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<mods:title id="FBD7F24E7A4F9FCA664EFD1F151CD5F4">Phylogenetic systematics of the keratin-feeding genus Polononcus Burmeister, 1876 (Coleoptera: Scarabaeoidea: Trogidae)</mods:title>
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<mods:namePart id="3C0A7C5C0A464C0FEB41D444F13A8015">Vinícius</mods:namePart>
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<mods:namePart id="F6A379BCB32BCED21EB8B0E018B41B48">Costa-Silva</mods:namePart>
<mods:affiliation id="1457C79013E3BB7080F31BA58636B4ED">Laboratório de Entomologia Integrativa, Departamento de Biologia Animal, Instituto de Biologia, Universidade de Campinas (UNICAMP), 13083 - 862, Campinas, São Paulo, Brazil &amp; Laboratório de Scarabaeoidologia, Instituto de Biociências, Universidade Federal de Mato Grosso (UFMT), 78060 - 900, Cuiabá, Mato Grosso, Brazil &amp; Department of Zoology and Entomology, University of Pretoria, Private Bag X 20, Hatfield, 0028, South Africa</mods:affiliation>
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<mods:namePart id="DABC0BDFE9A7CF0432CCE5F6C220795E">Strümpher, Werner P.</mods:namePart>
<mods:affiliation id="7742218467640E23A59E93A605F6E4D7">Ditsong National Museum of Natural History, Department of Coleoptera, P. O. Box 413, Pretoria, 0001, South Africa</mods:affiliation>
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<mods:namePart id="3E7FE909EE5B078DD8C324D6A9E04DA1">Tyssen, Patricia J.</mods:namePart>
<mods:affiliation id="48756DBD3658AF375DE91FAE31DDEF1B">Laboratório de Entomologia Integrativa, Departamento de Biologia Animal, Instituto de Biologia, Universidade de Campinas (UNICAMP), 13083 - 862, Campinas, São Paulo, Brazil</mods:affiliation>
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<mods:namePart id="415890FCB78DD8A5741216F3872E5DAA">Vaz-de-Mello, Fernando Z.</mods:namePart>
<mods:affiliation id="DAB13E1EE815B3647AC4E76A835A88F2">Laboratório de Scarabaeoidologia, Instituto de Biociências, Universidade Federal de Mato Grosso (UFMT), 78060 - 900, Cuiabá, Mato Grosso, Brazil</mods:affiliation>
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<mods:date id="73F6F8CFF0905E25B56C6156C68B822B">2025</mods:date>
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<subSubSection id="68D08EFFFFA0CB028BA0FAD1AD752F0B" box="[898,1372,1366,1393]" pageId="15" pageNumber="16" type="nomenclature">
<paragraph id="2075DD74FFA0CB028BA0FAD1AD752F0B" blockId="15.[810,1460,1366,1962]" box="[898,1372,1366,1393]" pageId="15" pageNumber="16">
<emphasis id="12BE0166FFA0CB028BA0FAD1AD752F0B" bold="true" box="[898,1372,1366,1393]" pageId="15" pageNumber="16">
Phylogenetic relationships within
<taxonomicName id="E7CAA6F7FFA0CB028CC8FAD0AD752F0B" ID-CoL="63N92" authorityName="Burmeister" authorityYear="1876" box="[1258,1372,1366,1393]" class="Insecta" family="Trogidae" genus="Polynoncus" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFA0CB028CC8FAD0AD752F0B" bold="true" box="[1258,1372,1366,1393]" italics="true" pageId="15" pageNumber="16">Polynoncus</emphasis>
</taxonomicName>
</emphasis>
</paragraph>
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<subSubSection id="68D08EFFFFA0CB1B8B08FAFBA9702902" lastPageId="22" lastPageNumber="23" pageId="15" pageNumber="16" type="discussion">
<paragraph id="2075DD74FFA0CB028B08FAFBAC9A2D57" blockId="15.[810,1460,1366,1962]" pageId="15" pageNumber="16">
Under both phylogenetic analyses (EW and IW),
<taxonomicName id="E7CAA6F7FFA0CB028D3EFAFBADA02FEF" authorityName="Burmeister" authorityYear="1876" box="[1308,1417,1405,1429]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFA0CB028D3EFAFBADA02FEF" box="[1308,1417,1405,1429]" italics="true" pageId="15" pageNumber="16">Polononcus</emphasis>
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was recovered as monophyletic, corroborating the hypotheses of previous research on the genus (
<emphasis id="12BE0166FFA0CB028C71FA3BAC582FAE" box="[1107,1137,1469,1492]" italics="true" pageId="15" pageNumber="16">e.g.</emphasis>
Scholtz 1986, Scholtz and Peck 1990, Browne
<emphasis id="12BE0166FFA0CB028BE3FA5AABC22F89" box="[961,1003,1499,1523]" italics="true" pageId="15" pageNumber="16">et al</emphasis>
. 1993, Strümpher
<emphasis id="12BE0166FFA0CB028C8FFA5AACFF2F89" box="[1197,1238,1499,1523]" italics="true" pageId="15" pageNumber="16">et al</emphasis>
. 2014). Nonetheless, only the topology obtained from the IW model will be discussed here, as the resolution and level of support were beter solved than that of the EW model (
<figureCitation id="B8F1C1F1FFA0CB028C65F9BFACA22C2B" box="[1095,1163,1593,1617]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="15" pageNumber="16">Fig. 10</figureCitation>
).
<taxonomicName id="E7CAA6F7FFA0CB028C83F9BFAD272C2B" authorityName="Burmeister" authorityYear="1876" box="[1185,1294,1593,1617]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="16" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFA0CB028C83F9BFAD272C2B" box="[1185,1294,1593,1617]" italics="true" pageId="15" pageNumber="16">Polononcus</emphasis>
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monophyly was well supported (RBS = 92; SR = 71), with five synapomorphies (the last one homoplastic): presence of the one single seta in the sixth antennomere (3:0—
<figureCitation id="B8F1C1F1FFA0CB028C1AF911ACAB2CD5" box="[1080,1154,1687,1711]" captionStart="Figure 1" captionStartId="5.[113,178,1725,1749]" captionTargetBox="[114,1458,144,1697]" captionTargetId="figure-7@5.[114,1458,144,1697]" captionTargetPageId="5" captionText="Figure 1. Characters relative of antennae and mouthparts of Trogidae: antennae of: A, Polononcus gemmingeri; B, Omorgus suberosus; C, Trox plicatus. Labrum of:D, Polononcus juglans; E, Trox plicatus; F, Omorgus batesi in dorsal view.Mentum of: G, Polononcus argentinus; H, Trox variolatus in ventral view.Scale bar:0.5 mm." pageId="15" pageNumber="16">Fig. 1A</figureCitation>
); clypeus with apical portion downward abruptly (20:0—
<figureCitation id="B8F1C1F1FFA0CB028C76F931AC892CB5" box="[1108,1184,1719,1743]" captionStart="Figure 2" captionStartId="6.[129,194,1774,1798]" captionTargetBox="[131,1471,145,1744]" captionTargetId="figure-8@6.[129,1473,143,1746]" captionTargetPageId="6" captionText="Figure 2. Characters relative of the head and pronotum of Trogidae.Fronto-lateral view of: A, Polononcus hemisphaericus; B, Trox plicatus. Frontal view of head of: C, Polononcus bullatus; D, Polononcus juglans. Dorsal view of head and pronotum of: E, Polononcus vazdemelloi; F, Trox plicatus. Scale bar: 1 mm." pageId="15" pageNumber="16">Fig. 2A</figureCitation>
); aedeagus with a medial lobe and a dorsal portion surrounding the sof parts (83:0—
<figureCitation id="B8F1C1F1FFA0CB028DAFF950AB6F2D77" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="15" pageNumber="16">Fig. 9B</figureCitation>
); parameres not fully sclerotized (92:1—
<figureCitation id="B8F1C1F1FFA0CB028CD4F973AD172D77" box="[1270,1342,1781,1805]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="15" pageNumber="16">Fig. 9F</figureCitation>
); and with apical torsion inward (93:0—
<figureCitation id="B8F1C1F1FFA0CB028C7BF893AC8A2D57" box="[1113,1187,1813,1837]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="15" pageNumber="16">Fig. 9C</figureCitation>
).
</paragraph>
<paragraph id="2075DD74FFA0CB1D8B64F8B2A99D2DDA" blockId="15.[810,1460,1366,1962]" lastBlockId="16.[128,778,1614,1952]" lastPageId="16" lastPageNumber="17" pageId="15" pageNumber="16">
Te presence of one single seta in the sixth antennomere was first documented by Costa-Silva
<emphasis id="12BE0166FFA0CB028C9DF8D2ACC42D11" box="[1215,1261,1875,1899]" italics="true" pageId="15" pageNumber="16">et al</emphasis>
. (2024) and used as differential diagnosis for the
<taxonomicName id="E7CAA6F7FFA0CB028C5AF8F5AD692DF1" box="[1144,1344,1907,1931]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="15" pageNumber="16" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="12BE0166FFA0CB028C5AF8F5ACCE2DF1" box="[1144,1255,1907,1931]" italics="true" pageId="15" pageNumber="16">Polononcus</emphasis>
species.
</taxonomicName>
Te apical portion of the clypeus slopes downward abruptly (forming a straight angle with the front—
<figureCitation id="B8F1C1F1FFBFCB1D89E4F9C8AA3A2C1C" box="[454,531,1614,1638]" captionStart="Figure 2" captionStartId="6.[129,194,1774,1798]" captionTargetBox="[131,1471,145,1744]" captionTargetId="figure-8@6.[129,1473,143,1746]" captionTargetPageId="6" captionText="Figure 2. Characters relative of the head and pronotum of Trogidae.Fronto-lateral view of: A, Polononcus hemisphaericus; B, Trox plicatus. Frontal view of head of: C, Polononcus bullatus; D, Polononcus juglans. Dorsal view of head and pronotum of: E, Polononcus vazdemelloi; F, Trox plicatus. Scale bar: 1 mm." pageId="16" pageNumber="17">Fig. 2A</figureCitation>
) is historically used to separate
<taxonomicName id="E7CAA6F7FFBFCB1D88FCF9E8A9642CFC" authorityName="Burmeister" authorityYear="1876" box="[222,333,1646,1670]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBFCB1D88FCF9E8A9642CFC" box="[222,333,1646,1670]" italics="true" pageId="16" pageNumber="17">Polononcus</emphasis>
</taxonomicName>
from
<taxonomicName id="E7CAA6F7FFBFCB1D89B3F9E8A9C42CFF" authorityName="Erichson" authorityYear="1847" box="[401,493,1646,1669]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBFCB1D89B3F9E8A9C42CFF" box="[401,493,1646,1669]" italics="true" pageId="16" pageNumber="17">Omorgus</emphasis>
</taxonomicName>
in identification keys (e.g. Vaurie 1962, Scholtz 1986, 1990, Gómez 2008, Almeida and Mise 2009, Strümpher
<emphasis id="12BE0166FFBFCB1D895EF92BA9862CBE" box="[380,431,1708,1732]" italics="true" pageId="16" pageNumber="17">et al.</emphasis>
2016, Costa-Silva
<emphasis id="12BE0166FFBFCB1D8A5FF92BAA992CBE" box="[637,688,1708,1732]" italics="true" pageId="16" pageNumber="17">et al.</emphasis>
2024a). dHotman and Scholtz (1990) mentioned that parameres with inward apical torsion is a synapomorphy of
<taxonomicName id="E7CAA6F7FFBFCB1D8AAEF96DAB2F2D79" authorityName="Nikolajev" authorityYear="2005" box="[652,774,1771,1795]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="subFamily" subFamily="Omorginae">Omorginae</taxonomicName>
. However, in this study, this characteristic was observed just in
<taxonomicName id="E7CAA6F7FFBFCB1D88A2F8ACA96D2D38" box="[128,324,1834,1858]" pageId="16" pageNumber="17">
<emphasis id="12BE0166FFBFCB1D88A2F8ACA8C62D38" box="[128,239,1834,1858]" italics="true" pageId="16" pageNumber="17">Polononcus</emphasis>
species
</taxonomicName>
, indicating that parameres with apical torsion inward is a derived condition (apomorphic), while the primitive one (apex with straight shape) is found in
<taxonomicName id="E7CAA6F7FFBFCB1D8A8CF8EFAB232DFA" authorityName="Erichson" authorityYear="1847" box="[686,778,1897,1920]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBFCB1D8A8CF8EFAB232DFA" box="[686,778,1897,1920]" italics="true" pageId="16" pageNumber="17">Omorgus</emphasis>
</taxonomicName>
and other genera of
<taxonomicName id="E7CAA6F7FFBFCB1D8973F80EA9982DDA" authorityName="MacLeay" authorityYear="1819" box="[337,433,1928,1952]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="family">Trogidae</taxonomicName>
.
