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<document id="3BB43BA5B5245B7FBA5D320AD077D4AC" ID-DOI="10.37520/aemnp.2021.009" ID-ISSN="1804-6487" ID-Zenodo-Dep="5037330" ID-ZooBank="27EA8466-6B29-40F4-8236-DD49061C523B" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1624895377019" checkinUser="carolina" docAuthor="Liu, Tian-tian, Ono, Hiroki &amp; Maruyama, Munetoshi" docDate="2021" docId="03CB023BAC1AFF914CE7F8FE15C8F8F8" docLanguage="en" docName="ActaEntMusNatPra.61.1.163-201.pdf" docOrigin="Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 61 (1)" docSource="http://dx.doi.org/10.37520/aemnp.2021.009" docStyle="DocumentStyle:BA2C5C8540539042E893A65A182CEA85.8:ActaEntMusNatPra.2018-.journal_article" docStyleId="BA2C5C8540539042E893A65A182CEA85" docStyleName="ActaEntMusNatPra.2018-.journal_article" docStyleVersion="8" docTitle="Bryothinusa hokkaidensis Liu &amp; Ono &amp; Maruyama 2021, sp. nov." docType="treatment" docVersion="3" lastPageNumber="182" masterDocId="FFF27A43AC0AFF824F48FFFB175FFFCD" masterDocTitle="Revision of the intertidal rove beetle genus Bryothinusa from Japan (Coleoptera: Staphylinidae: Aleocharinae)" masterLastPageNumber="201" masterPageNumber="163" pageNumber="179" updateTime="1698977316832" updateUser="plazi" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:title id="2AF47AC4395C6F60CBCE9FAA445FD6E0">Revision of the intertidal rove beetle genus Bryothinusa from Japan (Coleoptera: Staphylinidae: Aleocharinae)</mods:title>
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<mods:namePart id="C666E4FF5BE817BD63AD6EA86D4EFFF8">Liu, Tian-tian</mods:namePart>
<mods:affiliation id="9F89F00B7CA248E5AC7A6538D18B4EB8">Entomological Laboratory, Graduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, Fukuoka, 819 - 0395, Japan &amp; The Kyushu University Museum, Hakozaki</mods:affiliation>
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<mods:namePart id="EFD23C3F7AA39286C631DAAA53E8A232">Ono, Hiroki</mods:namePart>
<mods:affiliation id="17E2E4FF31091AA70AB23767A1FA2066">Sounan, Minami-ku, Sagamihara-shi, Kanagawa, 252 - 0312, Japan</mods:affiliation>
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<mods:namePart id="CD12C1F4295A328FBEE6D29F1629043E">Maruyama, Munetoshi</mods:namePart>
<mods:affiliation id="F651A6E555AD22BE01975DC86C4482C5">The Kyushu University Museum, Hakozaki</mods:affiliation>
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<subSubSection id="C378E0A6AC1AFF924CE7F8FE1275F8D0" box="[943,1322,1797,1821]" pageId="16" pageNumber="179" type="nomenclature">
<paragraph id="8BDDB32DAC1AFF924CE7F8FE1275F8D0" blockId="16.[943,1322,1797,1821]" box="[943,1322,1797,1821]" pageId="16" pageNumber="179">
<heading id="D0950441AC1AFF924CE7F8FE1275F8D0" box="[943,1322,1797,1821]" centered="true" fontSize="10" level="2" pageId="16" pageNumber="179" reason="3">
<emphasis id="B9166F3FAC1AFF924CE7F8FE1275F8D0" bold="true" box="[943,1322,1797,1821]" pageId="16" pageNumber="179">
