" blockId="19.[189,1156,1114,1276]" pageId="19" pageNumber="962">
30. Genital capsule, phallobase, ventrobasal margin, shape: entirely (0); with median incision (1) ({figs 53
<emphasis id="B946FC05BC472A5E16C3FB9837D9FB36" box="[458,471,1148,1172]" italics="true" pageId="19" pageNumber="962">–</emphasis>
56}).
</paragraph>
<paragraph id="8B8D2017BC472A5E17B4FB7B3758FB5E" blockId="19.[189,1156,1114,1276]" pageId="19" pageNumber="962">
31. Internal sac, medial sclerite, form: wider and strongly curved (0) ({figs 57
<emphasis id="B946FC05BC472A5E134FFB7B325DFB15" box="[1094,1107,1183,1207]" italics="true" pageId="19" pageNumber="962">–</emphasis>
58}); wider and only slightly curved (1) ({figs 59
<emphasis id="B946FC05BC472A5E15C2FB2534D6FB7B" box="[715,728,1217,1241]" italics="true" pageId="19" pageNumber="962">–</emphasis>
60}); very fine and strongly curved (2) ({fig. 61}).
</paragraph>
<paragraph id="8B8D2017BC472A5E17A9FADE3402FAF1" blockId="19.[160,524,1338,1363]" box="[160,524,1338,1363]" pageId="19" pageNumber="962">
<heading id="D0C5977BBC472A5E17A9FADE3402FAF1" box="[160,524,1338,1363]" fontSize="11" level="3" pageId="19" pageNumber="962" reason="5">
<emphasis id="B946FC05BC472A5E17A9FADE3402FAF1" bold="true" box="[160,524,1338,1363]" inLineHeading="true" pageId="19" pageNumber="962" reason="3">
<emphasis id="B946FC05BC472A5E17A9FADE378FFAF1" bold="true" box="[160,385,1338,1363]" italics="true" pageId="19" pageNumber="962">The monophyly of</emphasis>
<taxonomicName id="4C325B94BC472A5E1683FADE3402FAF1" authorityName="Brulle" authorityYear="1837" box="[394,524,1338,1363]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="962" phylum="Arthropoda" rank="genus">Gromphas</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC472A5E17A9FA8B37E2F91D" blockId="19.[160,1156,1391,1727]" pageId="19" pageNumber="962">
In our analysis, the monophyly of
<taxonomicName id="4C325B94BC472A5E153AFA8B34AAFA25" authorityName="Brulle" authorityYear="1837" box="[563,676,1391,1415]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="962" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC472A5E153AFA8B34AAFA25" box="[563,676,1391,1415]" italics="true" pageId="19" pageNumber="962">Gromphas</emphasis>
</taxonomicName>
was strongly supported by eight uncontroverted synapomorphies: outer sides of pterothorax (mesepisternum, metepisternum and sides of metasternum) entirely glabrous (char. 11
<emphasis id="B946FC05BC472A5E140CFA50351CFA6E" box="[773,786,1460,1484]" italics="true" pageId="19" pageNumber="962">–</emphasis>
1;
<figureCitation id="13093C92BC472A5E1426FA5135ABFA6F" box="[815,933,1461,1485]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="19" pageNumber="962">Figures 1B</figureCitation>
,
<figureCitation id="13093C92BC472A5E14BDFA5135DCFA6F" box="[948,978,1461,1485]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="19" pageNumber="962">7A</figureCitation>
), metepisternal tab absent (12-1;
<figureCitation id="13093C92BC472A5E1662FA3337D7FA4D" box="[363,473,1495,1519]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="19" pageNumber="962">Figure 7A</figureCitation>
), sides of anteromedian angle of metasternum with granulose tegument (13-1); {figs 49
<emphasis id="B946FC05BC472A5E16E1FA1E37FBF9B0" box="[488,501,1530,1554]" italics="true" pageId="19" pageNumber="962">–</emphasis>
52}), centre of metasternum densely punctate (15-1), inner apical angle of male protibiae with a tiny tubercle (21-1; {fig. 23b}), elytral striae without basal fossae (24-1), epipleura strongly curved and wide from base of elytron to metacoxa (28-1), and ventrobasal margin of phallobase with median incision (30-1; {figs 53
<emphasis id="B946FC05BC472A5E134FF986325DF9D8" box="[1094,1107,1634,1658]" italics="true" pageId="19" pageNumber="962">–</emphasis>
56}). The high support values (bootstrap 99, decay index 7) also give us great confidence in the monophyly of
<taxonomicName id="4C325B94BC472A5E167FF94337E9F91D" authorityName="Brulle" authorityYear="1837" box="[374,487,1703,1727]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="962" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC472A5E167FF94337E9F91D" box="[374,487,1703,1727]" italics="true" pageId="19" pageNumber="962">Gromphas</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8B8D2017BC402A5917C9FF6A34C4FBAB" blockId="20.[160,1156,142,1483]" pageId="20" pageNumber="963">
<bibRefCitation id="EFA35DE6BC402A5917C9FF6A377BFF04" author="Edmonds WD" box="[192,373,142,166]" pageId="20" pageNumber="963" pagination="731 - 874" refId="ref14833" refString="Edmonds WD. 1972. Comparative skeletal morphology, systematics and evolution of the Phanaeine dung beetles (Coleoptera: Scarabaeidae). Univ Kansas Sci Bull. 49: 731 - 874." type="journal article" year="1972">Edmonds (1972)</bibRefCitation>
and
<bibRefCitation id="EFA35DE6BC402A5916A0FF6A3476FF04" author="Philips TK &amp; Edmonds WD &amp; Scholtz CH" box="[425,632,142,166]" pageId="20" pageNumber="963" pagination="43 - 63" refId="ref15926" refString="Philips TK, Edmonds WD, Scholtz CH. 2004. A phylogenetic analysis of the New World tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae): hypothesis on relationship and origins. Insect Syst Evol. 35: 43 - 63." type="journal article" year="2004">Philips et al. (2004)</bibRefCitation>
stated that the metepisternal tab was absent in
<taxonomicName id="4C325B94BC402A5917A9FF55371FFF6B" authorityName="Brulle" authorityYear="1837" box="[160,273,177,201]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917A9FF55371FFF6B" box="[160,273,177,201]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC402A591643FF5537B8FF6B" authorityName="Bates" authorityYear="1870" box="[330,438,177,201]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A591643FF5537B8FF6B" box="[330,438,177,201]" italics="true" pageId="20" pageNumber="963">Oruscatus</emphasis>
</taxonomicName>
; the first author considered the presence the metepisternal tab as one of the defining characteristics of the phanaeines when these two genera are excluded. However, as we observed in
<bibRefCitation id="EFA35DE6BC402A59155EFF1235C9FEAC" author="Cupello M &amp; Vaz-de-Mello FZ" box="[599,967,246,270]" pageId="20" pageNumber="963" pagination="439 - 482" refId="ref14473" 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>
, the two species of
<taxonomicName id="4C325B94BC402A5917B6FEFD3725FE93" authorityName="Bates" authorityYear="1870" box="[191,299,281,305]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917B6FEFD3725FE93" box="[191,299,281,305]" italics="true" pageId="20" pageNumber="963">Oruscatus</emphasis>
</taxonomicName>
have, in fact, a short tab in their metepisterna covering the margin of the elytral epipleura, which has an evident depression at this point to receive the tab, in the same way as in Phanaeina (
<figureCitation id="13093C92BC402A5916C4FEBA3438FED4" box="[461,566,350,374]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="20" pageNumber="963">Figure 7B</figureCitation>
). We observed this same short metepisternal tab and the respective epipleural depression in a series of other dung beetle genera, including species of
<taxonomicName id="4C325B94BC402A59162DFE40342FFE1E" authority="Geoffroy, 1762" authorityName="Geoffroy" authorityYear="1762" box="[292,545,419,444]" class="Insecta" family="Scarabaeidae" genus="Copris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59162DFE403769FE1E" box="[292,359,420,444]" italics="true" pageId="20" pageNumber="963">Copris</emphasis>
<bibRefCitation id="EFA35DE6BC402A59167FFE40342FFE1E" author="Geoffroy EL" box="[374,545,419,444]" pageId="20" pageNumber="963" refId="ref15141" refString="Geoffroy EL. 1762. Histoire abregee des Insectes qui se trouvent aux environs de Paris; Dans laquelle ces Animaux sont ranges suivant un ordre methodique. Tome Premier. Paris: Durant." type="book" year="1762">Geoffroy, 1762</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C325B94BC402A59153CFE40355DFE1E" authority="Hope, 1838" authorityName="Hope" authorityYear="1838" box="[565,851,420,444]" class="Insecta" family="Scarabaeidae" genus="Dichotomius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59153CFE4034B0FE1E" box="[565,702,420,444]" italics="true" pageId="20" pageNumber="963">Dichotomius</emphasis>
<bibRefCitation id="EFA35DE6BC402A5915C5FE40355DFE1E" author="Hope FW" box="[716,851,420,444]" pageId="20" pageNumber="963" pagination="312 - 326" refId="ref15376" refString="Hope FW. 1838. Observations on the Lamellicorns of Olivier. Entomol Mag. 5: 312 - 326." type="journal article" year="1838">Hope, 1838</bibRefCitation>
</taxonomicName>
(
<figureCitation id="13093C92BC402A591460FE4035D3FE1E" box="[873,989,420,444]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="20" pageNumber="963">Figure 7D</figureCitation>
),
<taxonomicName id="4C325B94BC402A5914F0FE40376DFE7C" authority="Burmeister, 1846" authorityName="Burmeister" authorityYear="1846" class="Insecta" family="Scarabaeidae" genus="Chalcocopris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5914F0FE40328DFE1E" box="[1017,1155,420,444]" italics="true" pageId="20" pageNumber="963">Chalcocopris</emphasis>
<bibRefCitation id="EFA35DE6BC402A5917A9FE22376DFE7C" author="Burmeister H" box="[160,355,454,478]" pageId="20" pageNumber="963" pagination="1838 - 1846" refId="ref14272" refString="Burmeister H, editor. 1846. Genera quaedam insectorum iconibus illustravit et descripsit: Copris, Heliocopris, Pelidnota, Trichius, Clastocnemis, Trigonopeltastes. In: 1838 - 1846. Genera quaedam insectorum iconibus illustravit et descripsit. Volumen I. Continet tabulas XL. Berlin [&quot; Berolini &quot;]: Privately published, unpaginated." type="book chapter" year="1846">Burmeister, 1846</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C325B94BC402A59167CFE2234C9FE7C" authority="Burmeister, 1846" authorityName="Burmeister" authorityYear="1846" box="[373,711,454,478]" class="Insecta" family="Scarabaeidae" genus="Homocopris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59167CFE2237F9FE7C" box="[373,503,454,478]" italics="true" pageId="20" pageNumber="963">Homocopris</emphasis>
