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<mods:title id="B4F9343661815C99CFA9BEA20996B35D">A review of Thysanarthria with description of seven new species and comments on its relationship to Chaetarthria (Hydrophilidae: Chaetarthriini)</mods:title>
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<mods:namePart id="860D151C820E90BD4FAF56DDE9441515">Fikáček, Martin</mods:namePart>
<mods:affiliation id="1A7F1F4EBD7CB2CB9D1C171F2D28E6D0">Department of Entomology, National Museum, Cirkusová 1740, CZ- 19100 Praha 9 Horní Počernice, Czech Republic &amp; Department of Zoology, Faculty of Science, Charles University, Viničná 7, CZ- 12844 Praha 2, Czech Republic</mods:affiliation>
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<mods:namePart id="11988356DD96DEF141E1D427D5A91F14">Liu, Hsing-Che</mods:namePart>
<mods:affiliation id="864CBE19FC05BDC3B70C74740A395BAA">Department of Environmental Engineering and Management, Chaoyang University of Technology, No. 168, Jifeng E. Rd., Taichung City 413, Taiwan</mods:affiliation>
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<paragraph id="8BE536D37A2BFFBBFE9B04EE364CB2D4" blockId="5.[274,637,1478,1501]" box="[274,637,1478,1501]" pageId="5" pageNumber="234">
<heading id="D0AD81BF7A2BFFBBFE9B04EE364CB2D4" box="[274,637,1478,1501]" centered="true" fontSize="10" level="2" pageId="5" pageNumber="234" reason="3">
<taxonomicName id="4C5A4D507A2BFFBBFE9B04EE364CB2D4" ID-CoL="8X7X2" authority="Orchymont, 1926" authorityName="Orchymnont" authorityYear="1926" box="[274,637,1478,1501]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFE9B04EE364CB2D4" bold="true" box="[274,637,1478,1501]" pageId="5" pageNumber="234">
<emphasis id="B92EEAC17A2BFFBBFE9B04EE3580B2D4" bold="true" box="[274,433,1478,1501]" italics="true" pageId="5" pageNumber="234">Thysanarthria</emphasis>
<bibRefCitation id="EFCB4B227A2BFFBBFE3E04EE364CB2D4" author="ORCHYMONT A. d'" box="[439,637,1478,1501]" journalOrPublisher="Bulletin et Annales de la Societe Entomologique de Belgique" pageId="5" pageNumber="234" pagination="195 - 196" part="66" refId="ref18492" refString="ORCHYMONT A. d' 1926 a: Note sur Thysanarthria n. g. Bulletin et Annales de la Societe Entomologique de Belgique 66: 195 - 196." title="Note sur Thysanarthria n. g." type="journal article" year="1926">Orchymont, 1926</bibRefCitation>
</emphasis>
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<paragraph id="8BE536D37A2BFFBBFF1E04C43627B113" blockId="5.[151,760,1516,2071]" pageId="5" pageNumber="234">
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<taxonomicName id="4C5A4D507A2BFFBBFF1E04C435EDB2F6" ID-CoL="8X7X2" authority="Orchymont, 1926 a: 195" authorityName="Orchymnont" authorityPageNumber="195" authorityYear="1926" box="[151,476,1516,1535]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFF1E04C43521B2F6" box="[151,272,1516,1535]" italics="true" pageId="5" pageNumber="234">Thysanarthria</emphasis>
<treatmentCitation id="0AFB10C27A2BFFBBFE9D04C435EDB2F6" author="ORCHYMONT A." box="[276,476,1516,1535]" page="195" pageId="5" pageNumber="234" year="1926">
<bibRefCitation id="EFCB4B227A2BFFBBFE9D04C435EDB2F6" author="ORCHYMONT A." box="[276,476,1516,1535]" firstAuthor="Orchymont" page="195" pageId="5" pageNumber="234" pagination="195 - 196" refId="ref18492" refString="ORCHYMONT A. d' 1926 a: Note sur Thysanarthria n. g. Bulletin et Annales de la Societe Entomologique de Belgique 66: 195 - 196." type="journal article" year="1926" yearSuffix="a">Orchymont, 1926a:195</bibRefCitation>
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.
<typeStatus id="54E188717A2BFFBBFE6A04C4363CB2F6" box="[483,525,1516,1535]" pageId="5" pageNumber="234">Type</typeStatus>
species:
<taxonomicName id="4C5A4D507A2BFFBBFDDE04C43574B113" ID-CoL="6MJVW" authority="Regimbart, 1903" authorityName="Regimbart" authorityYear="1903" class="Insecta" family="Hydrophilidae" genus="Hydrobius" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="atriceps">
<emphasis id="B92EEAC17A2BFFBBFDDE04C436C9B2F6" box="[599,760,1516,1535]" italics="true" pageId="5" pageNumber="234">Hydrobius atriceps</emphasis>
<treatmentCitation id="0AFB10C27A2BFFBBFF3C072F3574B113" author="REGIMBART M." box="[181,325,1543,1562]" pageId="5" pageNumber="234" year="1903">
<bibRefCitation id="EFCB4B227A2BFFBBFF3C072F3574B113" author="Regimbart" box="[181,325,1543,1562]" firstAuthor="Regimbart" pageId="5" pageNumber="234" pagination="1 - 51" refId="ref18592" refString="REGIMBART M. 1903: Coleopteres aquatiques (Haliplidae, Dytiscidae, Gyrinidae et Hydrophilidae) recueillis dans le Sud de Madagascar par M. Ch. Alluaud (Juillet 1900 - mai 1901). Annales de la Societe Entomologique de France 72: 1 - 51." type="journal article" year="1903">Régimbart, 1903</bibRefCitation>
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by original designation.
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</paragraph>
<paragraph id="8BE536D37A2BFFBBFF1E070A36DAB146" blockId="5.[151,760,1516,2071]" pageId="5" pageNumber="234">
<treatmentCitationGroup id="AB4A11FD7A2BFFBBFF1E070A36DAB146" pageId="5" pageNumber="234">
=
<treatmentCitation id="0AFB10C27A2BFFBBFF22070A363EB13C" author="CHIESA A." box="[171,527,1570,1589]" page="276" pageId="5" pageNumber="234" year="1967">
<taxonomicName id="4C5A4D507A2BFFBBFF22070A363EB13C" ID-CoL="8XKCK" authority="Chiesa, 1967: 276" authorityName="Chiesa" authorityPageNumber="276" authorityYear="1967" box="[171,527,1570,1589]" class="Insecta" family="Hydrophilidae" genus="Chaetarthriomorphus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFF22070A3554B13C" box="[171,357,1570,1589]" italics="true" pageId="5" pageNumber="234">Chaetarthriomorphus</emphasis>
<bibRefCitation id="EFCB4B227A2BFFBBFEE4070A363EB13C" author="Chiesa" box="[365,527,1570,1589]" firstAuthor="Chiesa" page="276" pageId="5" pageNumber="234" pagination="275 - 277" refId="ref17771" refString="CHIESA A. 1967: Compimento di una revisione di Hydrophilidae del Afghanistan (Coleoptera: Hydrophilidae). Annales Historico-Naturales Musei Nationalis Hungarici 59: 275 - 277." type="journal article" year="1967">Chiesa, 1967: 276</bibRefCitation>
</taxonomicName>
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.
