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<mods:title id="3410DB8A26FFB555D483704F3C55EB5C">Morphological phylogenetics provide new insights into the classification and evolution of fossil soldier beetles from Mid-Cretaceous Burmese amber (Coleoptera: Cantharidae)</mods:title>
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<mods:namePart id="D91541B1083AF4F5C3ADF54CBECB36CD">Hsiao, Yun</mods:namePart>
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<mods:namePart id="9F697FA7C6D9AC469703E4FF96673EE9">Li, Yun</mods:namePart>
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<mods:namePart id="617EC413D2A87BA14FB9C921E65130FF">Ren, Dong</mods:namePart>
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<treatment id="03E3640FFF8FFFEFD0AA2E94ABB98D4A" ID-DOI="http://doi.org/10.5281/zenodo.5753244" ID-GBIF-Taxon="190877384" ID-Zenodo-Dep="5753244" LSID="urn:lsid:zoobank.org:act:FAB17477-85E4-4D7D-95C9-895FFCD9E9B5" httpUri="http://treatment.plazi.org/id/03E3640FFF8FFFEFD0AA2E94ABB98D4A" lastPageNumber="1281" pageId="10" pageNumber="1281">
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<paragraph id="8BF5D519FF8FFFEFD0AA2E94A9DA89ED" blockId="10.[145,762,953,1006]" pageId="10" pageNumber="1281">
<taxonomicName id="4C4AAE9AFF8FFFEFD0AA2E94A9DA89ED" authority="Hsiao &amp; Li &amp; Ren &amp; Pang, 2021" authorityName="Hsiao &amp; Li &amp; Ren &amp; Pang" authorityYear="2021" class="Insecta" family="Cantharidae" genus="Palaeocantharis" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="1281" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B93E090BFF8FFFEFD0AA2E94A85B89CD" box="[145,334,953,974]" italics="true" pageId="10" pageNumber="1281">Palaeocantharis</emphasis>
<taxonomicNameLabel id="A20DB470FF8FFFEFD16D2E97A8AA89CC" box="[342,447,954,975]" pageId="10" pageNumber="1281" rank="genus">gen. nov.</taxonomicNameLabel>
;
<emphasis id="B93E090BFF8FFFEFD1F02E94AB7D89CD" box="[459,616,953,974]" italics="true" pageId="10" pageNumber="1281">Sanaungulus</emphasis>
Fanti
<emphasis id="B93E090BFF8FFFEFD2832E97ABE689CD" box="[696,755,953,975]" italics="true" pageId="10" pageNumber="1281">et al.</emphasis>
, 2018.
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C3508692FF8FFFEFD0AA293AABB98D4A" pageId="10" pageNumber="1281" type="description">
<paragraph id="8BF5D519FF8FFFEFD0AA293AABB98D4A" blockId="10.[145,762,1047,1866]" pageId="10" pageNumber="1281">
<emphasis id="B93E090BFF8FFFEFD0AA293AA8BA8E2F" box="[145,431,1047,1068]" italics="true" pageId="10" pageNumber="1281">General characteristics:</emphasis>
Body small (2.66.0 mm); head prognathous to slightly deflexed (
<figureCitation id="1371C99CFF8FFFEFD2B42918A9B68E69" captionStart="Figure 4" captionStartId="10.[809,890,1041,1063]" captionTargetBox="[813,1421,195,1001]" captionTargetId="figure-719@10.[813,1421,195,1001]" captionTargetPageId="10" captionText="Figure 4. Details of body parts of Cretocantharis veda: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments; F, tarsus of left hind leg. Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.5753266" httpUri="https://zenodo.org/record/5753266/files/figure.png" pageId="10" pageNumber="1281">Figs 4A, B</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD08C2979A8158E69" box="[183,256,1108,1130]" captionStart="Figure 6" captionStartId="13.[163,241,1157,1179]" captionTargetBox="[403,1203,195,1117]" captionTargetId="figure-285@13.[403,1203,195,1117]" captionTargetPageId="13" captionText="Figure 6. Details of body parts of Palaeocantharis panna: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments of abdomen, with gonostyli indicated by a black arrow; F, tarsi of hind legs. Scale bars: 1.0 mm (A, B, D); 0.5 mm (C, E, F)." figureDoi="http://doi.org/10.5281/zenodo.5753272" httpUri="https://zenodo.org/record/5753272/files/figure.png" pageId="10" pageNumber="1281">6A, B</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD12F2979A84B8E69" box="[276,350,1108,1130]" captionStart="Figure 8" captionStartId="15.