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<document id="F5F9316DBF95CAFBC0173028017CBE36" ID-CLB-Dataset="2773" ID-DOI="10.11646/zootaxa.5087.3.3" ID-GBIF-Dataset="ffd046ce-58d2-48e5-bdb4-310e24a02b4c" ID-ISSN="1175-5326" ID-Zenodo-Dep="5826878" ID-ZooBank="95583A37-2CD1-4C16-8902-512215B6CF9E" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1641539179045" checkinUser="plazi" docAuthor="Grebennikov, Vasily V." docDate="2022" docId="D71C072CFFCDFFB3E0C42E97FE3E04AB" docLanguage="en" docName="zootaxa.5087.3.3.pdf" docOrigin="Zootaxa 5087 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Melanophilharmostes Paulian 1968" docType="treatment" docVersion="5" lastPageNumber="445" masterDocId="2B257F54FFC9FFB6E0532810FF9B0505" masterDocTitle="Phylogenetic placement of a new Melanophilharmostes Paulian, 1968 pill scarab (Coleoptera: Hybosoridae: Ceratocanthinae) from Cameroon: molecular results decipher misleading morphology" masterLastPageNumber="450" masterPageNumber="441" pageNumber="445" updateTime="1699296579684" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
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<mods:title id="F1F632D71EBFAEF66837714AD6B97A2A">Phylogenetic placement of a new Melanophilharmostes Paulian, 1968 pill scarab (Coleoptera: Hybosoridae: Ceratocanthinae) from Cameroon: molecular results decipher misleading morphology</mods:title>
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<treatment id="D71C072CFFCDFFB3E0C42E97FE3E04AB" ID-DOI="http://doi.org/10.5281/zenodo.5826876" ID-GBIF-Taxon="191924648" ID-Zenodo-Dep="5826876" LSID="urn:lsid:plazi:treatment:D71C072CFFCDFFB3E0C42E97FE3E04AB" httpUri="http://treatment.plazi.org/id/D71C072CFFCDFFB3E0C42E97FE3E04AB" lastPageId="5" lastPageNumber="445" pageId="4" pageNumber="445">
<subSubSection id="17AFE5B1FFCDFFB2E0C42E97FDCB03A7" box="[151,592,1671,1698]" pageId="4" pageNumber="445" type="nomenclature">
<paragraph id="5F0AB63AFFCDFFB2E0C42E97FDCB03A7" blockId="4.[151,592,1671,1698]" box="[151,592,1671,1698]" pageId="4" pageNumber="445">
<heading id="04420156FFCDFFB2E0C42E97FDCB03A7" bold="true" box="[151,592,1671,1698]" fontSize="11" level="1" pageId="4" pageNumber="445" reason="1">
<taxonomicName id="98B5CDB9FFCDFFB2E0C42E97FDCB03A7" ID-CoL="5NHF" authority="Paulian, 1968" authorityName="Paulian" authorityYear="1968" box="[151,592,1671,1698]" class="Insecta" family="Hybosoridae" genus="Melanophilharmostes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="445" phylum="Arthropoda" rank="genus">
<emphasis id="6DC16A28FFCDFFB2E0C42E97FDCB03A7" bold="true" box="[151,592,1671,1698]" pageId="4" pageNumber="445">
<emphasis id="6DC16A28FFCDFFB2E0C42E97FE0403A7" bold="true" box="[151,415,1671,1698]" italics="true" pageId="4" pageNumber="445">Melanophilharmostes</emphasis>
Paulian, 1968
</emphasis>
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="17AFE5B1FFCDFFB3E0C42EC0FE3E04AB" lastPageId="5" lastPageNumber="446" pageId="4" pageNumber="445" type="type_taxon">
<paragraph id="5F0AB63AFFCDFFB2E0C42EC0FB7E03EF" blockId="4.[151,1436,1744,2023]" box="[151,1253,1744,1771]" pageId="4" pageNumber="445">
<emphasis id="6DC16A28FFCDFFB2E0C42EC0FEAF03EF" bold="true" box="[151,308,1744,1770]" pageId="4" pageNumber="445">
<typeStatus id="800E0898FFCDFFB2E0C42EC0FF4903EF" box="[151,210,1744,1770]" pageId="4" pageNumber="445">Type</typeStatus>
species:
</emphasis>
<taxonomicName id="98B5CDB9FFCDFFB2E16F2EC0FC4B03EF" ID-CoL="4G5ZR" authority="Paulian, 1968" authorityName="Paulian" authorityYear="1968" box="[316,976,1744,1771]" class="Insecta" family="Hybosoridae" genus="Philharmostes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="445" phylum="Arthropoda" rank="species" species="zicsii" subGenus="Melanophilharmostes">
<emphasis id="6DC16A28FFCDFFB2E16F2EC0FE4503EF" box="[316,478,1744,1770]" italics="true" pageId="4" pageNumber="445">Philharmostes</emphasis>
(
<emphasis id="6DC16A28FFCDFFB2E1BD2EC0FD7903EF" box="[494,738,1744,1770]" italics="true" pageId="4" pageNumber="445">Melanophilharmostes</emphasis>
)
<emphasis id="6DC16A28FFCDFFB2E2A12EC1FCB703EF" box="[754,812,1745,1770]" italics="true" pageId="4" pageNumber="445">zicsii</emphasis>
Paulian, 1968
</taxonomicName>
, by original designation.