<typeStatus id="54E188717A2BFFBBFD90070A3672B13C" box="[537,579,1570,1589]" pageId="5" pageNumber="234">Type</typeStatus>
species:
<taxonomicName id="4C5A4D507A2BFFBBFD1F070A3514B146" ID-CoL="5XNTC" authority="Chiesa, 1967" authorityName="Chiesa" authorityYear="1967" class="Insecta" family="Hydrophilidae" genus="Chaetarthriomorphus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="sulcatus">
<emphasis id="B92EEAC17A2BFFBBFD1F070A36C9B13C" box="[662,760,1570,1589]" italics="true" pageId="5" pageNumber="234">C. sulcatus</emphasis>
<bibRefCitation id="EFCB4B227A2BFFBBFF3C07143514B146" author="Chiesa" box="[181,293,1596,1615]" firstAuthor="Chiesa" pageId="5" pageNumber="234" pagination="275 - 277" refId="ref17771" refString="CHIESA A. 1967: Compimento di una revisione di Hydrophilidae del Afghanistan (Coleoptera: Hydrophilidae). Annales Historico-Naturales Musei Nationalis Hungarici 59: 275 - 277." type="journal article" year="1967">Chiesa, 1967</bibRefCitation>
</taxonomicName>
by monotypy. Synonymized by
<treatmentCitation id="0AFB10C27A2BFFBBFDB3071436D6B146" author="HANSEN M." box="[570,743,1596,1615]" page="126" pageId="5" pageNumber="234" year="1991">
<bibRefCitation id="EFCB4B227A2BFFBBFDB3071436D6B146" author="HANSEN" box="[570,743,1596,1615]" firstAuthor="HANSEN" page="126" pageId="5" pageNumber="234" pagination="1 - 368" refId="ref18021" refString="HANSEN M. 1991: The hydrophiloid beetles. Phylogeny, classification and a revision of the genera (Coleoptera, Hydrophiloidea). Biologiske Skrifter 40: 1 - 368." type="journal article" year="1991">HANSEN (1991: 126)</bibRefCitation>
</treatmentCitation>
.
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</paragraph>
</subSubSection>
<subSubSection id="C34065587A2BFFBBFF1E07483097B23C" pageId="5" pageNumber="234" type="diagnosis">
<paragraph id="8BE536D37A2BFFBBFF1E0748301BB41C" blockId="5.[151,760,1516,2071]" lastBlockId="5.[828,1437,190,2069]" pageId="5" pageNumber="234">
<emphasis id="B92EEAC17A2BFFBBFF1E0748353AB17E" bold="true" box="[151,267,1632,1655]" pageId="5" pageNumber="234">Diagnosis.</emphasis>
The genus can be recognized from other co-occurring genera of the
<taxonomicName id="4C5A4D507A2BFFBBFEF807A83638B19E" authorityName="Latreille" authorityYear="1802" box="[369,521,1664,1687]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="family">Hydrophilidae</taxonomicName>
based on the following combination of characters: body small to medium sized (
<quantity id="4CA29B367A2BFFBBFF1607E83515B1DE" box="[159,292,1728,1751]" metricMagnitude="-3" metricUnit="m" metricValue="1.8" metricValueMax="2.2" metricValueMin="1.4" pageId="5" pageNumber="234" unit="mm" value="1.8" valueMax="2.2" valueMin="1.4">1.42.2 mm</quantity>
); head black, pronotum and elytra yellowish to pale brown in most species, uniformly brown in the remaining ones (
<figureCitation id="13612A567A2BFFBBFEBD06283542B01E" box="[308,371,1792,1815]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Figs 1</figureCitation>
AD); head with large exposed well sclerotized labrum (e.g.,
<figureCitation id="13612A567A2BFFBBFE2D060835C6B03E" box="[420,503,1824,1847]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Fig. 1B</figureCitation>
); antenna with 9 antennomeres, scape very long, pedicel bulbose, antennomeres 35 very small, cupule and three-segment antennal club pubescent (
<figureCitation id="13612A567A2BFFBBFE9806A83556B09E" box="[273,359,1920,1943]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3D</figureCitation>
); maxillary palpomere 4 basally with row of many peg-like setae (
<figureCitation id="13612A567A2BFFBBFE4E0688362BB0BE" box="[455,538,1952,1975]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3E</figureCitation>
); mentum projecting anteromedially, with row of setae along anterior margin (
<figureCitation id="13612A567A2BFFBBFF1606C834C3B0FE" box="[159,242,2016,2039]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3A</figureCitation>
); gular sutures contiguous (
<figureCitation id="13612A567A2BFFBBFD8406C8366CB0FE" box="[525,605,2016,2039]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3C</figureCitation>
); mesoventrite distinctly divided from anepisterna by sutures, sutures widely separated on anterior margin of mesothorax (
<figureCitation id="13612A567A2BFFBBFAF80196376CB7FC" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3B</figureCitation>
); mesoventrite flat except of small semicircular elevation posteromesally (
<figureCitation id="13612A567A2BFFBBFB9E01D63058B61C" box="[1047,1129,254,277]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3B</figureCitation>
); metaventrite short, sparsely pubescent only mesally and anterolaterally (
<figureCitation id="13612A567A2BFFBBFB760036317EB63C" box="[1279,1359,286,309]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3B</figureCitation>
); elytra with 10 sharply impressed striae (
<figureCitation id="13612A567A2BFFBBFB18001630FCB65C" box="[1169,1229,318,341]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Figs 1</figureCitation>
AD); whole dorsal surface covered by sparsely arranged setae which are trifid basally with a long median projection (
<figureCitation id="13612A567A2BFFBBFB6600563178B69C" box="[1263,1353,382,405]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Figs 1C</figureCitation>
,
<figureCitation id="13612A567A2BFFBBFAD100563157B69C" box="[1368,1382,382,405]" captionStart="Fig" captionStartId="3.[151,182,1959,1978]" captionTargetBox="[157,1426,396,1946]" captionTargetId="figure-14@3.[151,1436,395,1949]" captionTargetPageId="3" captionText="Fig. 2. Morphological differences among Thysanarthria and principal groups of Chaetarthria. AZ male genitalia and surrounding sclerites; ae details of elytral pubescence. A Chaetarthria similis Wollaston, 1864; BG, c Chaetarthria seminulum (Herbst, 1797); HM, b Thysanarthria championi (Knisch, 1924); NS, d Chaetarthria panda Orchymont, 1939; TZ, e Chaetarthria malickyi Hebauer, 1995; a Thysanarthria atriceps (Régimbart, 1903). AD, KM, QS, XZ aedeagus in dorsal, lateral and ventral view; E, J, P, V male sternite 9; F, H, N, T male sternite 8; G, I, O, U male tergite 8. Figures ae are all to the same scale." figureDoi="http://doi.org/10.5281/zenodo.4488924" httpUri="https://zenodo.org/record/4488924/files/figure.png" pageId="5" pageNumber="234">2</figureCitation>
