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<document id="B54B58683328552AAF971102DB2E11A8" ID-DOI="10.11646/zootaxa.3716.2.8" ID-GBIF-Dataset="703c8417-4446-4e68-9ff2-537f42339b5a" ID-ISSN="1175-5326" ID-Zenodo-Dep="221980" ID-ZooBank="D975C77F-83DB-48C6-A698-84A24665D838" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460681307907" checkinUser="plazi" docAuthor="Fikáček, Martin, Maruyama, Munetoshi, Vondráček, Dominik &amp; Short, Andrew E. Z." docDate="2013" docId="038087C28A37143DFF76FC52FAE7F89D" docLanguage="en" docName="zt03716p288.pdf" docOrigin="Zootaxa 3716 (2)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Chimaerocyon sumatranus Fikáček, Maruyama, Vondráček &amp; Short, 2013, sp. nov." docType="treatment" docVersion="8" lastPageNumber="288" masterDocId="FFB9FFBA8A3F1436FFE1FFBDFFBAFFC3" masterDocTitle="Chimaerocyon gen. nov., a morphologically aberrant myrmecophilous genus of water scavenger beetle (Coleoptera: Hydrophilidae: Sphaeridiinae)" masterLastPageNumber="288" masterPageNumber="277" pageNumber="285" updateTime="1698350497756" updateUser="plazi">
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<mods:title id="1F1AAE09DA817F0B69DE437029E370FE">Chimaerocyon gen. nov., a morphologically aberrant myrmecophilous genus of water scavenger beetle (Coleoptera: Hydrophilidae: Sphaeridiinae)</mods:title>
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<mods:namePart id="89BAE48842EDF56571225072DC326E9C">Fikáček, Martin</mods:namePart>
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<mods:namePart id="23854BB22E2AA1850472206923313113">Maruyama, Munetoshi</mods:namePart>
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<mods:namePart id="31F05845DA2CA4B3EA02A87F82D7E390">Vondráček, Dominik</mods:namePart>
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<mods:namePart id="35E1EC94EEFC7800D246537E8B0EBBC3">Short, Andrew E. Z.</mods:namePart>
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<mods:title id="98BE01CFDBD8EC5D8B9ABD33AC6FA90D">Zootaxa</mods:title>
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<mods:date id="3C2CF191D33A3BF57387258ABB61B116">2013</mods:date>
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<mods:number id="A6751A3DC9459BA952EEC6D1F1FB84B1">3716</mods:number>
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<mods:identifier id="C0E1D3188F3C05B19715962E34EA0770" type="DOI">10.11646/zootaxa.3716.2.8</mods:identifier>
<mods:identifier id="560737EAA607BCA4D73182D5B6FBA5BA" type="GBIF-Dataset">703c8417-4446-4e68-9ff2-537f42339b5a</mods:identifier>
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<treatment id="038087C28A37143DFF76FC52FAE7F89D" ID-DOI="http://doi.org/10.5281/zenodo.6149749" ID-GBIF-Taxon="119579974" ID-Zenodo-Dep="6149749" LSID="urn:lsid:plazi:treatment:038087C28A37143DFF76FC52FAE7F89D" httpUri="http://treatment.plazi.org/id/038087C28A37143DFF76FC52FAE7F89D" lastPageId="11" lastPageNumber="288" pageId="8" pageNumber="285">
<subSubSection id="C333655F8A37143EFF76FC52FE2AFBE8" pageId="8" pageNumber="285" type="nomenclature">
<paragraph id="8B9636D48A37143EFF76FC52FDFEFBC9" blockId="8.[151,580,1007,1067]" box="[151,580,1007,1034]" pageId="8" pageNumber="285">
<heading id="D0DE81B88A37143EFF76FC52FDFEFBC9" bold="true" box="[151,580,1007,1034]" fontSize="11" level="1" pageId="8" pageNumber="285" reason="1">
<emphasis id="B95DEAC68A37143EFF76FC52FDFEFBC9" bold="true" box="[151,580,1007,1034]" pageId="8" pageNumber="285">
<taxonomicName id="4C294D578A37143EFF76FC52FE65FBCA" ID-CoL="69RWM" box="[151,479,1007,1033]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="285" phylum="Arthropoda" rank="species" species="sumatranus" status="sp. nov.">
<emphasis id="B95DEAC68A37143EFF76FC52FE65FBCA" bold="true" box="[151,479,1007,1033]" italics="true" pageId="8" pageNumber="285">Chimaerocyon sumatranus</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A26E57BD8A37143EFE07FC4DFDFEFBC9" box="[486,580,1008,1034]" pageId="8" pageNumber="285" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8B9636D48A37143EFF76FBAFFE2AFBE8" blockId="8.[151,580,1007,1067]" box="[151,400,1042,1067]" pageId="8" pageNumber="285">
(
<figureCitation id="13122A518A37143EFF7EFBAFFEB1FBE8" box="[159,267,1042,1067]" captionStart="FIGURES 1 5" captionStartId="4.[151,264,1928,1950]" captionTargetBox="[171,1417,240,1878]" captionTargetId="figure@4.[151,1436,195,1906]" captionTargetPageId="4" captionText="FIGURES 1 5. Habitus of Chimaerocyon species. 1 3 — male paratype of Chimaerocyon shimadai (1 — dorsal view, 2 — lateral view, 3 — frontal view). 4 5 — holotype of C. sumatranus (4 — dorsal view, 5 — frontal view)." httpUri="https://zenodo.org/record/221981/files/figure.png" pageId="8" pageNumber="285">Figs. 45</figureCitation>
,
