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<document id="6A1BE675D2EC287F8A6171E11AE4EE28" ID-DOI="10.13133/2284-4880/547" ID-ISSN="2284-4880" ID-Zenodo-Dep="12762398" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1721239357453" checkinUser="felipe" docAuthor="Grebennikov, Vasily V." docDate="2021" docId="03A5143AFF9EFFD3FC83B261FEB7C3C1" docLanguage="en" docName="FragEntomol.53.2.283-298.pdf" docOrigin="Fragmenta entomologica 53 (2)" docSource="http://dx.doi.org/10.13133/2284-4880/547" docStyle="DocumentStyle:8499FF4308F512386C0E43DC290A16BD.1:FragEntomol.2021-.journal_article.0cover" docStyleId="8499FF4308F512386C0E43DC290A16BD" docStyleName="FragEntomol.2021-.journal_article.0cover" docStyleVersion="1" docTitle="Balleriodes Grebennikov, 2021, gen. nov." docType="treatment" docVersion="2" lastPageNumber="290" masterDocId="FF9C6C42FF9BFFD4FFD5B035FFE6C12D" masterDocTitle="Phylogenetic placement of a new paleoendemic pill scarab from the Udzungwa Mountains, Tanzania, triggers biogeographic interpretations (Coleoptera: Hybosoridae, Ceratocanthinae)" masterLastPageNumber="298" masterPageNumber="283" pageNumber="288" updateTime="1721239891135" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-4.0">
<mods:mods id="8456D6FB7DBAD2393A96D44CCBC2E6DB" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="C2AA1AFA6B9F0170A5B24B46732BE31D">
<mods:title id="C5127D78293D18418418FA80A7EDD839">Phylogenetic placement of a new paleoendemic pill scarab from the Udzungwa Mountains, Tanzania, triggers biogeographic interpretations (Coleoptera: Hybosoridae, Ceratocanthinae)</mods:title>
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<mods:name id="9FDA2B7B0A986F97EA5174D49243F573" type="personal">
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<mods:namePart id="57830FC5996819B5184500A0A23571F0">Grebennikov, Vasily V.</mods:namePart>
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<mods:typeOfResource id="C3ABC74D801C6026C422535D8C17AE9D">text</mods:typeOfResource>
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<mods:title id="A25A98F12A7FC7DB18CDC6A29D35D348">Fragmenta entomologica</mods:title>
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<mods:date id="F22ED09A4864D972F036E3E107621362">2021</mods:date>
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<mods:number id="651FAB1720015A78B6B3D1466C128629">2021-11-30</mods:number>
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<mods:url id="4EFDF47F7A4FACB7C229515F7419FCBA">http://dx.doi.org/10.13133/2284-4880/547</mods:url>
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<mods:classification id="0F64E915BA3EB1F5137E72456991A384">journal article</mods:classification>
<mods:identifier id="BFAB3111081004768C42A5E6B95C26A4" type="DOI">10.13133/2284-4880/547</mods:identifier>
<mods:identifier id="CAB0FBDC8DA06879CD0D0E8930F842F4" type="ISSN">2284-4880</mods:identifier>
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<treatment id="03A5143AFF9EFFD3FC83B261FEB7C3C1" LSID="urn:lsid:plazi:treatment:03A5143AFF9EFFD3FC83B261FEB7C3C1" httpUri="http://treatment.plazi.org/id/03A5143AFF9EFFD3FC83B261FEB7C3C1" lastPageId="7" lastPageNumber="290" pageId="5" pageNumber="288">
<subSubSection id="C316F6A7FF9EFFD1FC83B261FBD8C341" box="[854,1086,596,620]" pageId="5" pageNumber="288" type="nomenclature">