<bibRefCitation id="EFA35DE6BC402A59150DFE2234C9FE7C" author="Burmeister H" box="[516,711,454,478]" pageId="20" pageNumber="963" pagination="1838 - 1846" refId="ref14272" refString="Burmeister H, editor. 1846. Genera quaedam insectorum iconibus illustravit et descripsit: Copris, Heliocopris, Pelidnota, Trichius, Clastocnemis, Trigonopeltastes. In: 1838 - 1846. Genera quaedam insectorum iconibus illustravit et descripsit. Volumen I. Continet tabulas XL. Berlin [&quot; Berolini &quot;]: Privately published, unpaginated." type="book chapter" year="1846">Burmeister, 1846</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C325B94BC402A5915D0FE2235FCFE7C" authority="Erichson, 1847" authorityName="Erichson" authorityYear="1847" box="[729,1010,454,478]" class="Insecta" family="Scarabaeidae" genus="Ontherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5915D0FE223532FE7C" box="[729,828,454,478]" italics="true" pageId="20" pageNumber="963">Ontherus</emphasis>
<bibRefCitation id="EFA35DE6BC402A591440FE2235FCFE7C" author="Erichson WF" box="[841,1010,454,478]" pageId="20" pageNumber="963" pagination="552 - 950" refId="ref14975" refString="Erichson WF, editor. 1847. Scarabaeides. In: Naturgeschichte der Insecten Deutschlands. Berlin: Nicolaische Buchhandlung; p. 552 - 950." type="book chapter" year="1847">Erichson, 1847</bibRefCitation>
</taxonomicName>
,
<taxonomicName id="4C325B94BC402A59130DFE22374DFDA3" authority="Erichson, 1847" authorityName="Erichson" authorityYear="1847" class="Insecta" family="Scarabaeidae" genus="Canthidium" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59130DFE22328DFE7C" box="[1028,1155,454,478]" italics="true" pageId="20" pageNumber="963">Canthidium</emphasis>
<bibRefCitation id="EFA35DE6BC402A5917A9FE0D374DFDA3" author="Erichson WF" box="[160,323,489,513]" pageId="20" pageNumber="963" pagination="552 - 950" refId="ref14975" refString="Erichson WF, editor. 1847. Scarabaeides. In: Naturgeschichte der Insecten Deutschlands. Berlin: Nicolaische Buchhandlung; p. 552 - 950." type="book chapter" year="1847">Erichson, 1847</bibRefCitation>
</taxonomicName>
, and
<taxonomicName id="4C325B94BC402A59168BFE0D34F2FDA3" authority="Pereira &amp; Martinez, 1960" authorityName="Pereira &amp; Martinez" authorityYear="1960" box="[386,764,489,513]" class="Insecta" family="Scarabaeidae" genus="Isocopris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59168BFE0D37EEFDA3" box="[386,480,489,513]" italics="true" pageId="20" pageNumber="963">Isocopris</emphasis>
<bibRefCitation id="EFA35DE6BC402A5916EEFE0D34F2FDA3" author="Pereira FS &amp; Martinez A" box="[487,764,489,513]" pageId="20" pageNumber="963" pagination="37 - 55" refId="ref15839" refString="Pereira FS, Martinez A. 1960. Notas escarabeidologicas - II. Rev Bras Entomol. 9: 37 - 55." type="journal article" year="1960">Pereira &amp; Martínez, 1960</bibRefCitation>
</taxonomicName>
, in the tribe
<taxonomicName id="4C325B94BC402A591485FE0D35EEFDA3" box="[908,992,489,513]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="tribe" tribe="Coprini">Coprini</taxonomicName>
,
<taxonomicName id="4C325B94BC402A5914E3FE0D328DFDA3" authorityName="Orbigny" authorityYear="1915" box="[1002,1155,489,513]" class="Insecta" family="Scarabaeidae" genus="Helictopleurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5914E3FE0D328DFDA3" box="[1002,1155,489,513]" italics="true" pageId="20" pageNumber="963">Helictopleurus</emphasis>
</taxonomicName>
d
<emphasis id="B946FC05BC402A5917A6FDEF36BAFD81" box="[175,180,523,547]" italics="true" pageId="20" pageNumber="963">’</emphasis>
<bibRefCitation id="EFA35DE6BC402A5917BCFDE8375BFD86" author="Orbigny H" box="[181,341,524,548]" pageId="20" pageNumber="963" pagination="402 - 434" refId="ref14583" refString="d ' Orbigny H. 1915. Synopsis d ' un genre nouveau d ' Oniticellides [Scarabaeidae Coprini] special a Madagascar. Ann Soc Entomol Fr. 84: 402 - 434." type="journal article" year="1915">Orbigny, 1915</bibRefCitation>
,
<taxonomicName id="4C325B94BC402A59166AFDE83468FD86" authority="Reitter, 1893" authorityName="Reitter" authorityYear="1893" box="[355,614,524,548]" class="Insecta" family="Scarabaeidae" genus="Liatongus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59166AFDE837C3FD86" box="[355,461,524,548]" italics="true" pageId="20" pageNumber="963">Liatongus</emphasis>
<bibRefCitation id="EFA35DE6BC402A5916DEFDE83468FD86" author="Reitter E" box="[471,614,524,548]" pageId="20" pageNumber="963" pagination="141 - 262" refId="ref16073" refString="Reitter E. 1893. Bestimmungs-Tabelle der Lucaniden und coprophagen Lamellicornen des pal ӕarctischen Faunengebietes. Verh Naturforsch Ver Brunn. 30: 141 - 262." type="journal article" year="1893">Reitter, 1893</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C325B94BC402A5915AAFDE835D4FD86" authority="Janssens, 1953" authorityName="Janssens" authorityYear="1953" box="[675,986,524,548]" class="Insecta" family="Scarabaeidae" genus="Euoniticellus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5915AAFDE83524FD86" box="[675,810,524,548]" italics="true" pageId="20" pageNumber="963">Euoniticellus</emphasis>
<bibRefCitation id="EFA35DE6BC402A59143BFDE835D4FD86" author="Janssens A" box="[818,986,524,548]" pageId="20" pageNumber="963" pagination="1 - 118" refId="ref15494" refString="Janssens A. 1953. Oniticellini (Coleoptera Lamellicornia). In Exploration du Parc National de l ' Upemba, Mission G. F. de Witte. Inst Parc Nat Congo Belg. 11: 1 - 118." type="journal article" year="1953">Janssens, 1953</bibRefCitation>
</taxonomicName>
, in Oniticellini, and
<taxonomicName id="4C325B94BC402A5917DDFDCA37B0FDE4" authority="Mulsant, 1842" authorityName="Mulsant" authorityYear="1842" box="[212,446,558,582]" class="Insecta" family="Scarabaeidae" genus="Bubas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917DDFDCA3718FDE4" box="[212,278,558,582]" italics="true" pageId="20" pageNumber="963">Bubas</emphasis>
<bibRefCitation id="EFA35DE6BC402A591616FDCA37B0FDE4" author="Mulsant ME" box="[287,446,558,582]" pageId="20" pageNumber="963" refId="ref15676" refString="Mulsant ME. 1842. Histoire naturelle des Coleopteres de France. Lamellicornes. Paris: Maison." type="book" year="1842">Mulsant, 1842</bibRefCitation>
</taxonomicName>
, in Onitini; in
<taxonomicName id="4C325B94BC402A59156DFDCA34DAFDE4" authorityName="Burmeister" authorityYear="1873" box="[612,724,558,582]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="tribe" tribe="Eucraniini">Eucraniini</taxonomicName>
, the putative sister tribe of
<taxonomicName id="4C325B94BC402A59131CFDCA328DFDE4" box="[1045,1155,558,582]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
(
<bibRefCitation id="EFA35DE6BC402A5917AFFDB53786FDCB" author="Philips TK &amp; Edmonds WD &amp; Scholtz CH" box="[166,392,593,617]" pageId="20" pageNumber="963" pagination="43 - 63" refId="ref15926" refString="Philips TK, Edmonds WD, Scholtz CH. 2004. A phylogenetic analysis of the New World tribe Phanaeini (Coleoptera: Scarabaeidae: Scarabaeinae): hypothesis on relationship and origins. Insect Syst Evol. 35: 43 - 63." type="journal article" year="2004">Philips et al. 2004</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC402A591697FDB534DEFDCB" author="Tarasov S &amp; Genier F" box="[414,720,593,617]" pageId="20" pageNumber="963" pagination="1 - 86" refId="ref16197" refString="Tarasov S, Genier F. 2015. Innovative bayesian and parsimony of dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) enhanced by ontology-based partitioning of morphological characters. Plos One. 10: 1 - 86." type="journal article" year="2015">Tarasov and Génier 2015</bibRefCitation>
), we have not seen this structure. Nonetheless, the metepisternal tab seen in Phanaeina is unique among the dung beetles by its great development, being much more curved and longer than in any other group observed by us (
<figureCitation id="13093C92BC402A591652FD5D37C6FD73" box="[347,456,697,721]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="20" pageNumber="963">Figure 7C</figureCitation>
). Therefore, we consider that this unique form, rather than its mere presence, is one of the defining synapomorphies of Phanaeina. In its turn,
<taxonomicName id="4C325B94BC402A59131AFD3F328AFD51" authorityName="Brulle" authorityYear="1837" box="[1043,1156,731,755]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59131AFD3F328AFD51" box="[1043,1156,731,755]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
is exclusive among the
<taxonomicName id="4C325B94BC402A5916AAFD1A341FFCB4" box="[419,529,766,790]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
in that it has no trace of a metepisternal tab (
<figureCitation id="13093C92BC402A591302FD1A3278FCB4" box="[1035,1142,766,790]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="20" pageNumber="963">Figure 7A</figureCitation>
). The epipleura, however, has a perceptible vestigial depression at the same point as in
<taxonomicName id="4C325B94BC402A5917A9FCA73702FCF9" authorityName="Bates" authorityYear="1870" box="[160,268,835,859]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917A9FCA73702FCF9" box="[160,268,835,859]" italics="true" pageId="20" pageNumber="963">Oruscatus</emphasis>
</taxonomicName>
and the other
<taxonomicName id="4C325B94BC402A5916BFFCA73429FCF9" box="[438,551,835,859]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
, indicating again that a metepisternal tab could have been present in the ancestors of the
<taxonomicName id="4C325B94BC402A59154FFC8234B9FCDC" authorityName="Brulle" authorityYear="1837" box="[582,695,870,894]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A59154FFC8234B9FCDC" box="[582,695,870,894]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
.