</paragraph>
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<paragraph id="2075DD74FFBFCB1D88A0FA48ADE82C64" blockId="16.[129,1473,1486,1567]" pageId="16" pageNumber="17">
<emphasis id="12BE0166FFBFCB1D88A0FA48A8F32F9C" bold="true" box="[130,218,1486,1510]" pageId="16" pageNumber="17">Figure 8.</emphasis>
Characters relative of the elytra and legs of
<taxonomicName id="E7CAA6F7FFBFCB1D8A53FA48AAFF2F9C" authorityName="Burmeister" authorityYear="1876" box="[625,726,1486,1510]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBFCB1D8A53FA48AAFF2F9C" box="[625,726,1486,1510]" italics="true" pageId="16" pageNumber="17">Polononcus</emphasis>
</taxonomicName>
. Half lef dorsal habitus of: A,
<emphasis id="12BE0166FFBFCB1D8BD6FA48ACC62F9C" box="[1012,1263,1486,1510]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="E7CAA6F7FFBFCB1D8BD6FA48AC702F9C" authorityName="Burmeister" authorityYear="1876" box="[1012,1113,1486,1510]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">Polononcus</taxonomicName>
pseudogemmifer
</emphasis>
; B,
<taxonomicName id="E7CAA6F7FFBFCB1D8D30FA48A8E42C78" baseAuthorityName="Vaurie" baseAuthorityYear="1962" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="neuquen">
<emphasis id="12BE0166FFBFCB1D8D30FA48A8E42C78" italics="true" pageId="16" pageNumber="17">Polononcus neuquen</emphasis>
</taxonomicName>
. C, lateral habitus of
<taxonomicName id="E7CAA6F7FFBFCB1D89B3FA6CAA032C78" baseAuthorityName="Guerin-Meneville" baseAuthorityYear="1830" box="[401,554,1514,1538]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="aeger">
<emphasis id="12BE0166FFBFCB1D89B3FA6CAA032C78" box="[401,554,1514,1538]" italics="true" pageId="16" pageNumber="17">Polononcus aeger</emphasis>
</taxonomicName>
. Ventral view of lef mesotibia in: D,
<taxonomicName id="E7CAA6F7FFBFCB1D8BA6FA6CAC682C78" baseAuthorityName="Erichson" baseAuthorityYear="1847" box="[900,1089,1514,1538]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="peruanus">
<emphasis id="12BE0166FFBFCB1D8BA6FA6CAC682C78" box="[900,1089,1514,1538]" italics="true" pageId="16" pageNumber="17">Polononcus peruanus</emphasis>
</taxonomicName>
; E,
<emphasis id="12BE0166FFBFCB1D8C46FA6CAD372C78" box="[1124,1310,1514,1538]" italics="true" pageId="16" pageNumber="17">
<taxonomicName id="E7CAA6F7FFBFCB1D8C46FA6CACE02C78" authorityName="Burmeister" authorityYear="1876" box="[1124,1225,1514,1538]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">Polononcus</taxonomicName>
<taxonomicName id="E7CAA6F7FFBFCB1D8CECFA6DAD372C78" authorityName="Dieguez" authorityYear="2019" box="[1230,1310,1515,1538]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="cropticus">cropticus</taxonomicName>
</emphasis>
. Dorsal view of right foretibia in: F,
<taxonomicName id="E7CAA6F7FFBFCB1D8919F980A9DF2C64" authorityName="Pitino" authorityYear="1987" box="[315,502,1542,1566]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="furcillifer">
<emphasis id="12BE0166FFBFCB1D8919F980A9DF2C64" box="[315,502,1542,1566]" italics="true" pageId="16" pageNumber="17">Polononcus furcillifer</emphasis>
</taxonomicName>
; G,
<taxonomicName id="E7CAA6F7FFBFCB1D8A3FF980AAC42C65" baseAuthorityName="Blanchard" baseAuthorityYear="1847" box="[541,749,1542,1567]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="gemmiferus">
<emphasis id="12BE0166FFBFCB1D8A3FF980AAC42C65" box="[541,749,1542,1567]" italics="true" pageId="16" pageNumber="17">Polononcus gemmiferus</emphasis>
</taxonomicName>
. Lateral view of right foretibia in: H,
<taxonomicName id="E7CAA6F7FFBFCB1D8C6BF980AD302C65" baseAuthorityName="Blanchard" baseAuthorityYear="1847" box="[1097,1305,1542,1567]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="gemmiferus">
<emphasis id="12BE0166FFBFCB1D8C6BF980AD302C65" box="[1097,1305,1542,1567]" italics="true" pageId="16" pageNumber="17">Polononcus gemmiferus</emphasis>
</taxonomicName>
. Scale bar: 1 mm.
</paragraph>
</caption>
<paragraph id="2075DD74FFBFCB1F8B77F9C8AA9C287B" blockId="16.[825,1475,1614,1952]" lastBlockId="18.[128,778,144,1954]" lastPageId="18" lastPageNumber="19" pageId="16" pageNumber="17">
Within the
<taxonomicName id="E7CAA6F7FFBFCB1D8BF1F9C8AC692C1C" authorityName="Burmeister" authorityYear="1876" box="[979,1088,1614,1638]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBFCB1D8BF1F9C8AC692C1C" box="[979,1088,1614,1638]" italics="true" pageId="16" pageNumber="17">Polononcus</emphasis>
</taxonomicName>
clade, three lineages were retrieved and will be hereafer treated as
<taxonomicName id="E7CAA6F7FFBFCB1D8CB8F9E8ACC42CFC" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1178,1261,1646,1670]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBFCB1D8CB8F9E8ACC42CFC" box="[1178,1261,1646,1670]" italics="true" pageId="16" pageNumber="17">pedestris</emphasis>
</taxonomicName>
(A),
<taxonomicName id="E7CAA6F7FFBFCB1D8D13F9E8ADA42CFC" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[1329,1421,1646,1670]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBFCB1D8D13F9E8ADA42CFC" box="[1329,1421,1646,1670]" italics="true" pageId="16" pageNumber="17">pilularius</emphasis>
</taxonomicName>
(B),
<taxonomicName id="E7CAA6F7FFBFCB1D8B1BF90BABB12CDF" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[825,920,1677,1701]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBFCB1D8B1BF90BABB12CDF" box="[825,920,1677,1701]" italics="true" pageId="16" pageNumber="17">brevicollis</emphasis>
</taxonomicName>
clades (C), and their respective subclades (e.g. B1, B2, C1, etc.). Te
<taxonomicName id="E7CAA6F7FFBFCB1D8BD2F92AAC6A2CBE" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1008,1091,1708,1732]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBFCB1D8BD2F92AAC6A2CBE" box="[1008,1091,1708,1732]" italics="true" pageId="16" pageNumber="17">pedestris</emphasis>
</taxonomicName>
clade presented a low branch support (RBS = 8) with three synapomorphies (all homoplastic) (
<figureCitation id="B8F1C1F1FFBFCB1D8DBEF94AAB7A2D79" captionStart="Figure 11" captionStartId="20.[128,193,1106,1130]" captionTargetBox="[131,1471,149,1077]" captionTargetId="figure-506@20.[129,1473,144,1079]" captionTargetPageId="20" captionText="Figure 11. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and pedestris clade.(Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies. Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus pedestris." pageId="16" pageNumber="17">Fig. 11</figureCitation>
): antero-basal constriction with shallow shape (52:1), basal tubercles and medial edge of pronotum not connected (55:1—
<figureCitation id="B8F1C1F1FFBFCB1D8B1BF8ACABAC2D38" box="[825,901,1834,1858]" captionStart="Figure 6" captionStartId="13.[113,178,1839,1863]" captionTargetBox="[175,1397,147,1809]" captionTargetId="figure-6@13.[173,1399,144,1811]" captionTargetPageId="13" captionText="Figure 6. Characters relative of the pronotum and hypomeron of Polononcus. Dorsal view of pronotum in: A, Polononcus chilensis; B, Polononcus gemmiferus; C, Polononcus argentinus; D, Polononcus patagonicus. Hypemeron of: E, Polononcus patagonicus F,Polononcus gemmifer. Scale bar: 1 mm." pageId="16" pageNumber="17">Fig. 6C</figureCitation>
), and the presence of a longitudinal sulcus in the inner face of protibial (78:0). Into this clade,
<taxonomicName id="E7CAA6F7FFBFCB1D8CC2F8CFAD802D1B" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1248,1449,1865,1889]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBFCB1D8CC2F8CFAD802D1B" box="[1248,1449,1865,1889]" italics="true" pageId="16" pageNumber="17">Polononcus pedestris</emphasis>
</taxonomicName>
is retrieved as a sister-group of
<taxonomicName id="E7CAA6F7FFBFCB1D8C42F8EFACDA2DFA" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[1120,1267,1896,1920]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="ecuadorensis">
<emphasis id="12BE0166FFBFCB1D8C42F8EFAC472DFA" box="[1120,1134,1897,1920]" italics="true" pageId="16" pageNumber="17">P</emphasis>
.
<emphasis id="12BE0166FFBFCB1D8C58F8EEACDA2DFA" box="[1146,1267,1896,1920]" italics="true" pageId="16" pageNumber="17">ecuadorensis</emphasis>
</taxonomicName>
+ (
<taxonomicName id="E7CAA6F7FFBFCB1D8D35F8EFADBC2DFA" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1303,1429,1896,1920]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="tenebrosus">
<emphasis id="12BE0166FFBFCB1D8D35F8EFAD0C2DFA" box="[1303,1317,1897,1920]" italics="true" pageId="16" pageNumber="17">P</emphasis>
.
<emphasis id="12BE0166FFBFCB1D8D13F8EEADBC2DFA" box="[1329,1429,1896,1920]" italics="true" pageId="16" pageNumber="17">tenebrosus</emphasis>
</taxonomicName>
+
<taxonomicName id="E7CAA6F7FFBFCB1D8D8CF8EFAB432DDA" baseAuthorityName="Harold" baseAuthorityYear="1872" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="sallei">
<emphasis id="12BE0166FFBFCB1D8D8CF8EFAD952DFA" box="[1454,1468,1897,1920]" italics="true" pageId="16" pageNumber="17">P</emphasis>
.
<emphasis id="12BE0166FFBFCB1D8B1BF80EAB432DDA" box="[825,874,1928,1952]" italics="true" pageId="16" pageNumber="17">sallei</emphasis>
</taxonomicName>
). Tis last subclade (
<taxonomicName id="E7CAA6F7FFBFCB1D8C68F80EACEA2DDA" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[1098,1219,1928,1952]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="ecuadorensis">
<emphasis id="12BE0166FFBFCB1D8C68F80EACEA2DDA" box="[1098,1219,1928,1952]" italics="true" pageId="16" pageNumber="17">ecuadorensis</emphasis>
</taxonomicName>
+ (
<taxonomicName id="E7CAA6F7FFBFCB1D8CC5F80EAD622DDA" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1255,1355,1928,1952]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="tenebrosus">
<emphasis id="12BE0166FFBFCB1D8CC5F80EAD622DDA" box="[1255,1355,1928,1952]" italics="true" pageId="16" pageNumber="17">tenebrosus</emphasis>
</taxonomicName>
<taxonomicName id="E7CAA6F7FFBFCB1D8D72F80EADBC2DDA" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1360,1429,1928,1952]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="17" phylum="Arthropoda" rank="species" species="sallei">
+
<emphasis id="12BE0166FFBFCB1D8D46F80EADBC2DDA" box="[1380,1429,1928,1952]" italics="true" pageId="16" pageNumber="17">sallei</emphasis>
</taxonomicName>
)) is supported by three synapomorphies (the last two being homoplastic) with low branch support (RBS = 8): dorsal surface of a median lobe with a blade-like shape (88:1—
<figureCitation id="B8F1C1F1FFBDCB1F8A60FF49AA832A9C" box="[578,682,207,231]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9G, H</figureCitation>
); apex of the median lobe with a spoon-like shape in lateral view (89:1—
<figureCitation id="B8F1C1F1FFBDCB1F88A3FE8BA8ED2B5F" box="[129,196,269,293]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9I</figureCitation>
); and the inner margin of parameres curve near of the base (96:0—
<figureCitation id="B8F1C1F1FFBDCB1F8928FEABA97E2B3F" box="[266,343,301,325]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9D</figureCitation>
). Even with markable external diagnostic characteristic to separate these three species (see diffential diagnostic in: Costa-Silva
<emphasis id="12BE0166FFBDCB1F8946FEEAA9BD2BF9" box="[356,404,363,387]" italics="true" pageId="18" pageNumber="19">et al.</emphasis>
2024a), the spoon-like shape of the median lobe of those three species as a synapomorphy, and the shared geographical distribution from the highest regions of the north-west of the Andean Mountains (
<collectingCountry id="58DD9DE4FFBDCB1F8A1DFE4FAA5B2B9B" box="[575,626,457,481]" name="Peru" pageId="18" pageNumber="19">Peru</collectingCountry>
and
<collectingCountry id="58DD9DE4FFBDCB1F8A8AFE4FAB2B2B9B" box="[680,770,457,481]" name="Ecuador" pageId="18" pageNumber="19">Ecuador</collectingCountry>
) suggest an evolutive relationship between these species.
