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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.743.23029" ID-GBIF-Dataset="1e5fbbb7-a293-466f-b354-9eb70c8befe7" ID-PMC="PMC5904386" ID-Pensoft-Pub="1313-2970-743-67" ID-PubMed="29670437" ID-ZBK="4D31DBBF07B1479C90489C9207B5103D" ModsDocAuthor="" ModsDocDate="2018" ModsDocID="1313-2970-743-67" ModsDocOrigin="ZooKeys 743" ModsDocTitle="Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae)" checkinTime="1521107529766" checkinUser="pensoft" docAuthor="Kohlmann, Bert, Arriaga-Jimenez, Alfonsina &amp; Roes, Matthias" docDate="2018" docId="355A9EEBAD9D2DDA6EACCCBF6A512440" docLanguage="en" docName="ZooKeys 743: 67-93" docOrigin="ZooKeys 743" docSource="http://dx.doi.org/10.3897/zookeys.743.23029" docTitle="Phanaeus (Notiophanaeus) dionysius Kohlmann, Arriaga-Jimenez &amp; Roes, sp. n." docType="treatment" docUuid="B4F5DBB5-6802-44F0-87FF-199895A96C80" docUuidSource="ZooBank" docVersion="5" lastPageNumber="73" masterDocId="FF813C19FFB8FFF8FFB7FFACFF851225" masterDocTitle="Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae)" masterLastPageNumber="93" masterPageNumber="67" pageNumber="69" updateTime="1668165570053" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Dung beetle vicariant speciation in the mountains of Oaxaca, Mexico, with a description of a new species of Phanaeus (Coleoptera, Geotrupidae, Scarabaeidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kohlmann, Bert</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Arriaga-Jimenez, Alfonsina</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Roes, Matthias</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2018</mods:date>
<mods:detail type="volume">
<mods:number>743</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>67</mods:start>
<mods:end>93</mods:end>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.743.23029</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.743.23029</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-743-67</mods:identifier>
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<mods:identifier type="ZooBank">4D31DBBF07B1479C90489C9207B5103D</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="142283989" LSID="urn:lsid:zoobank.org:act:B4F5DBB5-6802-44F0-87FF-199895A96C80" httpUri="http://treatment.plazi.org/id/355A9EEBAD9D2DDA6EACCCBF6A512440" lastPageId="6" lastPageNumber="73" pageId="2" pageNumber="69">
<subSubSection pageId="2" pageNumber="69" type="nomenclature">
<paragraph pageId="2" pageNumber="69">
<taxonomicName LSID="http://zoobank.org/B4F5DBB5-6802-44F0-87FF-199895A96C80" authority="Kohlmann, Arriaga-Jimenez &amp; Roes" class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus (Notiophanaeus) dionysius" order="Coleoptera" pageId="2" pageNumber="69" phylum="Arthropoda" rank="species" species="dionysius" subGenus="Notiophanaeus">
Phanaeus (Notiophanaeus) dionysius Kohlmann,
<normalizedToken originalValue="Arriaga-Jiménez">Arriaga-Jimenez</normalizedToken>
&amp;
<normalizedToken originalValue="Rös">Roes</normalizedToken>
</taxonomicName>
<taxonomicNameLabel pageId="2" pageNumber="69">sp. n.</taxonomicNameLabel>
Figures 1, 2, 3, 4, 5, 6
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="69" type="type material">
<paragraph pageId="2" pageNumber="69">Type material.</paragraph>
<paragraph pageId="2" pageNumber="69">
Holotype male, pinned, with genitalia in a separate microvial. Original label:
<normalizedToken originalValue="“México">&quot;Mexico</normalizedToken>
. La Mesita San Pablo Etla. Oaxaca.