</paragraph>
<paragraph id="5F0AB63AFFCDFFB2E0942EE4FD5E0237" blockId="4.[151,1436,1744,2023]" pageId="4" pageNumber="445">
<emphasis id="6DC16A28FFCDFFB2E0942EE4FDA0020B" bold="true" box="[199,571,1780,1806]" pageId="4" pageNumber="445">Composition and distribution:</emphasis>
17 named species (including the one described below) from sub-Saharan Africa and the Gulf of
<collectingCountry id="27A2F6AAFFCDFFB2E1C42F08FE730237" box="[407,488,1816,1842]" name="Guinea-Bissau" pageId="4" pageNumber="445">Guinea</collectingCountry>
island of
<collectingCountry id="27A2F6AAFFCDFFB2E2062F08FD5A0237" box="[597,705,1816,1842]" name="Equatorial Guinea" pageId="4" pageNumber="445">Annobón</collectingCountry>
.
</paragraph>
<paragraph id="5F0AB63AFFCDFFB3E0942F2CFE3E04AB" blockId="4.[151,1436,1744,2023]" lastBlockId="5.[151,1437,151,430]" lastPageId="5" lastPageNumber="446" pageId="4" pageNumber="445">
<emphasis id="6DC16A28FFCDFFB2E0942F2CFC230253" bold="true" box="[199,952,1852,1878]" pageId="4" pageNumber="445">Monophyly, sister group, and morphological synapomorphies:</emphasis>
the genus formed a moderately supported clade in both the morphology-based (Ballerio 2016) and DNA-based (
<figureCitation id="C78EAABFFFCDFFB2E3942F70FB97027E" box="[967,1036,1888,1915]" captionStart="FIGURE 5" captionStartId="8.[151,250,1591,1615]" captionTargetBox="[151,1431,464,1568]" captionTargetId="figure-149@8.[151,1435,453,1568]" captionTargetPageId="8" captionText="FIGURE 5. Maximum likelihood tree of Ceratocanthinae pill scarabs (Coleoptera: Hybosoridae) reconstructed by RAxML from the three-fragment concatenated matrix. Non-target clades are collapsed (see Grebennikov 2021 for their details). Except for the aberrant position of the Melanophilharmostes terminal 9604, M. tuber Grebennikov, new species, is inside monophyletic Melanophilharmostes, sister to monophyletic Pseudopterorthochaetes. Both genera are inside the monophyletic Synarmostes group of four genera. Habitus images are to scale; small black arrows indicate imaged specimens." figureDoi="http://doi.org/10.5281/zenodo.5826890" httpUri="https://zenodo.org/record/5826890/files/figure.png" pageId="4" pageNumber="445">Fig. 5</figureCitation>
) analyses. Its three morphological synapomorphies (Ballerio 2016) are (1) dorsal body surface with setae, (2) larger elytral punctures are “large horseshoe-shaped”, and (3) elytra with a lateral carina. In both analysis the genus formed either moderately (Ballerio 2016) or weakly-supported (
<figureCitation id="C78EAABFFFCDFFB2E1B02FDCFDB202E2" box="[483,553,1996,2023]" captionStart="FIGURE 5" captionStartId="8.[151,250,1591,1615]" captionTargetBox="[151,1431,464,1568]" captionTargetId="figure-149@8.[151,1435,453,1568]" captionTargetPageId="8" captionText="FIGURE 5. Maximum likelihood tree of Ceratocanthinae pill scarabs (Coleoptera: Hybosoridae) reconstructed by RAxML from the three-fragment concatenated matrix. Non-target clades are collapsed (see Grebennikov 2021 for their details). Except for the aberrant position of the Melanophilharmostes terminal 9604, M. tuber Grebennikov, new species, is inside monophyletic Melanophilharmostes, sister to monophyletic Pseudopterorthochaetes. Both genera are inside the monophyletic Synarmostes group of four genera. Habitus images are to scale; small black arrows indicate imaged specimens." figureDoi="http://doi.org/10.5281/zenodo.5826890" httpUri="https://zenodo.org/record/5826890/files/figure.png" pageId="4" pageNumber="445">Fig. 5</figureCitation>
) sister group to the genus
<taxonomicName id="98B5CDB9FFCDFFB2E33F2FDCFBE002E3" ID-CoL="63QQY" baseAuthorityName="Ballerio" baseAuthorityYear="2016" box="[876,1147,1996,2022]" class="Insecta" family="Hybosoridae" genus="Pseudopterorthochaetes" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="445" phylum="Arthropoda" rank="genus">
<emphasis id="6DC16A28FFCDFFB2E33F2FDCFBE002E3" box="[876,1147,1996,2022]" italics="true" pageId="4" pageNumber="445">Pseudopterorthochaetes</emphasis>
</taxonomicName>
; the only morphological synapomorphy of both genera is that the dorsal puncture on the head is shallow (not deep; Ballerio 2016). Two hind wing characters seem to additionally support the