</paragraph>
<caption id="74B58DFCFFBECB1C8853F8B4AA0F2DE4" pageId="17" pageNumber="18" startId="17.[113,178,1842,1866]" targetBox="[200,1372,147,1810]" targetPageId="17" targetType="figure">
<paragraph id="2075DD74FFBECB1C8853F8B4AA0F2DE4" blockId="17.[113,1427,1842,1950]" pageId="17" pageNumber="18">
<emphasis id="12BE0166FFBECB1C8853F8B4A8E02D30" bold="true" box="[113,201,1842,1866]" pageId="17" pageNumber="18">Figure 9.</emphasis>
Characters relative of male genitalia of
<taxonomicName id="E7CAA6F7FFBECB1C8A1CF8B4AABC2D30" authorityName="MacLeay" authorityYear="1819" box="[574,661,1842,1866]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="family">Trogidae</taxonomicName>
. Median lobe (with parameres removed, of: A,
<taxonomicName id="E7CAA6F7FFBECB1C8C6DF8B5ACD52D30" baseAuthorityName="Fabricius" baseAuthorityYear="1775" box="[1103,1276,1842,1866]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="suberosus">
<emphasis id="12BE0166FFBECB1C8C6DF8B5ACD52D30" box="[1103,1276,1842,1866]" italics="true" pageId="17" pageNumber="18">Omorgus suberosus</emphasis>
</taxonomicName>
; B,
<taxonomicName id="E7CAA6F7FFBECB1C8D3DF8B4A8F52D1C" authorityName="Huchet and Costa-Silva" authorityYear="2018" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="vazdemelloi">
<emphasis id="12BE0166FFBECB1C8D3DF8B4A8F52D1C" italics="true" pageId="17" pageNumber="18">Polononcus vazdemelloi</emphasis>
</taxonomicName>
. Aedeagus habitus of: C,
<taxonomicName id="E7CAA6F7FFBECB1C89E9F8C8AAAF2D1C" authorityName="Pitino" authorityYear="1987" box="[459,646,1870,1894]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="furcillifer">
<emphasis id="12BE0166FFBECB1C89E9F8C8AAAF2D1C" box="[459,646,1870,1894]" italics="true" pageId="17" pageNumber="18">Polononcus furcillifer</emphasis>
</taxonomicName>
; D,
<taxonomicName id="E7CAA6F7FFBECB1C8A8EF8C8AB422D1C" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[684,875,1870,1894]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBECB1C8A8EF8C8AB422D1C" box="[684,875,1870,1894]" italics="true" pageId="17" pageNumber="18">Polononcus pilularius</emphasis>
</taxonomicName>
; E,
<taxonomicName id="E7CAA6F7FFBECB1C8BADF8C8AC6E2D1C" authorityName="Pitino" authorityYear="1987" box="[911,1095,1870,1894]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="patriciae">
<emphasis id="12BE0166FFBECB1C8BADF8C8AC6E2D1C" box="[911,1095,1870,1894]" italics="true" pageId="17" pageNumber="18">Polononcus patriciae</emphasis>
</taxonomicName>
; F,
<taxonomicName id="E7CAA6F7FFBECB1C8C4AF8C9ACDC2D1C" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[1128,1269,1870,1894]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="batesi">
<emphasis id="12BE0166FFBECB1C8C4AF8C9ACDC2D1C" box="[1128,1269,1870,1894]" italics="true" pageId="17" pageNumber="18">Omorgus batesi</emphasis>
</taxonomicName>
. Dorsal (G) and (H) lateral views of
<taxonomicName id="E7CAA6F7FFBECB1C890FF8ECAA2E2DF8" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[301,519,1898,1922]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="ecuadorensis">
<emphasis id="12BE0166FFBECB1C890FF8ECAA2E2DF8" box="[301,519,1898,1922]" italics="true" pageId="17" pageNumber="18">Polononcus ecuadorensis</emphasis>
</taxonomicName>
. I, lateral view of
<taxonomicName id="E7CAA6F7FFBECB1C8A88F8ECAB492DF8" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[682,864,1898,1922]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="neuquen">
<emphasis id="12BE0166FFBECB1C8A88F8ECAB492DF8" box="[682,864,1898,1922]" italics="true" pageId="17" pageNumber="18">Polononcus neuquen</emphasis>
</taxonomicName>
. Dorsal (J) and (K) lateral views of
<taxonomicName id="E7CAA6F7FFBECB1C8C91F8ECAD632DF8" baseAuthorityName="Fabricius" baseAuthorityYear="1781" box="[1203,1354,1898,1922]" class="Insecta" family="Trogidae" genus="Phoberus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="luridus">
<emphasis id="12BE0166FFBECB1C8C91F8ECAD632DF8" box="[1203,1354,1898,1922]" italics="true" pageId="17" pageNumber="18">Phoberus luridus</emphasis>
</taxonomicName>
. Dorsal view of L,
<taxonomicName id="E7CAA6F7FFBECB1C88F0F801A9572DE4" baseAuthorityName="Olivier" baseAuthorityYear="1789" box="[210,382,1926,1950]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="species" species="squalidus">
<emphasis id="12BE0166FFBECB1C88F0F801A9572DE4" box="[210,382,1926,1950]" italics="true" pageId="17" pageNumber="18">Omorgus squalidus</emphasis>
</taxonomicName>
. Scale bar: 1 mm.
</paragraph>
</caption>
<paragraph id="2075DD74FFBDCB1F88BEFD8EA90F2940" blockId="18.[128,778,144,1954]" pageId="18" pageNumber="19">
Te
<taxonomicName id="E7CAA6F7FFBDCB1F88EBFD8EA90C285A" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[201,293,520,544]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBDCB1F88EBFD8EA90C285A" box="[201,293,520,544]" italics="true" pageId="18" pageNumber="19">pilularius</emphasis>
</taxonomicName>
clade (B) is beter supported than the
<taxonomicName id="E7CAA6F7FFBDCB1F8A94FD8EAB20285A" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[694,777,520,544]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBDCB1F8A94FD8EAB20285A" box="[694,777,520,544]" italics="true" pageId="18" pageNumber="19">pedestris</emphasis>
</taxonomicName>
clade (A) and includes all species with asymmetrical male genitalia (RBS = 49; SR = 35) (
<figureCitation id="B8F1C1F1FFBDCB1F89B9FDC1A9C92825" box="[411,480,583,607]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="18" pageNumber="19">Fig. 10</figureCitation>
). Four synapomorphies, two of which were homoplastic (characters 79 and 90), supported this clade (
<figureCitation id="B8F1C1F1FFBDCB1F88D0FD03A91128E7" box="[242,312,645,669]" captionStart="Figure 12" captionStartId="21.[113,178,903,927]" captionTargetBox="[125,1456,146,873]" captionTargetId="figure-6@21.[114,1458,144,875]" captionTargetPageId="21" captionText="Figure 12. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and pilularius clade.(Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies. Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus vazdemelloi." pageId="18" pageNumber="19">Fig. 12</figureCitation>
): a right mandible with a raised swelling parallel with outer margin (17:0—
<figureCitation id="B8F1C1F1FFBDCB1F899EFD23AA2828C7" box="[444,513,677,701]" captionStart="Figure 3" captionStartId="7.[113,178,1835,1859]" captionTargetBox="[321,1252,143,1807]" captionTargetId="figure-7@7.[321,1252,143,1807]" captionTargetPageId="7" captionText="Figure 3. Characters relative of mouthparts of Trogidae.Maxila, in dorsal view, of: A, Polononcus argentinus; B, Trox luridus. Dorsal view of right mandible of: C, Polononcus gemmingeri; D, Trox variolatus. Lateral view of right mandible of:E, Polononcus gemmingeri; F, Trox variolatus. Scale bar: 0.5 mm." pageId="18" pageNumber="19">Fig. 3E</figureCitation>
), protibiae without an apical tooth in the dorsal surface (79:1), dorsal portion of median lobe with asymmetrical lateral margins at the middle (85:1—
<figureCitation id="B8F1C1F1FFBDCB1F88A3FC85A8E52961" box="[129,204,771,795]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9D</figureCitation>
), and phallobase with a quarter of total aedeagus length (90:2—
<figureCitation id="B8F1C1F1FFBDCB1F88F7FCA4A93F2940" box="[213,278,802,826]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9I</figureCitation>
).
</paragraph>
<paragraph id="2075DD74FFBDCB1F88BEFCC4AA952FD7" blockId="18.[128,778,144,1954]" pageId="18" pageNumber="19">
Two subclades (B1 + B2—
<figureCitation id="B8F1C1F1FFBDCB1F8993FCC7A9DE2923" box="[433,503,833,857]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="18" pageNumber="19">Fig. 10</figureCitation>
) can be recognized within
<taxonomicName id="E7CAA6F7FFBDCB1F88A2FCE7A8CB2903" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[128,226,865,889]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBDCB1F88A4FCE7A8CB2903" box="[134,226,865,889]" italics="true" pageId="18" pageNumber="19">pilularius</emphasis>
</taxonomicName>
(B) clade. Te subclade B1 include the species
<taxonomicName id="E7CAA6F7FFBDCB1F88A2FC06A95F29E2" baseAuthorityName="Van Dyke" baseAuthorityYear="1953" box="[128,374,896,920]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="galapagoensis">
<emphasis id="12BE0166FFBDCB1F88A2FC06A95F29E2" box="[128,374,896,920]" italics="true" pageId="18" pageNumber="19">Polononcus galapagoensis</emphasis>
</taxonomicName>
and
<taxonomicName id="E7CAA6F7FFBDCB1F8987FC07AA1229E2" baseAuthorityName="Mutchler" baseAuthorityYear="1925" box="[421,571,896,920]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="seomourensis">
<emphasis id="12BE0166FFBDCB1F8987FC07A99A29E2" box="[421,435,897,920]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F899CFC07AA1229E2" box="[446,571,897,920]" italics="true" pageId="18" pageNumber="19">seomourensis</emphasis>
</taxonomicName>
, the only
<taxonomicName id="E7CAA6F7FFBDCB1F8ABFFC06A8E029CD" pageId="18" pageNumber="19">
<emphasis id="12BE0166FFBDCB1F8ABFFC06AB2329E2" box="[669,778,896,920]" italics="true" pageId="18" pageNumber="19">Polononcus</emphasis>
species
</taxonomicName>
to occur in the
<collectingRegion id="E20E1396FFBDCB1F894EFC19AA0D29CD" box="[364,548,927,951]" country="Ecuador" name="Galapagos" pageId="18" pageNumber="19">Galapagos Islands</collectingRegion>
(Vaurie 1962, Scholtz 1990b, Costa-Silva
<emphasis id="12BE0166FFBDCB1F8976FC39A9A329AD" box="[340,394,959,983]" italics="true" pageId="18" pageNumber="19">et al.</emphasis>
2024a). Considering the restricted geographical distribution of both species (both being confined to the
<collectingRegion id="E20E1396FFBDCB1F88E8FC7BA9A32E6F" box="[202,394,1021,1045]" country="Ecuador" name="Galapagos" pageId="18" pageNumber="19">Galapagos Islands</collectingRegion>
), Vaurie (1962) and Scholtz (1986, 1990b) hypothesized that both species share a common ancestor. Te highest branch support of our results (RBS = 100; SR = 34) corroborates this hypothesis (
<figureCitation id="B8F1C1F1FFBDCB1F8A02FBDDAA4E2E09" box="[544,615,1115,1139]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="18" pageNumber="19">Fig. 10</figureCitation>
). Furthermore,
<taxonomicName id="E7CAA6F7FFBDCB1F88A3FBFDA9A52EE9" baseAuthorityName="Van Dyke" baseAuthorityYear="1953" box="[129,396,1147,1171]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="galapagoensis">
<emphasis id="12BE0166FFBDCB1F88A3FBFDA9A52EE9" box="[129,396,1147,1171]" italics="true" pageId="18" pageNumber="19">Polononcus galapagoensis</emphasis>
</taxonomicName>
presents reduced membranous wings, suggesting that the fully winged
<taxonomicName id="E7CAA6F7FFBDCB1F8A39FB1DAA9D2EC8" baseAuthorityName="Mutchler" baseAuthorityYear="1925" box="[539,692,1178,1202]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="seomourensis">
<emphasis id="12BE0166FFBDCB1F8A39FB1DAA002EC8" box="[539,553,1179,1202]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8A15FB1DAA9D2EC8" box="[567,692,1179,1202]" italics="true" pageId="18" pageNumber="19">seomourensis</emphasis>
</taxonomicName>
is more primitive and responsible for the dispersion of these species among islands (Costa-Silva
<emphasis id="12BE0166FFBDCB1F8980FB5FA9FA2E8B" box="[418,467,1241,1265]" italics="true" pageId="18" pageNumber="19">et al.</emphasis>
2024a), while the wingless
<taxonomicName id="E7CAA6F7FFBDCB1F8AD7FB5FA92C2F6A" baseAuthorityName="Van Dyke" baseAuthorityYear="1953" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="galapagoensis">
<emphasis id="12BE0166FFBDCB1F8AD7FB5FAB2A2E8A" box="[757,771,1241,1264]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F88A3FB7EA92C2F6A" box="[129,261,1272,1296]" italics="true" pageId="18" pageNumber="19">galapagoensis</emphasis>
</taxonomicName>
is more derived and has lost its flight capacity due to environmental pressures [i.e. restricted areas (islands, mountains), wind, temperature, etc.; for more details, see:
<collectingRegion id="E20E1396FFBDCB1F8ABBFAB1AB202F35" box="[665,777,1335,1359]" country="United Kingdom" name="Darlington" pageId="18" pageNumber="19">Darlington</collectingRegion>
(1943), Scholtz (1981)]. Probably as a result of loss of flight capacity,
<taxonomicName id="E7CAA6F7FFBDCB1F8898FAF0A97F2FF7" baseAuthorityName="Van Dyke" baseAuthorityYear="1953" box="[186,342,1397,1421]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="galapagoensis">
<emphasis id="12BE0166FFBDCB1F8898FAF0A8E12FF7" box="[186,200,1398,1421]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F88F0FAF3A97F2FF7" box="[210,342,1397,1421]" italics="true" pageId="18" pageNumber="19">galapagoensis</emphasis>
</taxonomicName>
is found restricted to the small and isolated northern islands of
<collectingRegion id="E20E1396FFBDCB1F8965FA13A9982FD7" box="[327,433,1429,1453]" country="Ecuador" name="Galapagos" pageId="18" pageNumber="19">Galapagos</collectingRegion>
(Costa-Silva
<emphasis id="12BE0166FFBDCB1F8A18FA13AA402FD7" box="[570,617,1429,1453]" italics="true" pageId="18" pageNumber="19">et al.</emphasis>
2024a).