</paragraph>
<paragraph pageId="2" pageNumber="69">
23-VI-17, coprotrampa,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164999">17°9'54&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.73833">96°44'18&quot;W</geoCoordinate>
, bosque de Encino, 1976 m, Arriaga A. and Arenas A. Col.&quot; &quot;HOLOTYPE/
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="2" pageNumber="69" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
Kohlmann,
<normalizedToken originalValue="Arriaga-Jiménez">Arriaga-Jimenez</normalizedToken>
,
<normalizedToken originalValue="Rös">Roes</normalizedToken>
[red printed label]&quot;. Allotype female: &quot;Mexico. La Mesita San Pablo Etla. Oaxaca. 23-VI-17, coprotrampa,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164999">17°9'54&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.73861">96°44'19&quot;W</geoCoordinate>
, bosque de Encino, 1976 m, Arriaga A. and Arenas A. Col.&quot;
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="69" type="materials_examined">
<paragraph pageId="2" pageNumber="69">Other material.</paragraph>
<paragraph pageId="2" pageNumber="69">
(5 males, 5 females). Paratypes: &quot;Mexico. Reserva Comunitaria San Pablo Etla. Oaxaca. 27-IV-17, coprotrampa,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164722">17°9'53&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.738884">96°44'20&quot;W</geoCoordinate>
, bosque de Encino, 1974 m, Arriaga A. and Arenas A. Col.&quot; (1 males, 2 females) (CMN, CEMT, CPFA); &quot;Mexico. La Mesita San Pablo Etla. Oaxaca. 23-VI-17, coprotrampa,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164999">17°9'54&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.73833">96°44'18&quot;W</geoCoordinate>
, bosque de Encino, 1976 m, Arriaga A. and Arenas A. Col. (2 males, 2 females) (CMN, CEMT, IEXA, CPFA); 14-VII-17,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164999">17°9'54&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.74833">96°44'54&quot;W</geoCoordinate>
, 1954 m, (1 male) (IEXA); 27-IV-17,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.17111">17°10'16&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.73055">96°43'50&quot;W</geoCoordinate>
, 2219 (1 male) (JB). 23-VI-17,
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164999">17°09'54&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.73861">96°44'19&quot;W</geoCoordinate>
, 1976 m, (1 female) (JB).
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="69" type="type locality">
<paragraph pageId="2" pageNumber="69">Type locality.</paragraph>
<paragraph pageId="2" pageNumber="69">
La Mesita San Pablo Etla (
<geoCoordinate direction="north" orientation="latitude" precision="15" value="17.164999">17°9'54&quot;N</geoCoordinate>
,
<geoCoordinate direction="west" orientation="longitude" precision="15" value="-96.73861">96°44'19&quot;W</geoCoordinate>
, 1976 m), Oaxaca, Mexico.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="69" type="type deposition">
<paragraph pageId="2" pageNumber="69">Type deposition.</paragraph>
<paragraph pageId="2" pageNumber="69">
<normalizedToken originalValue="Colección">Coleccion</normalizedToken>
<normalizedToken originalValue="Entomológica">Entomologica</normalizedToken>
IEXA, Instituto de
<normalizedToken originalValue="Ecología">Ecologia</normalizedToken>
, Xalapa, Mexico.