<figureCitation id="13122A518A37143EFEF7FBAEFE8EFBE8" box="[278,308,1043,1067]" captionStart="FIGURES 6 14" captionStartId="5.[151,264,1470,1492]" captionTargetBox="[151,1436,205,1448]" captionTargetId="figure@5.[151,1436,193,1449]" captionTargetPageId="5" captionText="FIGURES 6 14. Morphological details of Chimaerocyon shimadai (6 13) and C. sumatranus (14). 6 — ventral view, 7 — egg drawn to scale with Fig. 6, 8 — aedeagus of the holotype, 9 — male sternite 8 of the holotype, 10 — male sternite 9, 11 — hind wing of the holotype, 12 14 — abdominal apex (12 — male holotype of C. shimadai, 13 — female paratype of C. shimadai, 14 — female holotype of C. sumatranus). Abbreviations (numbers refer to characters in Fig. 32): msuc — male suckers on maxilla, elpr — elevated median portion of prosternum, wepi — wide epipleuron posteriorly, shtars — shortened tarsus, abdem — abdominal emargination." httpUri="https://zenodo.org/record/221982/files/figure.png" pageId="8" pageNumber="285">14</figureCitation>
,
<figureCitation id="13122A518A37143EFEA1FBAEFEE4FBE8" box="[320,350,1043,1067]" captionStart="FIGURES 15 22" captionStartId="6.[151,264,1567,1589]" captionTargetBox="[151,1436,193,1545]" captionTargetId="figure@6.[151,1436,193,1546]" captionTargetPageId="6" captionText="FIGURES 15 22. Detailed morphology of Chimaerocyon shimadai (15 21) and C. sumatranus (22). 15 — head in dorsal view, 16 — head in ventral view, 17 — prothorax and head in ventral view, 18 — clypeus in lateral view, 19, 22 — mentum, 20 — profurca, 21 — elevated mesal portion of prosternum. Abbreviations (numbers refer to those used in Fig. 32): cly — clypeus, msuc — male sucker on maxilla." httpUri="https://zenodo.org/record/221983/files/figure.png" pageId="8" pageNumber="285">22</figureCitation>
,
<figureCitation id="13122A518A37143EFE8BFBAEFE3DFBE8" box="[362,391,1043,1067]" captionStart="FIGURES 23 30" captionStartId="7.[151,264,1910,1932]" captionTargetBox="[151,1436,482,1887]" captionTargetId="figure@7.[151,1436,481,1888]" captionTargetPageId="7" captionText="FIGURES 23 30. Detailed morphology of Chimaerocyon shimadai (23 26, 28 30) and C. sumatranus (27). 23 — meso- and metathorax in ventral view, 24 — mesoventrite in lateral view, 26 — median portions of meso- and metaventrite in ventral view, 26 27 — right elytral apex, 28 — metatarsus, 29 — metatarsal claws, 30 — detail of apical portion of elytral series 1 2." httpUri="https://zenodo.org/record/221984/files/figure.png" pageId="8" pageNumber="285">27</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF76FBE7FA9EFBB1" box="[151,1316,1114,1139]" pageId="8" pageNumber="285" type="materials_examined">
<paragraph id="8B9636D48A37143EFF76FBE7FA9EFBB1" blockId="8.[151,1437,1114,1823]" box="[151,1316,1114,1139]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF76FBE7FEFFFBB0" bold="true" box="[151,325,1114,1139]" pageId="8" pageNumber="285">
<typeStatus id="549288768A37143EFF76FBE7FF68FBB0" box="[151,210,1114,1139]" pageId="8" pageNumber="285">Type</typeStatus>
material.
</emphasis>
<typeStatus id="549288768A37143EFEADFBE7FE03FBB0" box="[332,441,1114,1139]" pageId="8" pageNumber="285" type="holotype">Holotype</typeStatus>
: female (ZMHB): NORD-SUMATRA
<quantity id="4CD19B318A37143EFC8CFBE6FC78FBB1" box="[877,962,1115,1139]" metricMagnitude="3" metricUnit="m" metricValue="1.3" pageId="8" pageNumber="285" unit="m" value="1300.0">1300 m</quantity>
/ Brastagi /
<date id="FF9710148A37143EFBAAFBE6FB01FBB0" box="[1099,1211,1115,1139]" pageId="8" pageNumber="285" value="1972-08-17">17.8.1972</date>
ERBER.
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF26FBC3FEFDF9E0" pageId="8" pageNumber="285" type="description">
<paragraph id="8B9636D48A37143EFF26FBC3FEFDF9E0" blockId="8.[151,1437,1114,1823]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF26FBC3FEE3FB54" bold="true" box="[199,345,1150,1175]" pageId="8" pageNumber="285">Description.</emphasis>
Body length 2.5 mm, body width 1.8 mm. Body widest slightly behind elytral base, gradually and arcuately narrowing posteriad; elytra evenly convex in anterior view. Coloration uniformly reddish brown. Head with dense punctation consisting of crescent-like to semicircular punctures, without microsculpture on interstices; punctation on pronotum and elytra as dense as that on head, consisting of slightly finer punctures; pronotal punctures transverse, scar-like, elytral punctures dot-like. Eyes divided by 10.0× the width of one eye in dorsal view. Mentum with sparse and fine punctation. Elytron with remnants of five striae posteromesally situated in moderately deep depression; lateral portion of elytra moderately explanate. Mesoventral plate rhomboid, slightly longer than wide, with slightly concave margins. Median portion of metaventrite uniformly pubescent in female. Abdominal ventrites with dense and rather coarse setiferous punctation, without longer and denser pubescence around apical emargination in female. Abdominal ventrite 5 widely emarginate apically, with posterior margin of emargination weakly convex in female. Metatarsus short, with metatarsoreme 1 ca. 3.0× as long as tarsomere 2. Male unknown.