</paragraph>
<paragraph id="2075DD74FFBDCB1F88BEFA32AC6F287B" blockId="18.[128,778,144,1954]" lastBlockId="18.[825,1475,144,1954]" pageId="18" pageNumber="19">
Te subclade B2 (RBR = 12; SR = 21) (
<figureCitation id="B8F1C1F1FFBDCB1F8A7EFA32AA802FB6" box="[604,681,1460,1484]" captionStart="Figure 12" captionStartId="21.[113,178,903,927]" captionTargetBox="[125,1456,146,873]" captionTargetId="figure-6@21.[114,1458,144,875]" captionTargetPageId="21" captionText="Figure 12. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and pilularius clade.(Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies. Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus vazdemelloi." pageId="18" pageNumber="19">Fig. 12</figureCitation>
) is supported in the presence of the apical callus in the inner margin of protibiae (80:0—
<figureCitation id="B8F1C1F1FFBDCB1F897AFA75A98B2C71" box="[344,418,1523,1547]" captionStart="Figure 8" captionStartId="16.[130,195,1486,1510]" captionTargetBox="[131,1471,146,1456]" captionTargetId="figure-232@16.[129,1473,144,1458]" captionTargetPageId="16" captionText="Figure 8.Characters relative of the elytra and legs of Polononcus. Half lef dorsal habitus of: A, Polononcus pseudogemmifer; B, Polononcus neuquen. C, lateral habitus of Polononcus aeger. Ventral view of lef mesotibia in: D, Polononcus peruanus; E, Polononcus cropticus. Dorsal view of right foretibia in: F, Polononcus furcillifer; G, Polononcus gemmiferus. Lateral view of right foretibia in: H, Polononcus gemmiferus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 8F</figureCitation>
). Tis characteristic is mentioned here for the first time in the literature and can be used to distinguish this group from the remaining species of
<taxonomicName id="E7CAA6F7FFBDCB1F8AB5F9B7AB2D2C33" authorityName="Burmeister" authorityYear="1876" box="[663,772,1585,1609]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBDCB1F8AB5F9B7AB2D2C33" box="[663,772,1585,1609]" italics="true" pageId="18" pageNumber="19">Polononcus</emphasis>
</taxonomicName>
. Also, two homoplasies with phylogenetic signal were identified here: the length of AA1+2 venation is a quarter of AA3+4 length (29:1—
<figureCitation id="B8F1C1F1FFBDCB1F88F7F909A9372CDD" box="[213,286,1679,1703]" captionStart="Figure 5" captionStartId="11.[113,178,1717,1741]" captionTargetBox="[226,1346,144,1689]" captionTargetId="figure-6@11.[226,1346,144,1689]" captionTargetPageId="11" captionText="Figure 5. Characters relative of the dorsal view of lef anterior wing of:A, Polononcus vazdemelloi; B, Polononcus hemisphaericus; C, Polononcus bullatus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 5A</figureCitation>
) and the inner margin of the parameres, near to the base, form a curve of ~90° (96:0—
<figureCitation id="B8F1C1F1FFBDCB1F8A2FF929AA712CBD" box="[525,600,1711,1735]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9D</figureCitation>
). All species that composed the subclade B2 are insular, and most of them occur within tropical regions (
<collectingCountry id="58DD9DE4FFBDCB1F895BF96BA9902D7F" box="[377,441,1773,1797]" name="Brazil" pageId="18" pageNumber="19">Brazil</collectingCountry>
,
<collectingCountry id="58DD9DE4FFBDCB1F89E6F968AA092D7C" box="[452,544,1774,1798]" name="Paraguay" pageId="18" pageNumber="19">Paraguay</collectingCountry>
,
<collectingCountry id="58DD9DE4FFBDCB1F8A09F96BAA5C2D7F" box="[555,629,1773,1797]" name="Bolivia" pageId="18" pageNumber="19">Bolivia</collectingCountry>
,
<collectingCountry id="58DD9DE4FFBDCB1F8A5DF968AAF12D7C" box="[639,728,1774,1798]" name="Uruguay" pageId="18" pageNumber="19">Uruguay</collectingCountry>
, and north-east
<collectingCountry id="58DD9DE4FFBDCB1F88D9F88BA94F2D5F" box="[251,358,1805,1829]" name="Argentina" pageId="18" pageNumber="19">Argentina</collectingCountry>
).
<taxonomicName id="E7CAA6F7FFBDCB1F89A3F88BAA462D5E" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[385,623,1805,1829]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="gemmingeri">
<emphasis id="12BE0166FFBDCB1F89A3F88BAA462D5E" box="[385,623,1805,1829]" italics="true" pageId="18" pageNumber="19">Polononcus gemmingeri</emphasis>
</taxonomicName>
+
<taxonomicName id="E7CAA6F7FFBDCB1F8AAAF88BAB202D5F" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[648,777,1805,1829]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBDCB1F8AAAF88BAABF2D5E" box="[648,662,1805,1828]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8A8FF88BAB202D5F" box="[685,777,1805,1829]" italics="true" pageId="18" pageNumber="19">pilularius</emphasis>
</taxonomicName>
clade (RBS = 3; SR = 35) is recognized as a sister-clade of all remaining species, distinguished by an asymmetrical superior lobe of the aedeagus (
<figureCitation id="B8F1C1F1FFBDCB1F8909F8EDA9462DF9" box="[299,367,1899,1923]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="18" pageNumber="19">Fig. 10</figureCitation>
). Te subclade B3 (RBS = 79; SR = 58) is a monophyletic and well-supported group with the lateral margin of pronotum bulbous located in front of the central portion (50:1—
<figureCitation id="B8F1C1F1FFBDCB1F8BE3FF29AC252ABD" box="[961,1036,175,199]" captionStart="Figure 4" captionStartId="9.[113,178,1839,1863]" captionTargetBox="[187,1385,146,1809]" captionTargetId="figure-6@9.[185,1387,144,1811]" captionTargetPageId="9" captionText="Figure 4. Characters relative of the head, pronotum and metasternum of Polononcus. Dorsal view of head and pronotum in: A, Polononcus bullatus; B, Polononcus hemisphaericus. Ventral view of body in: C, Polononcus patagonicus; D, Polononcus neuquen. Dorsal view of pronotum in: E, Polononcus furcillifer; F, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 4E</figureCitation>
), and symmetrical lateral margins at the middle of the superior lobe of aedeagus (85:0—
<figureCitation id="B8F1C1F1FFBDCB1F8D0AFF49AD592A9D" box="[1320,1392,207,231]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9E</figureCitation>
). Tese characteristics may also be used to distinguish the species group morphologically (Vaurie 1962) and are included in a dichotomous key for species identification or differential diagnosis (i.e. Pitino 1987, Scholtz 1990b, Gómez 2008, Costa-Silva and Diéguez 2020, Costa-Silva
<emphasis id="12BE0166FFBDCB1F8C72FEEAAC532BF9" box="[1104,1146,363,387]" italics="true" pageId="18" pageNumber="19">et al</emphasis>
. 2024a). Even without a formal hypothesis of homology, the arrangements of the terminals present in the identification keys presented by Scholtz (1990b) and, more recently, by Costa-Silva
<emphasis id="12BE0166FFBDCB1F8C54FE4CAC812B9B" box="[1142,1192,457,481]" italics="true" pageId="18" pageNumber="19">et al.</emphasis>
(2024a) are similar to the topologies presented here.
</paragraph>
<paragraph id="2075DD74FFBDCB1F8B77FD8EAB5F29AD" blockId="18.[825,1475,144,1954]" pageId="18" pageNumber="19">
Te
<taxonomicName id="E7CAA6F7FFBDCB1F8BA3FD8EABC9285A" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[897,992,520,544]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBDCB1F8BA3FD8EABC9285A" box="[897,992,520,544]" italics="true" pageId="18" pageNumber="19">brevicollis</emphasis>
</taxonomicName>
clade (C) (
<figureCitation id="B8F1C1F1FFBDCB1F8C7AFD8EACB7285A" box="[1112,1182,520,544]" captionStart="Figure 13" captionStartId="21.[112,177,1849,1873]" captionTargetBox="[129,1457,1045,1820]" captionTargetId="figure-65@21.[113,1457,1045,1820]" captionTargetPageId="21" captionText="Figure 13. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and brevicollis clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies.Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus hemisphaericus." pageId="18" pageNumber="19">Fig. 13</figureCitation>
) is composed of species distributed across
<collectingCountry id="58DD9DE4FFBDCB1F8BFFFDA1AC3E2845" box="[989,1047,551,575]" name="Chile" pageId="18" pageNumber="19">Chile</collectingCountry>
,
<collectingCountry id="58DD9DE4FFBDCB1F8C01FDA1ACA22845" box="[1059,1163,551,575]" name="Argentina" pageId="18" pageNumber="19">Argentina</collectingCountry>
, and
<collectingCountry id="58DD9DE4FFBDCB1F8CE5FDAEAD09283A" box="[1223,1312,552,576]" name="Uruguay" pageId="18" pageNumber="19">Uruguay</collectingCountry>
(
<figureCitation id="B8F1C1F1FFBDCB1F8D16FDA1AD552845" box="[1332,1404,551,575]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="18" pageNumber="19">Fig. 10</figureCitation>
). Tis clade is supported with two synapomorphies (all homoplastic) (RBS = 26): pronotum with a surface of medial edges glabrous (49:0—
<figureCitation id="B8F1C1F1FFBDCB1F8BACFD03ABF528E7" box="[910,988,645,669]" captionStart="Figure 4" captionStartId="9.[113,178,1839,1863]" captionTargetBox="[187,1385,146,1809]" captionTargetId="figure-6@9.[185,1387,144,1811]" captionTargetPageId="9" captionText="Figure 4. Characters relative of the head, pronotum and metasternum of Polononcus. Dorsal view of head and pronotum in: A, Polononcus bullatus; B, Polononcus hemisphaericus. Ventral view of body in: C, Polononcus patagonicus; D, Polononcus neuquen. Dorsal view of pronotum in: E, Polononcus furcillifer; F, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 4A</figureCitation>
) and odd-numbered tubercles of elytra with elongate shape (71:1—
<figureCitation id="B8F1C1F1FFBDCB1F8C0EFD23AC5128C7" box="[1068,1144,677,701]" captionStart="Figure 7" captionStartId="14.[129,194,1839,1863]" captionTargetBox="[220,1382,144,1808]" captionTargetId="figure-7@14.[218,1384,143,1810]" captionTargetPageId="14" captionText="Figure 7. Characters relative of the scutellum and elytra of Trogidae. Scutellum of: A, Polononcus bifurcatus; B, Polononcus bullatus; C, Omorgus suberosus. D, lef elytron of Polononcus pedestris; E, right elytron of Polononcus diffluens. Dorsal habitus of: F, Polononcus pampeanus; G, Polononcus longitarsis. Lateral view of: H, Polononcus chilensis; I, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 7G</figureCitation>
). With few exceptions, the species that compose clade C are the same as those cited by Haaf (1953) and Vaurie (1962) as
<taxonomicName id="E7CAA6F7FFBDCB1F8C53FD65ACFC2881" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[1137,1237,739,763]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBDCB1F8C54FD65ACFC2881" box="[1142,1237,739,763]" italics="true" pageId="18" pageNumber="19">brevicollis</emphasis>
</taxonomicName>
-group in their studies of the global and South American
<taxonomicName id="E7CAA6F7FFBDCB1F8CB0FC85ACC42961" authorityName="MacLeay" authorityYear="1819" box="[1170,1261,771,795]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Trogidae</taxonomicName>
species, respectively. Even so, the diagnostic characteristics used by those authors to recognize the
<taxonomicName id="E7CAA6F7FFBDCB1F8BECFCC7AC042923" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[974,1069,833,857]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBDCB1F8BECFCC7AC042923" box="[974,1069,833,857]" italics="true" pageId="18" pageNumber="19">brevicollis</emphasis>
</taxonomicName>
-group [i.e. presence of humeral callus (70:0—
<figureCitation id="B8F1C1F1FFBDCB1F8BACFCE7ABF12903" box="[910,984,865,889]" captionStart="Figure 7" captionStartId="14.[129,194,1839,1863]" captionTargetBox="[220,1382,144,1808]" captionTargetId="figure-7@14.[218,1384,143,1810]" captionTargetPageId="14" captionText="Figure 7. Characters relative of the scutellum and elytra of Trogidae. Scutellum of: A, Polononcus bifurcatus; B, Polononcus bullatus; C, Omorgus suberosus. D, lef elytron of Polononcus pedestris; E, right elytron of Polononcus diffluens. Dorsal habitus of: F, Polononcus pampeanus; G, Polononcus longitarsis. Lateral view of: H, Polononcus chilensis; I, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 7G</figureCitation>
); margins of elytra subparallel (74:1—
<figureCitation id="B8F1C1F1FFBDCB1F8D4EFCE7AD9E2903" box="[1388,1463,865,889]" captionStart="Figure 7" captionStartId="14.[129,194,1839,1863]" captionTargetBox="[220,1382,144,1808]" captionTargetId="figure-7@14.[218,1384,143,1810]" captionTargetPageId="14" captionText="Figure 7. Characters relative of the scutellum and elytra of Trogidae. Scutellum of: A, Polononcus bifurcatus; B, Polononcus bullatus; C, Omorgus suberosus. D, lef elytron of Polononcus pedestris; E, right elytron of Polononcus diffluens. Dorsal habitus of: F, Polononcus pampeanus; G, Polononcus longitarsis. Lateral view of: H, Polononcus chilensis; I, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 7G</figureCitation>
) and metasternum as wide as long (rhomboidal) (42:0—
<figureCitation id="B8F1C1F1FFBDCB1F8DBEFC06AB7029CD" captionStart="Figure 4" captionStartId="9.[113,178,1839,1863]" captionTargetBox="[187,1385,146,1809]" captionTargetId="figure-6@9.[185,1387,144,1811]" captionTargetPageId="9" captionText="Figure 4. Characters relative of the head, pronotum and metasternum of Polononcus. Dorsal view of head and pronotum in: A, Polononcus bullatus; B, Polononcus hemisphaericus. Ventral view of body in: C, Polononcus patagonicus; D, Polononcus neuquen. Dorsal view of pronotum in: E, Polononcus furcillifer; F, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 4D</figureCitation>
)] were not retrieved as a synapomorphy of this group in this study.