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="69" type="diagnosis">
<paragraph pageId="2" pageNumber="69">Diagnosis.</paragraph>
<paragraph pageId="2" pageNumber="69">Distinctly granulate male pronotal disk; sagittal furrow present on the female pronotum; unmodified sutural margin of the elytra; pygidium longer than wide. Its basal border forming a small indentation medially, usually all-black color.</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="70" pageId="2" pageNumber="69" type="description">
<paragraph pageId="2" pageNumber="69">Description.</paragraph>
<paragraph lastPageId="3" lastPageNumber="70" pageId="2" pageNumber="69">
Holotype. Major male (Fig. 1
<normalizedToken originalValue="ab">a-b</normalizedToken>
). Length: 16.5 mm. Humeral width: 10. 9 mm. Body appearing dull shiny black with a faint blue luster to the unaided eye. Magnification reveals faint greenish cast along the ocular, pronotal, and elytral borders, on the abdominal surface and underside of femora rugose. Clypeus with two conspicuous median teeth; surface and frons bearing long, slender horn strongly curved over the pronotum. Pronotum with large, flat triangular disk (Fig. 1a), with a well-developed, small callosity on each side near anterior margin and with postero-lateral angles projecting caudally; lateral portions faintly asperate, with distinct punctures present only behind lateral fossae (
<normalizedToken originalValue="×">x</normalizedToken>
20); flat triangular surface disk densely, evenly, and coarsely granulate; granules extending onto posterolateral angles and becoming eroded near and along disk borders. Pronotum with obsolete basal fossae; anterolateral angles subquadrate, distinctly upturned and surface behind angles concave; pronotal midline present, faintly developed, more evident on anterior half; weak punctures along posterior pronotal margin; pronotal surface shagreen. Elytral striae fine, with small but well-defined punctures separated at regular intervals; intervals broad and faintly convex, evenly and faintly shiny, covered with minute punctures (
<normalizedToken originalValue="×">x</normalizedToken>
20); surface shagreen. Pygidium black with shagreen surface and obsolete punctures, glabrous; pygidium wider than long (Fig. 2a); basal pygidial margin forming a small triangular tooth medially (Fig. 2b); pygidial margin with a green cast.
<pageBreakToken pageId="3" pageNumber="70" start="start">Protibia</pageBreakToken>
quadridentate. Lamella copulatrix as in Figure 3a; aedeagus similar to the
<taxonomicName lsidName="Ph. endymion" pageId="3" pageNumber="70" rank="species" species="endymion">Ph. endymion</taxonomicName>
species group (Fig. 3b, d).
</paragraph>
<caption pageId="3" pageNumber="70">
<paragraph pageId="3" pageNumber="70">
Figure 1. Major male habitus in a dorsal and b lateral view of
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="3" pageNumber="70" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n., and c dorsal view of
<taxonomicName lsidName="Ph. zapotecus" pageId="3" pageNumber="70" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
. Scale bar= 5mm.
</paragraph>
</caption>
<caption pageId="3" pageNumber="70">
<paragraph pageId="3" pageNumber="70">
Figure 2. Pygidia of a
<taxonomicName lsidName="Ph. dyionysius" pageId="3" pageNumber="70" rank="species" species="dyionysius">Ph. dyionysius</taxonomicName>
sp. n. b detail, arrow points to the triangle that forms the keel c
<taxonomicName lsidName="Ph. zapotecus" pageId="3" pageNumber="70" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
. Scale bar= 1mm.
</paragraph>
</caption>
<caption pageId="3" pageNumber="70">
<paragraph pageId="3" pageNumber="70">
Figure 3. a Lamella copulatrix and b aedeagus of
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="3" pageNumber="70" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n. c aedeagus of
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus zapotecus" order="Coleoptera" pageId="3" pageNumber="70" phylum="Arthropoda" rank="species" species="zapotecus">Phanaeus zapotecus</taxonomicName>
d parameres of
<taxonomicName lsidName="Ph. dionysius" pageId="3" pageNumber="70" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
sp. n. e parameres of
<taxonomicName lsidName="Ph. zapotecus" pageId="3" pageNumber="70" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
. Scale bar= 1mm.