[163,241,1245,1267]" captionTargetBox="[403,1203,195,1205]" captionTargetId="figure-356@15.[403,1203,195,1205]" captionTargetPageId="15" captionText="Figure 8. Details of body parts of Hukawngichthyurus maha: A, head, ventral-lateral view; B, ditto, dorsal view; C, antenna indicated by a black arrow; D, pronotum and elytra; E, apical tergite; F, caudal segments, with the process of aedeagus indicated by a black arrow; G, tarsus of left mid leg, with the tibial spurs indicated by a black arrow. Scale bars: 0.2 mm (A, B, E, F, G); 0.5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.5753276" httpUri="https://zenodo.org/record/5753276/files/figure.png" pageId="10" pageNumber="1281">8A, B</figureCitation>
); frontoclypeal suture absent; eyes large, hemispherically prominent; apical maxillary and labial palpomeres bilaterally symmetric, securiform; mandibles usually toothed (
<figureCitation id="1371C99CFF8FFFEFD0A029E2A9E98EE6" box="[155,252,1231,1253]" captionStart="Figure 6" captionStartId="13.[163,241,1157,1179]" captionTargetBox="[403,1203,195,1117]" captionTargetId="figure-285@13.[403,1203,195,1117]" captionTargetPageId="13" captionText="Figure 6. Details of body parts of Palaeocantharis panna: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments of abdomen, with gonostyli indicated by a black arrow; F, tarsi of hind legs. Scale bars: 1.0 mm (A, B, D); 0.5 mm (C, E, F)." figureDoi="http://doi.org/10.5281/zenodo.5753272" httpUri="https://zenodo.org/record/5753272/files/figure.png" pageId="10" pageNumber="1281">Figs 6A</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD13529E2A8418EE7" box="[270,340,1231,1252]" captionStart="Figure 8" captionStartId="15.[163,241,1245,1267]" captionTargetBox="[403,1203,195,1205]" captionTargetId="figure-356@15.[403,1203,195,1205]" captionTargetPageId="15" captionText="Figure 8. Details of body parts of Hukawngichthyurus maha: A, head, ventral-lateral view; B, ditto, dorsal view; C, antenna indicated by a black arrow; D, pronotum and elytra; E, apical tergite; F, caudal segments, with the process of aedeagus indicated by a black arrow; G, tarsus of left mid leg, with the tibial spurs indicated by a black arrow. Scale bars: 0.2 mm (A, B, E, F, G); 0.5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.5753276" httpUri="https://zenodo.org/record/5753276/files/figure.png" pageId="10" pageNumber="1281">8A, B</figureCitation>
), sometimes simple (
<figureCitation id="1371C99CFF8FFFEFD25329E2ABFD8EE7" box="[616,744,1231,1253]" captionStart="Figure 4" captionStartId="10.[809,890,1041,1063]" captionTargetBox="[813,1421,195,1001]" captionTargetId="figure-719@10.[813,1421,195,1001]" captionTargetPageId="10" captionText="Figure 4. Details of body parts of Cretocantharis veda: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments; F, tarsus of left hind leg. Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.5753266" httpUri="https://zenodo.org/record/5753266/files/figure.png" pageId="10" pageNumber="1281">Fig. 4A, B</figureCitation>
); gular sutures close, nearly confluent (Supporting Information,
<figureCitation id="1371C99CFF8FFFEFD1092821A8AC8F22" box="[306,441,1292,1314]" captionStart="Figure 1" captionStartId="2.[145,224,1158,1180]" captionTargetBox="[147,1428,197,1117]" captionTargetId="figure-292@2.[145,1428,195,1118]" captionTargetPageId="2" captionText="Figure 1. The most-parsimonious tree inferred from the implied weighting (K = 10.0) (tree length = 92 steps; consistency index = 77; retention index = 87). Black circles indicate synapomorphies; white circles indicate parallelisms or reversals; character numbers are given above circles, and character states below circles. Image courtesy of Stephen Marshall (Cantharinae), Hua-Te Fang (Silini), Zhenhua Liu (Chauliognathinae), Reiner Richter (Dysmorphocerinae), E. Christina Butler (Tytthonyxini) and Chih-Wen Hsu (Malthininae)." figureDoi="http://doi.org/10.5281/zenodo.5753258" httpUri="https://zenodo.org/record/5753258/files/figure.png" pageId="10" pageNumber="1281">Fig. S1AC</figureCitation>