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF26F993FD05F985" box="[199,703,1582,1607]" pageId="8" pageNumber="285" type="diagnosis">
<paragraph id="8B9636D48A37143EFF26F993FD05F985" blockId="8.[151,1437,1114,1823]" box="[199,703,1582,1607]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF26F993FEFBF984" bold="true" box="[199,321,1582,1607]" pageId="8" pageNumber="285">Diagnosis.</emphasis>
See the diagnosis of
<emphasis id="B95DEAC68A37143EFDD1F992FD05F985" box="[560,703,1582,1606]" italics="true" pageId="8" pageNumber="285">
<taxonomicName id="4C294D578A37143EFDD1F992FD00F985" box="[560,698,1582,1606]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="285" phylum="Arthropoda" rank="species" species="shimadai">C. shimadai</taxonomicName>
.
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF26F9EFFE8CF94C" pageId="8" pageNumber="285" type="etymology">
<paragraph id="8B9636D48A37143EFF26F9EFFE8CF94C" blockId="8.[151,1437,1114,1823]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF26F9EFFEF4F9A8" bold="true" box="[199,334,1618,1643]" pageId="8" pageNumber="285">Etymology.</emphasis>
The species name is derived from the name of Sumatra Island where the only known specimen was collected.
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF26F927FCA8F939" pageId="8" pageNumber="285" type="biology_ecology">
<paragraph id="8B9636D48A37143EFF26F927FCA8F939" blockId="8.[151,1437,1114,1823]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF26F927FE9DF970" bold="true" box="[199,295,1690,1715]" pageId="8" pageNumber="285">Biology.</emphasis>
Unknown. The specimen was collected by prof. J. Erber (Institut für Ekologie, Technische Universität Berlin) who is working on the neurophysiology of social hymenopterans (especially bees). Hence, it cannot be excluded that the specimen was also collected from ants.
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF26F8BBFC55F8DC" box="[199,1007,1798,1823]" pageId="8" pageNumber="285" type="distribution">
<paragraph id="8B9636D48A37143EFF26F8BBFC55F8DC" blockId="8.[151,1437,1114,1823]" box="[199,1007,1798,1823]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF26F8BBFEDAF8DC" bold="true" box="[199,352,1798,1823]" pageId="8" pageNumber="285">Distribution.</emphasis>
Known only from the
<typeStatus id="549288768A37143EFD81F8BAFD2AF8DC" box="[608,656,1799,1823]" pageId="8" pageNumber="285">type</typeStatus>
locality in Sumatra,
<collectingCountry id="F33E76448A37143EFC9BF8BBFC51F8DC" box="[890,1003,1798,1823]" name="Indonesia" pageId="8" pageNumber="285">Indonesia</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143EFF76F8D0FEAFF845" box="[151,277,1901,1926]" pageId="8" pageNumber="285" type="discussion">
<paragraph id="8B9636D48A37143EFF76F8D0FEAFF845" blockId="8.[151,277,1901,1926]" box="[151,277,1901,1926]" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF76F8D0FEAFF845" bold="true" box="[151,277,1901,1926]" pageId="8" pageNumber="285">Discussion</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A37143DFF76F808FB4FFCD6" lastPageId="11" lastPageNumber="288" pageId="8" pageNumber="285" type="biology_ecology">
<paragraph id="8B9636D48A37143FFF76F808FD53FE44" blockId="8.[151,1436,1972,2034]" lastBlockId="9.[151,1436,151,1977]" lastPageId="9" lastPageNumber="286" pageId="8" pageNumber="285">
<emphasis id="B95DEAC68A37143EFF76F808FE7DF80D" bold="true" box="[151,455,1972,1998]" pageId="8" pageNumber="285">
Biology of
<taxonomicName id="4C294D578A37143EFEF7F809FE7AF80E" box="[278,448,1972,1997]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="285" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A37143EFEF7F809FE7AF80E" bold="true" box="[278,448,1972,1997]" italics="true" pageId="8" pageNumber="285">Chimaerocyon</emphasis>
</taxonomicName>
.