<bibRefCitation id="EFA35DE6BC402A5915CDFC82357AFCDC" author="Edmonds WD" box="[708,884,870,894]" pageId="20" pageNumber="963" pagination="731 - 874" refId="ref14833" refString="Edmonds WD. 1972. Comparative skeletal morphology, systematics and evolution of the Phanaeine dung beetles (Coleoptera: Scarabaeidae). Univ Kansas Sci Bull. 49: 731 - 874." type="journal article" year="1972">Edmonds (1972</bibRefCitation>
, pp. 814
<emphasis id="B946FC05BC402A5914DEFC8235EAFCDC" box="[983,996,870,894]" italics="true" pageId="20" pageNumber="963">–</emphasis>
815) hypothesized that the metepisternal tab anchors the elytra, maintaining
<emphasis id="B946FC05BC402A59147EFC6D3573FC03" box="[887,893,905,929]" italics="true" pageId="20" pageNumber="963">‘</emphasis>
close elytral appression along the elytral suture by restricting lateral slippage of the elytra
<emphasis id="B946FC05BC402A5914A0FC4F35A1FC61" box="[937,943,939,963]" italics="true" pageId="20" pageNumber="963">’</emphasis>
. Giving assistance during digging, probably this is the same role that the metepisternal tab has in those other dung beetles, which are fossorial and paracoprid.
</paragraph>
<paragraph id="8B8D2017BC402A5917C9FBF73512FA69" blockId="20.[160,1156,142,1483]" pageId="20" pageNumber="963">
The presence of only four protarsomeres, rather than five, is also unique to
<taxonomicName id="4C325B94BC402A5917A9FBD2371BFBEC" authorityName="Brulle" authorityYear="1837" box="[160,277,1078,1102]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917A9FBD2371BFBEC" box="[160,277,1078,1102]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
, at least among the
<taxonomicName id="4C325B94BC402A591527FBD234AAFBEC" box="[558,676,1078,1102]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
. In this tribe, protarsi, if present, are found only in females, but
<emphasis id="B946FC05BC402A5916F7FBBD340AFBD3" box="[510,516,1113,1137]" italics="true" pageId="20" pageNumber="963">‘</emphasis>
are always reduced in size and clawless
<emphasis id="B946FC05BC402A5914F3FBBD320EFBD3" box="[1018,1024,1113,1137]" italics="true" pageId="20" pageNumber="963">’</emphasis>
(
<bibRefCitation id="EFA35DE6BC402A59131CFBBD36D7FB31" author="Edmonds WD" pageId="20" pageNumber="963" pagination="731 - 874" refId="ref14833" refString="Edmonds WD. 1972. Comparative skeletal morphology, systematics and evolution of the Phanaeine dung beetles (Coleoptera: Scarabaeidae). Univ Kansas Sci Bull. 49: 731 - 874." type="journal article" year="1972">Edmonds 1972</bibRefCitation>
, p. 770). In some groups, protarsi are completely absent.
<taxonomicName id="4C325B94BC402A5914E5FB9F324BFB31" authorityName="Harold" authorityYear="1868" box="[1004,1093,1147,1171]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5914E5FB9F324BFB31" box="[1004,1093,1147,1171]" italics="true" pageId="20" pageNumber="963">Bolbites</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC402A5917A9FB7A371FFB14" authorityName="Bates" authorityYear="1870" box="[160,273,1182,1206]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917A9FB7A371FFB14" box="[160,273,1182,1206]" italics="true" pageId="20" pageNumber="963">Oruscatus</emphasis>
</taxonomicName>
are included in this latter case and, in consequence, it is difficult to say if the condition seen in
<taxonomicName id="4C325B94BC402A5916D6FB25345AFB7B" authorityName="Brulle" authorityYear="1837" box="[479,596,1217,1241]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5916D6FB25345AFB7B" box="[479,596,1217,1241]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
(i.e. protarsi with four tarsomeres) was the ancestral condition of
<taxonomicName id="4C325B94BC402A5916B1FB073456FB59" box="[440,600,1251,1275]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
and then
<taxonomicName id="4C325B94BC402A5915E8FB07355CFB59" authorityName="Bates" authorityYear="1870" box="[737,850,1251,1275]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5915E8FB07355CFB59" box="[737,850,1251,1275]" italics="true" pageId="20" pageNumber="963">Oruscatus</emphasis>
</taxonomicName>
took a step further and lost completely the protarsi, or if the condition in the last common ancestor was the five-articulated protarsi and then the reduction occurred in
<taxonomicName id="4C325B94BC402A5914DEFACD3246FAE3" authorityName="Bates" authorityYear="1870" box="[983,1096,1321,1345]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5914DEFACD3246FAE3" box="[983,1096,1321,1345]" italics="true" pageId="20" pageNumber="963">Oruscatus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC402A5917A9FAAF371BFAC1" authorityName="Brulle" authorityYear="1837" box="[160,277,1355,1379]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5917A9FAAF371BFAC1" box="[160,277,1355,1379]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
independently. So, in the first case, protarsi with four tarsomeres would be a synapomorphy of
<taxonomicName id="4C325B94BC402A591512FA8A34B0FA24" box="[539,702,1390,1414]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
, whereas in the latter case it would be a synapomorphy of
<taxonomicName id="4C325B94BC402A5916B6FA75343AFA0B" authorityName="Brulle" authorityYear="1837" box="[447,564,1425,1449]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A5916B6FA75343AFA0B" box="[447,564,1425,1449]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
. Given these difficulties in the ambiguity of this character, we chose not to include it in the analysis.
</paragraph>
<paragraph id="8B8D2017BC402A5917A9F9EE3561F981" blockId="20.[160,879,1546,1571]" box="[160,879,1546,1571]" pageId="20" pageNumber="963">
<heading id="D0C5977BBC402A5917A9F9EE3561F981" box="[160,879,1546,1571]" fontSize="11" level="3" pageId="20" pageNumber="963" reason="5">
<emphasis id="B946FC05BC402A5917A9F9EE3561F981" bold="true" box="[160,879,1546,1571]" inLineHeading="true" pageId="20" pageNumber="963" reason="3">
<emphasis id="B946FC05BC402A5917A9F9EE34EAF981" bold="true" box="[160,740,1546,1571]" italics="true" pageId="20" pageNumber="963">The phylogenetic relationship of the species of</emphasis>
<taxonomicName id="4C325B94BC402A5915E4F9EE3561F981" authorityName="Brulle" authorityYear="1837" box="[749,879,1546,1571]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">Gromphas</taxonomicName>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B8D2017BC402A5817A9F9DB340AFDA3" blockId="20.[160,1156,1599,1727]" lastBlockId="21.[160,1156,142,1726]" lastPageId="21" lastPageNumber="964" pageId="20" pageNumber="963">
The six species of
<taxonomicName id="4C325B94BC402A591652F9DB37C9F9F5" authorityName="Brulle" authorityYear="1837" box="[347,455,1599,1623]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC402A591652F9DB37C9F9F5" box="[347,455,1599,1623]" italics="true" pageId="20" pageNumber="963">Gromphas</emphasis>
</taxonomicName>
are divided into two main clades, one including
<taxonomicName id="4C325B94BC402A5914CEF9DB325AF9F5" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[967,1108,1599,1623]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC402A5914CEF9DB325AF9F5" box="[967,1108,1599,1623]" italics="true" pageId="20" pageNumber="963">G. aeruginosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC402A5917A9F9863719F9D8" authorityName="Waterhouse" authorityYear="1891" box="[160,279,1634,1658]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="B946FC05BC402A5917A9F9863719F9D8" box="[160,279,1634,1658]" italics="true" pageId="20" pageNumber="963">G. lemoinei</emphasis>
</taxonomicName>
(clade A in
<figureCitation id="13093C92BC402A5916AAF98637F0F9D8" box="[419,510,1634,1658]" captionStart="Figure 6" captionStartId="16.[160,225,923,946]" captionTargetBox="[163,1153,142,892]" captionTargetId="figure@16.[163,1153,142,892]" captionTargetPageId="16" captionText="Figure 6. Phylogeny of Gromphas. Above: Cladogram showing the phylogenetic relationship of the species of Gromphas and the evolution of their characters. Black circles indicate uncontroverted synapormorphies; white circles with black margin, controverted synapomorphies; red circles, ambiguous uncontroverted synapomorphies; and white circles with red margin, ambiguous controverted synapomorphies. Below: Support values given for each clade are decay index (above branch) and bootstrap (below branch)." figureDoi="http://doi.org/10.5281/zenodo.3990455" httpUri="https://zenodo.org/record/3990455/files/figure.png" pageId="20" pageNumber="963">Figure 6</figureCitation>
), and other including the topology (
<taxonomicName id="4C325B94BC402A591493F9863209F9D8" authorityName="Blanchard" authorityYear="1846" box="[922,1031,1634,1658]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="B946FC05BC402A591493F9863209F9D8" box="[922,1031,1634,1658]" italics="true" pageId="20" pageNumber="963">G. dichroa</emphasis>
</taxonomicName>
(
<taxonomicName id="4C325B94BC402A591310F986328DF9D8" authorityName="Harold" authorityYear="1869" box="[1049,1155,1634,1658]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC402A591310F986328DF9D8" box="[1049,1155,1634,1658]" italics="true" pageId="20" pageNumber="963">G. inermis</emphasis>
</taxonomicName>
(
<taxonomicName id="4C325B94BC402A5917AEF961373BF93F" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[167,309,1669,1693]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="B946FC05BC402A5917AEF961373BF93F" box="[167,309,1669,1693]" italics="true" pageId="20" pageNumber="963">G. amazonica</emphasis>
</taxonomicName>
+
<taxonomicName id="4C325B94BC402A591658F96137A1F93F" box="[337,431,1669,1693]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="963" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC402A591658F96137A1F93F" box="[337,431,1669,1693]" italics="true" pageId="20" pageNumber="963">G. jardim</emphasis>
</taxonomicName>
))) (clade B). The first clade is supported by 10 synapomorphies (four uncontroverted, two controverted by homoplasies, and four ambiguous optimized manually): genae smooth adjacent to the eyes (char. 4
<emphasis id="B946FC05BC412A581573FF6A3489FF04" box="[634,647,142,166]" italics="true" pageId="21" pageNumber="964">–</emphasis>
1; ambiguous and homoplastic with
<taxonomicName id="4C325B94BC412A58131EFF6A3270FF04" authorityName="Bates" authorityYear="1870" box="[1047,1150,142,166]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A58131EFF6A3270FF04" box="[1047,1150,142,166]" italics="true" pageId="21" pageNumber="964">Oruscatus</emphasis>
</taxonomicName>
; {figs 24, 25}), cephalic projection developed as a horn flattened anteroposteriorly (5-2; ambiguous; {figs 29, 30}), centre of pronotum with a mid-longitudinal line of smooth and glossy tegument (7-1), presence of pronotal prominence (9-1; {figs 1
<emphasis id="B946FC05BC412A581487FF123595FEAC" box="[910,923,246,270]" italics="true" pageId="21" pageNumber="964">–</emphasis>
11}), ventral carina of protibiae simple in both sexes (16-1; {fig. 22}), protibiae with apical tuft of setae without any sexual dimorphism (19-1), protibiae very broad in both sexes (20-1), elytral striae carinulate (25-1; homoplastic with clade D; {fig. 37}), pygidium without basal margin (29-2; ambiguous and homoplastic with clade C; {fig. 36}), and medial sclerite of internal sac wider and strongly curved (30-0; ambiguous; {figs 57, 58}). The values of support are high (bootstrap 99, decay index 6) and give confidence to our hypothesis that these two species are closely related (
<bibRefCitation id="EFA35DE6BC412A5817AFFE0D37F6FDA3" author="Cupello M &amp; Vaz-de-Mello FZ" box="[166,504,489,513]" pageId="21" pageNumber="964" pagination="439 - 482" refId="ref14473" 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>
).