); antennae 11-segmented. Pronotum simple, not elaborately sculptured, without lateral glandular pores (
<figureCitation id="1371C99CFF8FFFEFD2112864AB928F5D" box="[554,647,1353,1375]" captionStart="Figure 4" captionStartId="10.[809,890,1041,1063]" captionTargetBox="[813,1421,195,1001]" captionTargetId="figure-719@10.[813,1421,195,1001]" captionTargetPageId="10" captionText="Figure 4. Details of body parts of Cretocantharis veda: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments; F, tarsus of left hind leg. Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.5753266" httpUri="https://zenodo.org/record/5753266/files/figure.png" pageId="10" pageNumber="1281">Figs 4C</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD2AC2864ABA28F5C" box="[663,695,1353,1375]" captionStart="Figure 6" captionStartId="13.[163,241,1157,1179]" captionTargetBox="[403,1203,195,1117]" captionTargetId="figure-285@13.[403,1203,195,1117]" captionTargetPageId="13" captionText="Figure 6. Details of body parts of Palaeocantharis panna: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments of abdomen, with gonostyli indicated by a black arrow; F, tarsi of hind legs. Scale bars: 1.0 mm (A, B, D); 0.5 mm (C, E, F)." figureDoi="http://doi.org/10.5281/zenodo.5753272" httpUri="https://zenodo.org/record/5753272/files/figure.png" pageId="10" pageNumber="1281">6C</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD2FC2864ABFD8F5D" box="[711,744,1353,1374]" captionStart="Figure 8" captionStartId="15.[163,241,1245,1267]" captionTargetBox="[403,1203,195,1205]" captionTargetId="figure-356@15.[403,1203,195,1205]" captionTargetPageId="15" captionText="Figure 8. Details of body parts of Hukawngichthyurus maha: A, head, ventral-lateral view; B, ditto, dorsal view; C, antenna indicated by a black arrow; D, pronotum and elytra; E, apical tergite; F, caudal segments, with the process of aedeagus indicated by a black arrow; G, tarsus of left mid leg, with the tibial spurs indicated by a black arrow. Scale bars: 0.2 mm (A, B, E, F, G); 0.5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.5753276" httpUri="https://zenodo.org/record/5753276/files/figure.png" pageId="10" pageNumber="1281">8D</figureCitation>
); mesopreepisternum contiguous; metaventrite with midline long, complete or near so, and branched. Elytra entirely covering abdomen (
<figureCitation id="1371C99CFF8FFFEFD2422888ABD28FB8" box="[633,711,1445,1467]" captionStart="Figure 5" captionStartId="12.[145,227,793,815]" captionTargetBox="[149,757,195,753]" captionTargetId="figure-713@12.[149,757,195,753]" captionTargetPageId="12" captionText="Figure 5. Habitus of Palaeocantharis panna: A, dorsal view; B, ventral-lateral view. Scale bars: 2.0 mm." figureDoi="http://doi.org/10.5281/zenodo.5753270" httpUri="https://zenodo.org/record/5753270/files/figure.png" pageId="10" pageNumber="1281">Fig. 5</figureCitation>
) or shortened in various degree, with abdomen and hindwings partially exposed (