ab); profemora pubescent in basal half (
<figureCitation id="13612A567A2BFFBBFB3800B63134B6BC" box="[1201,1285,414,437]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3C</figureCitation>
); mesofemora pubescent anterobasally (
<figureCitation id="13612A567A2BFFBBFBC8009630A3B6DC" box="[1089,1170,446,469]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3B</figureCitation>
); metafemora bare except on anterobasal margins (
<figureCitation id="13612A567A2BFFBBFBC300F63091B6FC" box="[1098,1184,478,501]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3B</figureCitation>
); tarsi rather short and stout, metatarsus with all tarsomeres c. equal in length (
<figureCitation id="13612A567A2BFFBBFAFB00D6376AB53C" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Fig. 3F</figureCitation>
); abdomen with 5 ventrites, basal two bearing shallow cavity covered by long setae arising from base of ventrite 1, holding whitish gelatinous substance (
<figureCitation id="13612A567A2BFFBBFA8403763164B57C" box="[1293,1365,606,629]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="5" pageNumber="234">Figs 3</figureCitation>
GH); ventrite 12 with median carina; male abdominal sternite 8 with narrow median projection (
<figureCitation id="13612A567A2BFFBBFB2003B630CEB5BC" box="[1193,1279,670,693]" captionStart="Fig" captionStartId="3.[151,182,1959,1978]" captionTargetBox="[157,1426,396,1946]" captionTargetId="figure-14@3.[151,1436,395,1949]" captionTargetPageId="3" captionText="Fig. 2. Morphological differences among Thysanarthria and principal groups of Chaetarthria. AZ male genitalia and surrounding sclerites; ae details of elytral pubescence. A Chaetarthria similis Wollaston, 1864; BG, c Chaetarthria seminulum (Herbst, 1797); HM, b Thysanarthria championi (Knisch, 1924); NS, d Chaetarthria panda Orchymont, 1939; TZ, e Chaetarthria malickyi Hebauer, 1995; a Thysanarthria atriceps (Régimbart, 1903). AD, KM, QS, XZ aedeagus in dorsal, lateral and ventral view; E, J, P, V male sternite 9; F, H, N, T male sternite 8; G, I, O, U male tergite 8. Figures ae are all to the same scale." figureDoi="http://doi.org/10.5281/zenodo.4488924" httpUri="https://zenodo.org/record/4488924/files/figure.png" pageId="5" pageNumber="234">Fig. 2H</figureCitation>
); male sternite 9 V-shaped medially (
<figureCitation id="13612A567A2BFFBBFBAB0396305CB5DC" box="[1058,1133,702,725]" captionStart="Fig" captionStartId="3.[151,182,1959,1978]" captionTargetBox="[157,1426,396,1946]" captionTargetId="figure-14@3.[151,1436,395,1949]" captionTargetPageId="3" captionText="Fig. 2. Morphological differences among Thysanarthria and principal groups of Chaetarthria. AZ male genitalia and surrounding sclerites; ae details of elytral pubescence. A Chaetarthria similis Wollaston, 1864; BG, c Chaetarthria seminulum (Herbst, 1797); HM, b Thysanarthria championi (Knisch, 1924); NS, d Chaetarthria panda Orchymont, 1939; TZ, e Chaetarthria malickyi Hebauer, 1995; a Thysanarthria atriceps (Régimbart, 1903). AD, KM, QS, XZ aedeagus in dorsal, lateral and ventral view; E, J, P, V male sternite 9; F, H, N, T male sternite 8; G, I, O, U male tergite 8. Figures ae are all to the same scale." figureDoi="http://doi.org/10.5281/zenodo.4488924" httpUri="https://zenodo.org/record/4488924/files/figure.png" pageId="5" pageNumber="234">Fig. 2J</figureCitation>
); aedeagus with long tubular phallobase, base of median lobe not reaching deeply into phallobase (
<figureCitation id="13612A567A2BFFBBFC3503D6302FB41C" box="[956,1054,766,789]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStart-4="Fig" captionStart-5="Fig" captionStartId-0="8.[151,182,1941,1960]" captionStartId-1="10.[887,918,1799,1818]" captionStartId-2="12.[151,182,1980,1999]" captionStartId-3="14.[151,182,1334,1353]" captionStartId-4="16.[151,182,1976,1995]" captionStartId-5="18.[151,182,1467,1486]" captionTargetBox-0="[165,1398,197,1882]" captionTargetBox-1="[178,1439,395,2038]" captionTargetBox-2="[155,1405,208,1868]" captionTargetBox-3="[177,1428,199,1315]" captionTargetBox-4="[204,1420,202,1944]" captionTargetBox-5="[178,1432,202,1437]" captionTargetId-0="figure-13@8.[157,1431,189,1911]" captionTargetId-1="figure-13@10.[175,1413,386,2041]" captionTargetId-2="figure-13@12.[153,1435,189,1873]" captionTargetId-3="figure-13@14.[151,1436,186,1319]" captionTargetId-4="figure-108@16.[162,1439,189,1956]" captionTargetId-5="figure-13@18.[151,1436,186,1450]" captionTargetPageId-0="8" captionTargetPageId-1="10" captionTargetPageId-2="12" captionTargetPageId-3="14" captionTargetPageId-4="16" captionTargetPageId-5="18" captionText-0="Fig. 4. Male genitalia of Thysanarthria species. AE T. atriceps (Régimbart, 1903) from Zambia: 20 km W of Gwanda, coll. NMPC (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view); FJ T. atriceps from Republic of South Africa: 8 km NEE of Stanford, coll. NMPC (FH whole aedeagus in dorsal, lateral and ventral view; IJ detail of median lobe and parameres in dorsal and ventral view); KO T. brittoni Balfour-Browne, 1951, holotype (KM whole aedeagus in dorsal, lateral and ventral view; NO detail of median lobe and parameres in dorsal and ventral view)." captionText-1="Fig. 5. Male genitalia of Thysanarthria species, holotypes.AE T.brincki Hebauer, 2001 (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view); FJ T. cardamona sp. nov. (FH whole aedeagus in dorsal, lateral and ventral view; IJ detail of median lobe and parameres in dorsal and ventral view); KM T. bengalensis Hebauer, 2001 (K photo of the aedeagus before re-mounting; L whole aedeagus in ventral view; M whole aedeagus in lateral view)." captionText-2="Fig. 6. Male genitalia of Thysanarthria species, holotypes (AI, KO) and paratype (J). AE T.bifida sp. nov. (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view). FJ T. trifida sp. nov. (FH whole aedeagus of the holotype in dorsal, lateral and ventral view; I details of parameres of the holotype; J same of the paratype). KO T. ceylonensis Hebauer, 2001 (KM whole aedeagus in dorsal, lateral and ventral view; NO details of median lobe and parameres in dorsal and ventral view)." captionText-3="Fig. 7. Male genitalia of Thysanarthria championi (Knisch, 1924).AE lectotype of Chaetarthria championi Knisch, 1924 (AC whole aedeagus in dorsal, lateral and ventral view; D detail of median lobe and parameres in dorsal view; E same after KOH treatment, with fully everted internal sac). FJ lectotype of Chaetarthriomorphus sulcatus Chiesa, 1967 (FH whole aedeagus in dorsal, lateral and ventral view; IJ details of parameres and median lobe in dorsal and ventral view)." captionText-4="Fig. 8. Male genitalia of Thysanarthria species, holotypes.AE T. chui sp. nov.(AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view). FJ T. hongsonensis Hebauer, 2001 (FH whole aedeagus in dorsal, lateral and ventral view; IJ details of parameres and median lobe in dorsal and ventral view). KO T. madurensis Hebauer, 2001 (KM whole aedeagus in dorsal, lateral and ventral view; NO details of median lobe and parameres in dorsal and ventral view)." captionText-5="Fig. 9. Male genitalia of Thysanarthria species, holotypes. AE T. saurahana sp. nov. (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view). FJ T. siamensis Hebauer, 2001 (FH whole aedeagus in dorsal, lateral and ventral view; IJ details of parameres and median lobe in dorsal and ventral view)." figureDoi-0="http://doi.org/10.5281/zenodo.4488930" figureDoi-1="http://doi.org/10.5281/zenodo.4488932" figureDoi-2="http://doi.org/10.5281/zenodo.4488934" figureDoi-3="http://doi.org/10.5281/zenodo.4488938" figureDoi-4="http://doi.org/10.5281/zenodo.4488940" figureDoi-5="http://doi.org/10.5281/zenodo.4488942" httpUri-0="https://zenodo.org/record/4488930/files/figure.png" httpUri-1="https://zenodo.org/record/4488932/files/figure.png" httpUri-2="https://zenodo.org/record/4488934/files/figure.png" httpUri-3="https://zenodo.org/record/4488938/files/figure.png" httpUri-4="https://zenodo.org/record/4488940/files/figure.png" httpUri-5="https://zenodo.org/record/4488942/files/figure.png" pageId="5" pageNumber="234">Figs 49</figureCitation>
).
</paragraph>
<paragraph id="8BE536D37A2BFFBBFCB502363097B23C" blockId="5.[828,1437,190,2069]" pageId="5" pageNumber="234">
<emphasis id="B92EEAC17A2BFFBBFCB502363007B43C" bold="true" box="[828,1078,798,821]" pageId="5" pageNumber="234">Differential diagnosis.</emphasis>
The base of abdomen with series of long setae covering a gelatinous substance and antenna with bulbous pedicel distinguish
<taxonomicName id="4C5A4D507A2BFFBBFB2D0276310CB47C" authorityName="Orchymnont" authorityYear="1926" box="[1188,1341,862,885]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFB2D0276310CB47C" box="[1188,1341,862,885]" italics="true" pageId="5" pageNumber="234">Thysanarthria</emphasis>
</taxonomicName>
from all other non-chaetarthriine genera. Within
<taxonomicName id="4C5A4D507A2BFFBBFB5402563142B49C" authorityName="Bedel" authorityYear="1881" box="[1245,1395,894,917]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="tribe" tribe="Chaetarthriini">Chaetarthriini</taxonomicName>
, the well sclerotized and widely exposed labrum differentiates it from
<taxonomicName id="4C5A4D507A2BFFBBFC0E02963087B4DC" authority="Pandelle, 1876" authorityName="Pandelle" authorityYear="1876" box="[903,1206,958,981]" class="Magnoliopsida" family="Campanulaceae" genus="Hemisphaera" higherTaxonomySource="CoL" kingdom="Plantae" order="Asterales" pageId="5" pageNumber="234" phylum="Tracheophyta" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFC0E02963024B4DC" box="[903,1045,958,981]" italics="true" pageId="5" pageNumber="234">Hemisphaera</emphasis>
Pandellé, 1876
</taxonomicName>
(which is also smaller and has more depressed body: see
<bibRefCitation id="EFCB4B227A2BFFBBFB2102F63145B4FC" author="FIKACEK" box="[1192,1396,990,1013]" etAl="et al." firstAuthor="FIKACEK" pageId="5" pageNumber="234" pagination="107 - 130" refId="ref17935" refString="FIKACEK M., DELGADO J. A. &amp; GENTILI E. 2012: The hydrophiloid beetles of Socotra Island (Coleoptera: Georissidae, Hydrophilidae). Acta Entomologica Musei Nationalis Pragae 52 (supplementum): 107 - 130." type="journal article" year="2012">FIKÁČEK et al. 2012</bibRefCitation>
,
<bibRefCitation id="EFCB4B227A2BFFBBFAF702F6379EB31C" author="JIA" etAl="et al." firstAuthor="JIA" pageId="5" pageNumber="234" pagination="301 - 306" refId="ref18294" refString="JIA F., FIKACEK M. &amp; SONG K. 2013: Hemisphaera orientalis new species, the first species of Hemisphaera from the Oriental Region (Coleoptera: Hydrophilidae: Chaetarthriini). Journal of the Kansas Entomological Society 86: 301 - 306." type="journal article" year="2013">JIA et al. 2013</bibRefCitation>
) which can co-occur with
<emphasis id="B92EEAC17A2BFFBBFB4602D6315FB31C" box="[1231,1390,1022,1045]" italics="true" pageId="5" pageNumber="234">
<taxonomicName id="4C5A4D507A2BFFBBFB4602D6315BB31C" authorityName="Orchymnont" authorityYear="1926" box="[1231,1386,1022,1045]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">Thysanarthria</taxonomicName>
,
</emphasis>
and from the Neotropical genus
<taxonomicName id="4C5A4D507A2BFFBBFBF9053631ADB33C" authority="Spangler, 1986" authorityName="Spangler" authorityYear="1986" box="[1136,1436,1054,1077]" class="Insecta" family="Hydrophilidae" genus="Guyanobius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFBF9053630C0B33C" box="[1136,1265,1054,1077]" italics="true" pageId="5" pageNumber="234">Guyanobius</emphasis>
<bibRefCitation id="EFCB4B227A2BFFBBFB73053631ADB33C" author="Spangler" box="[1274,1436,1054,1077]" firstAuthor="Spangler" pageId="5" pageNumber="234" pagination="509 - 516" refId="ref18794" refString="SPANGLER P. J. 1986: Three new species of water scavenger beetles of the genus Chaetarthria from South America (Coleoptera: Hydrophilidae). Proceedings of the Biological Society of Washington 99: 509 - 516." type="journal article" year="1986">Spangler, 1986</bibRefCitation>
</taxonomicName>
(see
<bibRefCitation id="EFCB4B227A2BFFBBFCFB051630A6B35C" author="GUSTAFSON &amp; SHORT" box="[882,1175,1086,1109]" firstAuthor="GUSTAFSON" pageId="5" pageNumber="234" pagination="245 - 258" refId="ref17978" refString="GUSTAFSON G. T. &amp; SHORT A. E. Z. 2010: Revision of the Neotropical water scavenger beetle genus Guyanobius Spangler, 1986 (Coleoptera: Hydrophilidae: Chaetarthriini). Aquatic Insects 32: 245 - 258." type="journal article" year="2010">GUSTAFSON &amp; SHORT 2010</bibRefCitation>
).