<taxonomicName id="98B5CDB9FFCCFFB3E34628ABFB9205D0" ID-CoL="5NHF" authorityName="Paulian" authorityYear="1968" box="[789,1033,187,213]" class="Insecta" family="Hybosoridae" genus="Melanophilharmostes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="446" phylum="Arthropoda" rank="genus">
<emphasis id="6DC16A28FFCCFFB3E34628ABFB9205D0" box="[789,1033,187,213]" italics="true" pageId="5" pageNumber="446">Melanophilharmostes</emphasis>
</taxonomicName>
and
<taxonomicName id="98B5CDB9FFCCFFB3E41428ABFACD05D0" ID-CoL="63QQY" baseAuthorityName="Ballerio" baseAuthorityYear="2016" box="[1095,1366,187,213]" class="Insecta" family="Hybosoridae" genus="Pseudopterorthochaetes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="446" phylum="Arthropoda" rank="genus">
<emphasis id="6DC16A28FFCCFFB3E41428ABFACD05D0" box="[1095,1366,187,213]" italics="true" pageId="5" pageNumber="446">Pseudopterorthochaetes</emphasis>
</taxonomicName>
clade (presence of distal fork in vein CuA and presence of the distal expansion of vein MP 1+2; characters 15/1 and 16/1, respectively, in Ballerio 2016; see also
<figureCitation id="C78EAABFFFCCFFB3E2C42913FD6A0418" box="[663,753,259,285]" captionStart="FIGURE 2" captionStartId="4.[151,250,1530,1554]" captionTargetBox="[151,1435,183,1505]" captionTargetId="figure-32@4.[151,1435,181,1505]" captionTargetPageId="4" captionText="FIGURE 2. Melanophilharmostes tuber Grebennikov,new species, holotype, habitus (AC; right mesothoracic andmetathoracic legs removed), hind wing (D), antenna (E), and head (FG)." figureDoi="http://doi.org/10.5281/zenodo.5826884" httpUri="https://zenodo.org/record/5826884/files/figure.png" pageId="5" pageNumber="446">Fig. 2D</figureCitation>
); however, ambiguity remains due to the widespread loss of hind wings within this clade. The genus
<taxonomicName id="98B5CDB9FFCCFFB3E2352937FCEE0444" ID-CoL="63QQY" baseAuthorityName="Ballerio" baseAuthorityYear="2016" box="[614,885,295,321]" class="Insecta" family="Hybosoridae" genus="Pseudopterorthochaetes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="446" phylum="Arthropoda" rank="genus">
<emphasis id="6DC16A28FFCCFFB3E2352937FCEE0444" box="[614,885,295,321]" italics="true" pageId="5" pageNumber="446">Pseudopterorthochaetes</emphasis>
</taxonomicName>
was recovered either as a moderately-supported (Ballerio 2016) or strongly-supported clade (
<figureCitation id="C78EAABFFFCCFFB3E2D5295BFD5C0460" box="[646,711,330,357]" captionStart="FIGURE 5" captionStartId="8.[151,250,1591,1615]" captionTargetBox="[151,1431,464,1568]" captionTargetId="figure-149@8.[151,1435,453,1568]" captionTargetPageId="8" captionText="FIGURE 5. Maximum likelihood tree of Ceratocanthinae pill scarabs (Coleoptera: Hybosoridae) reconstructed by RAxML from the three-fragment concatenated matrix. Non-target clades are collapsed (see Grebennikov 2021 for their details). Except for the aberrant position of the Melanophilharmostes terminal 9604, M. tuber Grebennikov, new species, is inside monophyletic Melanophilharmostes, sister to monophyletic Pseudopterorthochaetes. Both genera are inside the monophyletic Synarmostes group of four genera. Habitus images are to scale; small black arrows indicate imaged specimens." figureDoi="http://doi.org/10.5281/zenodo.5826890" httpUri="https://zenodo.org/record/5826890/files/figure.png" pageId="5" pageNumber="446">Fig. 5</figureCitation>
) with a single morphological autapomorphy: the inner mesotibial apical spur in males is parallel to the tibial axis (not bent inwards at a right angle, as in
<taxonomicName id="98B5CDB9FFCCFFB3E42B297FFAF7048C" ID-CoL="5NHF" authorityName="Paulian" authorityYear="1968" box="[1144,1388,367,393]" class="Insecta" family="Hybosoridae" genus="Melanophilharmostes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="446" phylum="Arthropoda" rank="genus">
<emphasis id="6DC16A28FFCCFFB3E42B297FFAF7048C" box="[1144,1388,367,393]" italics="true" pageId="5" pageNumber="446">Melanophilharmostes</emphasis>
</taxonomicName>
and most other pill scarabs).
</paragraph>
</subSubSection>
</treatment>
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