</paragraph>
</caption>
<paragraph pageId="3" pageNumber="70">Female. Allotype (Fig. 4). Length: 16.3 mm. Humeral width: 10 mm. Body faintly shining black. Head with low, narrow trituberculate carina. Pronotum with a faint green lustre, evenly and densely covered with punctures, punctures becoming fainter on middle of disk; surface shagreen; with raised anteromedian trituberculate tumosity near anterior margin, tubercles equal in size and set in a more-or-less straight, transverse line; disk with distinct mid-longitudinal furrow, extending forward from posterior margin to about middle of disk, furrow more strongly sculptured than adjacent surface of disk. Pygidium with faint to distinct fine, sparse punctures.</paragraph>
<caption pageId="3" pageNumber="70">
<paragraph pageId="3" pageNumber="70">
Figure 4. Female habitus of a dorsal and b lateral view of
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="3" pageNumber="70" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n. Scale bar= 5mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="3" pageNumber="70" type="variation">
<paragraph pageId="3" pageNumber="70">Variations.</paragraph>
<paragraph pageId="3" pageNumber="70">Length: 12.6-18.7 mm. Humeral width: 7.9-11.1 mm. Pronotal disk of males may vary from black without reflections to having a green or red lustre. Minor male (Fig. 5): Similar to major male, except the cephalic horn is smaller and the posterolateral angles of the pronotum are reduced.</paragraph>
<caption pageId="3" pageNumber="70">
<paragraph pageId="3" pageNumber="70">
Figure 5. Dorsal habitus of a minor male of
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="3" pageNumber="70" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp.n. Scale bar= 5mm.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="4" pageNumber="71" type="etymology">
<paragraph pageId="4" pageNumber="71">
<pageBreakToken pageId="4" pageNumber="71" start="start">Etymology</pageBreakToken>
.
</paragraph>
<paragraph pageId="4" pageNumber="71">
Due to the fact that this species has been collected in association with
<taxonomicName lsidName="Ph. damocles" pageId="4" pageNumber="71" rank="species" species="damocles">Ph. damocles</taxonomicName>
Harold, this new species (a noun in the nominative singular) is named after Dionysius II of Syracuse, one of the main characters alluded to in the moral anecdote of the &quot;Sword of Damocles&quot;.
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="73" pageId="5" pageNumber="72" type="distribution">
<paragraph pageId="5" pageNumber="72">
<pageBreakToken pageId="5" pageNumber="72" start="start">Distribution</pageBreakToken>
and ecology.
</paragraph>
<paragraph pageId="5" pageNumber="72">
So far, this species is only known from San Pablo Etla in the Sierra Norte (Sierra de
<normalizedToken originalValue="Ixtlán">Ixtlan</normalizedToken>
) in Oaxaca (Fig. 6), along the internal dry slope facing the Oaxaca Valley. It has been collected from altitudes of 1950 m to 2250 m. The dry deciduous oak forest where
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n. was found is characterised by trees between five and ten meters tall. Abundant oak species are
<taxonomicName class="Magnoliopsida" family="Fagaceae" genus="Quercus" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Quercus laeta" order="Fagales" pageId="5" pageNumber="72" phylum="Tracheophyta" rank="species" species="laeta">Quercus laeta</taxonomicName>
Liebm. and
<taxonomicName lsidName="Q. laurina" pageId="5" pageNumber="72" rank="species" species="laurina">Q. laurina</taxonomicName>
Humb. and Bonpl., predominant species of this ecosystem, ranging from 1800 m to 2400 m altitude (Fig. 7). Other species dominating this forest in the sampling site are
<taxonomicName lsidName="Q. glaucoides" pageId="5" pageNumber="72" rank="species" species="glaucoides">Q. glaucoides</taxonomicName>
Mart. and Gal.,
<taxonomicName lsidName="Q. liebmannii" pageId="5" pageNumber="72" rank="species" species="liebmannii">Q. liebmannii</taxonomicName>
Oersted.,
<taxonomicName lsidName="Q. rugosa" pageId="5" pageNumber="72" rank="species" species="rugosa">Q. rugosa</taxonomicName>
<normalizedToken originalValue="Née">Nee</normalizedToken>
, and
<taxonomicName lsidName="Q. castanea" pageId="5" pageNumber="72" rank="species" species="castanea">Q. castanea</taxonomicName>
<normalizedToken originalValue="Née">Nee</normalizedToken>
, also found at higher or lower altitudes (J. Williams, CIIDIR-Oaxaca, pers. comm.,
<bibRefCitation pageId="5" pageNumber="72">
<normalizedToken originalValue="Valencia-Ávalos">Valencia-Avalos</normalizedToken>
and Nixon 2004
</bibRefCitation>
). This dry deciduous oak forest shows a strong seasonality, when most trees lose their leaves for around four to five months between December and May. This new species is found next to the Oaxaca Metropolitan Area in a voluntary protected area, the San Pablo Etla Community Reserve &quot;La Mesita&quot;.