</paragraph>
<paragraph id="2075DD74FFBDCB1F8B77FC58ABC42D5E" blockId="18.[825,1475,144,1954]" pageId="18" pageNumber="19">
Within clade C, the species
<taxonomicName id="E7CAA6F7FFBDCB1F8C56FC58AD11298C" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[1140,1336,990,1014]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="diffluens">
<emphasis id="12BE0166FFBDCB1F8C56FC58AD11298C" box="[1140,1336,990,1014]" italics="true" pageId="18" pageNumber="19">Polononcus diffluens</emphasis>
</taxonomicName>
+
<taxonomicName id="E7CAA6F7FFBDCB1F8D73FC59ADEB298C" authorityName="Dieguez" authorityYear="2019" box="[1361,1474,990,1014]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="cropticus">
<emphasis id="12BE0166FFBDCB1F8D73FC59AD76298C" box="[1361,1375,991,1014]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8D4EFC59ADEB298C" box="[1388,1474,991,1014]" italics="true" pageId="18" pageNumber="19">cropticus</emphasis>
</taxonomicName>
are retrieved as a sister-clade of the other species of the group (
<figureCitation id="B8F1C1F1FFBDCB1F8B66FB9BABA72E4F" box="[836,910,1053,1077]" captionStart="Figure 13" captionStartId="21.[112,177,1849,1873]" captionTargetBox="[129,1457,1045,1820]" captionTargetId="figure-65@21.[113,1457,1045,1820]" captionTargetPageId="21" captionText="Figure 13. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and brevicollis clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies.Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus hemisphaericus." pageId="18" pageNumber="19">Fig. 13</figureCitation>
). Te dichotomy of
<taxonomicName id="E7CAA6F7FFBDCB1F8C56FB9BACCB2E4F" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[1140,1250,1053,1077]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="diffluens">
<emphasis id="12BE0166FFBDCB1F8C56FB9BACAB2E4E" box="[1140,1154,1053,1076]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8CB0FB9BACCB2E4F" box="[1170,1250,1053,1077]" italics="true" pageId="18" pageNumber="19">diffluens</emphasis>
</taxonomicName>
+
<taxonomicName id="E7CAA6F7FFBDCB1F8CDEFB9BAD582E4E" authorityName="Dieguez" authorityYear="2019" box="[1276,1393,1053,1076]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="cropticus">
<emphasis id="12BE0166FFBDCB1F8CDEFB9BAD232E4E" box="[1276,1290,1053,1076]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8D39FB9BAD582E4E" box="[1307,1393,1053,1076]" italics="true" pageId="18" pageNumber="19">cropticus</emphasis>
</taxonomicName>
is supported with four synapomorphies, the first three being homoplastic: shape of anterior margin of labrum symmetric (7:1
<figureCitation id="B8F1C1F1FFBDCB1F8DBEFBDDAB702EE9" captionStart="Figure 1" captionStartId="5.[113,178,1725,1749]" captionTargetBox="[114,1458,144,1697]" captionTargetId="figure-7@5.[114,1458,144,1697]" captionTargetPageId="5" captionText="Figure 1. Characters relative of antennae and mouthparts of Trogidae: antennae of: A, Polononcus gemmingeri; B, Omorgus suberosus; C, Trox plicatus. Labrum of:D, Polononcus juglans; E, Trox plicatus; F, Omorgus batesi in dorsal view.Mentum of: G, Polononcus argentinus; H, Trox variolatus in ventral view.Scale bar:0.5 mm." pageId="18" pageNumber="19">Fig. 1D</figureCitation>
); basal tubercles of pronotum rudimentary (44:0—
<figureCitation id="B8F1C1F1FFBDCB1F8D48FBFDAD862EE9" box="[1386,1455,1147,1171]" captionStart="Figure 4" captionStartId="9.[113,178,1839,1863]" captionTargetBox="[187,1385,146,1809]" captionTargetId="figure-6@9.[185,1387,144,1811]" captionTargetPageId="9" captionText="Figure 4. Characters relative of the head, pronotum and metasternum of Polononcus. Dorsal view of head and pronotum in: A, Polononcus bullatus; B, Polononcus hemisphaericus. Ventral view of body in: C, Polononcus patagonicus; D, Polononcus neuquen. Dorsal view of pronotum in: E, Polononcus furcillifer; F, Polononcus neuquen. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 4B</figureCitation>
); absence of elytral tubercles (64:1); and dorsal portion of median lobe simple, without ornaments and symmetric (87:0—
<figureCitation id="B8F1C1F1FFBDCB1F8DBEFB3FAB7D2E8B" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9F</figureCitation>
). According to Diéguez (2019),
<taxonomicName id="E7CAA6F7FFBDCB1F8C8DFB5FAD332E8B" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[1199,1306,1241,1265]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="diffluens">
<emphasis id="12BE0166FFBDCB1F8C8DFB5FAC942E8A" box="[1199,1213,1241,1264]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8CE8FB5FAD332E8B" box="[1226,1306,1241,1265]" italics="true" pageId="18" pageNumber="19">diffluens</emphasis>
</taxonomicName>
and
<taxonomicName id="E7CAA6F7FFBDCB1F8D72FB5FADEB2E8A" authorityName="Dieguez" authorityYear="2019" box="[1360,1474,1241,1264]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="cropticus">
<emphasis id="12BE0166FFBDCB1F8D72FB5FAD772E8A" box="[1360,1374,1241,1264]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8D4EFB5FADEB2E8A" box="[1388,1474,1241,1264]" italics="true" pageId="18" pageNumber="19">cropticus</emphasis>
</taxonomicName>
present a similar morphology that can be easily confused at first glance. Moreover, both species share a morphological characteristic not typical for
<taxonomicName id="E7CAA6F7FFBDCB1F8C24FAB1ACEE2F35" box="[1030,1223,1335,1359]" pageId="18" pageNumber="19">
<emphasis id="12BE0166FFBDCB1F8C24FAB1AC5A2F35" box="[1030,1139,1335,1359]" italics="true" pageId="18" pageNumber="19">Polononcus</emphasis>
species
</taxonomicName>
: male genitalia are compact and simple, and there is an absence of elytral tubercles. As mentioned by Vaurie (1962), the male genitalia of
<taxonomicName id="E7CAA6F7FFBDCB1F8D77FAF3ABAB2FD6" pageId="18" pageNumber="19">
<emphasis id="12BE0166FFBDCB1F8D77FAF3ADEB2FF7" box="[1365,1474,1397,1421]" italics="true" pageId="18" pageNumber="19">Polononcus</emphasis>
species
</taxonomicName>
are rather complex, with a dorsal (usually asymmetrical and variable) portion surrounding the sof parts (also see: Pitino 1987). Tis complexity of the aedeagus shape was treated as a synapomorphy of
<taxonomicName id="E7CAA6F7FFBDCB1F8BDAFA74AC4C2C70" authorityName="Burmeister" authorityYear="1876" box="[1016,1125,1522,1546]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBDCB1F8BDAFA74AC4C2C70" box="[1016,1125,1522,1546]" italics="true" pageId="18" pageNumber="19">Polononcus</emphasis>
</taxonomicName>
by Scholtz (1986). However, the male genitalia of
<taxonomicName id="E7CAA6F7FFBDCB1F8BCBF995AC7A2C50" baseAuthorityName="Vaurie" baseAuthorityYear="1962" box="[1001,1107,1554,1578]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="diffluens">
<emphasis id="12BE0166FFBDCB1F8BCBF995ABDE2C50" box="[1001,1015,1555,1578]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8C21F994AC7A2C50" box="[1027,1107,1554,1578]" italics="true" pageId="18" pageNumber="19">diffluens</emphasis>
</taxonomicName>
and
<taxonomicName id="E7CAA6F7FFBDCB1F8CA5F995ACDE2C50" authorityName="Dieguez" authorityYear="2019" box="[1159,1271,1554,1578]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="species" species="cropticus">
<emphasis id="12BE0166FFBDCB1F8CA5F995ACBC2C50" box="[1159,1173,1555,1578]" italics="true" pageId="18" pageNumber="19">P</emphasis>
.
<emphasis id="12BE0166FFBDCB1F8C83F995ACDE2C50" box="[1185,1271,1555,1578]" italics="true" pageId="18" pageNumber="19">cropticus</emphasis>
</taxonomicName>
lack the ornaments and bizarre structures on the dorsal portions, instead assuming a compact and simple shape typical of those found in
<taxonomicName id="E7CAA6F7FFBDCB1F8D4AF9D7ABAC2CF2" pageId="18" pageNumber="19">
<emphasis id="12BE0166FFBDCB1F8D4AF9D7ADEB2C12" box="[1384,1474,1617,1640]" italics="true" pageId="18" pageNumber="19">Omorgus</emphasis>
species
</taxonomicName>
; a plesiomorphic condition suggested by Scholtz (1986). Te hypothesis of homology of this character showed a great phylogenetic signal that both species evolved from a common ancestor, and the lack of ornaments in the dorsal portion of aedeagus was result of a secondary lost by phylogenetic reversion (or atavism).
</paragraph>
<paragraph id="2075DD74FFBDCB1E8B77F8AAAA8B2D18" blockId="18.[825,1475,144,1954]" lastBlockId="19.[113,764,1553,1953]" lastPageId="19" lastPageNumber="20" pageId="18" pageNumber="19">
Te subclade C1 (RBS = 63) is supported with two synapomorphies (all homoplastic): presence of a longitudinal sulcus in the inner face of the protibiae (78:0); and apical part of median lobe wide in dorsal view (86:1—
<figureCitation id="B8F1C1F1FFBDCB1F8CF6F80CAD362DD8" box="[1236,1311,1930,1954]" captionStart="Figure 9" captionStartId="17.[113,178,1842,1866]" captionTargetBox="[200,1372,147,1810]" captionTargetId="figure-6@17.[197,1375,144,1814]" captionTargetPageId="17" captionText="Figure 9. Characters relative of male genitalia of Trogidae.Median lobe (with parameres removed, of: A, Omorgus suberosus; B, Polononcus vazdemelloi. Aedeagus habitus of: C, Polononcus furcillifer; D, Polononcus pilularius; E, Polononcus patriciae; F, Omorgus batesi. Dorsal (G) and (H) lateral views of Polononcus ecuadorensis. I, lateral view of Polononcus neuquen. Dorsal (J) and (K) lateral views of Phoberus luridus. Dorsal view of L, Omorgus squalidus. Scale bar: 1 mm." pageId="18" pageNumber="19">Fig. 9E</figureCitation>
). Two lineages of C1 are recognized (
<figureCitation id="B8F1C1F1FFBCCB1E8942F997A9842C53" box="[352,429,1553,1577]" captionStart="Figure 10" captionStartId="19.[114,179,1425,1449]" captionTargetBox="[116,1457,145,1395]" captionTargetId="figure-304@19.[114,1458,144,1397]" captionTargetPageId="19" captionText="Figure 10. Hypothetical reconstruction of the relationships of Polononcus based on the strict consensus tree under implied weight (k = 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively.Te scheme showsthree lineages (pedestris, pilularius, and brevicollis clades) retrieved in the analysis and the geographical distribution in the South America region." pageId="19" pageNumber="20">Figs 10</figureCitation>
,
<figureCitation id="B8F1C1F1FFBCCB1E8999F997A9FC2C53" box="[443,469,1553,1577]" captionStart="Figure 13" captionStartId="21.[112,177,1849,1873]" captionTargetBox="[129,1457,1045,1820]" captionTargetId="figure-65@21.[113,1457,1045,1820]" captionTargetPageId="21" captionText="Figure 13. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and brevicollis clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies.Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus hemisphaericus." pageId="19" pageNumber="20">13</figureCitation>
): subclade
<taxonomicName id="E7CAA6F7FFBCCB1E8A73F994AAE42C53" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[593,717,1553,1577]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBCCB1E8A73F994AA762C53" box="[593,607,1554,1577]" italics="true" pageId="19" pageNumber="20">P</emphasis>
.
