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<document id="7B82F6E8FE9D4A5FDB2EC87181285DC5" ID-CLB-Dataset="22582" ID-DOI="10.3897/zookeys.34.342" ID-GBIF-Dataset="0981ff15-3ede-42a5-9475-980c31a88b2d" ID-ISSN="1313–2970" ID-Zenodo-Dep="576603" IM.metadata_requiresApprovalFor="plazi" IM.taxonomicNames_requiresApprovalFor="plazi" checkinTime="1588014235105" checkinUser="plazi" docAuthor="Krell, Frank &amp; Philips, T." docDate="2010" docId="03AE87BC330F152BDB8DFC6D784DDEB3" docLanguage="en" docName="ZK_article_2137.pdf" docOrigin="ZooKeys 34 (34)" docStyle="DocumentStylede.uka.ipd.idaho.easyIO.settings.Settings@359066bc" docStyleName="zookeys.2008.journal_article" docTitle="Haroldius Boucomont 1914" docType="treatment" docVersion="6" lastPageNumber="46" masterDocId="FF97FFC4330E152EDB01FFB27C44DD7B" masterDocTitle="Formicdubius Philips &amp; Scholtz from South Africa, a junior synonym of Haroldius Boucomont, and a survey of the trichomes in the African species (Coleoptera, Scarabaeidae, Onthophagini)" masterLastPageNumber="48" masterPageNumber="41" pageNumber="42" updateTime="1698837357546" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-3.0">
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<mods:title id="F910E147DD5DB1105D040630F98864F6">Formicdubius Philips &amp; Scholtz from South Africa, a junior synonym of Haroldius Boucomont, and a survey of the trichomes in the African species (Coleoptera, Scarabaeidae, Onthophagini)</mods:title>
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<mods:namePart id="4392B58568E58A344BB883A00790A95E">Krell, Frank</mods:namePart>
<mods:affiliation id="EFA7455C9620A4FA8C95010956E1803B">Denver Museum of Nature &amp; Science, Denver, United States of America</mods:affiliation>
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<mods:namePart id="3FB43FBEBBB79B8020BABA2E3E32ECEF">Philips, T.</mods:namePart>
<mods:affiliation id="8634CDFE42F328B1971777B7610D2E0B">Western Kentucky University, Bowling Green ,, United States of America</mods:affiliation>
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<mods:date id="FE29112E94E842DFA1A32077019C7E6F">2010</mods:date>
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<mods:identifier id="7C7D3BFB9B3E96B3B39A40FC3953BEE4" type="DOI">10.3897/zookeys.34.342</mods:identifier>
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<treatment id="03AE87BC330F152BDB8DFC6D784DDEB3" ID-DOI="http://doi.org/10.5281/zenodo.3789796" ID-GBIF-Taxon="163812682" ID-Zenodo-Dep="3789796" LSID="urn:lsid:plazi:treatment:03AE87BC330F152BDB8DFC6D784DDEB3" httpUri="http://treatment.plazi.org/id/03AE87BC330F152BDB8DFC6D784DDEB3" lastPageId="5" lastPageNumber="46" pageId="1" pageNumber="42">
<subSubSection id="C31D6521330F152FDB8DFC6D7FA7D853" pageId="1" pageNumber="42" type="nomenclature">
<paragraph id="8BB836AA330F152FDB8DFC6D7D9ADE81" blockId="1.[140,478,991,1018]" box="[140,478,991,1018]" pageId="1" pageNumber="42">
<heading id="D0F081C6330F152FDB8DFC6D7D9ADE81" box="[140,478,991,1018]" fontSize="11" level="2" pageId="1" pageNumber="42" reason="3">
<emphasis id="B973EAB8330F152FDB8DFC6D7D9ADE81" bold="true" box="[140,478,991,1018]" pageId="1" pageNumber="42">
African
<taxonomicName id="4C074D29330F152FDBF9FC6D7D30DE81" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[248,372,991,1018]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330F152FDBF9FC6D7D30DE81" bold="true" box="[248,372,991,1018]" italics="true" pageId="1" pageNumber="42">Haroldius</emphasis>
</taxonomicName>
species
</emphasis>
</heading>
</paragraph>
<paragraph id="8BB836AA330F152FDB8DFBA57FA7D853" blockId="1.[140,1108,1047,1671]" pageId="1" pageNumber="42">
All African species of
<taxonomicName id="4C074D29330F152FDA8DFBA57DBCD94A" authorityName="Boucomont" authorityYear="1914" box="[396,504,1047,1073]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330F152FDA8DFBA57DBCD94A" box="[396,504,1047,1073]" italics="true" pageId="1" pageNumber="42">Haroldius</emphasis>
</taxonomicName>
appear to have trichomes on the mesepimeron, the pronotal base, and the elytral base. Trichomes appear as distinct and relatively tight clusters of setae while others are looser or even single rows of thick setae such as those on the elytral base. Without microsections we are unable to decide whether those might be proper trichomes associated with glands or mechanoreceptors. However, as further evidence of a glandular association, longitudinal grooves at the pronotal base are also present that may facilitate the spreading of allomone secretions onto the pronotal surface, increasing the attractiveness of these beetles to their host ants.