<paragraph id="8BB3A52CFF9EFFD1FC83B261FBD8C341" blockId="5.[854,1086,596,620]" box="[854,1086,596,620]" pageId="5" pageNumber="288">
<heading id="D0FB1240FF9EFFD1FC83B261FBD8C341" bold="true" box="[854,1086,596,620]" centered="true" fontSize="10" level="2" pageId="5" pageNumber="288" reason="3">
<emphasis id="B978793EFF9EFFD1FC83B261FBD8C341" bold="true" box="[854,1086,596,620]" pageId="5" pageNumber="288">
<taxonomicName id="4C0CDEAFFF9EFFD1FC83B261FC34C341" ID-CoL="8SSNK" authorityName="Grebennikov" authorityYear="2021" box="[854,978,596,620]" class="Insecta" family="Hybosoridae" genus="Balleriodes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="288" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B978793EFF9EFFD1FC83B261FC34C341" bold="true" box="[854,978,596,620]" italics="true" pageId="5" pageNumber="288">Balleriodes</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A24BC445FF9EFFD1FC0CB261FBD8C341" box="[985,1086,596,620]" pageId="5" pageNumber="288" rank="genus">gen. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C316F6A7FF9EFFD1FC83B2A1FBE6C3E1" pageId="5" pageNumber="288" type="type_taxon">
<paragraph id="8BB3A52CFF9EFFD1FC83B2A1FAFAC381" blockId="5.[854,1308,660,716]" box="[854,1308,660,684]" pageId="5" pageNumber="288">
<emphasis id="B978793EFF9EFFD1FC83B2A1FC01C381" bold="true" box="[854,999,660,684]" pageId="5" pageNumber="288">
<typeStatus id="54B71B8EFF9EFFD1FC83B2A1FC6BC381" box="[854,909,660,684]" pageId="5" pageNumber="288">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="4C0CDEAFFF9EFFD1FC38B2A1FB59C381" ID-CoL="8SSQY" authorityName="Grebennikov" authorityYear="2021" box="[1005,1215,660,684]" class="Insecta" family="Hybosoridae" genus="Balleriodes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="288" phylum="Arthropoda" rank="species" species="sphaera" status="sp. nov.">
<emphasis id="B978793EFF9EFFD1FC38B2A1FB59C381" box="[1005,1215,660,684]" italics="true" pageId="5" pageNumber="288">Balleriodes sphaera</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A24BC445FF9EFFD1FB13B2A0FAFFC381" box="[1222,1305,661,684]" pageId="5" pageNumber="288" rank="species">sp. nov.</taxonomicNameLabel>
,
</paragraph>
<paragraph id="8BB3A52CFF9EFFD1FC83B281FBE6C3E1" blockId="5.[854,1308,660,716]" box="[854,1024,692,716]" pageId="5" pageNumber="288">here designated.</paragraph>
</subSubSection>
<subSubSection id="C316F6A7FF9EFFD3FC83B2C1FDCEC361" lastPageId="7" lastPageNumber="290" pageId="5" pageNumber="288" type="diagnosis">
<paragraph id="8BB3A52CFF9EFFD3FC83B2C1FE9DC0C1" blockId="5.[814,1438,756,1932]" lastBlockId="7.[135,758,180,748]" lastPageId="7" lastPageNumber="290" pageId="5" pageNumber="288">
<emphasis id="B978793EFF9EFFD1FC83B2C1FC2CC221" bold="true" box="[854,970,756,780]" pageId="5" pageNumber="288">Diagnosis.</emphasis>
The only known species of the genus can be immediately recognized among all pill scarabs (including most similar
<taxonomicName id="4C0CDEAFFF9EFFD1FC3CB300FB8BC261" authorityName="Germar" authorityYear="1843" box="[1001,1133,821,844]" class="Insecta" family="Hybosoridae" genus="Synarmostes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B978793EFF9EFFD1FC3CB300FB8BC261" box="[1001,1133,821,844]" italics="true" pageId="5" pageNumber="288">Synarmostes</emphasis>