<emphasis id="12BE0166FFBCCB1E8A4CF997AAE42C53" box="[622,717,1553,1577]" italics="true" pageId="19" pageNumber="20">brevicollis</emphasis>
</taxonomicName>
+
<taxonomicName id="E7CAA6F7FFBCCB1E8AC4F994A8FD2C32" authorityName="Scholtz" authorityYear="1990" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="gibberosus">
<emphasis id="12BE0166FFBCCB1E8AC4F994AADD2C53" box="[742,756,1554,1577]" italics="true" pageId="19" pageNumber="20">P</emphasis>
.
<emphasis id="12BE0166FFBCCB1E8853F9B6A8FD2C32" box="[113,212,1584,1608]" italics="true" pageId="19" pageNumber="20">gibberosus</emphasis>
</taxonomicName>
+ (C2 + C3). All species of these subclades are restricted to
<collectingCountry id="58DD9DE4FFBCCB1E88CAF9D6A9642C12" box="[232,333,1616,1640]" name="Argentina" pageId="19" pageNumber="20">Argentina</collectingCountry>
and/or
<collectingCountry id="58DD9DE4FFBCCB1E898EF9D6A9CD2C12" box="[428,484,1616,1640]" name="Chile" pageId="19" pageNumber="20">Chile</collectingCountry>
(Diéguez 2008; also see: Costa-Silva
<emphasis id="12BE0166FFBCCB1E88CEF9F6A9322CFD" box="[236,283,1647,1671]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2024a). Both
<taxonomicName id="E7CAA6F7FFBCCB1E898FF9F6AA0F2CFD" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[429,550,1647,1671]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBCCB1E898FF9F6A9922CFD" box="[429,443,1648,1671]" italics="true" pageId="19" pageNumber="20">P</emphasis>
.
<emphasis id="12BE0166FFBCCB1E89E5F9E9AA0F2CFD" box="[455,550,1647,1671]" italics="true" pageId="19" pageNumber="20">brevicollis</emphasis>
</taxonomicName>
and
<taxonomicName id="E7CAA6F7FFBCCB1E8A7AF9F6AAFC2CFD" authorityName="Scholtz" authorityYear="1990" box="[600,725,1647,1671]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="gibberosus">
<emphasis id="12BE0166FFBCCB1E8A7AF9F6AA4F2CFD" box="[600,614,1648,1671]" italics="true" pageId="19" pageNumber="20">P</emphasis>
.
<emphasis id="12BE0166FFBCCB1E8A50F9E9AAFC2CFD" box="[626,725,1647,1671]" italics="true" pageId="19" pageNumber="20">gibberosus</emphasis>
</taxonomicName>
are morphological related species as discussed by Scholtz (1990) and Costa-Silva
<emphasis id="12BE0166FFBCCB1E8938F928A9622CBC" box="[282,331,1710,1734]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
(2024a). Te lineage C2 + C3 (RBS = 8) is supported with three synapomorphies, the last two of which are homoplastic: fourth maxillary palpomere with a sinuosity in the ventral margin near the apex (12:0—
<figureCitation id="B8F1C1F1FFBCCB1E8A3FF88AAA412D5E" box="[541,616,1804,1828]" captionStart="Figure 3" captionStartId="7.[113,178,1835,1859]" captionTargetBox="[321,1252,143,1807]" captionTargetId="figure-7@7.[321,1252,143,1807]" captionTargetPageId="7" captionText="Figure 3. Characters relative of mouthparts of Trogidae.Maxila, in dorsal view, of: A, Polononcus argentinus; B, Trox luridus. Dorsal view of right mandible of: C, Polononcus gemmingeri; D, Trox variolatus. Lateral view of right mandible of:E, Polononcus gemmingeri; F, Trox variolatus. Scale bar: 0.5 mm." pageId="19" pageNumber="20">Fig. 3A</figureCitation>
); basal tubercles of pronotum rudimentary (44:0—
<figureCitation id="B8F1C1F1FFBCCB1E8A20F8ADAA602D39" box="[514,585,1835,1859]" captionStart="Figure 4" captionStartId="9.[113,178,1839,1863]" captionTargetBox="[187,1385,146,1809]" captionTargetId="figure-6@9.[185,1387,144,1811]" captionTargetPageId="9" captionText="Figure 4. Characters relative of the head, pronotum and metasternum of Polononcus. Dorsal view of head and pronotum in: A, Polononcus bullatus; B, Polononcus hemisphaericus. Ventral view of body in: C, Polononcus patagonicus; D, Polononcus neuquen. Dorsal view of pronotum in: E, Polononcus furcillifer; F, Polononcus neuquen. Scale bar: 1 mm." pageId="19" pageNumber="20">Fig. 4B</figureCitation>
); and absence of punctuation in the surface of tubercles (69:1—
<figureCitation id="B8F1C1F1FFBCCB1E8A73F8CCAABB2D18" box="[593,658,1866,1890]" captionStart="Figure 7" captionStartId="14.[129,194,1839,1863]" captionTargetBox="[220,1382,144,1808]" captionTargetId="figure-7@14.[218,1384,143,1810]" captionTargetPageId="14" captionText="Figure 7. Characters relative of the scutellum and elytra of Trogidae. Scutellum of: A, Polononcus bifurcatus; B, Polononcus bullatus; C, Omorgus suberosus. D, lef elytron of Polononcus pedestris; E, right elytron of Polononcus diffluens. Dorsal habitus of: F, Polononcus pampeanus; G, Polononcus longitarsis. Lateral view of: H, Polononcus chilensis; I, Polononcus neuquen. Scale bar: 1 mm." pageId="19" pageNumber="20">Fig. 7I</figureCitation>
).
</paragraph>
<caption id="74B58DFCFFBCCB1E8850FA17ADAB2F9B" pageId="19" pageNumber="20" startId="19.[114,179,1425,1449]" targetBox="[116,1457,145,1395]" targetPageId="19" targetType="figure">
<paragraph id="2075DD74FFBCCB1E8850FA17ADAB2F9B" blockId="19.[113,1448,1425,1505]" pageId="19" pageNumber="20">
<emphasis id="12BE0166FFBCCB1E8850FA17A8FF2FD3" bold="true" box="[114,214,1425,1449]" pageId="19" pageNumber="20">Figure 10.</emphasis>
Hypothetical reconstruction of the relationships of
<taxonomicName id="E7CAA6F7FFBCCB1E8A9CFA17AB0A2FD3" authorityName="Burmeister" authorityYear="1876" box="[702,803,1425,1449]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBCCB1E8A9CFA17AB0A2FD3" box="[702,803,1425,1449]" italics="true" pageId="19" pageNumber="20">Polononcus</emphasis>
</taxonomicName>
based on the strict consensus tree under implied weight (
<emphasis id="12BE0166FFBCCB1E8D63FA17AD652FD3" box="[1345,1356,1425,1449]" italics="true" pageId="19" pageNumber="20">k</emphasis>
= 2762). Values of clade supported obtained by RBS and SR analyses are given above and below the branches, respectively. Te scheme shows three lineages (
<taxonomicName id="E7CAA6F7FFBCCB1E88EEFA4FA9302F9B" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[204,281,1481,1505]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBCCB1E88EEFA4FA9302F9B" box="[204,281,1481,1505]" italics="true" pageId="19" pageNumber="20">pedestris</emphasis>
</taxonomicName>
,
<taxonomicName id="E7CAA6F7FFBCCB1E8901FA4FA9512F9B" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[291,376,1481,1505]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBCCB1E8901FA4FA9512F9B" box="[291,376,1481,1505]" italics="true" pageId="19" pageNumber="20">pilularius</emphasis>
</taxonomicName>
, and
<taxonomicName id="E7CAA6F7FFBCCB1E8988FA4FAA2B2F9B" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[426,514,1481,1505]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBCCB1E8988FA4FAA2B2F9B" box="[426,514,1481,1505]" italics="true" pageId="19" pageNumber="20">brevicollis</emphasis>
</taxonomicName>
clades) retrieved in the analysis and the geographical distribution in the South America region.
</paragraph>
</caption>
<paragraph id="2075DD74FFBCCB1988AFF8ECAB2C2F33" blockId="19.[113,764,1553,1953]" lastBlockId="20.[128,778,1234,1948]" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20">
Te subclade C2 (RBS = 100; SR = 26) is a well-supported clade of Argentinean species (with the exception of
<taxonomicName id="E7CAA6F7FFBCCB1E8A52F80CAADD2DDB" authorityName="Pitino" authorityYear="1987" box="[624,756,1929,1953]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="20" phylum="Arthropoda" rank="species" species="burmeisteri">
<emphasis id="12BE0166FFBCCB1E8A52F80CAA572DDB" box="[624,638,1930,1953]" italics="true" pageId="19" pageNumber="20">P</emphasis>
.
<emphasis id="12BE0166FFBCCB1E8AA5F80FAADD2DDB" box="[647,756,1929,1953]" italics="true" pageId="19" pageNumber="20">burmeisteri</emphasis>
</taxonomicName>
, which also occurs in
<collectingCountry id="58DD9DE4FFBCCB1E8BDEF997AC612C53" box="[1020,1096,1553,1577]" name="Bolivia" pageId="19" pageNumber="20">Bolivia</collectingCountry>
—see: Costa-Silva
<emphasis id="12BE0166FFBCCB1E8D26F994AD1A2C53" box="[1284,1331,1553,1577]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2024a) with three synapomorphies (all homoplastic): AA1+2 venation of wings with a quarter of AA3+4 length (29:1—
<figureCitation id="B8F1C1F1FFBCCB1E8D3CF9D6AD422C1D" box="[1310,1387,1615,1640]" captionStart="Figure 5" captionStartId="11.[113,178,1717,1741]" captionTargetBox="[226,1346,144,1689]" captionTargetId="figure-6@11.[226,1346,144,1689]" captionTargetPageId="11" captionText="Figure 5. Characters relative of the dorsal view of lef anterior wing of:A, Polononcus vazdemelloi; B, Polononcus hemisphaericus; C, Polononcus bullatus. Scale bar: 1 mm." pageId="19" pageNumber="20">Fig. 5A</figureCitation>
); posterior margin of pronotum with constriction (notched) near the angles (54:0—
<figureCitation id="B8F1C1F1FFBCCB1E8BE4F908AC3B2CDC" box="[966,1042,1678,1702]" captionStart="Figure 6" captionStartId="13.[113,178,1839,1863]" captionTargetBox="[175,1397,147,1809]" captionTargetId="figure-6@13.[173,1399,144,1811]" captionTargetPageId="13" captionText="Figure 6. Characters relative of the pronotum and hypomeron of Polononcus. Dorsal view of pronotum in: A, Polononcus chilensis; B, Polononcus gemmiferus; C, Polononcus argentinus; D, Polononcus patagonicus. Hypemeron of: E, Polononcus patagonicus F,Polononcus gemmifer. Scale bar: 1 mm." pageId="19" pageNumber="20">Fig. 6A</figureCitation>
); and surface of hypomeron with setae (57:0—
<figureCitation id="B8F1C1F1FFBCCB1E8B5CF928ABED2CBF" box="[894,964,1709,1734]" captionStart="Figure 6" captionStartId="13.[113,178,1839,1863]" captionTargetBox="[175,1397,147,1809]" captionTargetId="figure-6@13.[173,1399,144,1811]" captionTargetPageId="13" captionText="Figure 6. Characters relative of the pronotum and hypomeron of Polononcus. Dorsal view of pronotum in: A, Polononcus chilensis; B, Polononcus gemmiferus; C, Polononcus argentinus; D, Polononcus patagonicus. Hypemeron of: E, Polononcus patagonicus F,Polononcus gemmifer. Scale bar: 1 mm." pageId="19" pageNumber="20">Fig. 6F</figureCitation>
). Tis group of species (C2) is also supported by the taxonomic history proposed by several authors, where the similarity of morphological characteristic put all species together in the identification key and/or differential diagnosis (i.e. Vaurie 1962, Scholtz 1990b, Costa-Silva and Diéguez 2020, Costa-Silva
<emphasis id="12BE0166FFBCCB1E8B08F8CDAB7D2D18" box="[810,852,1866,1890]" italics="true" pageId="19" pageNumber="20">et al</emphasis>
. 2024a). In both dichotomous keys for
<taxonomicName id="E7CAA6F7FFBCCB1E8CD3F8CCAD992D18" box="[1265,1456,1866,1890]" pageId="19" pageNumber="20">
<emphasis id="12BE0166FFBCCB1E8CD3F8CCAD772D18" box="[1265,1374,1866,1890]" italics="true" pageId="19" pageNumber="20">Polononcus</emphasis>
species
</taxonomicName>
, one carried out by Vaurie (1962) and the other by Costa-Silva
<emphasis id="12BE0166FFBCCB1E8D80F8ECAB6A2DDB" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
(2024a), the authors used the presence of a tooth in the apex of the outer margin of the mesotibia to separate this group from other congeneric species. However, even though it is a good taxonomical characteristic for identification purposes, this character was retrieved as homoplastic (also present in
<taxonomicName id="E7CAA6F7FFBBCB198AA0FAB7AADD2F32" baseAuthorityName="Erichson" baseAuthorityYear="1847" box="[642,756,1329,1352]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="peruanus">
<emphasis id="12BE0166FFBBCB198AA0FAB7AAB92F32" box="[642,656,1329,1352]" italics="true" pageId="20" pageNumber="21">P</emphasis>
.