</paragraph>
<paragraph id="8B8D2017BC412A5817C9FDE8344BF9B3" blockId="21.[160,1156,142,1726]" pageId="21" pageNumber="964">
One of the most interesting features of this clade is the reduced sexual dimorphism. In
<taxonomicName id="4C325B94BC412A5817A9FDCA3739FDE4" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[160,311,558,582]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC412A5817A9FDCA3739FDE4" box="[160,311,558,582]" italics="true" pageId="21" pageNumber="964">G. aeruginosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC412A581666FDCA37E7FDE4" authorityName="Waterhouse" authorityYear="1891" box="[367,489,558,582]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="B946FC05BC412A581666FDCA37E7FDE4" box="[367,489,558,582]" italics="true" pageId="21" pageNumber="964">G. lemoinei</emphasis>
</taxonomicName>
, the ventral carina of protibiae is simple in both sexes ({fig. 22}), whereas it is ornamented with tubercles or teeth in
<taxonomicName id="4C325B94BC412A581438FDB53593FDCB" authorityName="Bates" authorityYear="1870" box="[817,925,593,617]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A581438FDB53593FDCB" box="[817,925,593,617]" italics="true" pageId="21" pageNumber="964">Oruscatus</emphasis>
</taxonomicName>
(
<figureCitation id="13093C92BC412A5814A4FDB53217FDCB" box="[941,1049,593,617]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="21" pageNumber="964">Figure 8B</figureCitation>
),
<taxonomicName id="4C325B94BC412A581326FDB5328DFDCB" authorityName="Harold" authorityYear="1868" box="[1071,1155,593,617]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A581326FDB5328DFDCB" box="[1071,1155,593,617]" italics="true" pageId="21" pageNumber="964">Bolbites</emphasis>
</taxonomicName>
(
<figureCitation id="13093C92BC412A5817A1FD97371FFD29" box="[168,273,627,651]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="21" pageNumber="964">Figure 8C</figureCitation>
) and in three of the other four
<taxonomicName id="4C325B94BC412A58156DFD9734DBFD29" authorityName="Brulle" authorityYear="1837" box="[612,725,627,651]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A58156DFD9734DBFD29" box="[612,725,627,651]" italics="true" pageId="21" pageNumber="964">Gromphas</emphasis>
</taxonomicName>
(
<taxonomicName id="4C325B94BC412A5815E8FD97354DFD29" box="[737,835,627,651]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC412A5815E8FD97354DFD29" box="[737,835,627,651]" italics="true" pageId="21" pageNumber="964">G. jardim</emphasis>
</taxonomicName>
is the exception) (
<figureCitation id="13093C92BC412A581305FD973278FD29" box="[1036,1142,627,651]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="21" pageNumber="964">Figure 8A</figureCitation>
). In the other Phanaeina, this carina is also simple in both sexes. Other reductions in sexual dimorphism are the protibiae very broad and the protibial tuft of setae of same length in both sexes. In the other
<taxonomicName id="4C325B94BC412A5816CBFD3F343DFD51" authorityName="Brulle" authorityYear="1837" box="[450,563,731,755]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A5816CBFD3F343DFD51" box="[450,563,731,755]" italics="true" pageId="21" pageNumber="964">Gromphas</emphasis>
</taxonomicName>
(and
<taxonomicName id="4C325B94BC412A581573FD3F34E8FD51" authorityName="Bates" authorityYear="1870" box="[634,742,731,755]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A581573FD3F34E8FD51" box="[634,742,731,755]" italics="true" pageId="21" pageNumber="964">Oruscatus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC412A58142FFD3F3574FD51" authorityName="Harold" authorityYear="1868" box="[806,890,731,755]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A58142FFD3F3574FD51" box="[806,890,731,755]" italics="true" pageId="21" pageNumber="964">Bolbites</emphasis>
</taxonomicName>
), males have narrower protibiae and the tuft of setae is much longer and denser in males than in females (
<figureCitation id="13093C92BC412A5817A1FCC5370DFC9B" box="[168,259,801,825]" captionStart="Figure 2" captionStartId="5.[160,225,933,956]" captionTargetBox="[178,1148,161,902]" captionTargetId="figure@5.[163,1153,142,902]" captionTargetPageId="5" captionText="Figure 2. Protibia of Gromphas jardim sp. nov. (A, B) Male protibia: (A) Ventral view, (B) Dorsal view. Red arrow indicates apical tubercle modified in a tapered spur. (C) Female protibia. Note that the apical tuft of setae is much denser and longer in male than in female." figureDoi="http://doi.org/10.5281/zenodo.3990445" httpUri="https://zenodo.org/record/3990445/files/figure.png" pageId="21" pageNumber="964">Figure 2</figureCitation>
, {fig. 23}). The presence of a pronotal prominence is also a remarkable characteristic of this clade, being absent in the other
<taxonomicName id="4C325B94BC412A581593FCA73539FCF9" box="[666,823,835,859]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
; nonetheless, as is typical for these two species, there is no sexual difference in this feature. In Phanaeina, in general, males have very elaborate ornamentation on head and pronotum, and it has been known since
<bibRefCitation id="EFA35DE6BC412A5817EAFC4F3775FC61" author="Darwin C" box="[227,379,939,963]" pageId="21" pageNumber="964" refId="ref14778" refString="Darwin C. 1859. On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. London: John Murray." type="book" year="1859">Darwin (1859</bibRefCitation>
,
<bibRefCitation id="EFA35DE6BC412A581683FC4F37CFFC61" author="Darwin C" box="[394,449,939,963]" pageId="21" pageNumber="964" refId="ref14811" refString="Darwin C. 1871. The descent of man, and selection in relation to sex. London: John Murray." type="book" year="1871">1871</bibRefCitation>
) that these structures play a key role in the sexual access to females by males and, therefore, are under constant pressure from sexual selection (see, for example,
<bibRefCitation id="EFA35DE6BC412A581635FC1537D3FBAB" author="Otronen M" box="[316,477,1009,1033]" pageId="21" pageNumber="964" pagination="741 - 748" refId="ref15770" refString="Otronen M. 1988. Intra- and intersexual interactions at breeding burrows in the horned beetle, Coprophanaeus ensifer. Anim Behav. 36: 741 - 748." type="journal article" year="1988">Otronen 1988</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC412A5816E7FC1534A3FBAB" author="Rasmussen JL" box="[494,685,1009,1033]" pageId="21" pageNumber="964" pagination="67 - 82" refId="ref16033" refString="Rasmussen JL. 1994. The influence of horn and body size on the reproductive behavior of the horned rainbow scarab beetle Phanaeus difformis (Coleoptera: Scarabaeidae). J Insect Behav. 7: 67 - 82." type="journal article" year="1994">Rasmussen 1994</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC412A5815B7FC153556FBAB" author="Escobar F" box="[702,856,1009,1033]" pageId="21" pageNumber="964" pagination="307 - 310" refId="ref15002" refString="Escobar F. 2003. Seleccion de habitat y comportamiento sexual de Sulcophanaeus velutinus (Murray 1856) (Coleoptera: Scarabaeidae: Scarabaeinae) en un bosque de montana en Colombia. Acta Zool Mex (NS). 90: 307 - 310." type="journal article" year="2003">Escobar 2003</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC412A581460FC153270FBAB" author="Emlen DJ &amp; Philips TK" box="[873,1150,1009,1033]" pageId="21" pageNumber="964" pagination="47 - 56" refId="ref14933" refString="Emlen DJ, Philips TK. 2006. Phylogenetic evidence for an association between tunneling behavior and the evolution of horns in dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopt Soc Monogr. 5: 47 - 56." type="journal article" year="2006">Emlen and Philips 2006</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC412A5817A9FBF737CDFB89" author="Rowland JM &amp; Emlen DJ" box="[160,451,1043,1067]" pageId="21" pageNumber="964" pagination="773 - 776" refId="ref16100" refString="Rowland JM, Emlen DJ. 2009. Two thresholds, three male forms result in facultative male trimorphism in beetles. Science. 323: 773 - 776." type="journal article" year="2009">Rowland and Emlen 2009</bibRefCitation>
). But what would be the reason for these structures being as developed in females as in males in this clade? A few other groups of
<taxonomicName id="4C325B94BC412A5814AFFBD2321AFBEC" box="[934,1044,1078,1102]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="tribe" tribe="Phanaeini">Phanaeini</taxonomicName>
also have females with ornamentation greatly developed, as in males, including those of the group lancifer of subgenus
<taxonomicName id="4C325B94BC412A581691FB9F3431FB31" authorityName="d'Olsoufieff" authorityYear="1924" box="[408,575,1147,1171]" class="Insecta" family="Scarabaeidae" genus="Megaphanaeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A581691FB9F3431FB31" box="[408,575,1147,1171]" italics="true" pageId="21" pageNumber="964">Megaphanaeus</emphasis>
</taxonomicName>
d
<emphasis id="B946FC05BC412A581553FB9F346EFB31" box="[602,608,1147,1171]" italics="true" pageId="21" pageNumber="964">’</emphasis>
<bibRefCitation id="EFA35DE6BC412A581569FB9F351DFB31" author="Olsoufieff G" box="[608,787,1147,1171]" pageId="21" pageNumber="964" pagination="4 - 201" refId="ref14554" refString="d ' Olsoufieff G. 1924. Les Phanaeides (Coleoptera - Lamellicornia), famille Scarabaeidae - tr. Coprini Insecta. 13: 4 - 201." type="journal article" year="1924">Olsoufieff, 1924</bibRefCitation>
, of
<taxonomicName id="4C325B94BC412A58144FFB9F35FFFB31" authorityName="D Olsoufieff" authorityYear="1924" box="[838,1009,1147,1171]" class="Insecta" family="Scarabaeidae" genus="Coprophanaeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A58144FFB9F35FFFB31" box="[838,1009,1147,1171]" italics="true" pageId="21" pageNumber="964">Coprophanaeus</emphasis>
</taxonomicName>
d
<emphasis id="B946FC05BC412A581304FB9F321DFB31" box="[1037,1043,1147,1171]" italics="true" pageId="21" pageNumber="964">’</emphasis>