</paragraph>
</subSubSection>
<subSubSection id="C31D6521330F152CDBB8FA827CB0DBF7" lastPageId="2" lastPageNumber="43" pageId="1" pageNumber="42" type="reference_group">
<paragraph id="8BB836AA330F152FDBB8FA827D22DBFC" blockId="1.[140,1108,1047,1671]" pageId="1" pageNumber="42">
<taxonomicName id="4C074D29330F152FDBB8FA827DF3D831" ID-CoL="6JHY9" authorityName="Philips &amp; Scholtz" authorityYear="2000" box="[185,439,1328,1354]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330F152FDBB8FA827DF3D831" box="[185,439,1328,1354]" italics="true" pageId="1" pageNumber="42">Formicdubius convexus</emphasis>
</taxonomicName>
(Figs 1–6) (and the very similar or conspecific
<taxonomicName id="4C074D29330F152FD8E9FA827DC4D815" ID-CoL="6LCGH" authority="Janssens, 1953" authorityName="Janssens" authorityYear="1953" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330F152FD8E9FA827C97D816" italics="true" pageId="1" pageNumber="42">Haroldius leleupi</emphasis>
<bibRefCitation id="EF964B5B330F152FDBD8FAE67DC4D815" author="Janssens A" box="[217,384,1363,1390]" pageId="1" pageNumber="42" pagination="344 - 347" refId="ref2860" refString="Janssens A (1953) Contribution a l'etude des Coleopteres Lamellicornes d'Afrique II. Especes nouvelles du genre Afroharoldius A. Janssens. Revue de Zoologie et de Botanique Africaines 47: 344 - 347." type="journal article" year="1953">Janssens, 1953</bibRefCitation>
</taxonomicName>
; Figs 11–13) can easily be distinguished from the remaining
<taxonomicName id="4C074D29330F152FDF26FAE67C91D8EA" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330F152FDF26FAE67C91D8EA" italics="true" pageId="1" pageNumber="42">Haroldius</emphasis>
</taxonomicName>
species by the presence of a distinct notch between the pronotum and the elytral base accommodating the pronotal trichomes (Figs 2, 12). In
<taxonomicName id="4C074D29330F152FD85CFA287D70D8AC" ID-CoL="3JRNG" authority="Janssens, 1949" authorityName="Janssens" authorityYear="1949" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="species" species="ennearthrus">
<emphasis id="B973EAB8330F152FD85CFA287810D8CF" box="[861,1108,1434,1460]" italics="true" pageId="1" pageNumber="42">Haroldius ennearthrus</emphasis>
Janssens, 1949
</taxonomicName>
(Figs 14–19), the basal margins of pronotum and elytra are straight with the pronotal margin slightly emarginate close to the edge (Figs 15, 18); a notch is missing, but a slight lateral depression towards the bases of the pronotum and elytra is visible.
<taxonomicName id="4C074D29330F152FDBE0F9957E56DB3A" ID-CoL="3JRNX" authority="Janssens, 1953" authorityName="Janssens" authorityYear="1953" box="[225,530,1574,1601]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="species" species="modestus">
<emphasis id="B973EAB8330F152FDBE0F9957D21DB3A" box="[225,357,1575,1601]" italics="true" pageId="1" pageNumber="42">H. modestus</emphasis>
<bibRefCitation id="EF964B5B330F152FDA6CF9957E56DB3A" author="Janssens A" box="[365,530,1574,1601]" pageId="1" pageNumber="42" pagination="344 - 347" refId="ref2860" refString="Janssens A (1953) Contribution a l'etude des Coleopteres Lamellicornes d'Afrique II. Especes nouvelles du genre Afroharoldius A. Janssens. Revue de Zoologie et de Botanique Africaines 47: 344 - 347." type="journal article" year="1953">Janssens, 1953</bibRefCitation>
</taxonomicName>
shares the general shape of the pronotal and elytral bases with
<taxonomicName id="4C074D29330F152FDA04F9F97DEDDB1F" ID-CoL="3JRNG" authorityName="Janssens" authorityYear="1949" box="[261,425,1610,1636]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="1" pageNumber="42" phylum="Arthropoda" rank="species" species="ennearthrus">
<emphasis id="B973EAB8330F152FDA04F9F97DEDDB1F" box="[261,425,1610,1636]" italics="true" pageId="1" pageNumber="42">H. ennearthrus</emphasis>
</taxonomicName>
but the pronotal margin is slightly more strongly emarginate laterally (Figs 8–9).
</paragraph>
<paragraph id="8BB836AA330C152CDB8DFA447CB0DBF7" blockId="2.[140,1108,1523,1676]" pageId="2" pageNumber="43">
<emphasis id="B973EAB8330C152CDB8DFA447D53DB77" bold="true" box="[140,279,1523,1550]" pageId="2" pageNumber="43">Figures Ι–6.</emphasis>
<taxonomicName id="4C074D29330C152CDA21FA447D8ADB77" ID-CoL="6LCGG" authorityName="Krell &amp; Philips" authorityYear="2010" baseAuthorityName="Philips &amp; Scholtz" baseAuthorityYear="2000" box="[288,462,1526,1548]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="2" pageNumber="43" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330C152CDA21FA447D8ADB77" box="[288,462,1526,1548]" italics="true" pageId="2" pageNumber="43">Haroldius convexus</emphasis>
</taxonomicName>
:
<emphasis id="B973EAB8330C152CDADDFA447DADDB77" bold="true" box="[476,489,1526,1548]" pageId="2" pageNumber="43">Ι</emphasis>
Dorsal habitus
<emphasis id="B973EAB8330C152CD984FA447ED6DB77" bold="true" box="[645,658,1526,1548]" pageId="2" pageNumber="43">2</emphasis>
Trichomes within cleft between prothorax and mesothorax and showing pronotal sulci
<emphasis id="B973EAB8330C152CDAF8F9A47E42DB57" bold="true" box="[505,518,1558,1580]" pageId="2" pageNumber="43">3</emphasis>
SEM (scanning electron microscope) view of pronotal trichome
<emphasis id="B973EAB8330C152CDB8DF9847CDDDB37" bold="true" box="[140,153,1590,1612]" pageId="2" pageNumber="43">4</emphasis>
SEM view of pronotal trichome showing possibly ant-gnawed tips of the setae
<emphasis id="B973EAB8330C152CD879F9847FC1DB37" bold="true" box="[888,901,1590,1612]" pageId="2" pageNumber="43">5</emphasis>
SEM view of possible trichomes on the elytral base
<emphasis id="B973EAB8330C152CDAA2F9E47DF4DB17" bold="true" box="[419,432,1622,1644]" pageId="2" pageNumber="43">6</emphasis>
SEM view of trichome on the mesepisternum and additional setal row proximally.