</taxonomicName>
) by the unique combination of the following easily observed characters: body black, non-metallic, without prominent pubescence or punctures (although sparse fine surface sculpturing is present), in conglobate position nearly circular in dorsal view and nearly globular in 3-dimensional shape. Most notably (and uniquely with allopatric
<taxonomicName id="4C0CDEAFFF9EFFD1FC23B3C0FB9DC521" authorityName="Germar" authorityYear="1843" box="[1014,1147,1013,1036]" class="Insecta" family="Hybosoridae" genus="Synarmostes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B978793EFF9EFFD1FC23B3C0FB9DC521" box="[1014,1147,1013,1036]" italics="true" pageId="5" pageNumber="288">Synarmostes</emphasis>
</taxonomicName>
), adults of the new genus have elytra with markedly developed longitudinal striation only in posterior halves. Adult beetles of the new genus additionally differ from those of
<taxonomicName id="4C0CDEAFFF9EFFD1FB42B460FAFAC541" authorityName="Germar" authorityYear="1843" box="[1175,1308,1109,1132]" class="Insecta" family="Hybosoridae" genus="Synarmostes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B978793EFF9EFFD1FB42B460FAFAC541" box="[1175,1308,1109,1132]" italics="true" pageId="5" pageNumber="288">Synarmostes</emphasis>
</taxonomicName>
in having a single apical outer spur on protibia apices (two in
<taxonomicName id="4C0CDEAFFF9EFFD1FA83B440FC9EC581" authorityName="Germar" authorityYear="1843" class="Insecta" family="Hybosoridae" genus="Synarmostes" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B978793EFF9EFFD1FA83B440FC9EC581" italics="true" pageId="5" pageNumber="288">Synarmostes</emphasis>
</taxonomicName>
), head lacking a transverse furrow on vertex, parameres with a basal projection, and relatively larger basal piece of male genitalia (
<collectingRegion id="49C86BCEFF9EFFD1FBF8B4E1FB99C5C1" box="[1069,1151,1236,1260]" country="Canada" name="Alberta" pageId="5" pageNumber="288">Alberto</collectingRegion>
Ballerio, personal communication). Additionally, the
<typeStatus id="54B71B8EFF9EFFD1FB83B4C0FB62C420" box="[1110,1156,1269,1293]" pageId="5" pageNumber="288">type</typeStatus>
species of this monotypic genus displays two characters seemingly unique within all of
<taxonomicName id="4C0CDEAFFF9EFFD1FC84B501FC1AC461" authorityName="Martinez" authorityYear="1968" box="[849,1020,1332,1356]" class="Insecta" family="Hybosoridae" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="284" phylum="Arthropoda" rank="subFamily" subFamily="Ceratocanthinae">Ceratocanthinae</taxonomicName>
(
<collectingRegion id="49C86BCEFF9EFFD1FBD8B501FBB9C461" box="[1037,1119,1332,1356]" country="Canada" name="Alberta" pageId="5" pageNumber="288">Alberto</collectingRegion>
Ballerio, personal communication): remarkably long and dense setae on the fore tarsi (arrow in
<figureCitation id="1337B9A9FF9EFFD1FC48B541FC0AC4A1" box="[925,1004,1396,1420]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Fig. 2L</figureCitation>
) and the transparent membrane adjacent to the ventral surface of parameres (arrows in
<figureCitation id="1337B9A9FF9EFFD1FAC2B5A1FA6CC481" box="[1303,1418,1428,1452]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Figs 2P–R</figureCitation>
).