<taxonomicName id="4C62C8AEAC1AFF924CE7F8FE1391F8D0" ID-CoL="8XW4J" authority="Liu &amp; Ono &amp; Maruyama, 2021" authorityName="Liu &amp; Ono &amp; Maruyama" authorityYear="2021" box="[943,1230,1797,1821]" class="Insecta" family="Staphylinidae" genus="Bryothinusa" kingdom="Animalia" order="Coleoptera" pageId="16" pageNumber="179" phylum="Arthropoda" rank="species" species="hokkaidensis" status="sp. nov.">
<emphasis id="B9166F3FAC1AFF924CE7F8FE1391F8D0" bold="true" box="[943,1230,1797,1821]" italics="true" pageId="16" pageNumber="179">Bryothinusa hokkaidensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A225D244AC1AFF924B9CF8FD1275F8D0" box="[1236,1322,1798,1821]" pageId="16" pageNumber="179" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C378E0A6AC1AFF924B49F8DF126FF89C" pageId="16" pageNumber="179" type="description">
<paragraph id="8BDDB32DAC1AFF924B49F8DF1388F8FA" blockId="16.[828,1437,1828,2049]" box="[1025,1239,1828,1847]" pageId="16" pageNumber="179">
(
<figureCitation id="1359AFA8AC1AFF924B4FF8DF131AF8FA" box="[1031,1093,1828,1847]" captionStart="Fig" captionStartId="2.[151,182,1972,1991]" captionTargetBox="[189,1405,205,1944]" captionTargetId="figure-13@2.[175,1411,189,1947]" captionTargetPageId="2" captionText="Fig. 1. Habitus of Bryothinusa spp. A B. algarum Sawada, 1971; B B. minuta (Sawada, 1955); C B. tsutsuii (Sawada, 1955); D B. koreana Ahn &amp; Jeon, 2004; E B. yoshigoui sp. nov.; F B. japonica Liu, Ono &amp; Maruyama, 2020; G B. sakishimana Sawada, 1991; H B. constricta sp. nov.; I B. hokkaidensis sp. nov.; J B. fulvipennis sp. nov.; K B. itsuroi sp. nov.; L B. aikoae sp. nov.; M B. okinawana sp. nov.; N B. nigra sp. nov.; O B. nakanei (Sawada, 1955); P B. moriguchii sp. nov.; Q B. gangjinensis Ahn &amp; Jeon, 2004. Scale bars: 1.00 mm." figureDoi="http://doi.org/10.5281/zenodo.5037332" httpUri="https://zenodo.org/record/5037332/files/figure.png" pageId="16" pageNumber="179">Figs 1I</figureCitation>
;
<figureCitation id="1359AFA8AC1AFF924B07F8DF133CF8FA" box="[1103,1123,1828,1847]" captionStart="Fig" captionStartId="4.[151,182,1978,1997]" captionTargetBox="[196,1391,190,1955]" captionTargetId="figure-13@4.[193,1394,189,1958]" captionTargetPageId="4" captionText="Fig. 2. Compound eyes of Bryothinusa spp.A B. algarum Sawada, 1971; B B. minuta (Sawada, 1955); C B. tsutsuii (Sawada, 1955); D B. koreana Ahn &amp; Jeon, 2004; E B. yoshigoui sp. nov.; F B. japonica Liu, Ono &amp; Maruyama, 2020; G B. sakishimana Sawada, 1991; H B. constricta sp. nov.; I B. hokkaidensis sp. nov.; J B. fulvipennis sp. nov.; K B. itsuroi sp. nov.; L B. aikoae sp. nov.; M B. okinawana sp. nov.; N B. nigra sp. nov.; O B. nakanei (Sawada, 1955); P B. moriguchii sp. nov.; Q B. gangjinensis Ahn &amp; Jeon, 2004. Scale bars: 0.05 mm." figureDoi="http://doi.org/10.5281/zenodo.5037334" httpUri="https://zenodo.org/record/5037334/files/figure.png" pageId="16" pageNumber="179">2I</figureCitation>
;
<figureCitation id="1359AFA8AC1AFF924B24F8DF13DDF8FA" box="[1132,1154,1828,1847]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="16" pageNumber="179">11</figureCitation>