<emphasis id="12BE0166FFBBCB198AB9FAB7AADD2F32" box="[667,756,1329,1352]" italics="true" pageId="20" pageNumber="21">peruanus</emphasis>
</taxonomicName>
).
</paragraph>
<caption id="74B58DFCFFBBCB1988A2FBD4AC192ED8" pageId="20" pageNumber="21" startId="20.[128,193,1106,1130]" targetBox="[131,1471,149,1077]" targetPageId="20" targetType="figure">
<paragraph id="2075DD74FFBBCB1988A2FBD4AC192ED8" blockId="20.[128,1441,1106,1187]" pageId="20" pageNumber="21">
<emphasis id="12BE0166FFBBCB1988A2FBD4A8CD2E11" bold="true" box="[128,228,1106,1131]" pageId="20" pageNumber="21">Figure 11.</emphasis>
Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and
<taxonomicName id="E7CAA6F7FFBBCB198924FBE9A9712EFD" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[262,344,1135,1159]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBBCB198929FBE9A9712EFD" box="[267,344,1135,1159]" italics="true" pageId="20" pageNumber="21">pedestris</emphasis>
</taxonomicName>
clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies. Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image:
<taxonomicName id="E7CAA6F7FFBBCB198B56FB0DAC022ED9" baseAuthorityName="Harold" baseAuthorityYear="1872" box="[884,1067,1163,1187]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="pedestris">
<emphasis id="12BE0166FFBBCB198B56FB0DAC022ED9" box="[884,1067,1163,1187]" italics="true" pageId="20" pageNumber="21">Polononcus pedestris</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="2075DD74FFBBCB1988BEFAD6AD5C2C39" blockId="20.[128,778,1234,1948]" lastBlockId="20.[825,1475,1234,1948]" pageId="20" pageNumber="21">
Te subclade C3 (RBS = 30) is supported with only one synapomorphy (
<figureCitation id="B8F1C1F1FFBBCB19890DFAE9A95E2FFD" box="[303,375,1391,1415]" captionStart="Figure 13" captionStartId="21.[112,177,1849,1873]" captionTargetBox="[129,1457,1045,1820]" captionTargetId="figure-65@21.[113,1457,1045,1820]" captionTargetPageId="21" captionText="Figure 13. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and brevicollis clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies.Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus hemisphaericus." pageId="20" pageNumber="21">Fig. 13</figureCitation>
): the interocular distance four or five times the length of the eyes (
<figureCitation id="B8F1C1F1FFBBCB1989EAFA08AA3E2FDD" box="[456,535,1422,1447]" captionStart="Figure 2" captionStartId="6.[129,194,1774,1798]" captionTargetBox="[131,1471,145,1744]" captionTargetId="figure-8@6.[129,1473,143,1746]" captionTargetPageId="6" captionText="Figure 2. Characters relative of the head and pronotum of Trogidae.Fronto-lateral view of: A, Polononcus hemisphaericus; B, Trox plicatus. Frontal view of head of: C, Polononcus bullatus; D, Polononcus juglans. Dorsal view of head and pronotum of: E, Polononcus vazdemelloi; F, Trox plicatus. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 2C</figureCitation>
). According to Vaurie (1962), this characteristic corresponds to the
<taxonomicName id="E7CAA6F7FFBBCB198A4BFA28AA952FBC" baseAuthorityName="Curtis" baseAuthorityYear="1844" box="[617,700,1454,1478]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bullatus">
<emphasis id="12BE0166FFBBCB198A4CFA28AA952FBC" box="[622,700,1454,1478]" italics="true" pageId="20" pageNumber="21">bullatus</emphasis>
</taxonomicName>
group, where the species are distinguished by the absence of a humeral callus and reduced or absent wings (as those from the subclade C4). However, the related species
<taxonomicName id="E7CAA6F7FFBBCB1989C8F98BAA762C5E" baseAuthorityName="Erichson" baseAuthorityYear="1847" box="[490,607,1548,1572]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="peruanus">
<emphasis id="12BE0166FFBBCB1989C8F98BA9D12C5E" box="[490,504,1549,1572]" italics="true" pageId="20" pageNumber="21">P</emphasis>
.
<emphasis id="12BE0166FFBBCB198A24F98BAA762C5E" box="[518,607,1549,1572]" italics="true" pageId="20" pageNumber="21">peruanus</emphasis>
</taxonomicName>
and
<taxonomicName id="E7CAA6F7FFBBCB198AB4F98BAB2D2C5E" baseAuthorityName="Gutierrez" baseAuthorityYear="1950" box="[662,772,1548,1572]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="aricensis">
<emphasis id="12BE0166FFBBCB198AB4F98BAA8D2C5E" box="[662,676,1549,1572]" italics="true" pageId="20" pageNumber="21">P</emphasis>
.
<emphasis id="12BE0166FFBBCB198A90F98BAB2D2C5E" box="[690,772,1549,1572]" italics="true" pageId="20" pageNumber="21">aricensis</emphasis>
</taxonomicName>
, both macropterous, also present small eyes, contradicting the diagnostic characteristic of Vaurie (1962) and Haaf (1953) for the
<taxonomicName id="E7CAA6F7FFBBCB1988F2F9ECA90D2CF8" baseAuthorityName="Curtis" baseAuthorityYear="1844" box="[208,292,1642,1666]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bullatus">
<emphasis id="12BE0166FFBBCB1988F4F9ECA90D2CF8" box="[214,292,1642,1666]" italics="true" pageId="20" pageNumber="21">bullatus</emphasis>
</taxonomicName>
group. Te true
<taxonomicName id="E7CAA6F7FFBBCB1989F8F9ECAA072CF8" baseAuthorityName="Curtis" baseAuthorityYear="1844" box="[474,558,1642,1666]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="bullatus">
<emphasis id="12BE0166FFBBCB1989C2F9ECAA072CF8" box="[480,558,1642,1666]" italics="true" pageId="20" pageNumber="21">bullatus</emphasis>
</taxonomicName>
group (
<emphasis id="12BE0166FFBBCB198AAEF9EDAA962CF8" box="[652,703,1643,1666]" italics="true" pageId="20" pageNumber="21">sensu</emphasis>
Vaurie 1962) is retrieved here as monophyletic under the subclade C4 (RBS = 99; SR = 100). Tis well-defined subclade is supported with 18 synapomorphies, being the characters 28, 30, 33, 35, 37, 38, 54, 58, 59, and 69, homoplastic (
<figureCitation id="B8F1C1F1FFBBCB1989D4F961AA152C85" box="[502,572,1767,1791]" captionStart="Figure 13" captionStartId="21.[112,177,1849,1873]" captionTargetBox="[129,1457,1045,1820]" captionTargetId="figure-65@21.[113,1457,1045,1820]" captionTargetPageId="21" captionText="Figure 13. Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and brevicollis clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies.Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image: Polononcus hemisphaericus." pageId="20" pageNumber="21">Fig. 13</figureCitation>
): vestigial posterior wings (25:2—
<figureCitation id="B8F1C1F1FFBBCB198934F880A9482D64" box="[278,353,1798,1822]" captionStart="Figure 5" captionStartId="11.[113,178,1717,1741]" captionTargetBox="[226,1346,144,1689]" captionTargetId="figure-6@11.[226,1346,144,1689]" captionTargetPageId="11" captionText="Figure 5. Characters relative of the dorsal view of lef anterior wing of:A, Polononcus vazdemelloi; B, Polononcus hemisphaericus; C, Polononcus bullatus. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 5C</figureCitation>
); the constriction between R3+4, R3, and R4 venation (pinch) absent (27:1); absence of AA1+2 venation (28:1); end of MP venation not fusing with any venations (30:1—
<figureCitation id="B8F1C1F1FFBBCB19893EF8E2A94F2D06" box="[284,358,1892,1916]" captionStart="Figure 5" captionStartId="11.[113,178,1717,1741]" captionTargetBox="[226,1346,144,1689]" captionTargetId="figure-6@11.[226,1346,144,1689]" captionTargetPageId="11" captionText="Figure 5. Characters relative of the dorsal view of lef anterior wing of:A, Polononcus vazdemelloi; B, Polononcus hemisphaericus; C, Polononcus bullatus. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 5B</figureCitation>
); absence of fusion of the R3+4 venation before the pinch (32:1); absence of the R3 venation (33:1); absence of the R4 venation (34:1); absence of the RP venation (35:1); absence of the RP2 venation (36:1); absence of the MP4 venation (37:1); absence of the fusion of R3+4 venation (38:1); shape of metadiscrimen area nearly twice as wide as long (42:0—
<figureCitation id="B8F1C1F1FFBBCB198C02FAD6AC472F12" box="[1056,1134,1360,1384]" captionStart="Figure 4" captionStartId="9.[113,178,1839,1863]" captionTargetBox="[187,1385,146,1809]" captionTargetId="figure-6@9.[185,1387,144,1811]" captionTargetPageId="9" captionText="Figure 4. Characters relative of the head, pronotum and metasternum of Polononcus. Dorsal view of head and pronotum in: A, Polononcus bullatus; B, Polononcus hemisphaericus. Ventral view of body in: C, Polononcus patagonicus; D, Polononcus neuquen. Dorsal view of pronotum in: E, Polononcus furcillifer; F, Polononcus neuquen. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 4C</figureCitation>
); posterior margin with a constriction (notched) near the angles (54:0—
<figureCitation id="B8F1C1F1FFBBCB198D20FAE9AD642FFD" box="[1282,1357,1391,1415]" captionStart="Figure 6" captionStartId="13.[113,178,1839,1863]" captionTargetBox="[175,1397,147,1809]" captionTargetId="figure-6@13.[173,1399,144,1811]" captionTargetPageId="13" captionText="Figure 6. Characters relative of the pronotum and hypomeron of Polononcus. Dorsal view of pronotum in: A, Polononcus chilensis; B, Polononcus gemmiferus; C, Polononcus argentinus; D, Polononcus patagonicus. Hypemeron of: E, Polononcus patagonicus F,Polononcus gemmifer. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 6A</figureCitation>
); presence of an emargination in the internal margin of anterior angle of proepisternum (58:0—
<figureCitation id="B8F1C1F1FFBBCB198C0EFA28AC5C2FBC" box="[1068,1141,1454,1478]" captionStart="Figure 6" captionStartId="13.[113,178,1839,1863]" captionTargetBox="[175,1397,147,1809]" captionTargetId="figure-6@13.[173,1399,144,1811]" captionTargetPageId="13" captionText="Figure 6. Characters relative of the pronotum and hypomeron of Polononcus. Dorsal view of pronotum in: A, Polononcus chilensis; B, Polononcus gemmiferus; C, Polononcus argentinus; D, Polononcus patagonicus. Hypemeron of: E, Polononcus patagonicus F,Polononcus gemmifer. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 6E</figureCitation>
); absence of a posterior margin projecting backward in obtuse lobe at the middle (59:1—
<figureCitation id="B8F1C1F1FFBBCB198DBEFA4BAB712C7E" captionStart="Figure 6" captionStartId="13.[113,178,1839,1863]" captionTargetBox="[175,1397,147,1809]" captionTargetId="figure-6@13.[173,1399,144,1811]" captionTargetPageId="13" captionText="Figure 6. Characters relative of the pronotum and hypomeron of Polononcus. Dorsal view of pronotum in: A, Polononcus chilensis; B, Polononcus gemmiferus; C, Polononcus argentinus; D, Polononcus patagonicus. Hypemeron of: E, Polononcus patagonicus F,Polononcus gemmifer. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 6C</figureCitation>
); presence of the punctuation in the surface of tubercles (69:0); absence of a humeral callus (70:1—
<figureCitation id="B8F1C1F1FFBBCB198CDAF98AAD172C59" box="[1272,1342,1548,1572]" captionStart="Figure 7" captionStartId="14.[129,194,1839,1863]" captionTargetBox="[220,1382,144,1808]" captionTargetId="figure-7@14.[218,1384,143,1810]" captionTargetPageId="14" captionText="Figure 7. Characters relative of the scutellum and elytra of Trogidae. Scutellum of: A, Polononcus bifurcatus; B, Polononcus bullatus; C, Omorgus suberosus. D, lef elytron of Polononcus pedestris; E, right elytron of Polononcus diffluens. Dorsal habitus of: F, Polononcus pampeanus; G, Polononcus longitarsis. Lateral view of: H, Polononcus chilensis; I, Polononcus neuquen. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 7F</figureCitation>
); and elytral with margins arcuate from apex to base (74:0—
<figureCitation id="B8F1C1F1FFBBCB198D3CF9ADAD4D2C39" box="[1310,1380,1579,1603]" captionStart="Figure 7" captionStartId="14.[129,194,1839,1863]" captionTargetBox="[220,1382,144,1808]" captionTargetId="figure-7@14.[218,1384,143,1810]" captionTargetPageId="14" captionText="Figure 7. Characters relative of the scutellum and elytra of Trogidae. Scutellum of: A, Polononcus bifurcatus; B, Polononcus bullatus; C, Omorgus suberosus. D, lef elytron of Polononcus pedestris; E, right elytron of Polononcus diffluens. Dorsal habitus of: F, Polononcus pampeanus; G, Polononcus longitarsis. Lateral view of: H, Polononcus chilensis; I, Polononcus neuquen. Scale bar: 1 mm." pageId="20" pageNumber="21">Fig. 7F</figureCitation>
).