<figureCitation id="1371C99CFF8FFFEFD21628CEAB948FFB" box="[557,641,1507,1529]" captionStart="Figure 3" captionStartId="10.[145,224,813,835]" captionTargetBox="[149,757,195,773]" captionTargetId="figure-700@10.[149,757,195,773]" captionTargetPageId="10" captionText="Figure 3. Habitus of Cretocantharis veda: A, dorsal view; B, ventral-lateral view. Scale bars: 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.5753264" httpUri="https://zenodo.org/record/5753264/files/figure.png" pageId="10" pageNumber="1281">Figs 3</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD2AC28CEABB08FFB" box="[663,677,1507,1528]" captionStart="Figure 7" captionStartId="14.[146,224,783,805]" captionTargetBox="[149,757,195,743]" captionTargetId="figure-687@14.[149,757,195,743]" captionTargetPageId="14" captionText="Figure 7. Habitus of Hukawngichthyurus maha: A, dorsal view; B, ventral-lateral view. Scale bars: 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.5753274" httpUri="https://zenodo.org/record/5753274/files/figure.png" pageId="10" pageNumber="1281">7</figureCitation>
); disc usually deeply punctate, sometimes forming coarse reticulation (
<figureCitation id="1371C99CFF8FFFEFD1012B0DA8898C36" box="[314,412,1568,1590]" captionStart="Figure 4" captionStartId="10.[809,890,1041,1063]" captionTargetBox="[813,1421,195,1001]" captionTargetId="figure-719@10.[813,1421,195,1001]" captionTargetPageId="10" captionText="Figure 4. Details of body parts of Cretocantharis veda: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments; F, tarsus of left hind leg. Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.5753266" httpUri="https://zenodo.org/record/5753266/files/figure.png" pageId="10" pageNumber="1281">Figs 4D</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD1952B0DA8C58C35" box="[430,464,1568,1590]" captionStart="Figure 6" captionStartId="13.[163,241,1157,1179]" captionTargetBox="[403,1203,195,1117]" captionTargetId="figure-285@13.[403,1203,195,1117]" captionTargetPageId="13" captionText="Figure 6. Details of body parts of Palaeocantharis panna: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments of abdomen, with gonostyli indicated by a black arrow; F, tarsi of hind legs. Scale bars: 1.0 mm (A, B, D); 0.5 mm (C, E, F)." figureDoi="http://doi.org/10.5281/zenodo.5753272" httpUri="https://zenodo.org/record/5753272/files/figure.png" pageId="10" pageNumber="1281">6D</figureCitation>
), or sometimes smooth (
<figureCitation id="1371C99CFF8FFFEFD0A02B12A9EE8C57" box="[155,251,1599,1621]" captionStart="Figure 8" captionStartId="15.[163,241,1245,1267]" captionTargetBox="[403,1203,195,1205]" captionTargetId="figure-356@15.[403,1203,195,1205]" captionTargetPageId="15" captionText="Figure 8. Details of body parts of Hukawngichthyurus maha: A, head, ventral-lateral view; B, ditto, dorsal view; C, antenna indicated by a black arrow; D, pronotum and elytra; E, apical tergite; F, caudal segments, with the process of aedeagus indicated by a black arrow; G, tarsus of left mid leg, with the tibial spurs indicated by a black arrow. Scale bars: 0.2 mm (A, B, E, F, G); 0.5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.5753276" httpUri="https://zenodo.org/record/5753276/files/figure.png" pageId="10" pageNumber="1281">Fig. 8D</figureCitation>
). Tibial spurs usually present, rarely absent. Abdomen with large lateral eversible glandular pores (Supporting Information,
<figureCitation id="1371C99CFF8FFFEFD2F32B51A9D58CB3" captionStart="Figure 1" captionStartId="2.[145,224,1158,1180]" captionTargetBox="[147,1428,197,1117]" captionTargetId="figure-292@2.[145,1428,195,1118]" captionTargetPageId="2" captionText="Figure 1. The most-parsimonious tree inferred from the implied weighting (K = 10.0) (tree length = 92 steps; consistency index = 77; retention index = 87). Black circles indicate synapomorphies; white circles indicate parallelisms or reversals; character numbers are given above circles, and character states below circles. Image courtesy of Stephen Marshall (Cantharinae), Hua-Te Fang (Silini), Zhenhua Liu (Chauliognathinae), Reiner Richter (Dysmorphocerinae), E. Christina Butler (Tytthonyxini) and Chih-Wen Hsu (Malthininae)." figureDoi="http://doi.org/10.5281/zenodo.5753258" httpUri="https://zenodo.org/record/5753258/files/figure.png" pageId="10" pageNumber="1281">Fig. S1L</figureCitation>