;
<figureCitation id="1359AFA8AC1AFF924BC4F8DF13EDF8FA" box="[1164,1202,1828,1847]" captionStart="Fig" captionStartId="34.[151,182,1955,1974]" captionTargetBox="[167,1408,193,1935]" captionTargetId="figure-13@34.[161,1426,189,1938]" captionTargetPageId="34" captionText="Fig. 21. Collection localities of Bryothinusa species in Japan: A B. koreana Ahn &amp; Jeon, 2004 and B. japonica Liu, Ono &amp; Maruyama, 2020 in Shima-nogita, Fukuoka; B B. aikoae sp. nov., B. fulvipennis sp. nov., B. japonica and B. nigra sp. nov. in Kuroto, Chiba; C B. hokkaidensis sp. nov. in Shunkunitai, Hokkaidô; D B. nigra sp. nov. and B. tsutsuii (Sawada, 1955) in Edateku-jima; E B. nakanei (Sawada, 1955) in Nagura-anparu, Ishigaki-jima; F B. gangjinensis Ahn &amp; Jeon, 2004 in Iki-no-matsubara, Fukuoka; G B. algarum Sawada, 1971, B. japonica and B. minuta (Sawada, 1955) in Sunosaki, Fukuoka." figureDoi="http://doi.org/10.5281/zenodo.5037374" httpUri="https://zenodo.org/record/5037374/files/figure.png" pageId="16" pageNumber="179">21C</figureCitation>
;
<figureCitation id="1359AFA8AC1AFF924BF3F8DF138EF8FA" box="[1211,1233,1828,1847]" captionStart="Fig" captionStartId="35.[150,181,1388,1407]" captionTargetBox="[150,1434,191,1370]" captionTargetId="figure-236@35.[150,1435,191,1371]" captionTargetPageId="35" captionText="Fig. 22. Map of examined specimens of Bryothinusa spp. from Japan: white inverted triangle B. algarum Sawada, 1971; black inverted triangle B. minuta (Sawada, 1955); white star B. tsutsuii (Sawada, 1955); black star B. koreana Ahn &amp; Jeon, 2004; black square B. yoshigoui sp. nov.; white triangle B. japonica Liu, Ono &amp; Maruyama, 2020; black diamond B. sakishimana Sawada, 1991; white diamond B. constricta sp. nov.; white square B. hokkaidensis sp. nov.; black triangle B. fulvipennis sp. nov.; double circle B. itsuroi sp. nov.; arrow B. aikoae sp. nov.; black hart B. okinawana sp. nov.; circled dot B. nigra sp. nov.; white circle B. nakanei (Sawada, 1955); white hart B. moriguchii sp. nov.; black circle B. gangjinensis Ahn &amp; Jeon, 2004." figureDoi="http://doi.org/10.5281/zenodo.5037376" httpUri="https://zenodo.org/record/5037376/files/figure.png" pageId="16" pageNumber="179">22</figureCitation>
)
</paragraph>
<paragraph id="8BDDB32DAC1AFF924CE0F8C5126FF89C" blockId="16.[828,1437,1828,2049]" box="[936,1328,1854,1873]" pageId="16" pageNumber="179">(Japanese name: Hokkai-nagisa-hanekakushi)</paragraph>
</subSubSection>
<subSubSection id="C378E0A6AC1AFF924C74F89312C2F7CC" pageId="16" pageNumber="179" type="materials_examined">
<paragraph id="8BDDB32DAC1AFF924C74F89312C2F7CC" blockId="16.[828,1437,1828,2049]" pageId="16" pageNumber="179">
<emphasis id="B9166F3FAC1AFF924C74F8931496F8B6" bold="true" box="[828,969,1896,1915]" pageId="16" pageNumber="179">Type material.</emphasis>
Hඈඅඈඍඒඉൾ: J,
<emphasis id="B9166F3FAC1AFF924B15F893127DF8B6" bold="true" box="[1117,1314,1896,1915]" pageId="16" pageNumber="179">