<taxonomicName id="4C5A4D507A2BFFBBFB2405163178B35C" authorityName="Orchymnont" authorityYear="1926" box="[1197,1353,1086,1109]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFB2405163178B35C" box="[1197,1353,1086,1109]" italics="true" pageId="5" pageNumber="234">Thysanarthria</emphasis>
</taxonomicName>
can be distinguished from all three groups of
<taxonomicName id="4C5A4D507A2BFFBBFB4B0576317AB37C" authorityName="Stephens" authorityYear="1835" box="[1218,1355,1118,1141]" class="Insecta" family="Hydrophilidae" genus="Chaetarthria" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFB4B0576317AB37C" box="[1218,1355,1118,1141]" italics="true" pageId="5" pageNumber="234">Chaetarthria</emphasis>
</taxonomicName>
defined above by the elytra with 10 sharply impressed striae (
<figureCitation id="13612A567A2BFFBBFAE405563774B3BC" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Figs 1</figureCitation>
AD). Most species of the genus are easy to recognize in the samples by their small body size and pale coloration of pronotum and elytra constrasting with the black head (this coloration is not present only in the Near East
<taxonomicName id="4C5A4D507A2BFFBBFABA05D631ACB21C" authority="Fikáček &amp; Liu, 2019" authorityName="Fikáček &amp; Liu" authorityYear="2019" box="[1331,1437,1278,1301]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="persica" status="sp. nov.">
<emphasis id="B92EEAC17A2BFFBBFABA05D631ACB21C" box="[1331,1437,1278,1301]" italics="true" pageId="5" pageNumber="234">T. persica</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21D57BA7A2BFFBBFCB5043637BFB23C" box="[828,910,1310,1333]" pageId="5" pageNumber="234" rank="species">sp. nov.</taxonomicNameLabel>
and
<taxonomicName id="4C5A4D507A2BFFBBFC480436300FB23C" authority="Fikáček &amp; Liu, 2019" authorityName="Fikáček &amp; Liu" authorityYear="2019" box="[961,1086,1310,1333]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="wadicola" status="sp. nov.">
<emphasis id="B92EEAC17A2BFFBBFC480436300FB23C" box="[961,1086,1310,1333]" italics="true" pageId="5" pageNumber="234">T. wadicola</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A21D57BA7A2BFFBBFBCD043630ABB23C" box="[1092,1178,1310,1333]" pageId="5" pageNumber="234" rank="species">sp. nov.</taxonomicNameLabel>
).
</paragraph>
</subSubSection>
<subSubSection id="C34065587A2BFFB9FCB504163655BF1C" lastPageId="7" lastPageNumber="236" pageId="5" pageNumber="234" type="discussion">
<paragraph id="8BE536D37A2BFFB9FCB5041635C9B5DC" blockId="5.[828,1437,190,2069]" lastBlockId="7.[151,760,190,2069]" lastPageId="7" lastPageNumber="236" pageId="5" pageNumber="234">
<emphasis id="B92EEAC17A2BFFBBFCB5041631ADB25C" bold="true" box="[828,1436,1342,1365]" pageId="5" pageNumber="234">Characters important for species-level identification.</emphasis>
All known species of
<taxonomicName id="4C5A4D507A2BFFBBFBA7047630F9B27C" authorityName="Orchymnont" authorityYear="1926" box="[1070,1224,1374,1397]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A2BFFBBFBA7047630F9B27C" box="[1070,1224,1374,1397]" italics="true" pageId="5" pageNumber="234">Thysanarthria</emphasis>
</taxonomicName>
are very similar to each other in most external characters, and their tiny size makes the observation of many characters very difficult. The only external characters are (1) the presence/absence of the microsculpture on the head and labrum, pronotum and elytra, which can be either strongly mesh-like, weak and granulate, or totally absent; and (2) the body shape which can be wider (
<figureCitation id="13612A567A2BFFBBFB920716305CB15C" box="[1051,1133,1598,1621]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Fig. 1A</figureCitation>
) or more elongate (
<figureCitation id="13612A567A2BFFBBFAC907163763B17C" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Figs 1B, D</figureCitation>
), but this is hard to compare in specimens which are not mounted in extended position on labels. Body coloration differs between species, with pronotum and elytra either uniformly yellowish (
<figureCitation id="13612A567A2BFFBBFBAB079630A6B1DC" box="[1058,1175,1726,1749]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Figs 1A, C</figureCitation>
) or partly darkened (e.g. pronotum in
<figureCitation id="13612A567A2BFFBBFC4C07F63027B1FC" box="[965,1046,1758,1781]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Fig. 1D</figureCitation>
) or uniformly dark brown (
<figureCitation id="13612A567A2BFFBBFAB207F631BFB1FC" box="[1339,1422,1758,1781]" captionStart="Fig" captionStartId="2.[151,182,2004,2023]" captionTargetBox="[165,1409,612,1979]" captionTargetId="figure-13@2.[151,1437,604,1996]" captionTargetPageId="2" captionText="Fig. 1. Habitus of the Thysanarthria species and related Old Word Chaetarthriini.A Thysanarthria bengalensis Hebauer, 2001, holotype; B T. persica sp. nov., paratype; C T. siamensis Hebauer, 2001, holotype; D T. hongsonensis Hebauer, 2001, holotype; E European Chaetarthria: C. seminulum (Herbst, 1797); F large Old World Chaetarthria: C. nigerrima (Blackburn, 1891)." figureDoi="http://doi.org/10.5281/zenodo.4488920" httpUri="https://zenodo.org/record/4488920/files/figure.png" pageId="5" pageNumber="234">Fig. 1B</figureCitation>
). However, examination of longer series of some species (
<taxonomicName id="4C5A4D507A2BFFBBFCCC063637E2B03C" baseAuthorityName="Knisch" baseAuthorityYear="1924" box="[837,979,1822,1845]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="championi">
<emphasis id="B92EEAC17A2BFFBBFCCC063637E2B03C" box="[837,979,1822,1845]" italics="true" pageId="5" pageNumber="234">T. championi</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5A4D507A2BFFBBFB80063630BFB03C" authorityName="Hebauer" authorityYear="2001" box="[1033,1166,1822,1845]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="siamensis">
<emphasis id="B92EEAC17A2BFFBBFB80063630BFB03C" box="[1033,1166,1822,1845]" italics="true" pageId="5" pageNumber="234">T. siamensis</emphasis>
</taxonomicName>
) revealed that the coloration can vary within a species, and hence is not always reliable for identification. The same is true for the dorsal body microsculpture which seems to vary in intensity, at least in
<taxonomicName id="4C5A4D507A2BFFBBFC0706B637C8B0BC" authorityName="Balfour-Browne" authorityYear="1951" box="[910,1017,1950,1973]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="234" phylum="Arthropoda" rank="species" species="brittoni">
<emphasis id="B92EEAC17A2BFFBBFC0706B637C8B0BC" box="[910,1017,1950,1973]" italics="true" pageId="5" pageNumber="234">T. brittoni</emphasis>
</taxonomicName>
(see under that species). The body size also differs between species, and the presence of specimens of different size in the same series may indicate the presence of multiple species. However, in species in which more specimens were available, the body size was revealed to vary to some extent as well, and the body size can be hence used as an additional character only. Therefore, examination of the male genitalia is necessary for reliable identification in all cases. Ideally, the genitalia should be examined under a medium magnification of the compound microscope, and attention should be paid also to the membranous structures on the apical part of the median lobe (including short, paired, subapical projections which are present only in some species, e.g.