</paragraph>
<caption pageId="5" pageNumber="72">
<paragraph pageId="5" pageNumber="72">
Figure 6. Map of the known distribution of
<taxonomicName class="Insecta" family="Geotrupidae" genus="Geotrupes" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Geotrupes pecki" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="pecki">Geotrupes pecki</taxonomicName>
,
<taxonomicName lsidName="G. viridiobscurus" pageId="5" pageNumber="72" rank="species" species="viridiobscurus">G. viridiobscurus</taxonomicName>
,
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n., and
<taxonomicName lsidName="Ph. zapotecus" pageId="5" pageNumber="72" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
. Orography of Oaxaca is shown, based on the Digital Elevation Model downloaded from INEGI (2017, http://www.inegi.org.mx). Grey area shows the limits of Mexico with the Gulf of Mexico in the North and the Pacific Ocean in the South.
</paragraph>
</caption>
<caption pageId="5" pageNumber="72">
<paragraph pageId="5" pageNumber="72">
Figure 7. Drone photographs of the dry oak forest where
<taxonomicName lsidName="Ph. dionysius" pageId="5" pageNumber="72" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
sp. n. was collected; a April 2017 and b August 2017.
</paragraph>
</caption>
<paragraph lastPageId="6" lastPageNumber="73" pageId="5" pageNumber="72">
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n. has been collected simultaneously in dung-baited traps with
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Canthidium" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Canthidium quercetorum" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="quercetorum">Canthidium quercetorum</taxonomicName>
Kohlmann,
<normalizedToken originalValue="Arriaga-Jiménez">Arriaga-Jimenez</normalizedToken>
and
<normalizedToken originalValue="Rös">Roes</normalizedToken>
,
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Canthon" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Canthon humectus" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="humectus">Canthon humectus</taxonomicName>
(Say),
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Copris" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Copris klugi" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="klugi">Copris klugi</taxonomicName>
Harold,
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Deltochilum" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Deltochilum mexicanum" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="mexicanum">Deltochilum mexicanum</taxonomicName>
Burmeister,
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Dichotomius" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Dichotomius colonicus" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="colonicus">Dichotomius colonicus</taxonomicName>
(Say),
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Onthophagus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Onthophagus anthracinus" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="anthracinus">Onthophagus near anthracinus</taxonomicName>
Harold,
<taxonomicName lsidName="O. aureofuscus" pageId="5" pageNumber="72" rank="species" species="aureofuscus">O. aureofuscus</taxonomicName>
Bates,
<taxonomicName lsidName="O. chevrolati" pageId="5" pageNumber="72" rank="subspecies" species="chevrolati" subspecies="retusus">O. chevrolati retusus</taxonomicName>
Harold,
<taxonomicName lsidName="O. mexicanus" pageId="5" pageNumber="72" rank="species" species="mexicanus">O. mexicanus</taxonomicName>
Bates,
<taxonomicName lsidName="O. zapotecus" pageId="5" pageNumber="72" rank="species" species="zapotecus">O. zapotecus</taxonomicName>
Zunino and Halffter and
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus damocles" order="Coleoptera" pageId="5" pageNumber="72" phylum="Arthropoda" rank="species" species="damocles">Phanaeus damocles</taxonomicName>
Harold, in the oak forest. Although its closest relative,
<taxonomicName lsidName="Ph. zapotecus" pageId="5" pageNumber="72" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
Edmonds, seems
<pageBreakToken pageId="6" pageNumber="73" start="start">to</pageBreakToken>
be a strictly mycetophagous species,
<taxonomicName lsidName="Ph. dionysius" pageId="6" pageNumber="73" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
has only been collected in dung, despite the presence of fungi-baited traps put in the forest. Interestingly, no big fleshy fungi (toadstools) were observed in this type of forest, only small
<normalizedToken originalValue="“clavitos”">&quot;clavitos&quot;</normalizedToken>
(
<taxonomicName class="Agaricomycetes" family="Lyophyllaceae" genus="Lyophyllum" higherTaxonomySource="CoL" kingdom="Fungi" lsidName="Lyophyllum" order="Agaricales" pageId="6" pageNumber="73" phylum="Basidiomycota" rank="genus">Lyophyllum</taxonomicName>
), which could probably explain why this species does not exploit fungi.