<collectingCountry id="F375F3BDAC1AFF924B15F89313F8F8B6" box="[1117,1191,1896,1915]" name="Japan" pageId="16" pageNumber="179">JAPAN</collectingCountry>
: Hඈκκൺൽත:
</emphasis>
Shunkunitai,
<geoCoordinate id="EE56D5EAAC1AFF924C74F87814CAF85B" box="[828,917,1923,1942]" degrees="43.2725" direction="north" orientation="latitude" pageId="16" pageNumber="179" precision="5" value="43.2725">43.2725 N</geoCoordinate>
,
<geoCoordinate id="EE56D5EAAC1AFF924CD4F87814ACF85B" box="[924,1011,1923,1942]" degrees="145.472" direction="east" orientation="longitude" pageId="16" pageNumber="179" precision="55" value="145.472">145.472 E</geoCoordinate>
, Nemuro-shi,
<collectingRegion id="49A67DCFAC1AFF924B2FF87813E3F85B" box="[1127,1212,1923,1942]" country="Japan" name="Hokkaido" pageId="16" pageNumber="179">Hokkaidô</collectingRegion>
,
<date id="FFDC95EDAC1AFF924B8BF8781241F85B" box="[1219,1310,1923,1942]" pageId="16" pageNumber="179" value="1999-09-11">11.ix.1999</date>
, M. Maruyama (
<collectionCode id="ED732BE8AC1AFF924C0AF8651427F87C" box="[834,888,1950,1969]" country="Ghana" lsid="urn:lsid:biocol.org:col:13924" name="Resource Management Support Center" pageId="16" pageNumber="179" type="Herbarium">KUM</collectionCode>
). Pൺ*©ൺඍඒඉൾඌ:
<emphasis id="B9166F3FAC1AFF924CA5F8651308F87C" bold="true" box="[1005,1111,1950,1969]" pageId="16" pageNumber="179">Hඈκκൺൽත:</emphasis>
3 JJ
<specimenCount id="9D6478A4AC1AFF924BDCF865139AF87D" box="[1172,1221,1943,1969]" pageId="16" pageNumber="179" type="female">4 ♀♀</specimenCount>
, 96 unsexed spec., same data as
<typeStatus id="54D90D8FAC1AFF924C30F843149DF806" box="[888,962,1976,1995]" pageId="16" pageNumber="179" type="holotype">holotype</typeStatus>
(
<collectionCode id="ED732BE8AC1AFF924C84F843135EF806" box="[972,1025,1976,1995]" country="Ghana" lsid="urn:lsid:biocol.org:col:13924" name="Resource Management Support Center" pageId="16" pageNumber="179" type="Herbarium">KUM</collectionCode>
,
<collectionCode id="ED732BE8AC1AFF924B4FF843131AF806" box="[1031,1093,1976,1995]" country="Czech Republic" httpUri="http://grbio.org/cool/hj12-2xm7" name="National Museum Prague" pageId="16" pageNumber="179">NMPC</collectionCode>
,
<collectionCode id="ED732BE8AC1AFF924B04F84313D0F806" box="[1100,1167,1976,1995]" pageId="16" pageNumber="179">BMNH</collectionCode>
);
<specimenCount id="9D6478A4AC1AFF924BD5F84313E4F806" box="[1181,1211,1970,1995]" pageId="16" pageNumber="179" type="female">1 ♀</specimenCount>
, 4 unsexed spec., Usu-chô, Date-shi,
<collectingRegion id="49A67DCFAC1AFF924CC5F82814BCF82B" box="[909,995,2003,2022]" country="Japan" name="Hokkaido" pageId="16" pageNumber="179">Hokkaidô</collectingRegion>
,
<date id="FFDC95EDAC1AFF924CA2F828131AF82B" box="[1002,1093,2003,2022]" pageId="16" pageNumber="179" value="2020-04-12">12.iv.2020</date>
,Y.Tasaku (
<collectionCode id="ED732BE8AC1AFF924BEFF8281382F82B" box="[1191,1245,2003,2022]" country="Ghana" lsid="urn:lsid:biocol.org:col:13924" name="Resource Management Support Center" pageId="16" pageNumber="179" type="Herbarium">KUM</collectionCode>
).