<emphasis id="B978793EFF9EFFD1FC83B581FC05C4E1" bold="true" box="[854,995,1460,1484]" pageId="5" pageNumber="288">Description.</emphasis>
Body black; without pubescence; without much microsculpture; non-metallic; in fully conglobate position visible part of legs (tibiae) and elytral apical halves often covered with dirt-like coating (
<figureCitation id="1337B9A9FF9EFFD1FAB9B621FCBAC761" captionStart-0="Fig" captionStart-1="Fig" captionStartId-0="8.[151,182,1720,1739]" captionStartId-1="9.[137,168,1724,1743]" captionTargetBox-0="[153,1455,164,1703]" captionTargetBox-1="[136,1436,169,1701]" captionTargetId-0="figure-101@8.[151,1465,161,1703]" captionTargetId-1="figure-87@9.[133,1447,166,1701]" captionTargetPageId-0="8" captionTargetPageId-1="9" captionText-0="Fig. 3 – Habitus of select ingroup and outgroup pill scarabs, dorsal view; images are to scale." captionText-1="Fig. 4 – Habitus of select ingroup and outgroup pill scarabs, left lateral view; images are to scale." figureDoi-0="http://doi.org/10.5281/zenodo.12762414" figureDoi-1="http://doi.org/10.5281/zenodo.12762418" httpUri-0="https://zenodo.org/record/12762414/files/figure.png" httpUri-1="https://zenodo.org/record/12762418/files/figure.png" pageId="5" pageNumber="288">Figs 3–4</figureCitation>
); neatly circular in dorsal view and nearly globular in 3-dimensional shape. Enrollment coaptations (ability to make a perfect ball) present. Head with interocular bridge (=“genal canthus”) completely subdividing compound eye into dorsal and ventral parts (
<figureCitation id="1337B9A9FF9EFFD1FBA6B681FB2AC7E1" box="[1139,1228,1716,1740]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Fig. 2A</figureCitation>
); dorsal eye small, ellipse-shaped, with major axis about four times greater than minor axis (
<figureCitation id="1337B9A9FF9EFFD1FC3BB6C1FBAEC621" box="[1006,1096,1780,1804]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Fig. 2A</figureCitation>
). Antennae with 10 antennomeres (
<figureCitation id="1337B9A9FF9EFFD1FCB9B721FC22C601" box="[876,964,1812,1836]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Fig. 2G</figureCitation>
). Mandibles in apical part without teeth (in addition to mandibular apex,
<figureCitation id="1337B9A9FF9EFFD1FB54B701FB13C661" box="[1153,1269,1844,1868]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Figs 2E–F</figureCitation>
). Maxillae with distal palpomere longer than combined length of both preceding ones (
<figureCitation id="1337B9A9FF9EFFD1FC4DB741FC0CC6A1" box="[920,1002,1908,1932]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="5" pageNumber="288">Fig. 2C</figureCitation>
). Pronotum evenly convex, sparsely, and shallowly punctured, its anterior and posterior margins not swollen (
<figureCitation id="1337B9A9FF9CFFD3FECBB0E1FE9EC1C1" box="[286,376,212,236]" captionStart="Fig" captionStartId="2.[151,183,1015,1034]" captionTargetBox="[151,1451,166,998]" captionTargetId="figure-428@2.[151,1451,164,998]" captionTargetPageId="2" captionText="Fig. 1 – A, Distribution of Balleriodes sphaera gen. et sp. nov. and potentially related and/or similar pill scarabs in Africa and Madagascar. B-F: Balleriodes sphaera gen. et sp. nov., holotype, habitus (B: left lateral, C: dorsal, D: ventral; E: anterior, F: posterior)." figureDoi="http://doi.org/10.5281/zenodo.12762406" httpUri="https://zenodo.org/record/12762406/files/figure.png" pageId="7" pageNumber="290">Fig. 1D</figureCitation>
), marginal bead complete. Elytra longitudinally striate on dorsal side in posterior half, and not striate in anterior half, with sparse shallow transverse comma-shaped punctures. Metathoracic wings absent. Protibiae with fine serration on distal third of outer side and with a larger apical outer tooth (
<figureCitation id="1337B9A9FF9CFFD3FDCEB141FD96C0A1" box="[539,624,372,396]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="7" pageNumber="290">Fig. 2L</figureCitation>
). Male genitalia with parameres almost symmetrical, basally with a projection (
<figureCitation id="1337B9A9FF9CFFD3FEDCB181FE65C0E1" box="[265,387,436,460]" captionStart="Fig" captionStartId="6.[151,182,1832,1851]" captionTargetBox="[183,1430,219,1714]" captionTargetId="figure-9@6.[148,1454,163,1801]" captionTargetPageId="6" captionText="Fig. 2 – Balleriodes sphaera gen. et sp. nov., paratypes (A-R, specimen 8969, S, specimen 8752), details. A-B, head, fronto-dorsal (A) and frontal (B); C, maxilla; D, labrum; E-F, mandibles; G, antenna; H: labium; I-J, elytron in ventral view showing phoretic mites (J, enlarged); K, prothorax, ventral view; L-N, fore, middle and hind left legs; O, pterothorax, ventral view; P-R, phallobase and parameres, right lateral (P), left lateral (Q) and dorsal (R) views; abdomen, with male genitalia in situ. Images are not to scale." figureDoi="http://doi.org/10.5281/zenodo.12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="7" pageNumber="290">Figs 2P–R</figureCitation>
). Female genitalia and sexual dimorphism not studied.