</paragraph>
</subSubSection>
<subSubSection id="C31D6521330D152DDBB8FF157D5DD806" pageId="3" pageNumber="44" type="description">
<paragraph id="8BB836AA330D152DDBB8FF157F0BDC0A" blockId="3.[140,1108,167,1284]" pageId="3" pageNumber="44">The trichomes on these species are located on the mesepisternum anterolaterally, the elytra anterolaterally (on the vertical surface facing the pronotum), and on the pronotal base posterolaterally. One should be aware that these setae may be damaged from the effects of gnawing by their host ants and the actual number of setae may vary in number more than we report due to our limited sample sizes. Regardless, the degree of development of these structures in each species is as follows:</paragraph>
<paragraph id="8BB836AA330D152DDBB8FEC87EF3DF59" blockId="3.[140,1108,167,1284]" pageId="3" pageNumber="44">
Specimens of
<taxonomicName id="4C074D29330D152DDA62FEC97E4BDCEF" authorityName="Janssens" authorityYear="1949" box="[355,527,378,404]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="ennearthrus">
<emphasis id="B973EAB8330D152DDA62FEC97E4BDCEF" box="[355,527,378,404]" italics="true" pageId="3" pageNumber="44">H. ennearthrus</emphasis>
</taxonomicName>
possess a large rounded cluster of setae on the mesepisternum (Fig. 19) and no additional setae proximally. They also appear to have two short and relatively thick setae on the elytral base. Three to five thick setae are present on the pronotum (Figs 16–18) as well as a row of similar aligned setae proximal to these, all of which may be trichomes (Fig. 16).
</paragraph>
<paragraph id="8BB836AA330D152DDBB8FD987FB3DFD5" blockId="3.[140,1108,167,1284]" pageId="3" pageNumber="44">
<taxonomicName id="4C074D29330D152DDBB8FD987DCBDF3F" box="[185,399,554,580]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="modestus">
<emphasis id="B973EAB8330D152DDBB8FD987DCBDF3F" box="[185,399,554,580]" italics="true" pageId="3" pageNumber="44">Haroldius modestus</emphasis>
</taxonomicName>
(Figs 7–10) also has a large, rounded cluster of setae on the mesepisternum (Fig. 10) and lacks any thick setae proximally. There are two to three thick setae on the elytral base. On the pronotum a brush-like trichome is well developed, consisting of six to sometimes ten or more, long, thick setae (Figs 8–9).
</paragraph>
<paragraph id="8BB836AA330D152DDBB8FD057D27DE91" blockId="3.[140,1108,167,1284]" pageId="3" pageNumber="44">
The third described
<taxonomicName id="4C074D29330D152DDAADFD057E5FDFAA" authorityName="Boucomont" authorityYear="1914" box="[428,539,695,721]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DDAADFD057E5FDFAA" box="[428,539,695,721]" italics="true" pageId="3" pageNumber="44">Haroldius</emphasis>
</taxonomicName>
species from Africa,
<taxonomicName id="4C074D29330D152DD81AFD057FC9DFAA" authorityName="Janssens" authorityYear="1953" box="[795,909,695,721]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330D152DD81AFD057FC9DFAA" box="[795,909,695,721]" italics="true" pageId="3" pageNumber="44">H. leleupi</emphasis>
</taxonomicName>
(Figs 11–13), is strikingly similar to
<taxonomicName id="4C074D29330D152DDA79FD687E3DDF8F" authorityName="Philips &amp; Scholtz" authorityYear="2000" box="[376,633,730,756]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330D152DDA79FD687E3DDF8F" box="[376,633,730,756]" italics="true" pageId="3" pageNumber="44">Formicdubius convexus</emphasis>
</taxonomicName>
(Figs 1–6). They both have a small cluster of stout setae on the mesepisternum (Figs 6, 13) and proximal to this are a few more thick and elongate setae arranged in a transverse row (Fig. 6). These same setae become more slender and slightly more widely separated the further from the mesepisternal trichome. Additionally, there are four to five short, thick setae at the base of the elytra near the lateral margin (Fig. 5). Both species also have a well developed trichome on the pronotal base consisting of 20 or more lobe shaped setae (Figs 3–4, 12–13).
</paragraph>
<paragraph id="8BB836AA330D152DDBB8FC467E51D87F" blockId="3.[140,1108,167,1284]" pageId="3" pageNumber="44">
We are unsure whether
<taxonomicName id="4C074D29330D152DDAD1FC467E7AD976" authorityName="Janssens" authorityYear="1953" box="[464,574,1011,1037]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330D152DDAD1FC467E7AD976" box="[464,574,1011,1037]" italics="true" pageId="3" pageNumber="44">H. leleupi</emphasis>
</taxonomicName>
and
<taxonomicName id="4C074D29330D152DD97CFC467EBCD976" authorityName="Philips &amp; Scholtz" authorityYear="2000" box="[637,760,1012,1037]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330D152DD97CFC467EBCD976" box="[637,760,1012,1037]" italics="true" pageId="3" pageNumber="44">F. convexus</emphasis>
</taxonomicName>
are distinct taxa.