<emphasis id="B9166F3FAC1AFF924BA1F8281263F82B" bold="true" box="[1257,1340,2003,2022]" pageId="16" pageNumber="179">KඒǕඌΗǕ:</emphasis>
<specimenCount id="9D6478A4AC1AFF924A76F8281204F828" box="[1342,1371,1996,2022]" pageId="16" pageNumber="179" type="female">1♀</specimenCount>
, Nokono-shima, Fukuoka-shi, Fukuoka-ken,
<date id="FFDC95EDAC1AFF924B36F8151385F7CC" box="[1150,1242,2030,2049]" pageId="16" pageNumber="179" value="2018-04-21">21.iv.2018</date>
,
<collectionCode id="ED732BE8AC1AFF924BA8F81513B1F7CC" box="[1248,1262,2030,2049]" name="Tavera, Department of Geology and Geophysics" pageId="16" pageNumber="179">T</collectionCode>
.Hashizume (
<collectionCode id="ED732BE8AC1AFF924A15F81512CCF7CC" box="[1373,1427,2030,2049]" country="Ghana" lsid="urn:lsid:biocol.org:col:13924" name="Resource Management Support Center" pageId="16" pageNumber="179" type="Herbarium">KUM</collectionCode>
).
</paragraph>
</subSubSection>
<caption id="DF1DE3A5AC1BFF934FDEF8091267F7ED" ID-DOI="http://doi.org/10.5281/zenodo.5037352" ID-Zenodo-Dep="5037352" httpUri="https://zenodo.org/record/5037352/files/figure.png" pageId="17" pageNumber="180" startId="17.[150,181,2034,2053]" targetBox="[153,1427,191,1935]" targetPageId="17">
<paragraph id="8BDDB32DAC1BFF934FDEF8091267F7ED" blockId="17.[150,1436,2034,2080]" pageId="17" pageNumber="180">
Fig. 10.
<taxonomicName id="4C62C8AEAC1BFF934F94F80916C2F7C9" authority="Liu &amp; Ono &amp; Maruyama, 2021" authorityName="Liu &amp; Ono &amp; Maruyama" authorityYear="2021" box="[220,413,2034,2053]" class="Insecta" family="Staphylinidae" genus="Bryothinusa" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="180" phylum="Arthropoda" rank="species" species="constricta" status="sp. nov.">
<emphasis id="B9166F3FAC1BFF934F94F80916C2F7C9" box="[220,413,2034,2053]" italics="true" pageId="17" pageNumber="180">Bryothinusa constricta</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A225D244AC1BFF934EEAF80916BBF7C8" box="[418,484,2034,2053]" pageId="17" pageNumber="180" rank="species">sp. nov.</taxonomicNameLabel>
A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm.
</paragraph>
</caption>
<caption id="DF1DE3A5AC18FF904FDFF80A1237F7D3" ID-DOI="http://doi.org/10.5281/zenodo.5037354" ID-Zenodo-Dep="5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="18" pageNumber="181" startId="18.[151,182,2033,2052]" targetBox="[155,1436,191,1952]" targetPageId="18">
<paragraph id="8BDDB32DAC18FF904FDFF80A1237F7D3" blockId="18.[151,1436,2032,2078]" pageId="18" pageNumber="181">
Fig. 11.
<taxonomicName id="4C62C8AEAC18FF904F95F80B16E6F7CE" authority="Liu &amp; Ono &amp; Maruyama, 2021" authorityName="Liu &amp; Ono &amp; Maruyama" authorityYear="2021" box="[221,441,2032,2051]" class="Insecta" family="Staphylinidae" genus="Bryothinusa" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="181" phylum="Arthropoda" rank="species" species="hokkaidensis" status="sp. nov.">
<emphasis id="B9166F3FAC18FF904F95F80B16E6F7CE" box="[221,441,2032,2051]" italics="true" pageId="18" pageNumber="181">Bryothinusa hokkaidensis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A225D244AC18FF904EF6F80A155EF7C9" box="[446,513,2033,2052]" pageId="18" pageNumber="181" rank="species">sp. nov.</taxonomicNameLabel>
A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm.