<figureCitation id="13612A567A28FFB8FAA800363152B63C" box="[1313,1379,286,309]" captionStart="Fig" captionStartId="10.[887,918,1799,1818]" captionTargetBox="[178,1439,395,2038]" captionTargetId="figure-13@10.[175,1413,386,2041]" captionTargetPageId="10" captionText="Fig. 5. Male genitalia of Thysanarthria species, holotypes.AE T.brincki Hebauer, 2001 (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view); FJ T. cardamona sp. nov. (FH whole aedeagus in dorsal, lateral and ventral view; IJ detail of median lobe and parameres in dorsal and ventral view); KM T. bengalensis Hebauer, 2001 (K photo of the aedeagus before re-mounting; L whole aedeagus in ventral view; M whole aedeagus in lateral view)." figureDoi="http://doi.org/10.5281/zenodo.4488932" httpUri="https://zenodo.org/record/4488932/files/figure.png" pageId="6" pageNumber="235">Figs 5</figureCitation>
DE, IJ,
<figureCitation id="13612A567A28FFB8FCE40016374BB65C" box="[877,890,318,341]" captionStart="Fig" captionStartId="12.[151,182,1980,1999]" captionTargetBox="[155,1405,208,1868]" captionTargetId="figure-13@12.[153,1435,189,1873]" captionTargetPageId="12" captionText="Fig. 6. Male genitalia of Thysanarthria species, holotypes (AI, KO) and paratype (J). AE T.bifida sp. nov. (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view). FJ T. trifida sp. nov. (FH whole aedeagus of the holotype in dorsal, lateral and ventral view; I details of parameres of the holotype; J same of the paratype). KO T. ceylonensis Hebauer, 2001 (KM whole aedeagus in dorsal, lateral and ventral view; NO details of median lobe and parameres in dorsal and ventral view)." figureDoi="http://doi.org/10.5281/zenodo.4488934" httpUri="https://zenodo.org/record/4488934/files/figure.png" pageId="6" pageNumber="235">6</figureCitation>
DE, IJ). The proportions of parameres may be uneasy to observe as they are partly affected by the position of the aedeagus, and the genitalia should be carefully observed under slightly different angles in case of doubts. The ratio of paramere length to phallobase length should be evaluated in lateral view, due to the strongly bent phallobase in many species. The examination of the material used for this study shows that especially the form of the median lobe is constant and diagnostic, whereas the shape of parameres may slightly vary. The apex of the median lobe is membranous in many species, even though usually rather constant in shape, and it seems that at least in some species it can include parts which are normally inverted and hence not easy to observe, and may sometimes get fully everted after the treatment in KOH (which however may distort other parts of the aedeagus; see e.g. 4J and 7E which show fully everted apical membranous parts). Since the observation of this part is difficult, we did not consider it for species diagnosis.
</paragraph>
<caption id="DF25665B7A28FFB8FF1E069A361EBF1C" ID-DOI="http://doi.org/10.5281/zenodo.4488928" ID-Zenodo-Dep="4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="6" pageNumber="235" startId="6.[151,182,1970,1989]" targetBox="[178,1407,394,1955]" targetPageId="6">
<paragraph id="8BE536D37A28FFB8FF1E069A361EBF1C" blockId="6.[151,1437,1970,2069]" pageId="6" pageNumber="235">
Fig. 3. Morphology of
<taxonomicName id="4C5A4D507A28FFB8FED1069A35E3B0CC" authorityName="Orchymnont" authorityYear="1926" box="[344,466,1970,1989]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="235" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A28FFB8FED1069A35E3B0CC" box="[344,466,1970,1989]" italics="true" pageId="6" pageNumber="235">Thysanarthria</emphasis>
</taxonomicName>
. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF
<taxonomicName id="4C5A4D507A28FFB8FBFA06CF34FABF1C" authority="(Knisch, 1924)" baseAuthorityName="Knisch" baseAuthorityYear="1924" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="235" phylum="Arthropoda" rank="species" species="championi">
<emphasis id="B92EEAC17A28FFB8FBFA06CF317CB0F3" box="[1139,1357,2023,2042]" italics="true" pageId="6" pageNumber="235">Thysanarthria championi</emphasis>
(
<bibRefCitation id="EFCB4B227A28FFB8FAD006CF34F4BF1C" author="KNISCH A." bookContentInfo="306 pp." editor="JUNK W. &amp; SCHENKLING S." journalOrPublisher="W. Junk, Berlin" pageId="6" pageNumber="235" refId="ref18391" refString="KNISCH A. 1924 a: Hydrophilidae. In: JUNK W. &amp; SCHENKLING S. (eds): Coleopterorum Catalogus. Vol. 14, part 79. W. Junk, Berlin, 306 pp." title="Hydrophilidae" type="book chapter" volumeTitle="Coleopterorum Catalogus. Vol. 14, part 79." year="1924">Knisch, 1924</bibRefCitation>
)
</taxonomicName>
; B, GH
<taxonomicName id="4C5A4D507A28FFB8FEA5092A361ABF1C" authority="(Regimbart, 1903)" baseAuthorityName="Regimbart" baseAuthorityYear="1903" box="[300,555,2050,2069]" class="Insecta" family="Hydrophilidae" genus="Hydrobius" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="235" phylum="Arthropoda" rank="species" species="atriceps">
<emphasis id="B92EEAC17A28FFB8FEA5092A35B7BF1C" box="[300,390,2050,2069]" italics="true" pageId="6" pageNumber="235">T. atriceps</emphasis>
(
<bibRefCitation id="EFCB4B227A28FFB8FE18092A3614BF1C" author="Regimbart" box="[401,549,2050,2069]" firstAuthor="Regimbart" pageId="6" pageNumber="235" pagination="1 - 51" refId="ref18592" refString="REGIMBART M. 1903: Coleopteres aquatiques (Haliplidae, Dytiscidae, Gyrinidae et Hydrophilidae) recueillis dans le Sud de Madagascar par M. Ch. Alluaud (Juillet 1900 - mai 1901). Annales de la Societe Entomologique de France 72: 1 - 51." type="journal article" year="1903">Régimbart, 1903</bibRefCitation>
)
</taxonomicName>
.