</paragraph>
<caption id="DF73F5A4FF9DFFD2FF42B71DFD39C6A6" ID-DOI="http://doi.org/10.5281/zenodo.12762410" ID-Zenodo-Dep="12762410" httpUri="https://zenodo.org/record/12762410/files/figure.png" pageId="6" pageNumber="289" startId="6.[151,182,1832,1851]" targetBox="[183,1430,219,1714]" targetPageId="6" targetType="figure">
<paragraph id="8BB3A52CFF9DFFD2FF42B71DFD39C6A6" blockId="6.[151,1452,1832,1931]" pageId="6" pageNumber="289">
<emphasis id="B978793EFF9DFFD2FF42B71DFF3DC616" bold="true" box="[151,219,1832,1851]" pageId="6" pageNumber="289">Fig. 2 –</emphasis>
<taxonomicName id="4C0CDEAFFF9DFFD2FF34B71DFE68C616" authorityName="Grebennikov" authorityYear="2021" box="[225,398,1832,1851]" class="Insecta" family="Hybosoridae" genus="Balleriodes" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="289" phylum="Arthropoda" rank="species" species="sphaera" status="gen. et sp. nov.">
<emphasis id="B978793EFF9DFFD2FF34B71DFE68C616" box="[225,398,1832,1851]" italics="true" pageId="6" pageNumber="289">Balleriodes sphaera</emphasis>
</taxonomicName>
<emphasis id="B978793EFF9DFFD2FE46B71DFDFFC616" bold="true" box="[403,537,1832,1851]" pageId="6" pageNumber="289">
<taxonomicNameLabel id="A24BC445FF9DFFD2FE46B71DFDFFC616" box="[403,537,1832,1851]" pageId="6" pageNumber="289" rank="species">gen. et sp. nov.</taxonomicNameLabel>
</emphasis>
, paratypes (
<emphasis id="B978793EFF9DFFD2FD57B71DFD4EC616" bold="true" box="[642,680,1832,1851]" pageId="6" pageNumber="289">A-R</emphasis>
, specimen 8969,
<emphasis id="B978793EFF9DFFD2FCEAB71DFCADC616" bold="true" box="[831,843,1832,1851]" pageId="6" pageNumber="289">S</emphasis>
, specimen 8752), details.
<emphasis id="B978793EFF9DFFD2FBFEB71DFBA9C616" bold="true" box="[1067,1103,1832,1851]" pageId="6" pageNumber="289">A-B</emphasis>
, head, fronto-dorsal (
<emphasis id="B978793EFF9DFFD2FADFB71DFAFFC616" bold="true" box="[1290,1305,1832,1851]" pageId="6" pageNumber="289">A</emphasis>
) and frontal (
<emphasis id="B978793EFF9DFFD2FA45B71DFA78C616" bold="true" box="[1424,1438,1832,1851]" pageId="6" pageNumber="289">B</emphasis>
);
<emphasis id="B978793EFF9DFFD2FF42B777FF40C678" bold="true" box="[151,166,1858,1877]" pageId="6" pageNumber="289">C</emphasis>
, maxilla;
<emphasis id="B978793EFF9DFFD2FF2BB776FEEBC67B" bold="true" box="[254,269,1859,1878]" pageId="6" pageNumber="289">D</emphasis>
, labrum;
<emphasis id="B978793EFF9DFFD2FEB4B776FE65C67B" bold="true" box="[353,387,1859,1878]" pageId="6" pageNumber="289">E-F</emphasis>
, mandibles;
<emphasis id="B978793EFF9DFFD2FE26B777FDE5C678" bold="true" box="[499,515,1858,1877]" pageId="6" pageNumber="289">G</emphasis>
, antenna;
<emphasis id="B978793EFF9DFFD2FD88B776FD8BC67B" bold="true" box="[605,621,1859,1878]" pageId="6" pageNumber="289">H</emphasis>
: labium;
<emphasis id="B978793EFF9DFFD2FD15B776FD3FC67B" bold="true" box="[704,729,1859,1878]" pageId="6" pageNumber="289">I-J</emphasis>
, elytron in ventral view showing phoretic mites (