<taxonomicName id="4C074D29330D152DD8CBFC46787DD976" authorityName="Janssens" authorityYear="1953" box="[970,1081,1011,1037]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330D152DD8CBFC46787DD976" box="[970,1081,1011,1037]" italics="true" pageId="3" pageNumber="44">H. leleupi</emphasis>
</taxonomicName>
is generally larger; the five specimens studied measure
<quantity id="4CFF9B4F330D152DD9D9FBA57FC7D94A" box="[728,899,1047,1073]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="2.2" metricValueMin="1.8" pageId="3" pageNumber="44" unit="mm" value="2.0" valueMax="2.2" valueMin="1.8">1.80–2.20 mm</quantity>
in length (average
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), whereas the 14 sampled
<typeStatus id="54BC8808330D152DD921FB887E1ED92F" box="[544,602,1082,1108]" pageId="3" pageNumber="44">types</typeStatus>
of
<taxonomicName id="4C074D29330D152DD97EFB897EB2D92F" authorityName="Philips &amp; Scholtz" authorityYear="2000" box="[639,758,1083,1108]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330D152DD97EFB897EB2D92F" box="[639,758,1083,1108]" italics="true" pageId="3" pageNumber="44">F. convexus</emphasis>
</taxonomicName>
measure
<quantity id="4CFF9B4F330D152DD860FB88784FD92F" box="[865,1035,1082,1108]" metricMagnitude="-3" metricUnit="m" metricValue="1.765" metricValueMax="1.9" metricValueMin="1.63" pageId="3" pageNumber="44" unit="mm" value="1.765" valueMax="1.9" valueMin="1.63">1.63–1.90 mm</quantity>
(average
<quantity id="4CFF9B4F330D152DDBB9FBEF7D61D903" box="[184,293,1117,1144]" metricMagnitude="-3" metricUnit="m" metricValue="1.78" pageId="3" pageNumber="44" unit="mm" value="1.78">1.78 mm</quantity>
).
<taxonomicName id="4C074D29330D152DDA39FBEF7DB6D90C" authorityName="Janssens" authorityYear="1953" box="[312,498,1117,1143]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330D152DDA39FBEF7DB6D90C" box="[312,498,1117,1143]" italics="true" pageId="3" pageNumber="44">Haroldius leleupi</emphasis>
</taxonomicName>
also has slightly stronger (more clearly defined) punctures on the disk of the pronotum and the elytral intervals and slightly deeper elytral striae, particularly near the suture. Additional material will be required from localities between the
<collectingCountry id="F310763A330D152DDA1AFB757D2CD99A" box="[283,360,1223,1249]" name="Democratic Republic of the Congo" pageId="3" pageNumber="44">Congo</collectingCountry>
and
<collectingCountry id="F310763A330D152DDAA0FB757E74D99A" box="[417,560,1223,1249]" name="South Africa" pageId="3" pageNumber="44">South Africa</collectingCountry>
to decide to what extent these minor differences indicate taxonomic differentiation.
</paragraph>
<paragraph id="8BB836AA330D152DDB8DFAD07D5DD806" blockId="3.[140,281,1378,1405]" box="[140,281,1378,1405]" pageId="3" pageNumber="44">
<emphasis id="B973EAB8330D152DDB8DFAD07D5DD806" bold="true" box="[140,281,1378,1405]" pageId="3" pageNumber="44">Synonymy</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C31D6521330D152ADB8DFA287E2CDB1F" lastPageId="4" lastPageNumber="45" pageId="3" pageNumber="44" type="reference_group">
<paragraph id="8BB836AA330D152DDB8DFA287ECBDB1F" blockId="3.[140,1108,1434,1636]" pageId="3" pageNumber="44">
<taxonomicName id="4C074D29330D152DDB8DFA287D0CD8CF" ID-CoL="6LCGH" authorityName="Janssens" authorityYear="1953" box="[140,328,1434,1460]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330D152DDB8DFA287D0CD8CF" box="[140,328,1434,1460]" italics="true" pageId="3" pageNumber="44">Haroldius leleupi</emphasis>
</taxonomicName>
and
<taxonomicName id="4C074D29330D152DDA85FA287EC5D8CF" ID-CoL="6JHY9" authorityName="Philips &amp; Scholtz" authorityYear="2000" box="[388,641,1434,1460]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330D152DDA85FA287EC5D8CF" box="[388,641,1434,1460]" italics="true" pageId="3" pageNumber="44">Formicdubius convexus</emphasis>
</taxonomicName>
are difficult to distinguish at the species level, but there is no doubt that
<taxonomicName id="4C074D29330D152DD90DFA0F7EE5D8AC" ID-CoL="63535" authorityName="Philips and Scholtz" authorityYear="2000" box="[524,673,1469,1495]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DD90DFA0F7EE5D8AC" box="[524,673,1469,1495]" italics="true" pageId="3" pageNumber="44">Formicdubius</emphasis>
</taxonomicName>
is identical with
<taxonomicName id="4C074D29330D152DD86FFA0F7F9ED8AC" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[878,986,1469,1495]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DD86FFA0F7F9ED8AC" box="[878,986,1469,1495]" italics="true" pageId="3" pageNumber="44">Haroldius</emphasis>
</taxonomicName>
at the generic level.