</paragraph>
</caption>
<paragraph id="8BE536D37A29FFB9FF3C03F6356EB4FC" blockId="7.[151,760,190,2069]" pageId="7" pageNumber="236">As male genitalia are the only reliable character for species identification, below we provide detailed illustrations of the genitalia with which new specimens to be identified should be compared. Once the candidate species is found based on genital morphology, the external characters mentioned above (body size and coloration, presence/ absence of the microsculpture) should be compared with the (re)descriptions provided below. No identification key is hence provided.</paragraph>
<paragraph id="8BE536D37A29FFB9FF1E02D636AFB27C" blockId="7.[151,760,190,2069]" pageId="7" pageNumber="236">
<emphasis id="B92EEAC17A29FFB9FF1E02D63577B31C" bold="true" box="[151,326,1022,1045]" pageId="7" pageNumber="236">Species groups.</emphasis>
The limited number of characters makes it difficult to group the species into supposedly monophyletic species groups. Based on the genital morphology, the African
<taxonomicName id="4C5A4D507A29FFB9FE9E05763623B37C" baseAuthorityName="Regimbart" baseAuthorityYear="1903" box="[279,530,1118,1141]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="atriceps">
<emphasis id="B92EEAC17A29FFB9FE9E05763623B37C" box="[279,530,1118,1141]" italics="true" pageId="7" pageNumber="236">Thysanarthria atriceps</emphasis>
</taxonomicName>
is very similar to the Arabian
<taxonomicName id="4C5A4D507A29FFB9FF7D05563551B39C" authorityName="Balfour-Browne" authorityYear="1951" box="[244,352,1150,1173]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="brittoni">
<emphasis id="B92EEAC17A29FFB9FF7D05563551B39C" box="[244,352,1150,1173]" italics="true" pageId="7" pageNumber="236">T. brittoni</emphasis>
</taxonomicName>
, and these two species seem to form a group of closely related species. The presence of subapical membranous lobes on the median lobe in
<taxonomicName id="4C5A4D507A29FFB9FD01059636C0B3DC" authorityName="Hebauer" authorityYear="2001" box="[648,753,1214,1237]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="brincki">
<emphasis id="B92EEAC17A29FFB9FD01059636C0B3DC" box="[648,753,1214,1237]" italics="true" pageId="7" pageNumber="236">T. brincki</emphasis>
</taxonomicName>
,
<taxonomicName id="4C5A4D507A29FFB9FF1E05F634DCB3FC" authorityName="Fikáček &amp; Liu" authorityYear="2019" box="[151,237,1246,1269]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="bifida">
<emphasis id="B92EEAC17A29FFB9FF1E05F634DCB3FC" box="[151,237,1246,1269]" italics="true" pageId="7" pageNumber="236">T. bifida</emphasis>
</taxonomicName>
,
<taxonomicName id="4C5A4D507A29FFB9FF7005F635BBB3FC" authorityName="Fikáček &amp; Liu" authorityYear="2019" box="[249,394,1246,1269]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="cardamona">
<emphasis id="B92EEAC17A29FFB9FF7005F635BBB3FC" box="[249,394,1246,1269]" italics="true" pageId="7" pageNumber="236">T. cardamona</emphasis>
</taxonomicName>
,
<taxonomicName id="4C5A4D507A29FFB9FE1F05F6361BB3FC" authorityName="Hebauer" authorityYear="2001" box="[406,554,1246,1269]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="madurensis">
<emphasis id="B92EEAC17A29FFB9FE1F05F6361BB3FC" box="[406,554,1246,1269]" italics="true" pageId="7" pageNumber="236">T. madurensis</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C5A4D507A29FFB9FDE805F6368DB3FC" authorityName="Fikáček &amp; Liu" authorityYear="2019" box="[609,700,1246,1269]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="trifida">
<emphasis id="B92EEAC17A29FFB9FDE805F6368DB3FC" box="[609,700,1246,1269]" italics="true" pageId="7" pageNumber="236">T. trifida</emphasis>
</taxonomicName>
(
<figureCitation id="13612A567A29FFB9FD4105F63492B21C" captionStart="Fig" captionStartId="10.[887,918,1799,1818]" captionTargetBox="[178,1439,395,2038]" captionTargetId="figure-13@10.[175,1413,386,2041]" captionTargetPageId="10" captionText="Fig. 5. Male genitalia of Thysanarthria species, holotypes.AE T.brincki Hebauer, 2001 (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view); FJ T. cardamona sp. nov. (FH whole aedeagus in dorsal, lateral and ventral view; IJ detail of median lobe and parameres in dorsal and ventral view); KM T. bengalensis Hebauer, 2001 (K photo of the aedeagus before re-mounting; L whole aedeagus in ventral view; M whole aedeagus in lateral view)." figureDoi="http://doi.org/10.5281/zenodo.4488932" httpUri="https://zenodo.org/record/4488932/files/figure.png" pageId="7" pageNumber="236">Figs 5</figureCitation>
DE, IJ;
<figureCitation id="13612A567A29FFB9FE9C05D63513B21C" box="[277,290,1278,1301]" captionStart="Fig" captionStartId="12.[151,182,1980,1999]" captionTargetBox="[155,1405,208,1868]" captionTargetId="figure-13@12.[153,1435,189,1873]" captionTargetPageId="12" captionText="Fig. 6. Male genitalia of Thysanarthria species, holotypes (AI, KO) and paratype (J). AE T.bifida sp. nov. (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view). FJ T. trifida sp. nov. (FH whole aedeagus of the holotype in dorsal, lateral and ventral view; I details of parameres of the holotype; J same of the paratype). KO T. ceylonensis Hebauer, 2001 (KM whole aedeagus in dorsal, lateral and ventral view; NO details of median lobe and parameres in dorsal and ventral view)." figureDoi="http://doi.org/10.5281/zenodo.4488934" httpUri="https://zenodo.org/record/4488934/files/figure.png" pageId="7" pageNumber="236">6</figureCitation>
DE, IJ;
<figureCitation id="13612A567A29FFB9FE1A05D635AFB21C" box="[403,414,1278,1301]" captionStart="Fig" captionStartId="16.[151,182,1976,1995]" captionTargetBox="[204,1420,202,1944]" captionTargetId="figure-108@16.[162,1439,189,1956]" captionTargetPageId="16" captionText="Fig. 8. Male genitalia of Thysanarthria species, holotypes.AE T. chui sp. nov.(AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view). FJ T. hongsonensis Hebauer, 2001 (FH whole aedeagus in dorsal, lateral and ventral view; IJ details of parameres and median lobe in dorsal and ventral view). KO T. madurensis Hebauer, 2001 (KM whole aedeagus in dorsal, lateral and ventral view; NO details of median lobe and parameres in dorsal and ventral view)." figureDoi="http://doi.org/10.5281/zenodo.4488940" httpUri="https://zenodo.org/record/4488940/files/figure.png" pageId="7" pageNumber="236">8</figureCitation>
NO) may also point to a close relationships.