<emphasis id="B978793EFF9DFFD2FB51B776FB69C67B" bold="true" box="[1156,1167,1859,1878]" pageId="6" pageNumber="289">J</emphasis>
, enlarged);
<emphasis id="B978793EFF9DFFD2FB22B776FAE1C67B" bold="true" box="[1271,1287,1859,1878]" pageId="6" pageNumber="289">K</emphasis>
, prothorax, ventral view;
<emphasis id="B978793EFF9DFFD2FF19B768FF17C65D" bold="true" box="[204,241,1885,1904]" pageId="6" pageNumber="289">L-N</emphasis>
, fore, middle and hind left legs;
<emphasis id="B978793EFF9DFFD2FDDFB768FDFCC65D" bold="true" box="[522,538,1885,1904]" pageId="6" pageNumber="289">O</emphasis>
, pterothorax, ventral view;
<emphasis id="B978793EFF9DFFD2FCDCB768FCCAC65D" bold="true" box="[777,812,1885,1904]" pageId="6" pageNumber="289">P-R</emphasis>
, phallobase and parameres, right lateral (
<emphasis id="B978793EFF9DFFD2FB5AB768FB7AC65D" bold="true" box="[1167,1180,1885,1904]" pageId="6" pageNumber="289">P</emphasis>
), left lateral (
<emphasis id="B978793EFF9DFFD2FAC7B768FAC5C65D" bold="true" box="[1298,1315,1885,1904]" pageId="6" pageNumber="289">Q</emphasis>
) and dorsal (
<emphasis id="B978793EFF9DFFD2FA40B768FA42C65D" bold="true" box="[1429,1444,1885,1904]" pageId="6" pageNumber="289">R</emphasis>
) views; abdomen, with male genitalia
<emphasis id="B978793EFF9DFFD2FE03B74DFDEDC6A7" box="[470,523,1912,1930]" italics="true" pageId="6" pageNumber="289">in situ</emphasis>
. Images are not to scale.
</paragraph>
</caption>
<paragraph id="8BB3A52CFF9CFFD3FF7BB1C1FDCEC361" blockId="7.[135,758,180,748]" pageId="7" pageNumber="290">
<emphasis id="B978793EFF9CFFD3FF7BB1C1FDBBC321" bold="true" box="[174,605,500,524]" pageId="7" pageNumber="290">Species composition and distribution.</emphasis>
This genus is monotypic and its known distribution is limited to the
<typeStatus id="54B71B8EFF9CFFD3FD1DB220FD10C300" box="[712,758,533,557]" pageId="7" pageNumber="290">type</typeStatus>
locality of its only species (see below).
</paragraph>
</subSubSection>
<subSubSection id="C316F6A7FF9CFFD3FF7BB261FEB7C3C1" pageId="7" pageNumber="290" type="etymology">
<paragraph id="8BB3A52CFF9CFFD3FF7BB261FEB7C3C1" blockId="7.[135,758,180,748]" pageId="7" pageNumber="290">
<emphasis id="B978793EFF9CFFD3FF7BB261FEC8C341" bold="true" box="[174,302,596,620]" pageId="7" pageNumber="290">Etymology.</emphasis>
The generic name is a patronymic, derived by adding the Latin suffix -
<emphasis id="B978793EFF9CFFD3FE79B241FE3BC3A1" box="[428,477,628,652]" italics="true" pageId="7" pageNumber="290">odes</emphasis>
to the family name of Alberto Ballerio (Brescia,
<collectingCountry id="F31BE5BCFF9CFFD3FE46B2A1FE2EC381" box="[403,456,660,684]" name="Italy" pageId="7" pageNumber="290">Italy</collectingCountry>
), in recognition of his role in triggering my interest in the evolution of pill scarabs. Gender masculine.
</paragraph>
</subSubSection>
</treatment>
</document>
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