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="73" type="taxonomic relationships">
<paragraph pageId="6" pageNumber="73">Taxonomic relationships.</paragraph>
<paragraph pageId="6" pageNumber="73">
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="6" pageNumber="73" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
sp. n. belongs to the
<taxonomicName lsidName="Ph. endymion" pageId="6" pageNumber="73" rank="species" species="endymion">Ph. endymion</taxonomicName>
species group and due to its close taxonomic similarity discussed below is postulated to be the sister species of
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
Edmonds, 2006.
<taxonomicName class="Insecta" family="Scarabaeidae" genus="Phanaeus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phanaeus dionysius" order="Coleoptera" pageId="6" pageNumber="73" phylum="Arthropoda" rank="species" species="dionysius">Phanaeus dionysius</taxonomicName>
will key out to
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
in the key of
<bibRefCitation author="Moctezuma, V" journalOrPublisher="ZooKeys" pageId="21" pageNumber="88" pagination="113 - 135" title="Two new species of the Phanaeusendymion group (Coleopter; Scarabaeidae, Scarabaeinae)." url="https://doi.org/10.3897/zookeys.702.14728" volume="702" year="2017">Moctezuma et al. (2017)</bibRefCitation>
and can be separated from it because it has long and slender pronotal posterolateral angles (Fig. 1a) whereas
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
has short and rounded posterolateral angles (Fig. 1c). The basal border of the pygidium in
<taxonomicName lsidName="Ph. dionysius" pageId="6" pageNumber="73" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
forms a small indentation at its middle (Fig. 2b), whereas it runs completely straight in
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
(Fig. 2c). Additionally, the apex of the parameres of
<taxonomicName lsidName="Ph. dionysius" pageId="6" pageNumber="73" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
sp. n. is more projected (Fig. 3b, d), than that from
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
(Fig. 3c, e). Moreover, the middle sinuation of the parameres in lateral view is much more pronounced in
<taxonomicName lsidName="Ph. dionysius" pageId="6" pageNumber="73" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
sp. n. (Fig. 3b) than in
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
(Fig. 3c).
</paragraph>
</subSubSection>
<subSubSection pageId="6" pageNumber="73" type="chorological affinities">
<paragraph pageId="6" pageNumber="73">Chorological affinities.</paragraph>
<paragraph pageId="6" pageNumber="73">
The known distribution of
<taxonomicName lsidName="Ph. dionysius" pageId="6" pageNumber="73" rank="species" species="dionysius">Ph. dionysius</taxonomicName>
sp. n. is relatively near to its closest relative,
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
, 90 km distance in a straight line, which is distributed in dry pine-oak and pine-oak-juniper forests on the internal slope of the Sierra Sur (Sierra de Tlaxiaco), going from 1850 m to 2150 m altitude. Interestingly, attempts at trying to collect
<taxonomicName lsidName="Ph. zapotecus" pageId="6" pageNumber="73" rank="species" species="zapotecus">Ph. zapotecus</taxonomicName>
in the environs of San
<normalizedToken originalValue="José">Jose</normalizedToken>
del
<normalizedToken originalValue="Pacífico">Pacifico</normalizedToken>
in the Sierra Sur (Sierra de
<normalizedToken originalValue="Miahuatlán">Miahuatlan</normalizedToken>
) with fungi-baited traps did not produce any results.
</paragraph>
</subSubSection>
</treatment>
</document>