</paragraph>
<paragraph id="2075DD74FFBBCB1B8B77F9CDAA3F2BB8" blockId="20.[825,1475,1234,1948]" lastBlockId="22.[129,778,144,889]" lastPageId="22" lastPageNumber="23" pageId="20" pageNumber="21">
Te last two characters (70 and 74) were also cited by Vaurie (1962), Scholtz (1990), Gómez (2008), and Costa-Silva
<emphasis id="12BE0166FFBBCB198DB0F9ECADEA2CF8" box="[1426,1475,1642,1666]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
(2024a) in their respective identification keys for
<taxonomicName id="E7CAA6F7FFBBCB198D77F90FABA12CBA" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="undetermined">
<emphasis id="12BE0166FFBBCB198D77F90FADEB2CDB" box="[1365,1474,1673,1697]" italics="true" pageId="20" pageNumber="21">Polononcus</emphasis>
species.
</taxonomicName>
However, these characters are not a true synapomorphy of
<taxonomicName id="E7CAA6F7FFBBCB198B7AF94EABEC2C9A" authorityName="Burmeister" authorityYear="1876" box="[856,965,1736,1760]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBBCB198B7AF94EABEC2C9A" box="[856,965,1736,1760]" italics="true" pageId="20" pageNumber="21">Polononcus</emphasis>
</taxonomicName>
when compared with other wingless species of
<taxonomicName id="E7CAA6F7FFBBCB198B1BF961ABBD2C85" authorityName="MacLeay" authorityYear="1819" box="[825,916,1767,1791]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="family">Trogidae</taxonomicName>
(Scholtz 1981, 2000, Browne
<emphasis id="12BE0166FFBBCB198CF6F96EACD62C85" box="[1236,1279,1767,1791]" italics="true" pageId="20" pageNumber="21">et al</emphasis>
. 1993). Groups of wingless species of the family are reported from different zoogeographical regions [e.g.
<taxonomicName id="E7CAA6F7FFBBCB198C66F8A0ACB72D47" authorityName="Erichson" authorityYear="1847" box="[1092,1182,1830,1853]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBBCB198C66F8A0ACB72D47" box="[1092,1182,1830,1853]" italics="true" pageId="20" pageNumber="21">Omorgus</emphasis>
</taxonomicName>
(
<emphasis id="12BE0166FFBBCB198C8DF8A0ACEB2D47" box="[1199,1218,1830,1853]" italics="true" pageId="20" pageNumber="21">O</emphasis>
.)
<emphasis id="12BE0166FFBBCB198CFAF8A0AD6A2D44" box="[1240,1347,1830,1854]" italics="true" pageId="20" pageNumber="21">pastillarius</emphasis>
(Blanchard, 1847) from South America;
<taxonomicName id="E7CAA6F7FFBBCB198C5BF8C3ACFA2D26" authorityName="Erichson" authorityYear="1847" box="[1145,1235,1861,1884]" class="Insecta" family="Trogidae" genus="Omorgus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFBBCB198C5BF8C3ACFA2D26" box="[1145,1235,1861,1884]" italics="true" pageId="20" pageNumber="21">Omorgus</emphasis>
</taxonomicName>
(
<emphasis id="12BE0166FFBBCB198CC9F8C3ACD72D26" box="[1259,1278,1861,1884]" italics="true" pageId="20" pageNumber="21">O</emphasis>
.)
<emphasis id="12BE0166FFBBCB198D39F8C3AD752D27" box="[1307,1372,1861,1885]" italics="true" pageId="20" pageNumber="21">dohrni</emphasis>
(Harold, 1872) from
<collectingCountry id="58DD9DE4FFBBCB198B96F8E2AC3A2D06" box="[948,1043,1892,1916]" name="Australia" pageId="20" pageNumber="21">Australia</collectingCountry>
;
<taxonomicName id="E7CAA6F7FFBBCB198C3EF8E2AD4C2D06" authority="(Haaf, 1954)" baseAuthorityName="Haaf" baseAuthorityYear="1954" box="[1052,1381,1892,1916]" class="Insecta" family="Trogidae" genus="Phoberus" kingdom="Animalia" order="Coleoptera" pageId="20" pageNumber="21" phylum="Arthropoda" rank="species" species="natalensis">
<emphasis id="12BE0166FFBBCB198C3EF8E2ACF32D06" box="[1052,1242,1892,1916]" italics="true" pageId="20" pageNumber="21">Phoberus natalensis</emphasis>
(Haaf, 1954)
</taxonomicName>
from the Afrotropical region], suggesting that loss of flight appeared many times during the evolution of the group. According to Scholtz (1981), the reduction of the wings can ofen be correlated with the locality and habit of the species (i.e. mountains, deserts, or islands; for additional details, see: Roff 1990). As these environments are stable and favourable for the group (e.g. there are readily available food sources), the species do not need to disperse to search for new habitats (Aukema 1990). Due to this selective pressure or occupation, some morphological structures of beetles wings were lost, termed atrophy (Fiori 1977, Wagner and Liebherr 1992, Browne
<emphasis id="12BE0166FFB9CB1B89BCFE2DA9EE2BB8" box="[414,455,426,450]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. 1993).
</paragraph>
<caption id="74B58DFCFFBACB188853FC01AC1729AC" pageId="21" pageNumber="22" startId="21.[113,178,903,927]" targetBox="[125,1456,146,873]" targetPageId="21" targetType="figure">
<paragraph id="2075DD74FFBACB188853FC01AC1729AC" blockId="21.[113,1434,903,983]" pageId="21" pageNumber="22">
<emphasis id="12BE0166FFBACB188853FC01A8FF29E5" bold="true" box="[113,214,903,927]" pageId="21" pageNumber="22">Figure 12.</emphasis>
Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and
<taxonomicName id="E7CAA6F7FFBACB1888D5FC25A97829C1" authorityName="Polononcus" baseAuthorityName="Germar" baseAuthorityYear="1823" box="[247,337,931,955]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="pilularius">
<emphasis id="12BE0166FFBACB1888DEFC25A97829C1" box="[252,337,931,955]" italics="true" pageId="21" pageNumber="22">pilularius</emphasis>
</taxonomicName>
clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies. Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image:
<taxonomicName id="E7CAA6F7FFBACB188B47FC39AC1029AD" authorityName="Huchet and Costa-Silva" authorityYear="2018" box="[869,1081,959,983]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="vazdemelloi">
<emphasis id="12BE0166FFBACB188B47FC39AC1029AD" box="[869,1081,959,983]" italics="true" pageId="21" pageNumber="22">Polononcus vazdemelloi</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<caption id="74B58DFCFFBACB188852F8BFAC722DF2" pageId="21" pageNumber="22" startId="21.[112,177,1849,1873]" targetBox="[129,1457,1045,1820]" targetPageId="21" targetType="figure">
<paragraph id="2075DD74FFBACB188852F8BFAC722DF2" blockId="21.[112,1435,1849,1929]" pageId="21" pageNumber="22">
<emphasis id="12BE0166FFBACB188852F8BFA8FD2D2B" bold="true" box="[112,212,1849,1873]" pageId="21" pageNumber="22">Figure 13.</emphasis>
Part of the strict consensus tree from two most parsimonious trees showing the phylogenetic relationships of species of the outgroup and
<taxonomicName id="E7CAA6F7FFBACB1888D7F8D3A97A2D17" baseAuthorityName="Eschscholtz" baseAuthorityYear="1822" box="[245,339,1877,1901]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="brevicollis">
<emphasis id="12BE0166FFBACB1888D9F8D3A97A2D17" box="[251,339,1877,1901]" italics="true" pageId="21" pageNumber="22">brevicollis</emphasis>
</taxonomicName>
clade. (Full circles indicate synapomorphies, while empty white circles indicate homoplastic synapomorphies. Black colour refers to state 0; grey refers to state 1; and blue refers to state 2. Image:
<taxonomicName id="E7CAA6F7FFBACB188B46F8F7AC7F2DF3" baseAuthorityName="Burmeister" baseAuthorityYear="1876" box="[868,1110,1905,1929]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="21" pageNumber="22" phylum="Arthropoda" rank="species" species="hemisphaericus">
<emphasis id="12BE0166FFBACB188B46F8F7AC7F2DF3" box="[868,1110,1905,1929]" italics="true" pageId="21" pageNumber="22">Polononcus hemisphaericus</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="2075DD74FFB9CB1B88BEFE4FA9702902" blockId="22.[129,778,144,889]" pageId="22" pageNumber="23">
As also discussed by Scholtz (1981), wingless species of
<taxonomicName id="E7CAA6F7FFB9CB1B88A3FE6FA8F5287B" authorityName="MacLeay" authorityYear="1819" box="[129,220,489,513]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Trogidae</taxonomicName>
are ofen reported within desert and arid regions (i.e. North America, Africa, and
<collectingCountry id="58DD9DE4FFB9CB1B89BBFD8EA9D3285A" box="[409,506,520,544]" name="Australia" pageId="22" pageNumber="23">Australia</collectingCountry>
). Tese reports match with the geographical distribution of the wingless
<taxonomicName id="E7CAA6F7FFB9CB1B8A61FDA1AA992845" authorityName="Burmeister" authorityYear="1876" box="[579,688,551,575]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFB9CB1B8A61FDA1AA992845" box="[579,688,551,575]" italics="true" pageId="22" pageNumber="23">Polononcus</emphasis>
</taxonomicName>
in South America, mainly in arid regions of
<collectingCountry id="58DD9DE4FFB9CB1B89C7FDC0AA632824" box="[485,586,582,606]" name="Argentina" pageId="22" pageNumber="23">Argentina</collectingCountry>
and
<collectingCountry id="58DD9DE4FFB9CB1B8A5FFDC0AA9C2824" box="[637,693,582,606]" name="Chile" pageId="22" pageNumber="23">Chile</collectingCountry>
(Costa-Silva
<emphasis id="12BE0166FFB9CB1B889EFDE1A8D92804" box="[188,240,614,638]" italics="true" pageId="22" pageNumber="23">et al.</emphasis>
2024a). Terefore, as occurs in other
<taxonomicName id="E7CAA6F7FFB9CB1B8AB5FDE0AB202804" box="[663,777,614,638]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
species, the wingless species of
<taxonomicName id="E7CAA6F7FFB9CB1B89F2FD03AA1428E7" authorityName="Burmeister" authorityYear="1876" box="[464,573,645,669]" class="Insecta" family="Trogidae" genus="Polononcus" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="genus">
<emphasis id="12BE0166FFB9CB1B89F2FD03AA1428E7" box="[464,573,645,669]" italics="true" pageId="22" pageNumber="23">Polononcus</emphasis>
</taxonomicName>
are ofen large (up to
<quantity id="E7327091FFB9CB1B88BFFD22A8C028C6" box="[157,233,676,700]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="22" pageNumber="23" unit="mm" value="1.5">1.5 mm</quantity>
), relative to the macropterous species (
<quantity id="E7327091FFB9CB1B8AA3FD22AAD728C6" box="[641,766,676,700]" metricMagnitude="-3" metricUnit="m" metricValue="1.05" metricValueMax="1.5" metricValueMin="0.6" pageId="22" pageNumber="23" unit="mm" value="1.05" valueMax="1.5" valueMin="0.6">0.61.5 mm</quantity>
) (Scholtz 1981). In addition, these species also present a short metasternum, reduced scutellum, absence of humeral callus, and small eyes, in accordance with all non-related species that live in similar regions of the world. For a comprehensive study about the flightless species of
<taxonomicName id="E7CAA6F7FFB9CB1B8991FCC7AA392923" authorityName="MacLeay" authorityYear="1819" box="[435,528,833,857]" class="Insecta" family="Trogidae" kingdom="Animalia" order="Coleoptera" pageId="22" pageNumber="23" phylum="Arthropoda" rank="family">Trogidae</taxonomicName>
, see Scholtz (1981) and Browne
<emphasis id="12BE0166FFB9CB1B88F4FCE7A8D72903" box="[214,254,865,889]" italics="true" pageId="22" pageNumber="23">et al</emphasis>
. (1993).
</paragraph>
</subSubSection>
</treatment>
</document>