<bibRefCitation id="EFA35DE6BC412A58131BFB9F36D9FB14" author="Olsoufieff G" pageId="21" pageNumber="964" pagination="4 - 201" refId="ref14554" refString="d ' Olsoufieff G. 1924. Les Phanaeides (Coleoptera - Lamellicornia), famille Scarabaeidae - tr. Coprini Insecta. 13: 4 - 201." type="journal article" year="1924">Olsoufieff, 1924</bibRefCitation>
, and, to a lesser extent, the group faunus of
<taxonomicName id="4C325B94BC412A5815B7FB7A356CFB14" authorityName="d'Olsoufieff" authorityYear="1924" box="[702,866,1182,1206]" class="Insecta" family="Scarabaeidae" genus="Sulcophanaeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A5815B7FB7A356CFB14" box="[702,866,1182,1206]" italics="true" pageId="21" pageNumber="964">Sulcophanaeus</emphasis>
</taxonomicName>
d
<emphasis id="B946FC05BC412A58147EFB7A3573FB14" box="[887,893,1182,1206]" italics="true" pageId="21" pageNumber="964">’</emphasis>
<bibRefCitation id="EFA35DE6BC412A581475FB7A3224FB14" author="Olsoufieff G" box="[892,1066,1182,1206]" pageId="21" pageNumber="964" pagination="4 - 201" refId="ref14554" refString="d ' Olsoufieff G. 1924. Les Phanaeides (Coleoptera - Lamellicornia), famille Scarabaeidae - tr. Coprini Insecta. 13: 4 - 201." type="journal article" year="1924">Olsoufieff, 1924</bibRefCitation>
, groups that include some of the largest dung beetles of the New World (
<bibRefCitation id="EFA35DE6BC412A58146EFB253201FB7B" author="Edmonds WD" box="[871,1039,1217,1241]" pageId="21" pageNumber="964" pagination="1 - 60" refId="ref14867" refString="Edmonds WD. 2000. Revision of the Neotropical dung beetle genus Sulcophanaeus (Coleoptera: Scarabaeidae: Scarabaeinae). Fol Heyrovskyana. 6: 1 - 60." type="journal article" year="2000">Edmonds 2000</bibRefCitation>
;
<bibRefCitation id="EFA35DE6BC412A581313FB25375AFB59" author="Edmonds WD &amp; Zidek J" pageId="21" pageNumber="964" pagination="1 - 111" refId="ref14897" 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">Edmonds and Zidek 2010</bibRefCitation>
).
<bibRefCitation id="EFA35DE6BC412A581662FB073417FB59" author="Otronen M" box="[363,537,1251,1275]" pageId="21" pageNumber="964" pagination="741 - 748" refId="ref15770" refString="Otronen M. 1988. Intra- and intersexual interactions at breeding burrows in the horned beetle, Coprophanaeus ensifer. Anim Behav. 36: 741 - 748." type="journal article" year="1988">Otronen (1988)</bibRefCitation>
supposed that females of
<taxonomicName id="4C325B94BC412A58145AFB073748FABC" authority="(Germar, 1821)" baseAuthorityName="Germar" baseAuthorityYear="1821" class="Insecta" family="Scarabaeidae" genus="Coprophanaeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="species" species="ensifer" subGenus="M.">
<emphasis id="B946FC05BC412A58145AFB07328DFB59" box="[851,1155,1251,1275]" italics="true" pageId="21" pageNumber="964">Coprophanaeus (M.) ensifer</emphasis>
(
<bibRefCitation id="EFA35DE6BC412A5817AFFAE23731FABC" author="Germar EF" box="[166,319,1286,1310]" pageId="21" pageNumber="964" pagination="107 - 183" refId="ref15176" refString="Germar EF. 1821. Neue exotische Kafer beschrieben von C. R. W. Widermann u. E. F. Germar. Mag Entomol. 4: 107 - 183." type="journal article" year="1821">Germar, 1821</bibRefCitation>
)
</taxonomicName>
need a developed armature to face combat against other females and so obtain enough food for their large larvae; larvae of smaller dung beetles, on the other hand, do not need large amounts of food and therefore females of these species do not need to face such fierce disputes. Not having such a large size, females of
<taxonomicName id="4C325B94BC412A5814E5FA8A328AFA24" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[1004,1156,1390,1414]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC412A5814E5FA8A328AFA24" box="[1004,1156,1390,1414]" italics="true" pageId="21" pageNumber="964">G. aeruginosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC412A5817DFFA75375DFA0B" authorityName="Waterhouse" authorityYear="1891" box="[214,339,1425,1449]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="B946FC05BC412A5817DFFA75375DFA0B" box="[214,339,1425,1449]" italics="true" pageId="21" pageNumber="964">G. lemoinei</emphasis>
</taxonomicName>
are unique in possessing well-developed ornamentation. Only with a greater knowledge of their biology will we be able to answer this question more firmly, but it is possible that some idiosyncrasy in their behaviour leads females to fight battles and so be in need of such armament.
</paragraph>
<paragraph id="8B8D2017BC412A5B17C9F9FF34DDFDA3" blockId="21.[160,1156,142,1726]" lastBlockId="22.[160,1157,142,1622]" lastPageId="22" lastPageNumber="965" pageId="21" pageNumber="964">
The second main lineage within
<taxonomicName id="4C325B94BC412A58154EF9FF34B2F991" authorityName="Brulle" authorityYear="1837" box="[583,700,1563,1587]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="964" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC412A58154EF9FF34B2F991" box="[583,700,1563,1587]" italics="true" pageId="21" pageNumber="964">Gromphas</emphasis>
</taxonomicName>
, clade B, is supported by five synapomorphies (three uncontroverted, two ambiguous): cephalic projection as a carina with base wider than apex (char. 5
<emphasis id="B946FC05BC412A5816F2F9843406F9DA" box="[507,520,1632,1656]" italics="true" pageId="21" pageNumber="964">–</emphasis>
1; ambiguous; {figs 31
<emphasis id="B946FC05BC412A581403F9843519F9DA" box="[778,791,1632,1656]" italics="true" pageId="21" pageNumber="964">–</emphasis>
33}), pronotum globular (10-1), ventral carina of male protibiae with a row of tubercles on basal half (17-2; ambiguous; {fig. 23b}), protibial spur strongly expanded at apex (18-1; {fig. 23}), and, on basal third or basal half, sheen and punctation of sutural margin of elytra extend onto first or second interstriae (27-1; {figs 13
<emphasis id="B946FC05BC422A5B1576FF553482FF6B" box="[639,652,177,201]" italics="true" pageId="22" pageNumber="965">–</emphasis>
15}). The values of support for this clade, however, are the lowest in our analysis (bootstrap 55, decay index 2). One interesting synapomorphy of this lineage is the form of the ornamentation of the ventral carina of male protibiae. In
<taxonomicName id="4C325B94BC422A5B168AFEFD37D2FE93" authorityName="Harold" authorityYear="1868" box="[387,476,281,305]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B168AFEFD37D2FE93" box="[387,476,281,305]" italics="true" pageId="22" pageNumber="965">Bolbites</emphasis>
</taxonomicName>
, this carina has few tiny, irregular teeth and one large central tooth (
<figureCitation id="13093C92BC422A5B1645FED837B0FEF6" box="[332,446,316,340]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="22" pageNumber="965">Figure 8C</figureCitation>
); in
<taxonomicName id="4C325B94BC422A5B16FCFED83468FEF6" authorityName="Bates" authorityYear="1870" box="[501,614,316,340]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B16FCFED83468FEF6" box="[501,614,316,340]" italics="true" pageId="22" pageNumber="965">Oruscatus</emphasis>
</taxonomicName>
, this ornamentation is modified into a single central tooth without any accessory teeth or tubercles (
<figureCitation id="13093C92BC422A5B145AFEBA35C9FED4" box="[851,967,350,374]" captionStart="Figure 8" captionStartId="18.[160,225,961,984]" captionTargetBox="[163,1153,142,930]" captionTargetId="figure@18.[163,1153,142,930]" captionTargetPageId="18" captionText="Figure 8. Lateral view of male protibia, ornamentation of ventral carina. (A) Gromphas inermis: a row of tubercles (indicated by red arrow), a synapomorphy of clade B of Gromphas. (B) Oruscatus davus: a single large tooth. (C) Bolbites onitoides: a large central tooth with minor lateral teeth." figureDoi="http://doi.org/10.5281/zenodo.3990461" httpUri="https://zenodo.org/record/3990461/files/figure.png" pageId="22" pageNumber="965">Figure 8B</figureCitation>
). In
<emphasis id="B946FC05BC422A5B130DFEBA371CFE3B" italics="true" pageId="22" pageNumber="965">
<taxonomicName id="4C325B94BC422A5B130DFEBA3271FED4" authorityName="Blanchard" authorityYear="1846" box="[1028,1151,350,374]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="dichroa">G. dichroa</taxonomicName>
,
<taxonomicName id="4C325B94BC422A5B17A9FE65371CFE3B" authorityName="Harold" authorityYear="1869" box="[160,274,385,409]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="inermis">G. inermis</taxonomicName>
</emphasis>
and
<taxonomicName id="4C325B94BC422A5B1659FE6537E3FE3B" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[336,493,385,409]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="B946FC05BC422A5B1659FE6537E3FE3B" box="[336,493,385,409]" italics="true" pageId="22" pageNumber="965">G. amazonica</emphasis>
</taxonomicName>
, the carina has a row of tiny, regular tubercles, with no apparent difference between these species ({fig. 23b}); in
<taxonomicName id="4C325B94BC422A5B144BFE4035A7FE1E" box="[834,937,420,444]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC422A5B144BFE4035A7FE1E" box="[834,937,420,444]" italics="true" pageId="22" pageNumber="965">G. jardim</emphasis>
</taxonomicName>
, as a controverted autapomorphy (char. 16
<emphasis id="B946FC05BC422A5B16B3FE2237C9FE7C" box="[442,455,454,478]" italics="true" pageId="22" pageNumber="965">–</emphasis>
1, homoplastic with clade A), this ornamentation is lost and the carina is simple and continuous (
<figureCitation id="13093C92BC422A5B155DFE0D34CBFDA3" box="[596,709,489,513]" captionStart="Figure 2" captionStartId="5.[160,225,933,956]" captionTargetBox="[178,1148,161,902]" captionTargetId="figure@5.[163,1153,142,902]" captionTargetPageId="5" captionText="Figure 2. Protibia of Gromphas jardim sp. nov. (A, B) Male protibia: (A) Ventral view, (B) Dorsal view. Red arrow indicates apical tubercle modified in a tapered spur. (C) Female protibia. Note that the apical tuft of setae is much denser and longer in male than in female." figureDoi="http://doi.org/10.5281/zenodo.3990445" httpUri="https://zenodo.org/record/3990445/files/figure.png" pageId="22" pageNumber="965">Figure 2A</figureCitation>
).