</emphasis>
Adults of
<taxonomicName id="4C294D578A37143EFDDFF808FCEAF80E" box="[574,848,1973,1997]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="8" pageNumber="285" phylum="Arthropoda" rank="species" species="shimadai">
<emphasis id="B95DEAC68A37143EFDDFF808FCEAF80E" box="[574,848,1973,1997]" italics="true" pageId="8" pageNumber="285">Chimaerocyon shimadai</emphasis>
</taxonomicName>
were found inside of ant host nests and appear to be highly morphologically modified due to myrmecophily, indicating it is an obligatorily inquiline species. Its limuloid body shape (
<figureCitation id="13122A518A36143FFE73FF2AFDBAFF73" box="[402,512,151,176]" captionStart="FIGURES 1 5" captionStartId="4.[151,264,1928,1950]" captionTargetBox="[171,1417,240,1878]" captionTargetId="figure@4.[151,1436,195,1906]" captionTargetPageId="4" captionText="FIGURES 1 5. Habitus of Chimaerocyon species. 1 3 — male paratype of Chimaerocyon shimadai (1 — dorsal view, 2 — lateral view, 3 — frontal view). 4 5 — holotype of C. sumatranus (4 — dorsal view, 5 — frontal view)." httpUri="https://zenodo.org/record/221981/files/figure.png" pageId="9" pageNumber="286">Figs. 15</figureCitation>
), large body size and numerous defensive structures (ability to roll up the body, covering of antennal bases by sides of clypeus, shield-like body with explanate lateral margins, extremely shortened tarsi) suggest that it may be occasionally attacked by the host ants (Hölldobler &amp; Wilson 1990). We did not discover any gland openings, and even a detailed examination of the apical portion of elytra, which possess a pair of shallow pits (
<figureCitation id="13122A518A36143FFE9DFE9AFE5DFE83" box="[380,487,295,320]" captionStart="FIGURES 1 5" captionStartId="4.[151,264,1928,1950]" captionTargetBox="[171,1417,240,1878]" captionTargetId="figure@4.[151,1436,195,1906]" captionTargetPageId="4" captionText="FIGURES 1 5. Habitus of Chimaerocyon species. 1 3 — male paratype of Chimaerocyon shimadai (1 — dorsal view, 2 — lateral view, 3 — frontal view). 4 5 — holotype of C. sumatranus (4 — dorsal view, 5 — frontal view)." httpUri="https://zenodo.org/record/221981/files/figure.png" pageId="9" pageNumber="286">Figs. 1, 4</figureCitation>
,
<figureCitation id="13122A518A36143FFE12FE9AFD87FE83" box="[499,573,295,320]" captionStart="FIGURES 23 30" captionStartId="7.[151,264,1910,1932]" captionTargetBox="[151,1436,482,1887]" captionTargetId="figure@7.[151,1436,481,1888]" captionTargetPageId="7" captionText="FIGURES 23 30. Detailed morphology of Chimaerocyon shimadai (23 26, 28 30) and C. sumatranus (27). 23 — meso- and metathorax in ventral view, 24 — mesoventrite in lateral view, 26 — median portions of meso- and metaventrite in ventral view, 26 27 — right elytral apex, 28 — metatarsus, 29 — metatarsal claws, 30 — detail of apical portion of elytral series 1 2." httpUri="https://zenodo.org/record/221984/files/figure.png" pageId="9" pageNumber="286">2627</figureCitation>
), did not reveal any secretory structures (
<figureCitation id="13122A518A36143FFBE9FE9AFBE6FE83" box="[1032,1116,295,320]" captionStart="FIGURES 23 30" captionStartId="7.[151,264,1910,1932]" captionTargetBox="[151,1436,482,1887]" captionTargetId="figure@7.[151,1436,481,1888]" captionTargetPageId="7" captionText="FIGURES 23 30. Detailed morphology of Chimaerocyon shimadai (23 26, 28 30) and C. sumatranus (27). 23 — meso- and metathorax in ventral view, 24 — mesoventrite in lateral view, 26 — median portions of meso- and metaventrite in ventral view, 26 27 — right elytral apex, 28 — metatarsus, 29 — metatarsal claws, 30 — detail of apical portion of elytral series 1 2." httpUri="https://zenodo.org/record/221984/files/figure.png" pageId="9" pageNumber="286">Fig. 30</figureCitation>
). We therefore conclude that
<taxonomicName id="4C294D578A36143FFF76FEF1FE81FEA7" box="[151,315,332,356]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFF76FEF1FE81FEA7" box="[151,315,332,356]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
is likely an obligatory synoekete (i.e. guest living unnoticed in the nest) or synechthran (i.e. guest attacked by the host when spotted) in
<taxonomicName id="4C294D578A36143FFDDFFECDFD18FE4B" box="[574,674,368,392]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFDDFFECDFD18FE4B" box="[574,674,368,392]" italics="true" pageId="9" pageNumber="286">Pheidole</emphasis>
</taxonomicName>
nests.
</paragraph>
<paragraph id="8B9636D48A36143FFF26FE29FC4FFD0E" blockId="9.[151,1436,151,1977]" pageId="9" pageNumber="286">
The host of
<taxonomicName id="4C294D578A36143FFEB8FE29FDD4FE6F" box="[345,622,404,428]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="species" species="shimadai">
<emphasis id="B95DEAC68A36143FFEB8FE29FDD4FE6F" box="[345,622,404,428]" italics="true" pageId="9" pageNumber="286">Chimaerocyon shimadai</emphasis>
</taxonomicName>
(i.e.,
<taxonomicName id="4C294D578A36143FFD56FE29FC7AFE6F" box="[695,960,404,428]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="species" species="singaporensis">
<emphasis id="B95DEAC68A36143FFD56FE29FC7AFE6F" box="[695,960,404,428]" italics="true" pageId="9" pageNumber="286">Pheidole singaporensis</emphasis>
</taxonomicName>
) is an unusual species of the genus that differs from other
<taxonomicName id="4C294D578A36143FFE86FE05FE71FE13" box="[359,459,440,464]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFE86FE05FE71FE13" box="[359,459,440,464]" italics="true" pageId="9" pageNumber="286">Pheidole</emphasis>
</taxonomicName>
by its very long legs and large body. In the past, the species was even placed together with few other
<taxonomicName id="4C294D578A36143FFEA3FE61FE1CFE37" box="[322,422,476,500]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFEA3FE61FE1CFE37" box="[322,422,476,500]" italics="true" pageId="9" pageNumber="286">Pheidole</emphasis>
</taxonomicName>
species in a separate genus,
<taxonomicName id="4C294D578A36143FFD09FE61FC31FE37" box="[744,907,476,500]" class="Insecta" family="Formicidae" genus="Ischnomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFD09FE61FC31FE37" box="[744,907,476,500]" italics="true" pageId="9" pageNumber="286">Ischnomyrmex</emphasis>
</taxonomicName>
. The presence of
<taxonomicName id="4C294D578A36143FFBB5FE61FB42FE37" box="[1108,1272,476,500]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFBB5FE61FB42FE37" box="[1108,1272,476,500]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
in the nests of this ant species is rather surprising, as no hydrophilids were found in numerous nests of other
<taxonomicName id="4C294D578A36143FFB3FFDBDFAF8FDDB" box="[1246,1346,512,536]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFB3FFDBDFAF8FDDB" box="[1246,1346,512,536]" italics="true" pageId="9" pageNumber="286">Pheidole</emphasis>
</taxonomicName>
species examined by the second author.