<taxonomicName id="4C5A4D507A29FFB9FEBA04363614B23C" authorityName="Hebauer" authorityYear="2001" box="[307,549,1310,1333]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="brincki">
<emphasis id="B92EEAC17A29FFB9FEBA04363614B23C" box="[307,549,1310,1333]" italics="true" pageId="7" pageNumber="236">Thysanarthria brincki</emphasis>
</taxonomicName>
and
<taxonomicName id="4C5A4D507A29FFB9FDE9043636C6B23C" authorityName="Fikáček &amp; Liu" authorityYear="2019" box="[608,759,1310,1333]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="species" species="cardamona">
<emphasis id="B92EEAC17A29FFB9FDE9043636C6B23C" box="[608,759,1310,1333]" italics="true" pageId="7" pageNumber="236">T. cardamona</emphasis>
</taxonomicName>
may form a clade within this group characterized by lateroapical spine on the paramere (
<figureCitation id="13612A567A29FFB9FD9404763650B27C" box="[541,609,1374,1397]" captionStart="Fig" captionStartId="10.[887,918,1799,1818]" captionTargetBox="[178,1439,395,2038]" captionTargetId="figure-13@10.[175,1413,386,2041]" captionTargetPageId="10" captionText="Fig. 5. Male genitalia of Thysanarthria species, holotypes.AE T.brincki Hebauer, 2001 (AC whole aedeagus in dorsal, lateral and ventral view; DE detail of median lobe and parameres in dorsal and ventral view); FJ T. cardamona sp. nov. (FH whole aedeagus in dorsal, lateral and ventral view; IJ detail of median lobe and parameres in dorsal and ventral view); KM T. bengalensis Hebauer, 2001 (K photo of the aedeagus before re-mounting; L whole aedeagus in ventral view; M whole aedeagus in lateral view)." figureDoi="http://doi.org/10.5281/zenodo.4488932" httpUri="https://zenodo.org/record/4488932/files/figure.png" pageId="7" pageNumber="236">Figs 5</figureCitation>
AJ).
</paragraph>
<paragraph id="8BE536D37A29FFB9FF1E04563655BF1C" blockId="7.[151,760,190,2069]" pageId="7" pageNumber="236">
<emphasis id="B92EEAC17A29FFB9FF1E045636FCB29C" bold="true" box="[151,717,1406,1429]" pageId="7" pageNumber="236">Function of the abdominal gelatinous substance.</emphasis>
All members of the tribe
<taxonomicName id="4C5A4D507A29FFB9FEF404B63624B2BC" authorityName="Bedel" authorityYear="1881" box="[381,533,1438,1461]" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="tribe" tribe="Chaetarthriini">Chaetarthriini</taxonomicName>
including
<taxonomicName id="4C5A4D507A29FFB9FD0104B634FAB2DC" authorityName="Orchymnont" authorityYear="1926" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A29FFB9FD0104B634FAB2DC" italics="true" pageId="7" pageNumber="236">Thysanarthria</emphasis>
</taxonomicName>
bear a series of long setae on the base of the abdomen which cover a shallow depression in ventrites 12 filled in by whitish gelatinous substance (
<figureCitation id="13612A567A29FFB9FDAD04D63658B11C" box="[548,617,1534,1557]" captionStart="Fig" captionStartId="6.[151,182,1970,1989]" captionTargetBox="[178,1407,394,1955]" captionTargetId="figure-13@6.[176,1412,393,1956]" captionTargetPageId="6" captionText="Fig. 3. Morphology of Thysanarthria. A mentum and maxillary palps; B meso- and metaventrite and meso- and metafemora; C head and prothorax in ventral view; D antenna; E ultimate maxillary palpomere with basal peg-like setae; F metatarsus; G ventral view with gelatinous substrance on the base of abdomen; H detail of abdomen with gelatinous substance in the cavity on ventrites 12. A, CF Thysanarthria championi (Knisch, 1924); B, GH T. atriceps (Régimbart, 1903)." figureDoi="http://doi.org/10.5281/zenodo.4488928" httpUri="https://zenodo.org/record/4488928/files/figure.png" pageId="7" pageNumber="236">Figs 3</figureCitation>
GH). When submerged,
<taxonomicName id="4C5A4D507A29FFB9FE9107363580B13C" authorityName="Orchymnont" authorityYear="1926" box="[280,433,1566,1589]" class="Insecta" family="Hydrophilidae" genus="Thysanarthria" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A29FFB9FE9107363580B13C" box="[280,433,1566,1589]" italics="true" pageId="7" pageNumber="236">Thysanarthria</emphasis>
</taxonomicName>
floats with dorsal body surface facing up, i.e. in the position usual for most other
<taxonomicName id="4C5A4D507A29FFB9FD20071634C1B17C" authorityName="Latreille" authorityYear="1802" class="Insecta" family="Hydrophilidae" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="family">Hydrophilidae</taxonomicName>
. The gelatinous substance hence does not serve to increase the buoyancy of the beetle as might be the case in the non-related genus
<taxonomicName id="4C5A4D507A29FFB9FE1407B63681B1BC" authority="Erichson, 1843" authorityName="Erichson" authorityYear="1843" box="[413,688,1694,1717]" class="Insecta" family="Hydrophilidae" genus="Amphiops" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="7" pageNumber="236" phylum="Arthropoda" rank="genus">
<emphasis id="B92EEAC17A29FFB9FE1407B63638B1BC" box="[413,521,1694,1717]" italics="true" pageId="7" pageNumber="236">Amphiops</emphasis>
Erichson, 1843
</taxonomicName>
which bears similar-looking gelatinous substance at the base of abdomen and is swimming in an upside-down position when submerged (Fikáček &amp;
<collectingRegion id="499EF8317A29FFB9FE5F07D63611B01C" box="[470,544,1790,1813]" country="United Kingdom" name="Angus" pageId="7" pageNumber="236">Angus</collectingRegion>
, pers. observ.). Moreover, the gelatinous substance does not interfere with the ventral air bubble of the submerged beetle: the bubble covers the whole ventral side of the beetle including the whole abdomen. Therefore, it seems that the substance cannot be functional in submerged beetle but may be an adaptation to its usual environment on the wet sand outside water. The gelatinous substance is sticky in alive specimens. Its function remains unknown.
</paragraph>
</subSubSection>
</treatment>
</document>