</paragraph>
<paragraph id="8B8D2017BC422A5B17C9FDE834EAFB59" blockId="22.[160,1157,142,1622]" pageId="22" pageNumber="965">
The next clade, clade C, is supported by five synapomorphies (two uncontroverted, three ambiguous): apical margin of clypeus evidently upturned (1-1), lateral margin of clypeus lobate (3-1;
<figureCitation id="13093C92BC422A5B169AFDB5340FFDCB" box="[403,513,593,617]" captionStart="Figure 1" captionStartId="4.[160,225,1578,1601]" captionTargetBox="[163,1152,143,1546]" captionTargetId="figure@4.[163,1153,142,1546]" captionTargetPageId="4" captionText="Figure 1. Gromphas jardim sp. nov. (A–D) Holotype. (A) Dorsal view. (B) Lateral view. (C) Aedeagus. From left to right, dorsal, lateral and ventral views. (D) Labels. (E) Female paratype from Bolivia. (F) Dorsal view of head (holotype). (G) Frontal view of cephalic projection (male paratype)." figureDoi="http://doi.org/10.5281/zenodo.3990443" httpUri="https://zenodo.org/record/3990443/files/figure.png" pageId="22" pageNumber="965">Figure 1F</figureCitation>
), cephalic carina emarginate apically (6-1; ambiguous;
<figureCitation id="13093C92BC422A5B17A9FD9736F3FD29" box="[160,253,627,651]" captionStart="Figure 7" captionStartId="17.[160,225,794,817]" captionTargetBox="[163,1153,142,763]" captionTargetId="figure@17.[163,1153,142,763]" captionTargetPageId="17" captionText="Figure 7. Metepisterna. (A) Gromphas jardim sp. nov.: gromphas is unique among the Phanaeini by not showing any trace of a metepisternal tab (MTab) covering the epipleural margin, a synapomorphy of this genus. (B) Oruscatus davus: Both species of Oruscatus have a small but evident MTab (indicated by red arrow). (C) Bolbites onitoides: The MTab of Bolbites is strongly developed and the dorsal margin of metepisternum is highly curved. This is the same form seen in the rest of Phanaeina and is a synapomorphy of this taxon. (D) Dichotomius fissus (Harold, 1867): MTab in a non-Phanaeini. This structure is present also in a series of other Coprini, as well as in at least some Oniticellini and Onitini." figureDoi="http://doi.org/10.5281/zenodo.3990459" httpUri="https://zenodo.org/record/3990459/files/figure.png" pageId="22" pageNumber="965">Figure 7</figureCitation>
), pygidium without basal margin (29-1; ambiguous and homoplastic with clade A; {fig. 35}), and medial sclerite of internal sac wider and only slightly curved (31-1; ambiguous) ({figs 59, 60}). The support values give us confidence in this hypothesis (bootstrap 76, decay index 2). This lineage shares with
<taxonomicName id="4C325B94BC422A5B1401FD3F3591FD51" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[776,927,731,755]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC422A5B1401FD3F3591FD51" box="[776,927,731,755]" italics="true" pageId="22" pageNumber="965">G. aeruginosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC422A5B14DEFD3F325FFD51" authorityName="Waterhouse" authorityYear="1891" box="[983,1105,731,755]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="B946FC05BC422A5B14DEFD3F325FFD51" box="[983,1105,731,755]" italics="true" pageId="22" pageNumber="965">G. lemoinei</emphasis>
</taxonomicName>
one important condition, the basal margin of pygidium completely absent ({fig. 36}).
<taxonomicName id="4C325B94BC422A5B17A9FCC53762FC9B" authorityName="Blanchard" authorityYear="1846" box="[160,364,801,825]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="B946FC05BC422A5B17A9FCC53762FC9B" box="[160,364,801,825]" italics="true" pageId="22" pageNumber="965">Gromphas dichroa</emphasis>
</taxonomicName>
, in its turn, possesses the basal margin complete as
<taxonomicName id="4C325B94BC422A5B14C4FCC5322FFC9B" authorityName="Harold" authorityYear="1868" box="[973,1057,801,825]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B14C4FCC5322FFC9B" box="[973,1057,801,825]" italics="true" pageId="22" pageNumber="965">Bolbites</emphasis>
</taxonomicName>
and the other Phanaeina ({fig. 35}). The two species of
<taxonomicName id="4C325B94BC422A5B15BCFCA7352FFCF9" authorityName="Bates" authorityYear="1870" box="[693,801,835,859]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B15BCFCA7352FFCF9" box="[693,801,835,859]" italics="true" pageId="22" pageNumber="965">Oruscatus</emphasis>
</taxonomicName>
apparently have an intermediate condition: the basal margin is present, but in the majority of specimens it is interrupted in the middle by the sulcus of propygidium. So, albeit complete in
<taxonomicName id="4C325B94BC422A5B17A9FC4F371CFC61" authorityName="Blanchard" authorityYear="1846" box="[160,274,939,963]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="B946FC05BC422A5B17A9FC4F371CFC61" box="[160,274,939,963]" italics="true" pageId="22" pageNumber="965">G. dichroa</emphasis>
</taxonomicName>
, we believe that there is a general tendency in
<taxonomicName id="4C325B94BC422A5B1437FC4F35D7FC61" box="[830,985,939,963]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
for the loss of this basal margin, perhaps linked to the great development of the propygidium (which has the length of the pygidium in
<taxonomicName id="4C325B94BC422A5B1531FC1534DBFBAB" box="[568,725,1009,1033]" class="Insecta" family="Scarabaeidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="subTribe" subTribe="Gromphadina">Gromphadina</taxonomicName>
, and is shorter than the pygidium in Phanaeina), and, therefore, we judge that the most likely scenario was the parallel and independent loss in both lineages rather than a unique loss in the ancestral
<taxonomicName id="4C325B94BC422A5B131AFBD2328AFBEC" authorityName="Brulle" authorityYear="1837" box="[1043,1156,1078,1102]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B131AFBD2328AFBEC" box="[1043,1156,1078,1102]" italics="true" pageId="22" pageNumber="965">Gromphas</emphasis>
</taxonomicName>
and a new origin in
<taxonomicName id="4C325B94BC422A5B168CFBBD37F8FBD3" authorityName="Blanchard" authorityYear="1846" box="[389,502,1113,1137]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="B946FC05BC422A5B168CFBBD37F8FBD3" box="[389,502,1113,1137]" italics="true" pageId="22" pageNumber="965">G. dichroa</emphasis>
</taxonomicName>
. The form of the medial sclerite of the internal sac of this lineage is very different to that of clade A and
<taxonomicName id="4C325B94BC422A5B15DDFB9F3549FB31" authorityName="Blanchard" authorityYear="1846" box="[724,839,1147,1171]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="B946FC05BC422A5B15DDFB9F3549FB31" box="[724,839,1147,1171]" italics="true" pageId="22" pageNumber="965">G. dichroa</emphasis>
</taxonomicName>
and, at same time, homogeneous between its species ({figs 59, 60}). As this sclerite is absent both in
<taxonomicName id="4C325B94BC422A5B14F5FB7A325EFB14" authorityName="Harold" authorityYear="1868" box="[1020,1104,1182,1206]" class="Insecta" family="Scarabaeidae" genus="Bolbites" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B14F5FB7A325EFB14" box="[1020,1104,1182,1206]" italics="true" pageId="22" pageNumber="965">Bolbites</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC422A5B17A9FB253702FB7B" authorityName="Bates" authorityYear="1870" box="[160,268,1217,1241]" class="Insecta" family="Scarabaeidae" genus="Oruscatus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC422A5B17A9FB253702FB7B" box="[160,268,1217,1241]" italics="true" pageId="22" pageNumber="965">Oruscatus</emphasis>
</taxonomicName>
, it was not possible to properly polarize this character and, given the topology of the tree, it remained ambiguous in our analysis.