<taxonomicName id="4C074D29330D152DDA15FA527DEDD881" ID-CoL="63535" authorityName="Philips and Scholtz" authorityYear="2000" box="[276,425,1504,1530]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DDA15FA527DEDD881" box="[276,425,1504,1530]" italics="true" pageId="3" pageNumber="44">Formicdubius</emphasis>
</taxonomicName>
fully matches
<bibRefCitation id="EF964B5B330D152DD958FA527F02D881" author="Boucomont A" box="[601,838,1504,1530]" pageId="3" pageNumber="44" pagination="238 - 350" refId="ref2682" refString="Boucomont A (1914) Les coprophages de l'Archipel Malais. Annales de la Societe Entomologique de France 83: 238 - 350." type="journal article" year="1914">Boucomont’s (1914)</bibRefCitation>
original description of
<taxonomicName id="4C074D29330D152DDB8DF9B17CBCDB66" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[140,248,1539,1565]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DDB8DF9B17CBCDB66" box="[140,248,1539,1565]" italics="true" pageId="3" pageNumber="44">Haroldius</emphasis>
</taxonomicName>
. It also agrees with the extended diagnosis of
<taxonomicName id="4C074D29330D152DD80FF9B17F3EDB66" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[782,890,1539,1565]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DD80FF9B17F3EDB66" box="[782,890,1539,1565]" italics="true" pageId="3" pageNumber="44">Haroldius</emphasis>
</taxonomicName>
recently published by
<bibRefCitation id="EF964B5B330D152DDBAFF9957E4ADB39" author="Krikken J &amp; Huijbregts J" box="[174,526,1575,1602]" pageId="3" pageNumber="44" pagination="167 - 187" refId="ref2896" refString="Krikken J, Huijbregts J (2006) Miniature scarabs of the genus Haroldius on Sulawesi, with notes on their relatives (Coleoptera: Scarabaeidae). Tijdschrift voor Entomologie 149: 167 - 187." type="journal article" year="2006">Krikken and Huijbregts (2006)</bibRefCitation>
with the exception that all African species currently subsumed under
<taxonomicName id="4C074D29330D152DDA4FF9F87DFEDB1F" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[334,442,1610,1636]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="44" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330D152DDA4FF9F87DFEDB1F" box="[334,442,1610,1636]" italics="true" pageId="3" pageNumber="44">Haroldius</emphasis>
</taxonomicName>
possess trichomes.
</paragraph>
<paragraph id="8BB836AA330A152ADB8DFA797E2CDB1F" blockId="4.[140,1108,1479,1636]" pageId="4" pageNumber="45">
<emphasis id="B973EAB8330A152ADB8DFA797D65D89A" bold="true" box="[140,289,1479,1506]" pageId="4" pageNumber="45">Figures 7–Ι3.</emphasis>
<taxonomicName id="4C074D29330A152ADA29FA797DC4D89B" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[296,384,1483,1504]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="45" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330A152ADA29FA797DC4D89B" box="[296,384,1483,1504]" italics="true" pageId="4" pageNumber="45">Haroldius</emphasis>
</taxonomicName>
species. Figures
<emphasis id="B973EAB8330A152AD918FA797E12D89A" bold="true" box="[537,598,1479,1506]" pageId="4" pageNumber="45">7–Ι0.</emphasis>
<taxonomicName id="4C074D29330A152AD95DFA7E7E8DD89B" ID-CoL="3JRNX" box="[604,713,1483,1505]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="45" phylum="Arthropoda" rank="species" species="modestus">
<emphasis id="B973EAB8330A152AD95DFA7E7E8DD89B" box="[604,713,1483,1505]" italics="true" pageId="4" pageNumber="45">H. modestus</emphasis>
</taxonomicName>
<emphasis id="B973EAB8330A152AD9CEFA797E98D89A" bold="true" box="[719,732,1483,1505]" pageId="4" pageNumber="45">7</emphasis>
Dorsal habitus
<emphasis id="B973EAB8330A152AD871FA797F39D89A" bold="true" box="[880,893,1483,1505]" pageId="4" pageNumber="45">8</emphasis>
Trichomes within cleft between prothorax and mesothorax and showing pronotal sulci
<emphasis id="B973EAB8330A152AD9EAFA597EBCDB7A" bold="true" box="[747,760,1515,1537]" pageId="4" pageNumber="45">9</emphasis>
Pronotal trichome
<emphasis id="B973EAB8330A152AD8B5FA597F8ADB7A" bold="true" box="[948,974,1515,1537]" pageId="4" pageNumber="45">Ι0</emphasis>
SEM view of trichomes on the pronotum and mesepisternum. Figures
<emphasis id="B973EAB8330A152AD9ADF9BC7EB2DB5F" bold="true" box="[684,758,1546,1573]" pageId="4" pageNumber="45">ΙΙ–Ι3.</emphasis>
<taxonomicName id="4C074D29330A152AD9FEF9BC7FDDDB58" ID-CoL="6LCGH" authorityName="Janssens" authorityYear="1953" box="[767,921,1550,1571]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="45" phylum="Arthropoda" rank="species" species="leleupi">
<emphasis id="B973EAB8330A152AD9FEF9BC7FDDDB58" box="[767,921,1550,1571]" italics="true" pageId="4" pageNumber="45">Haroldius leleupi</emphasis>
</taxonomicName>
:
<emphasis id="B973EAB8330A152AD8A6F9BC7F85DB5F" bold="true" box="[935,961,1550,1572]" pageId="4" pageNumber="45">ΙΙ</emphasis>
Dorsal habitus
<emphasis id="B973EAB8330A152ADB8DF99C7CE2DB3F" bold="true" box="[140,166,1582,1604]" pageId="4" pageNumber="45">Ι2</emphasis>
Trichomes within cleft between prothorax and mesothorax and showing pronotal sulci
<emphasis id="B973EAB8330A152AD8D2F99C7FA9DB3F" bold="true" box="[979,1005,1582,1604]" pageId="4" pageNumber="45">Ι3</emphasis>
SEM view of trichomes on the pronotum and mesepisternum.