<taxonomicName id="4C294D578A36143FFDE7FD99FCB7FDFF" box="[518,781,548,572]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="species" species="singaporensis">
<emphasis id="B95DEAC68A36143FFDE7FD99FCB7FDFF" box="[518,781,548,572]" italics="true" pageId="9" pageNumber="286">Pheidole singaporensis</emphasis>
</taxonomicName>
is a common species widely distributed from
<collectingCountry id="F33E76448A36143FFAFBFD99FAC4FDFE" box="[1306,1406,548,573]" name="Thailand" pageId="9" pageNumber="286">Thailand</collectingCountry>
to
<collectingCountry id="F33E76448A36143FFF76FDFAFEBBFDA3" box="[151,257,583,608]" name="Malaysia" pageId="9" pageNumber="286">Malaysia</collectingCountry>
,
<collectingCountry id="F33E76448A36143FFEEFFDFAFE3AFDA3" box="[270,384,583,608]" name="Singapore" pageId="9" pageNumber="286">Singapore</collectingCountry>
and
<collectingCountry id="F33E76448A36143FFE5DFDFAFD97FDA3" box="[444,557,583,608]" name="Indonesia" pageId="9" pageNumber="286">Indonesia</collectingCountry>
. Although the biology of
<taxonomicName id="4C294D578A36143FFCBDFDF5FBB9FDA3" box="[860,1027,584,608]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="species" species="sumatranus">
<emphasis id="B95DEAC68A36143FFCBDFDF5FBB9FDA3" box="[860,1027,584,608]" italics="true" pageId="9" pageNumber="286">C. sumatranus</emphasis>
</taxonomicName>
remains unknown, it could also be associated with
<taxonomicName id="4C294D578A36143FFEABFDD0FDBAFD47" box="[330,512,621,644]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="species" species="singaporensis">
<emphasis id="B95DEAC68A36143FFEABFDD0FDBAFD47" box="[330,512,621,644]" italics="true" pageId="9" pageNumber="286">P. singaporensis</emphasis>
</taxonomicName>
as the latter ant species also occurs in Sumatra. Further collecting effort should be focused on the nests of
<taxonomicName id="4C294D578A36143FFE43FD2CFDE3FD64" box="[418,601,656,680]" class="Insecta" family="Formicidae" genus="Pheidole" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="species" species="singaporensis">
<emphasis id="B95DEAC68A36143FFE43FD2CFDE3FD64" box="[418,601,656,680]" italics="true" pageId="9" pageNumber="286">P. singaporensis</emphasis>
</taxonomicName>
throughout its range as well as of the related species previously placed in
<emphasis id="B95DEAC68A36143FFF76FD09FEFBFD0F" box="[151,321,692,716]" italics="true" pageId="9" pageNumber="286">
<taxonomicName id="4C294D578A36143FFF76FD09FE87FD0F" box="[151,317,692,716]" class="Insecta" family="Formicidae" genus="Ischnomyrmex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Hymenoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">Ischnomyrmex</taxonomicName>
,
</emphasis>
as additional species of
<taxonomicName id="4C294D578A36143FFDB5FD09FD42FD0F" box="[596,760,692,716]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFDB5FD09FD42FD0F" box="[596,760,692,716]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
are likely to be found.
</paragraph>
<paragraph id="8B9636D48A36143FFF26FD6AFAC3FBD3" blockId="9.[151,1436,151,1977]" pageId="9" pageNumber="286">
<emphasis id="B95DEAC68A36143FFF26FD6AFD28FD33" bold="true" box="[199,658,727,752]" pageId="9" pageNumber="286">
Taxonomic position of
<taxonomicName id="4C294D578A36143FFE00FD6AFD30FD33" box="[481,650,727,752]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFE00FD6AFD30FD33" bold="true" box="[481,650,727,752]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
.
</emphasis>
Based on the initial morphological study of the above specimens,
<taxonomicName id="4C294D578A36143FFF76FD41FE81FCD7" box="[151,315,764,788]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFF76FD41FE81FCD7" box="[151,315,764,788]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
could be assigned to the subfamily
<taxonomicName id="4C294D578A36143FFD16FD41FC2BFCD6" box="[759,913,764,789]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
based on the combination of the following characters: (1) metaventrite completely fused with metanepisterna [outside
<taxonomicName id="4C294D578A36143FFBEFFCA2FB12FCFB" box="[1038,1192,799,824]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
present only in few highly derived genera of
<taxonomicName id="4C294D578A36143FFE54FCF9FDF9FC9E" box="[437,579,836,861]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Rygmodinae">Rygmodinae</taxonomicName>
and Amphiopini; see Short &amp; Fikáček 2013]; (2) posteromedian portion of metaventrite elevated into a mesoventral plate [characteristic for
<taxonomicName id="4C294D578A36143FFC7FFCDAFB81FC43" box="[926,1083,871,896]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
, infrequently present in other tribes]; (3) abdominal ventrite 1 carinate basally [characteristic for most
<taxonomicName id="4C294D578A36143FFC5FFC31FBE1FC66" box="[958,1115,908,933]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
, even though reversals exist; see Short &amp; Fikáček 2013], (4) metatarsomere 1 slightly longer than metatarsomere 2 [synapomorphy of
<taxonomicName id="4C294D578A36143FFF76FC69FE8EFC2E" box="[151,308,980,1005]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
, otherwise present only in three derived genera of the
<taxonomicName id="4C294D578A36143FFC77FC69FB90FC2E" box="[918,1066,980,1005]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Rygmodinae">Rygmodinae</taxonomicName>
], and (5) metathoracic wing with reduced anal veins [complete set of anal veins in present in all
<taxonomicName id="4C294D578A36143FFCB7FC4AFC5EFBD3" box="[854,996,1015,1040]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Rygmodinae">Rygmodinae</taxonomicName>
and many other hydrophilid tribes].