</paragraph>
<paragraph id="8B8D2017BC422A5B17C9FAE23219F9F4" blockId="22.[160,1157,142,1622]" pageId="22" pageNumber="965">
The last clade includes
<taxonomicName id="4C325B94BC422A5B16ECFAE23488FABC" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[485,646,1286,1310]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="B946FC05BC422A5B16ECFAE23488FABC" box="[485,646,1286,1310]" italics="true" pageId="22" pageNumber="965">G. amazonica</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC422A5B15C2FAE23539FABC" box="[715,823,1286,1310]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC422A5B15C2FAE23539FABC" box="[715,823,1286,1310]" italics="true" pageId="22" pageNumber="965">G. jardim</emphasis>
</taxonomicName>
, sharing four apomorphies (two uncontroverted and two controverted): centre of metasternum with sparse punctation (15-2; homoplastic with
<taxonomicName id="4C325B94BC422A5B1545FAA834E4FAC6" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[588,746,1356,1380]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC422A5B1545FAA834E4FAC6" box="[588,746,1356,1380]" italics="true" pageId="22" pageNumber="965">G. aeruginosa</emphasis>
</taxonomicName>
), tubercle of apical inner angle of protibiae modified in a strong spur (22-1;
<figureCitation id="13093C92BC422A5B15A9FA8A350CFA24" box="[672,770,1390,1414]" captionStart="Figure 2" captionStartId="5.[160,225,933,956]" captionTargetBox="[178,1148,161,902]" captionTargetId="figure@5.[163,1153,142,902]" captionTargetPageId="5" captionText="Figure 2. Protibia of Gromphas jardim sp. nov. (A, B) Male protibia: (A) Ventral view, (B) Dorsal view. Red arrow indicates apical tubercle modified in a tapered spur. (C) Female protibia. Note that the apical tuft of setae is much denser and longer in male than in female." figureDoi="http://doi.org/10.5281/zenodo.3990445" httpUri="https://zenodo.org/record/3990445/files/figure.png" pageId="22" pageNumber="965">Figure 2</figureCitation>
, {fig. 21b}), apical protarsomere with spiniform projection at apex (23-1; {fig. 45}), and elytral striae carinulate (25-1; homoplastic with clade
<taxonomicName id="4C325B94BC422A5B16B5FA503455FA6E" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[444,603,1460,1484]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC422A5B16B5FA503455FA6E" box="[444,603,1460,1484]" italics="true" pageId="22" pageNumber="965">G. aeruginosa</emphasis>
</taxonomicName>
+
<taxonomicName id="4C325B94BC422A5B1574FA5034F3FA6E" authorityName="Waterhouse" authorityYear="1891" box="[637,765,1460,1484]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="B946FC05BC422A5B1574FA5034F3FA6E" box="[637,765,1460,1484]" italics="true" pageId="22" pageNumber="965">G. lemoinei</emphasis>
</taxonomicName>
;{fig. 38}). The support values are also robust (bootstrap 78, decay index 3). The most remarkable synapomorphy of this lineage is the protibial tubercle developed into a strong spur, which has no parallel within the genus; the great differences between the two species regarding the form of this spur are discussed in the description of
<taxonomicName id="4C325B94BC422A5B145AF9DA35B3F9F4" box="[851,957,1598,1622]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="965" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC422A5B145AF9DA35B3F9F4" box="[851,957,1598,1622]" italics="true" pageId="22" pageNumber="965">G. jardim</emphasis>
</taxonomicName>
above.
</paragraph>
<paragraph id="8B8D2017BC432A5A17A9FF6A37EDFF0A" blockId="23.[160,483,142,168]" box="[160,483,142,168]" pageId="23" pageNumber="966">
<emphasis id="B946FC05BC432A5A17A9FF6A37EDFF0A" bold="true" box="[160,483,142,168]" pageId="23" pageNumber="966">Phylogenetic conclusion</emphasis>
</paragraph>
<paragraph id="8B8D2017BC432A5A17A9FF203257FC8B" blockId="23.[160,1157,196,1121]" pageId="23" pageNumber="966">
Through this phylogenetic analysis, we observed that features related to tegument [e.g. punctation (chars. 17 and 26) or carinulae of elytral striae (char. 25)], which are of a great importance for species identification, were the most error-prone characters for the construction of primary homologies and, so, led to more homoplasies (see
<bibRefCitation id="EFA35DE6BC432A5A1304FEC8375FFEC5" author="Nixon KC &amp; Carpenter JM" pageId="23" pageNumber="966" pagination="160 - 169" refId="ref15711" refString="Nixon KC, Carpenter JM. 2012. On homology. Cladistics. 28: 160 - 169." type="journal article" year="2012">Nixon and Carpenter 2012</bibRefCitation>
on homoplasy as error). The differences in pronotal granulation were also very difficult to codify and to include in our analysis because of their gradual variation between species. In fact, there is no pair of species of
<taxonomicName id="4C325B94BC432A5A14ADFE70321BFE0E" authorityName="Brulle" authorityYear="1837" box="[932,1045,404,428]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A14ADFE70321BFE0E" box="[932,1045,404,428]" italics="true" pageId="23" pageNumber="966">Gromphas</emphasis>
</taxonomicName>
with the same pattern of pronotal granulation. For the same reason, codifying the differences in the form of the anteromedian angle of metasternum was virtually impossible. Nonetheless, in this latter case, there is a clear homogeneity among the forms founded in
<emphasis id="B946FC05BC432A5A17B5FDFB37CDFD95" box="[188,451,543,567]" italics="true" pageId="23" pageNumber="966">
<taxonomicName id="4C325B94BC432A5A17B5FDFB375DFD95" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[188,339,543,567]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="amazonica">G. amazonica</taxonomicName>
,
<taxonomicName id="4C325B94BC432A5A1656FDFB37CDFD95" box="[351,451,543,567]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="jardim">G. jardim</taxonomicName>
</emphasis>
and
<taxonomicName id="4C325B94BC432A5A16F5FDFB3467FD95" authorityName="Harold" authorityYear="1869" box="[508,617,543,567]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC432A5A16F5FDFB3467FD95" box="[508,617,543,567]" italics="true" pageId="23" pageNumber="966">G. inermis</emphasis>
</taxonomicName>
({figs 50, 51}), on one side, and in
<taxonomicName id="4C325B94BC432A5A14E4FDFB328AFD95" baseAuthorityName="Perty" baseAuthorityYear="1830" box="[1005,1156,543,567]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="aeruginosa">
<emphasis id="B946FC05BC432A5A14E4FDFB328AFD95" box="[1005,1156,543,567]" italics="true" pageId="23" pageNumber="966">G. aeruginosa</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC432A5A17DEFDA5375BFDFB" authorityName="Waterhouse" authorityYear="1891" box="[215,341,577,601]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="lemoinei">
<emphasis id="B946FC05BC432A5A17DEFDA5375BFDFB" box="[215,341,577,601]" italics="true" pageId="23" pageNumber="966">G. lemoinei</emphasis>
</taxonomicName>
({fig. 52}), on the other. The form seen in
<taxonomicName id="4C325B94BC432A5A1453FDA535C0FDFB" authorityName="Blanchard" authorityYear="1846" box="[858,974,577,601]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="dichroa">
<emphasis id="B946FC05BC432A5A1453FDA535C0FDFB" box="[858,974,577,601]" italics="true" pageId="23" pageNumber="966">G. dichroa</emphasis>
</taxonomicName>
, which is high, narrow and truncate apically, is unique in the genus ({fig. 49}). Finally, the usual apomorphic absence of posterior pronotal fossae in
<taxonomicName id="4C325B94BC432A5A142EFD633592FD3D" authorityName="Harold" authorityYear="1869" box="[807,924,647,671]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC432A5A142EFD633592FD3D" box="[807,924,647,671]" italics="true" pageId="23" pageNumber="966">G. inermis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC432A5A14EFFD63328DFD3D" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[998,1155,647,671]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="B946FC05BC432A5A14EFFD63328DFD3D" box="[998,1155,647,671]" italics="true" pageId="23" pageNumber="966">G. amazonica</emphasis>
</taxonomicName>
appeared ambiguous in our analysis, with two possible scenarios: the loss of the fossae in the ancestral of clade C and its subsequent reappearance in
<taxonomicName id="4C325B94BC432A5A146DFD2835C9FD46" box="[868,967,716,740]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="jardim">
<emphasis id="B946FC05BC432A5A146DFD2835C9FD46" box="[868,967,716,740]" italics="true" pageId="23" pageNumber="966">G. jardim</emphasis>
</taxonomicName>
, or independent losses in
<taxonomicName id="4C325B94BC432A5A160CFD0B377FFCA5" authorityName="Harold" authorityYear="1869" box="[261,369,751,775]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="inermis">
<emphasis id="B946FC05BC432A5A160CFD0B377FFCA5" box="[261,369,751,775]" italics="true" pageId="23" pageNumber="966">G. inermis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C325B94BC432A5A16A1FD0B3433FCA5" authorityName=": Cupello and Vaz-de-Mello" authorityYear="2013" box="[424,573,751,775]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="species" species="amazonica">
<emphasis id="B946FC05BC432A5A16A1FD0B3433FCA5" box="[424,573,751,775]" italics="true" pageId="23" pageNumber="966">G. amazonica</emphasis>
</taxonomicName>
. In this case, we have no opinion on which scenario would be more likely and therefore we have not plotted this character on the tree.