</paragraph>
</subSubSection>
<subSubSection id="C31D6521330B152BDBB8FF157E1BDDBA" box="[185,607,167,193]" pageId="5" pageNumber="46" type="description">
<paragraph id="8BB836AA330B152BDBB8FF157E1BDDBA" blockId="5.[140,1108,167,968]" box="[185,607,167,193]" pageId="5" pageNumber="46">We propose the following synonymy:</paragraph>
</subSubSection>
<subSubSection id="C31D6521330B152BDBB8FF78784DDEB3" pageId="5" pageNumber="46" type="reference_group">
<paragraph id="8BB836AA330B152BDBB8FF787ECADC7C" blockId="5.[140,1108,167,968]" pageId="5" pageNumber="46">
<taxonomicName id="4C074D29330B152BDBB8FF787E47DD9F" ID-CoL="4TRW" authority="Boucomont, 1914" authorityName="Boucomont" authorityYear="1914" box="[185,515,202,228]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDBB8FF787D61DD9F" box="[185,293,202,228]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
<bibRefCitation id="EF964B5B330B152BDA2EFF787E47DD9F" author="Boucomont A" box="[303,515,202,228]" pageId="5" pageNumber="46" pagination="238 - 350" refId="ref2682" refString="Boucomont A (1914) Les coprophages de l'Archipel Malais. Annales de la Societe Entomologique de France 83: 238 - 350." type="journal article" year="1914">Boucomont, 1914</bibRefCitation>
</taxonomicName>
(
<typeStatus id="54BC8808330B152BD914FF787E03DD9F" box="[533,583,202,228]" pageId="5" pageNumber="46">type</typeStatus>
species by subsequent designation by
<bibRefCitation id="EF964B5B330B152BDF06FF787C8EDC7C" author="Arrow GJ" pageId="5" pageNumber="46" refId="ref2640" refString="Arrow GJ (1931) The Fauna of British India including Ceylon and Burma, Coleoptera, Lamellicornia, Part III. (Coprinae). Taylor and Francis, London, 428 pp., 13 pls, 1 map." type="book" year="1931">Arrow, 1931</bibRefCitation>
:
<taxonomicName id="4C074D29330B152BDBD4FF5F7EC2DC7C" ID-CoL="3JRP7" authority="Boucomont, 1914" authorityName="Boucomont" authorityYear="1914" box="[213,646,237,264]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="species" species="rugatulus">
<emphasis id="B973EAB8330B152BDBD4FF5F7DE9DC7C" box="[213,429,237,263]" italics="true" pageId="5" pageNumber="46">Haroldius rugatulus</emphasis>
<bibRefCitation id="EF964B5B330B152BDAB4FF5C7EC2DC7C" author="Boucomont A" box="[437,646,237,264]" pageId="5" pageNumber="46" pagination="238 - 350" refId="ref2682" refString="Boucomont A (1914) Les coprophages de l'Archipel Malais. Annales de la Societe Entomologique de France 83: 238 - 350." type="journal article" year="1914">Boucomont, 1914</bibRefCitation>
</taxonomicName>
)
</paragraph>
<paragraph id="8BB836AA330B152BDBB8FEA37E9FDC35" blockId="5.[140,1108,167,968]" pageId="5" pageNumber="46">
=
<taxonomicName id="4C074D29330B152BDBCEFEA27E3EDC51" ID-CoL="63535" authority="Philips &amp; Scholtz, 2000" authorityName="Philips &amp; Scholtz" authorityYear="2000" box="[207,634,272,298]" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDBCEFEA27D20DC51" box="[207,356,272,298]" italics="true" pageId="5" pageNumber="46">Formicdubius</emphasis>
<bibRefCitation id="EF964B5B330B152BDA6AFEA27E3EDC51" author="Philips TK &amp; Scholtz CH" box="[363,634,272,298]" pageId="5" pageNumber="46" pagination="227 - 231" refId="ref3062" refString="Philips TK, Scholtz CH (2000) A new genus and species of trichome-bearing dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) from South Africa. African Entomology 8: 227 - 231." type="journal article" year="2000">Philips &amp; Scholtz, 2000</bibRefCitation>
</taxonomicName>
(
<typeStatus id="54BC8808330B152BD988FEA27EFFDC51" box="[649,699,272,298]" pageId="5" pageNumber="46">type</typeStatus>
species by original designation:
<taxonomicName id="4C074D29330B152BDF24FEA37E39DC35" ID-CoL="6JHY9" authority="Philips &amp; Scholtz, 2000" authorityName="Philips &amp; Scholtz" authorityYear="2000" class="Insecta" family="Scarabaeidae" genus="Formicdubius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="species" species="convexus">
<emphasis id="B973EAB8330B152BDF24FEA37D25DC36" italics="true" pageId="5" pageNumber="46">Formicdubius convexus</emphasis>
<bibRefCitation id="EF964B5B330B152BDA69FE817E39DC35" author="Philips TK &amp; Scholtz CH" box="[360,637,307,334]" pageId="5" pageNumber="46" pagination="227 - 231" refId="ref3062" refString="Philips TK, Scholtz CH (2000) A new genus and species of trichome-bearing dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) from South Africa. African Entomology 8: 227 - 231." type="journal article" year="2000">Philips &amp; Scholtz, 2000</bibRefCitation>
</taxonomicName>
), syn. n.