</paragraph>
<paragraph id="8B9636D48A36143FFF26FBA1FA8EF9B6" blockId="9.[151,1436,151,1977]" pageId="9" pageNumber="286">
The taxonomic position of
<taxonomicName id="4C294D578A36143FFDEAFBA1FD15FBF7" box="[523,687,1052,1076]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFDEAFBA1FD15FBF7" box="[523,687,1052,1076]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
within
<taxonomicName id="4C294D578A36143FFCEFFBA1FC12FBF6" box="[782,936,1052,1077]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
was difficult to infer based on our initial morphological examination, as the taxon exhibits a mosaic of characters unique or typical for several different tribes. Most of the thoracic characters resemble representatives of the tribe
<taxonomicName id="4C294D578A36143FFC0FFBD9FBE2FBBE" box="[1006,1112,1124,1149]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Omicrini">Omicrini</taxonomicName>
: the antennal grooves on the prothoracic hypomeron and the grooves for reception of procoxae on the mesothorax are absent, the prosternum is largely reduced, the epipleuron is very wide throughout the elytral length, the tibiae are slightly bent outwards, and the tarsi have largely shortened tarsomeres. The sexually dimorphic emargination on abdominal ventrite 5 resembles a similar structure in some species of the omicrine genus
<taxonomicName id="4C294D578A36143FFC5DFB49FB9BFACF" box="[956,1057,1268,1292]" class="Insecta" family="Hydrophilidae" genus="Psalitrus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFC5DFB49FB9BFACF" box="[956,1057,1268,1292]" italics="true" pageId="9" pageNumber="286">Psalitrus</emphasis>
</taxonomicName>
. Few other characters, however, contradicted the assignment of
<taxonomicName id="4C294D578A36143FFE19FAA5FD26FAF3" box="[504,668,1304,1328]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFE19FAA5FD26FAF3" box="[504,668,1304,1328]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
to
<taxonomicName id="4C294D578A36143FFD20FAAAFC90FAF3" box="[705,810,1303,1328]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Omicrini">Omicrini</taxonomicName>
. The clypeus is not excised above the antennal bases (a character unique to
<taxonomicName id="4C294D578A36143FFE94FA81FE60FA96" box="[373,474,1340,1365]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Omicrini">Omicrini</taxonomicName>
and
<taxonomicName id="4C294D578A36143FFDF0FA81FD1DFA96" box="[529,679,1340,1365]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
) and lacks the anterolateral inclined lobes (a unique synapomorphy of
<taxonomicName id="4C294D578A36143FFF5FFAE2FE92FABB" box="[190,296,1375,1400]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Omicrini">Omicrini</taxonomicName>
), and the general morphology of the head resembles that found in the
<taxonomicName id="4C294D578A36143FFB4FFAE2FAE7FABB" box="[1198,1373,1375,1400]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Coelostomatini">Coelostomatini</taxonomicName>
. The representatives of the latter tribe are also characterized by a wide epipleuron throughout elytral length and by the absence of antennal grooves and grooves for reception of procoxae. The same is true for some Protosternini which are, however, all characterized by an arcuate ridge on the metaventrite, a character absent from
<taxonomicName id="4C294D578A36143FFB10FA71FA2FFA27" box="[1265,1429,1484,1508]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFB10FA71FA2FFA27" box="[1265,1429,1484,1508]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
. Based on all these characters,
<taxonomicName id="4C294D578A36143FFE1EFA4DFD19F9CB" box="[511,675,1520,1544]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFE1EFA4DFD19F9CB" box="[511,675,1520,1544]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
strongly differs from the
<taxonomicName id="4C294D578A36143FFC38FA52FBD7F9CB" box="[985,1133,1519,1544]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
and
<taxonomicName id="4C294D578A36143FFB4DFA52FA82F9CB" box="[1196,1336,1519,1544]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Sphaeridiini">Sphaeridiini</taxonomicName>
, both of which bear their own numerous unique morphological synapomorphies (Hansen 1991; Short &amp; Fikáček 2013). Hence, the initial study demonstrated that
<taxonomicName id="4C294D578A36143FFD9CF985FC9BF993" box="[637,801,1592,1616]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFD9CF985FC9BF993" box="[637,801,1592,1616]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
cannot be reliably assigned to any existing tribe of the
<taxonomicName id="4C294D578A36143FFF76F9E1FE8EF9B6" box="[151,308,1628,1653]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
, which provoked a more detailed study using scanning electron microscopy and DNA data.