); ventrite 7 of male with apical margin usually entire or concave (
<figureCitation id="1371C99CFF8FFFEFD14C2B94A8CD8CCD" box="[375,472,1721,1743]" captionStart="Figure 4" captionStartId="10.[809,890,1041,1063]" captionTargetBox="[813,1421,195,1001]" captionTargetId="figure-719@10.[813,1421,195,1001]" captionTargetPageId="10" captionText="Figure 4. Details of body parts of Cretocantharis veda: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments; F, tarsus of left hind leg. Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.5753266" httpUri="https://zenodo.org/record/5753266/files/figure.png" pageId="10" pageNumber="1281">Figs 4E</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD1DD2B94AB108CCD" box="[486,517,1721,1742]" captionStart="Figure 8" captionStartId="15.[163,241,1245,1267]" captionTargetBox="[403,1203,195,1205]" captionTargetId="figure-356@15.[403,1203,195,1205]" captionTargetPageId="15" captionText="Figure 8. Details of body parts of Hukawngichthyurus maha: A, head, ventral-lateral view; B, ditto, dorsal view; C, antenna indicated by a black arrow; D, pronotum and elytra; E, apical tergite; F, caudal segments, with the process of aedeagus indicated by a black arrow; G, tarsus of left mid leg, with the tibial spurs indicated by a black arrow. Scale bars: 0.2 mm (A, B, E, F, G); 0.5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.5753276" httpUri="https://zenodo.org/record/5753276/files/figure.png" pageId="10" pageNumber="1281">8F</figureCitation>
), sometimes deeply emarginate; ventrite 8 of male symmetrical, forming narrow, elongate process. Aedeagus symmetrical (
<figureCitation id="1371C99CFF8FFFEFD1002A38A88E8D29" box="[315,411,1813,1835]" captionStart="Figure 4" captionStartId="10.[809,890,1041,1063]" captionTargetBox="[813,1421,195,1001]" captionTargetId="figure-719@10.[813,1421,195,1001]" captionTargetPageId="10" captionText="Figure 4. Details of body parts of Cretocantharis veda: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments; F, tarsus of left hind leg. Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.5753266" httpUri="https://zenodo.org/record/5753266/files/figure.png" pageId="10" pageNumber="1281">Figs 4E</figureCitation>
,
<figureCitation id="1371C99CFF8FFFEFD1922A38A8DD8D29" box="[425,456,1813,1834]" captionStart="Figure 8" captionStartId="15.[163,241,1245,1267]" captionTargetBox="[403,1203,195,1205]" captionTargetId="figure-356@15.[403,1203,195,1205]" captionTargetPageId="15" captionText="Figure 8. Details of body parts of Hukawngichthyurus maha: A, head, ventral-lateral view; B, ditto, dorsal view; C, antenna indicated by a black arrow; D, pronotum and elytra; E, apical tergite; F, caudal segments, with the process of aedeagus indicated by a black arrow; G, tarsus of left mid leg, with the tibial spurs indicated by a black arrow. Scale bars: 0.2 mm (A, B, E, F, G); 0.5 mm (C, D)." figureDoi="http://doi.org/10.5281/zenodo.5753276" httpUri="https://zenodo.org/record/5753276/files/figure.png" pageId="10" pageNumber="1281">8F</figureCitation>
). Ovipositor with paired well-developed palpiform gonostyli (
<figureCitation id="1371C99CFF8FFFEFD27D2A19AB898D49" box="[582,668,1844,1866]" captionStart="Figure 6" captionStartId="13.[163,241,1157,1179]" captionTargetBox="[403,1203,195,1117]" captionTargetId="figure-285@13.[403,1203,195,1117]" captionTargetPageId="13" captionText="Figure 6. Details of body parts of Palaeocantharis panna: A, head, dorsal view; B, ditto, ventral-lateral view; C, pronotum; D, elytra; E, caudal segments of abdomen, with gonostyli indicated by a black arrow; F, tarsi of hind legs. Scale bars: 1.0 mm (A, B, D); 0.5 mm (C, E, F)." figureDoi="http://doi.org/10.5281/zenodo.5753272" httpUri="https://zenodo.org/record/5753272/files/figure.png" pageId="10" pageNumber="1281">Fig. 6E</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>