</paragraph>
<paragraph id="8BB836AA330B152BDBB8FEE57F2CDC0A" blockId="5.[140,1108,167,968]" box="[185,872,343,369]" pageId="5" pageNumber="46">Consequently, the following new combination is established:</paragraph>
<paragraph id="8BB836AA330B152BDBB8FEC87860DCEF" blockId="5.[140,1108,167,968]" box="[185,1060,378,404]" pageId="5" pageNumber="46">
<taxonomicName id="4C074D29330B152BDBB8FEC87FF0DCEF" ID-CoL="6LCGG" authority="(Philips &amp; Scholtz, 2000) Krell &amp; Philips, 2010" authorityName="Krell &amp; Philips" authorityYear="2010" baseAuthorityName="Philips &amp; Scholtz" baseAuthorityYear="2000" box="[185,948,378,404]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="species" species="convexus" status="comb. nov.">
<emphasis id="B973EAB8330B152BDBB8FEC87DCEDCEF" box="[185,394,378,404]" italics="true" pageId="5" pageNumber="46">Haroldius convexus</emphasis>
(
<bibRefCitation id="EF964B5B330B152BDA98FEC87EE8DCEF" author="Philips TK &amp; Scholtz CH" box="[409,684,378,404]" pageId="5" pageNumber="46" pagination="227 - 231" refId="ref3062" refString="Philips TK, Scholtz CH (2000) A new genus and species of trichome-bearing dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae) from South Africa. African Entomology 8: 227 - 231." type="journal article" year="2000">Philips &amp; Scholtz, 2000</bibRefCitation>
) Krell &amp; Philips, 2010
</taxonomicName>
,
<taxonomicNameLabel id="A24057C3330B152BD8C1FEC87860DCEF" box="[960,1060,378,404]" pageId="5" pageNumber="46" rank="species">comb. n.</taxonomicNameLabel>
</paragraph>
<paragraph id="8BB836AA330B152BDBB8FE2C784DDEB3" blockId="5.[140,1108,167,968]" pageId="5" pageNumber="46">
We refrain from determining the taxonomic status of the African species of the
<emphasis id="B973EAB8330B152BDB8DFE727DD6DCA1" box="[140,402,448,474]" italics="true" pageId="5" pageNumber="46">
<taxonomicName id="4C074D29330B152BDB8DFE727CB7DCA1" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[140,243,448,474]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">Haroldius</taxonomicName>
/
<taxonomicName id="4C074D29330B152BDBFFFE727DD6DCA1" ID-CoL="Q8C" authorityName="Janssens" authorityYear="1949" box="[254,402,448,474]" class="Insecta" family="Scarabaeidae" genus="Afroharoldius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">Afroharoldius</taxonomicName>
</emphasis>
group. All African
<taxonomicName id="4C074D29330B152BD96FFE727E9EDCA1" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[622,730,448,474]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD96FFE727E9EDCA1" box="[622,730,448,474]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
</taxonomicName>
species have more or less distinct trichomes which could be a synapomorphy justifying a genus or subgenus
<taxonomicName id="4C074D29330B152BD8E2FE517CFCDF5A" ID-CoL="Q8C" authorityName="Janssens" authorityYear="1949" class="Insecta" family="Scarabaeidae" genus="Afroharoldius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD8E2FE517CFCDF5A" italics="true" pageId="5" pageNumber="46">Afroharoldius</emphasis>
</taxonomicName>
. However, we neither know if trichomes (vestigial or distinctive) are common in Asian
<taxonomicName id="4C074D29330B152BDBD2FD987D7BDF3F" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[211,319,554,580]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDBD2FD987D7BDF3F" box="[211,319,554,580]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
</taxonomicName>
species, or whether the trichome-bearing species are the sister group of the trichome-less
<emphasis id="B973EAB8330B152BDA73FDFF7DA1DF1C" box="[370,485,589,615]" italics="true" pageId="5" pageNumber="46">
<taxonomicName id="4C074D29330B152BDA73FDFF7DA5DF1C" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[370,481,589,615]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">Haroldius</taxonomicName>
,
</emphasis>
or whether their exclusion from
<taxonomicName id="4C074D29330B152BD85AFDFF7F83DF1C" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[859,967,589,615]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD85AFDFF7F83DF1C" box="[859,967,589,615]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
</taxonomicName>
would leave
<taxonomicName id="4C074D29330B152BDB8DFDC27CBCDFF1" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[140,248,624,650]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDB8DFDC27CBCDFF1" box="[140,248,624,650]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
</taxonomicName>
or
<taxonomicName id="4C074D29330B152BDA21FDC27DF7DFF1" ID-CoL="Q8C" authorityName="Janssens" authorityYear="1949" box="[288,435,624,650]" class="Insecta" family="Scarabaeidae" genus="Afroharoldius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDA21FDC27DF7DFF1" box="[288,435,624,650]" italics="true" pageId="5" pageNumber="46">Afroharoldius</emphasis>
</taxonomicName>
paraphyletic. In
<taxonomicName id="4C074D29330B152BD974FDC2780BDFF1" ID-CoL="3JRN8" authority="Krikken &amp; Huijbregts" authorityName="Krikken &amp; Huijbregts" authorityYear="2009" box="[629,1103,624,650]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="species" species="brendelli">