</paragraph>
<paragraph id="8B9636D48A36143FFF26F9C2FE88F87A" blockId="9.[151,1436,151,1977]" pageId="9" pageNumber="286">
The examination by SEM revealed that the male maxilla of
<taxonomicName id="4C294D578A36143FFC8AF93DFBB5F95B" box="[875,1039,1664,1688]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFC8AF93DFBB5F95B" box="[875,1039,1664,1688]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
bears a sucking disc. The discs are unique within the
<taxonomicName id="4C294D578A36143FFE86F919FDB3F97E" box="[359,521,1700,1725]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="family">Hydrophilidae</taxonomicName>
for the clade
<taxonomicName id="4C294D578A36143FFD4BF919FC87F97E" box="[682,829,1700,1725]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
+
<taxonomicName id="4C294D578A36143FFCBEF919FB22F97F" authority="(Hansen 1991)" baseAuthorityName="Hansen" baseAuthorityYear="1991" box="[863,1176,1700,1725]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Sphaeridiini">Sphaeridiini (Hansen 1991)</taxonomicName>
and likely serve to fix the position of the male above the female during mating (Vogt 1968). Their presence in
<taxonomicName id="4C294D578A36143FFB21F975FADEF923" box="[1216,1380,1736,1760]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFB21F975FADEF923" box="[1216,1380,1736,1760]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
was unexpected, as the assignment to both
<taxonomicName id="4C294D578A36143FFD93F951FD41F8C6" box="[626,763,1772,1797]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Sphaeridiini">Sphaeridiini</taxonomicName>
and
<taxonomicName id="4C294D578A36143FFCDEF951FC69F8C6" box="[831,979,1772,1797]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
had been contradicted by most other morphological characters. Further detailed study of the metathoracic wing venation revealed an additional character suggesting placement within the
<taxonomicName id="4C294D578A36143FFD95F889FCB2F88E" box="[628,776,1844,1869]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
clade: the median spur arises subdistally of the M-Cu loop of the metathoracic wing, which is a unique synapomorphy of the tribe
<taxonomicName id="4C294D578A36143FFC3BF8EAFA9EF8B3" authority="(Hansen 1991)" baseAuthorityName="Hansen" baseAuthorityYear="1991" box="[986,1316,1879,1904]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini (Hansen 1991)</taxonomicName>
. Based on these morphological studies,
<taxonomicName id="4C294D578A36143FFE0BF8C1FD34F857" box="[490,654,1916,1940]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A36143FFE0BF8C1FD34F857" box="[490,654,1916,1940]" italics="true" pageId="9" pageNumber="286">Chimaerocyon</emphasis>
</taxonomicName>
was therefore hypothesized as a highly modified representative of
<taxonomicName id="4C294D578A36143FFF76F81DFE95F87A" box="[151,303,1952,1977]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="9" pageNumber="286" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
.
</paragraph>
<caption id="DF56665C8A35143CFF76F91DFDBAF8AD" httpUri="https://zenodo.org/record/221985/files/figure.png" pageId="10" pageNumber="287" targetBox="[151,1436,193,1674]" targetPageId="10">
<paragraph id="8B9636D48A35143CFF76F91DFDBAF8AD" blockId="10.[151,1436,1696,1902]" pageId="10" pageNumber="287">
<emphasis id="B95DEAC68A35143CFF76F91DFEECF975" bold="true" box="[151,342,1696,1718]" pageId="10" pageNumber="287">FIGURES 3137.</emphasis>
Phylogenetic position of
<taxonomicName id="4C294D578A35143CFD84F91DFD43F975" box="[613,761,1696,1718]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="287" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A35143CFD84F91DFD43F975" box="[613,761,1696,1718]" italics="true" pageId="10" pageNumber="287">Chimaerocyon</emphasis>
</taxonomicName>
. 31—Bayesian tree based on four genes concatenated (branches with posterior probability values). 32—mapping of relevant tribal-level characters on the tree (full circle = gain, crossed circle = loss). 3337—diagnostic tribal-level characters illustrated on selected representatives of the
<taxonomicName id="4C294D578A35143CFB93F960FABAF930" box="[1138,1280,1757,1779]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="10" pageNumber="287" phylum="Arthropoda" rank="subFamily" subFamily="Sphaeridiinae">Sphaeridiinae</taxonomicName>
. Abbreviations (numbers refer to characters in Fig. 32):
<emphasis id="B95DEAC68A35143CFDADF941FDC4F8D1" bold="true" box="[588,638,1788,1810]" pageId="10" pageNumber="287">angr</emphasis>
—antennal groove,
<emphasis id="B95DEAC68A35143CFCAFF941FC37F8D1" bold="true" box="[846,909,1788,1810]" pageId="10" pageNumber="287">comcl</emphasis>
—compact antennal club,
<emphasis id="B95DEAC68A35143CFB43F941FB75F8D1" bold="true" box="[1186,1231,1788,1810]" pageId="10" pageNumber="287">curt</emphasis>
—curved metatibia,
<emphasis id="B95DEAC68A35143CFF76F8A7FF79F8F3" bold="true" box="[151,195,1818,1840]" pageId="10" pageNumber="287">elpr</emphasis>
—elevated median portion of prosternum,
<emphasis id="B95DEAC68A35143CFD8EF8A7FD1CF8F3" bold="true" box="[623,678,1818,1840]" pageId="10" pageNumber="287">excly</emphasis>
—excised lateral portion of clypeus (exposed antennal bases),
<emphasis id="B95DEAC68A35143CFAFAF8A7FAFEF8F3" bold="true" box="[1307,1348,1818,1840]" pageId="10" pageNumber="287">llob</emphasis>
—lateral lobes of clypeus,
<emphasis id="B95DEAC68A35143CFEA8F884FE3BF88C" bold="true" box="[329,385,1849,1871]" pageId="10" pageNumber="287">msuc</emphasis>
—male sucker on maxilla,
<emphasis id="B95DEAC68A35143CFD72F884FD03F88C" bold="true" box="[659,697,1849,1871]" pageId="10" pageNumber="287">pgr</emphasis>
—grooves for reception of procoxae,
<emphasis id="B95DEAC68A35143CFBD6F884FBDCF88C" bold="true" box="[1079,1126,1849,1871]" pageId="10" pageNumber="287">pros</emphasis>
—well-developed prosternum,
<emphasis id="B95DEAC68A35143CFF76F8E5FF73F8AD" bold="true" box="[151,201,1880,1902]" pageId="10" pageNumber="287">wepi</emphasis>
—wide epipleuron posteriorly.