</paragraph>
<paragraph id="8B8D2017BC432A5A17C9FCD035D0FBC3" blockId="23.[160,1157,196,1121]" pageId="23" pageNumber="966">
We consider that the results of this phylogenetic analysis are important not only because they allows us to reconstruct the evolution of
<taxonomicName id="4C325B94BC432A5A1418FCB3358CFCCD" authorityName="Brulle" authorityYear="1837" box="[785,898,855,879]" class="Insecta" family="Scarabaeidae" genus="Gromphas" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A1418FCB3358CFCCD" box="[785,898,855,879]" italics="true" pageId="23" pageNumber="966">Gromphas</emphasis>
</taxonomicName>
, but, on a larger scale, because they represent a further contribution to the understanding of the evolution of the dung beetle fauna of the New World. Other American genera with published cladistic analyses of their species are
<taxonomicName id="4C325B94BC432A5A1545FC5B3549FC75" authority="Weber, 1801" authorityName="Weber" authorityYear="1801" box="[588,839,959,983]" class="Insecta" family="Scarabaeidae" genus="Ateuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A1545FC5B34A1FC75" box="[588,687,959,983]" italics="true" pageId="23" pageNumber="966">Ateuchus</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A15B1FC5B3549FC75" author="Weber F" box="[696,839,959,983]" pageId="23" pageNumber="966" refId="ref16312" refString="Weber F. 1801. Observationes entomologicae, continentes novorum quae condidit generum characters et nuper detectarum specierum descriptiones. Kiel [&quot; Kiliae &quot;], Bibliopolii Academici Novi." type="book" year="1801">Weber, 1801</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFA35DE6BC432A5A145FFC5B35F5FC75" author="Kohlman B" box="[854,1019,959,983]" pageId="23" pageNumber="966" pagination="3 - 81" refId="ref15535" refString="Kohlman B. 1984. Biosistematica de las especies norteamericanas del genero Ateuchus (Coleoptera: Scarabaeidae: Scarabaeinae). Fol Entomol Mex. 60: 3 - 81." type="journal article" year="1984">Kohlman 1984</bibRefCitation>
; only North American species),
<taxonomicName id="4C325B94BC432A5A1674FC05353BFC5B" authority="Erichson, 1847" authorityName="Erichson" authorityYear="1847" box="[381,821,993,1017]" class="Insecta" family="Scarabaeidae" genus="Ontherus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A1674FC0537EEFC5B" box="[381,480,993,1017]" italics="true" pageId="23" pageNumber="966">Ontherus</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A16E3FC05349EFC5B" author="Erichson WF" box="[490,656,993,1017]" pageId="23" pageNumber="966" pagination="552 - 950" refId="ref14975" refString="Erichson WF, editor. 1847. Scarabaeides. In: Naturgeschichte der Insecten Deutschlands. Berlin: Nicolaische Buchhandlung; p. 552 - 950." type="book chapter" year="1847">Erichson, 1847</bibRefCitation>
(
<bibRefCitation id="EFA35DE6BC432A5A1596FC053522FC5B" author="Genier F" box="[671,812,993,1017]" pageId="23" pageNumber="966" pagination="1 - 170" refId="ref15070" refString="Genier F. 1996. A revision of the Neotropical genus Ontherus Erichson (Coleoptera: Scarabaeidae, Scarabaeinae). Mem Entomol Soc Can. 170: 1 - 170." type="journal article" year="1996">Génier 1996</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C325B94BC432A5A144AFC053236FC5B" authority="Harold, 1869" authorityName="Harold" authorityYear="1869" box="[835,1080,993,1017]" class="Insecta" family="Scarabaeidae" genus="Bdelyrus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A144AFC053593FC5B" box="[835,925,993,1017]" italics="true" pageId="23" pageNumber="966">Bdelyrus</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A14AEFC053236FC5B" author="Harold E" box="[935,1080,993,1017]" pageId="23" pageNumber="966" pagination="46 - 70" refId="ref15433" refString="Harold E. 1869. Ueber coprophage Lamellicornien mit besonderer Berucksichtigung der Parises Sammlungen. Coleopterol Hefte. 5: 46 - 70." type="journal article" year="1869">Harold, 1869</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFA35DE6BC432A5A1341FC0536D9FBBE" author="Cook J" pageId="23" pageNumber="966" pagination="631 - 689" refId="ref14381" refString="Cook J. 1998. A revision of the Neotropical genus Bdelyrus Harold (Coleoptera: Scarabaeidae). Can Entomol. 130: 631 - 689." type="journal article" year="1998">Cook 1998</bibRefCitation>
,
<bibRefCitation id="EFA35DE6BC432A5A17E1FBE03711FBBE" author="Cook J" box="[232,287,1028,1052]" pageId="23" pageNumber="966" pagination="551 - 565" refId="ref14409" refString="Cook J. 2000. Four new species of Bdelyrus (Coleoptera: Scarabaeidae: Scarabaeinae) and a redescription of Bdelyrus lagopus. Can Entomol. 132: 551 - 565." type="journal article" year="2000">2000</bibRefCitation>
),
<taxonomicName id="4C325B94BC432A5A163EFBE03524FBBE" authority="Balthasar, 1942" authorityName="Balthasar" authorityYear="1942" box="[311,810,1028,1052]" class="Insecta" family="Scarabaeidae" genus="Cryptocanthon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A163EFBE037D6FBBE" box="[311,472,1028,1052]" italics="true" pageId="23" pageNumber="966">Cryptocanthon</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A16EDFBE0349CFBBE" author="Balthasar V" box="[484,658,1028,1052]" pageId="23" pageNumber="966" pagination="36 - 44" refId="ref14043" refString="Balthasar V. 1942. Neue Scarabaeiden aus Sud-Amerika. C asopis C esk Spol Ent. 39: 36 - 44." type="journal article" year="1942">Balthasar, 1942</bibRefCitation>
(
<bibRefCitation id="EFA35DE6BC432A5A15AAFBE0352FFBBE" author="Cook J" box="[675,801,1028,1052]" pageId="23" pageNumber="966" pagination="3 - 96" refId="ref14442" refString="Cook J. 2002. A revision of the Neotropical genus Cryptocanthon Balthasar (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopt Soc Monogr. 1: 3 - 96." type="journal article" year="2002">Cook 2002</bibRefCitation>
)
</taxonomicName>
,
<taxonomicName id="4C325B94BC432A5A1430FBE03243FBBE" authority="Erichson, 1847" authorityName="Erichson" authorityYear="1847" box="[825,1101,1028,1052]" class="Insecta" family="Scarabaeidae" genus="Scatimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A1430FBE03595FBBE" box="[825,923,1028,1052]" italics="true" pageId="23" pageNumber="966">Scatimus</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A14AFFBE03243FBBE" author="Erichson WF" box="[934,1101,1028,1052]" pageId="23" pageNumber="966" pagination="552 - 950" refId="ref14975" refString="Erichson WF, editor. 1847. Scarabaeides. In: Naturgeschichte der Insecten Deutschlands. Berlin: Nicolaische Buchhandlung; p. 552 - 950." type="book chapter" year="1847">Erichson, 1847</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C325B94BC432A5A17A9FBC3345FFB9D" authority="Genier and Kohlmann, 2003" authorityName="Genier and Kohlmann" authorityYear="2003" box="[160,593,1063,1087]" class="Insecta" family="Scarabaeidae" genus="Scatrichus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A17A9FBC33700FB9D" box="[160,270,1063,1087]" italics="true" pageId="23" pageNumber="966">Scatrichus</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A161FFBC3345FFB9D" author="Genier F &amp; Kohlmann B" box="[278,593,1063,1087]" pageId="23" pageNumber="966" pagination="57 - 111" refId="ref15102" refString="Genier F, Kohlmann B. 2003. Revision of the Neotropical dung beetle genera Scatimus Erichson and Scatrichus gen. nov. (Coleoptera: Scarabaeidae: Scarabaeinae). Fabreries. 28: 57 - 111." type="journal article" year="2003">Génier and Kohlmann, 2003</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFA35DE6BC432A5A1557FBC33598FB9D" author="Genier F &amp; Kohlmann B" box="[606,918,1063,1087]" pageId="23" pageNumber="966" pagination="57 - 111" refId="ref15102" refString="Genier F, Kohlmann B. 2003. Revision of the Neotropical dung beetle genera Scatimus Erichson and Scatrichus gen. nov. (Coleoptera: Scarabaeidae: Scarabaeinae). Fabreries. 28: 57 - 111." type="journal article" year="2003">Génier and Kohlmann 2003</bibRefCitation>
),
<taxonomicName id="4C325B94BC432A5A14A3FBC336D9FBC3" authority="MacLeay, 1819" authorityName="MacLeay" authorityYear="1819" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A14A3FBC3321AFB9D" box="[938,1044,1063,1087]" italics="true" pageId="23" pageNumber="966">Phanaeus</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A1315FBC336D9FBC3" author="MacLeay WS" pageId="23" pageNumber="966" refId="ref15566" refString="MacLeay WS. 1819. Hor ӕ entomologic ӕ: or essays on the annulose animals. Vol. I. Part I. London: Bagster." type="book" year="1819">MacLeay, 1819</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFA35DE6BC432A5A17ECFBAD3753FBC3" author="Price DL" box="[229,349,1097,1121]" pageId="23" pageNumber="966" pagination="1 - 18" refId="ref15970" refString="Price DL. 2007. A phylogenetic analysis of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae) based on morphological data. Insect Syst Evol. 38: 1 - 18." type="journal article" year="2007">Price 2007</bibRefCitation>
,
<bibRefCitation id="EFA35DE6BC432A5A1662FBAD37ACFBC3" author="Price DL" box="[363,418,1097,1121]" pageId="23" pageNumber="966" pagination="137 - 150" refId="ref16004" refString="Price DL. 2009. Phylogeny and biogeography of the dung beetle genus Phanaeus (Coleoptera: Scarabaeidae). Syst Entomol. 34: 137 - 150." type="journal article" year="2009">2009</bibRefCitation>
) and
<taxonomicName id="4C325B94BC432A5A16EFFBAD34FEFBC3" authority="Arrow, 1932" authorityName="Arrow" authorityYear="1932" box="[486,752,1097,1121]" class="Insecta" family="Scarabaeidae" genus="Zonocopris" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="23" pageNumber="966" phylum="Arthropoda" rank="genus">
<emphasis id="B946FC05BC432A5A16EFFBAD3450FBC3" box="[486,606,1097,1121]" italics="true" pageId="23" pageNumber="966">Zonocopris</emphasis>
<bibRefCitation id="EFA35DE6BC432A5A156EFBAD34FEFBC3" author="Arrow GJ" box="[615,752,1097,1121]" pageId="23" pageNumber="966" pagination="223 - 226" refId="ref14016" refString="Arrow GJ. 1932. New species of lamellicorn beetles (subfam. Coprinae) from South America. Stylops. 1: 223 - 226." type="journal article" year="1932">Arrow, 1932</bibRefCitation>
</taxonomicName>
(
<bibRefCitation id="EFA35DE6BC432A5A15F7FBAD35DCFBC3" author="Vaz-de-Mello FZ" box="[766,978,1097,1121]" pageId="23" pageNumber="966" pagination="231 - 239" refId="ref16236" refString="Vaz-de-Mello FZ. 2007. Revision and phylogeny of the dung beetle genus Zonocopris Arrow, 1932 (Coleoptera: Scarabaeidae: Scarabaeinae), a phoretic of land snails. Ann Soc Entomol Fr. 43: 231 - 239." type="journal article" year="2007">Vaz-de-Mello 2007</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>
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