<emphasis id="B973EAB8330B152BD974FDC27F0CDFF1" box="[629,840,624,650]" italics="true" pageId="5" pageNumber="46">Haroldius brendelli</emphasis>
Krikken &amp; Huijbregts
</taxonomicName>
, antehumeral elytral trichomes are described (
<bibRefCitation id="EF964B5B330B152BD998FD217FBFDFD5" author="Krikken J &amp; Huijbregts J" box="[665,1019,659,686]" pageId="5" pageNumber="46" pagination="257 - 266" refId="ref2933" refString="Krikken J, Huijbregts J (2009) A new Haroldius Boucomont and a new Tesserodon Hope from the Moluccas (Coleoptera: Scarabaeidae: Scarabaeinae). Koleopterologische Rundschau 79: 257 - 266." type="journal article" year="2009">Krikken and Huijbregts 2009</bibRefCitation>
). Based on the illustrations in
<bibRefCitation id="EF964B5B330B152BDA8CFD057EB2DFA9" author="Krikken J &amp; Huijbregts J" box="[397,758,695,722]" pageId="5" pageNumber="46" pagination="167 - 187" refId="ref2896" refString="Krikken J, Huijbregts J (2006) Miniature scarabs of the genus Haroldius on Sulawesi, with notes on their relatives (Coleoptera: Scarabaeidae). Tijdschrift voor Entomologie 149: 167 - 187." type="journal article" year="2006">Krikken and Huijbregts (2006)</bibRefCitation>
, it appears as though pronotal grooves are present on at least some Asian species and, moreover, trichomes seem to be present in
<taxonomicName id="4C074D29330B152BDA00FD4C7EC2DE6C" ID-CoL="3JRPB" authority="Krikken &amp; Huijbregts" authorityName="Krikken &amp; Huijbregts" authorityYear="2006" box="[257,646,765,791]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="species" species="tangkoko">
<emphasis id="B973EAB8330B152BDA00FD4C7DC2DE6C" box="[257,390,765,791]" italics="true" pageId="5" pageNumber="46">H. tangkoko</emphasis>
Krikken &amp; Huijbregts
</taxonomicName>
and
<taxonomicName id="4C074D29330B152BD9BCFD4C7817DE6C" ID-CoL="6LCGF" authority="Krikken &amp; Huijbregts" authorityName="Krikken &amp; Huijbregts" authorityYear="2006" box="[701,1107,765,791]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="species" species="cambeforti">
<emphasis id="B973EAB8330B152BD9BCFD4C7F17DE6C" box="[701,851,765,791]" italics="true" pageId="5" pageNumber="46">H. cambeforti</emphasis>
Krikken &amp; Huijbregts
</taxonomicName>
(see their figs 5 and 6). A revision and phylogenetic analysis of the whole group, including species of
<taxonomicName id="4C074D29330B152BDA5DFCF17E02DE26" ID-CoL="6MCM" authority="Paulian" authorityName="Paulian" authorityYear="1985" box="[348,582,835,861]" class="Insecta" family="Scarabaeidae" genus="Phaedotrogus" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDA5DFCF17DAEDE26" box="[348,490,835,861]" italics="true" pageId="5" pageNumber="46">Phaedotrogus</emphasis>
Paulian
</taxonomicName>
and probably
<taxonomicName id="4C074D29330B152BD9EDFCF67F92DE26" ID-CoL="63NZP" authority="Silvestri" authorityName="Silvestri" authorityYear="1924" box="[748,982,835,861]" class="Insecta" family="Scarabaeidae" genus="Ponerotrogus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD9EDFCF67F31DE26" box="[748,885,836,861]" italics="true" pageId="5" pageNumber="46">Ponerotrogus</emphasis>
Silvestri
</taxonomicName>
and
<taxonomicName id="4C074D29330B152BDF11FCF17D00DEFA" authority="Wasmann" authorityName="Wasmann" authorityYear="1918" class="Insecta" family="Scarabaeidae" genus="Cyclotrogus" higherTaxonomySource="CoL" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BDF11FCF17C88DEFB" italics="true" pageId="5" pageNumber="46">Cyclotrogus</emphasis>
Wasmann
</taxonomicName>
(both currently considered to be junior synonyms of
<taxonomicName id="4C074D29330B152BD8A7FCD57856DEFA" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[934,1042,871,897]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD8A7FCD57856DEFA" box="[934,1042,871,897]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
</taxonomicName>
), will probably be necessary to establish a sound genus-level classification. Currently we see no reason to change the status of
<taxonomicName id="4C074D29330B152BD904FC1F7EDCDEBC" ID-CoL="Q8C" authorityName="Janssens" authorityYear="1949" box="[517,664,941,967]" class="Insecta" family="Scarabaeidae" genus="Afroharoldius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD904FC1F7EDCDEBC" box="[517,664,941,967]" italics="true" pageId="5" pageNumber="46">Afroharoldius</emphasis>
</taxonomicName>
as junior synonym of
<taxonomicName id="4C074D29330B152BD897FC1F7846DEBC" ID-CoL="4TRW" authorityName="Boucomont" authorityYear="1914" box="[918,1026,941,967]" class="Insecta" family="Scarabaeidae" genus="Haroldius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="46" phylum="Arthropoda" rank="genus">
<emphasis id="B973EAB8330B152BD897FC1F7846DEBC" box="[918,1026,941,967]" italics="true" pageId="5" pageNumber="46">Haroldius</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>
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