</paragraph>
</caption>
<paragraph id="8B9636D48A34143DFF26FF2AFE3FFE6E" blockId="11.[151,1436,151,429]" pageId="11" pageNumber="288">
The molecular phylogenetic analysis based on four genes confirmed that
<taxonomicName id="4C294D578A34143DFBF5FF25FB02FF73" box="[1044,1208,152,176]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A34143DFBF5FF25FB02FF73" box="[1044,1208,152,176]" italics="true" pageId="11" pageNumber="288">Chimaerocyon</emphasis>
</taxonomicName>
is a member of the
<taxonomicName id="4C294D578A34143DFF76FF01FE91FF16" box="[151,299,188,213]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
deeply nested in the
<taxonomicName id="4C294D578A34143DFDF9FF01FDCCFF10" box="[536,630,188,211]" class="Insecta" family="Hydrophilidae" genus="Cercyon" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A34143DFDF9FF01FDCCFF10" box="[536,630,188,211]" italics="true" pageId="11" pageNumber="288">Cercyon</emphasis>
</taxonomicName>
-group. Further conclusions concerning its position within
<taxonomicName id="4C294D578A34143DFAE9FF01FA26FF16" box="[1288,1436,188,213]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
are premature, as our taxon sampling covers only a small portion of real taxonomic diversity of the tribe. The analysis confirmed that due to myrmecophilous habits,
<taxonomicName id="4C294D578A34143DFCEDFEB9FC0AFEDF" box="[780,944,260,284]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A34143DFCEDFEB9FC0AFEDF" box="[780,944,260,284]" italics="true" pageId="11" pageNumber="288">Chimaerocyon</emphasis>
</taxonomicName>
secondarily lost nearly all synapomorphies of the tribe
<taxonomicName id="4C294D578A34143DFEFBFE9AFE14FE83" box="[282,430,295,320]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
known to date, except the presence of suckers on male maxilla and the subdistal position of the M-Cu loop on hind wings. This strong morphological difference is in contrast to molecular data that do not reveal greater divergence of
<taxonomicName id="4C294D578A34143DFE08FECDFD37FE4B" box="[489,653,368,392]" class="Insecta" family="Hydrophilidae" genus="Chimaerocyon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="genus">
<emphasis id="B95DEAC68A34143DFE08FECDFD37FE4B" box="[489,653,368,392]" italics="true" pageId="11" pageNumber="288">Chimaerocyon</emphasis>
</taxonomicName>
from remaining
<taxonomicName id="4C294D578A34143DFCB7FED2FC50FE4B" box="[854,1002,367,392]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="tribe" tribe="Megasternini">Megasternini</taxonomicName>
than among morphologically similar megasternine genera.
</paragraph>
<paragraph id="8B9636D48A34143DFF76FE42FEC5FDDB" blockId="11.[151,383,511,536]" box="[151,383,511,536]" pageId="11" pageNumber="288">
<emphasis id="B95DEAC68A34143DFF76FE42FEC5FDDB" bold="true" box="[151,383,511,536]" pageId="11" pageNumber="288">Acknowledgements</emphasis>
</paragraph>
<paragraph id="8B9636D48A34143DFF76FDF5FB4FFCD6" blockId="11.[151,1436,583,789]" pageId="11" pageNumber="288">
We are indebted to Maxwell V. L. Barclay (The Natural History Museum, London,
<collectingCountry id="F33E76448A34143DFBDCFDF5FBDDFDA3" box="[1085,1127,584,608]" name="United Kingdom" pageId="11" pageNumber="288">UK</collectingCountry>
) for the possibility to study the
<typeStatus id="549288768A34143DFF29FDD0FEB9FD46" box="[200,259,621,645]" pageId="11" pageNumber="288">types</typeStatus>
of
<taxonomicName id="4C294D578A34143DFED8FDD0FDA0FD47" box="[313,538,620,644]" class="Insecta" family="Hydrophilidae" genus="Litrosurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="11" pageNumber="288" phylum="Arthropoda" rank="species" species="insolitus">
<emphasis id="B95DEAC68A34143DFED8FDD0FDA0FD47" box="[313,538,620,644]" italics="true" pageId="11" pageNumber="288">Litrosurus insolitus</emphasis>
</taxonomicName>
, to K. Taro Eldredge (Kansas University, Lawrence,
<collectingCountry id="F33E76448A34143DFB4CFDD1FB5DFD47" box="[1197,1255,620,644]" name="United States of America" pageId="11" pageNumber="288">USA</collectingCountry>
) for numerous discussions on myrmecophilous beetles and relevance of their morphological characters, and to two anonymous reviewers for their comments on the manuscript. The study was supported by the grant of the Czech Science Foundation (GAČR) P506/12/P096 and by the Ministry of Culture of the
<collectingCountry id="F33E76448A34143DFBE0FD6AFB0DFD33" box="[1025,1207,727,752]" name="Czech Republic" pageId="11" pageNumber="288">Czech Republic</collectingCountry>
(DKRVO 2013/12, National Museum, 00023272) to M. Fikáček, and by the grant SVV-2013-267 201 to D. Vondráček.
</paragraph>
</subSubSection>
<subSubSection id="C333655F8A34143DFF76FCDAFAE7F89D" pageId="11" pageNumber="288" type="reference_group">
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