</paragraph>
</caption>
<subSubSection id="C378E0A6AC19FF914FDFFF451592FA38" pageId="19" pageNumber="182" type="description">
<paragraph id="8BDDB32DAC19FF914FDFFF45160DFEB8" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
<emphasis id="B9166F3FAC19FF914FDFFF45167DFF18" bold="true" box="[151,290,190,213]" pageId="19" pageNumber="182">Description.</emphasis>
Body (
<figureCitation id="1359AFA8AC19FF914E3BFF4516E3FF18" box="[371,444,190,213]" captionStart="Fig" captionStartId="2.[151,182,1972,1991]" captionTargetBox="[189,1405,205,1944]" captionTargetId="figure-13@2.[175,1411,189,1947]" captionTargetPageId="2" captionText="Fig. 1. Habitus of Bryothinusa spp. A B. algarum Sawada, 1971; B B. minuta (Sawada, 1955); C B. tsutsuii (Sawada, 1955); D B. koreana Ahn &amp; Jeon, 2004; E B. yoshigoui sp. nov.; F B. japonica Liu, Ono &amp; Maruyama, 2020; G B. sakishimana Sawada, 1991; H B. constricta sp. nov.; I B. hokkaidensis sp. nov.; J B. fulvipennis sp. nov.; K B. itsuroi sp. nov.; L B. aikoae sp. nov.; M B. okinawana sp. nov.; N B. nigra sp. nov.; O B. nakanei (Sawada, 1955); P B. moriguchii sp. nov.; Q B. gangjinensis Ahn &amp; Jeon, 2004. Scale bars: 1.00 mm." figureDoi="http://doi.org/10.5281/zenodo.5037332" httpUri="https://zenodo.org/record/5037332/files/figure.png" pageId="19" pageNumber="182">Fig. 1I</figureCitation>
) small, about
<quantity id="4C9A1EC8AC19FF914D11FF451585FF18" box="[601,730,190,213]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="2.2" metricValueMin="1.8" pageId="19" pageNumber="182" unit="mm" value="2.0" valueMax="2.2" valueMin="1.8">1.82.2 mm</quantity>
in length, covered with minute setae. Abdomen almost dark brown and darker than head and pronotum; elytra lighter than pronotum; legs yellowish brown; antenna brown, similar coloured with mouthparts and legs, lightly paler than body colour.
</paragraph>
<paragraph id="8BDDB32DAC19FF914FFDFE85165AFDB8" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
Head evenly oval, wider than long, shorter than pronotum and elytra; eyes (
<figureCitation id="1359AFA8AC19FF914E80FE65154BFE78" box="[456,532,414,437]" captionStart="Fig" captionStartId="4.[151,182,1978,1997]" captionTargetBox="[196,1391,190,1955]" captionTargetId="figure-13@4.[193,1394,189,1958]" captionTargetPageId="4" captionText="Fig. 2. Compound eyes of Bryothinusa spp.A B. algarum Sawada, 1971; B B. minuta (Sawada, 1955); C B. tsutsuii (Sawada, 1955); D B. koreana Ahn &amp; Jeon, 2004; E B. yoshigoui sp. nov.; F B. japonica Liu, Ono &amp; Maruyama, 2020; G B. sakishimana Sawada, 1991; H B. constricta sp. nov.; I B. hokkaidensis sp. nov.; J B. fulvipennis sp. nov.; K B. itsuroi sp. nov.; L B. aikoae sp. nov.; M B. okinawana sp. nov.; N B. nigra sp. nov.; O B. nakanei (Sawada, 1955); P B. moriguchii sp. nov.; Q B. gangjinensis Ahn &amp; Jeon, 2004. Scale bars: 0.05 mm." figureDoi="http://doi.org/10.5281/zenodo.5037334" httpUri="https://zenodo.org/record/5037334/files/figure.png" pageId="19" pageNumber="182">Fig. 2I</figureCitation>
) large, setae present between about 2225 facets, slightly longer than tempora length; antenna brown, equals to total length of head and pronotum; segment I as long as II, almost 2.3 times longer than III, and as long as XI; antennal segments IIIIV subquadrate, VX almost same length and wider than long.
</paragraph>
<paragraph id="8BDDB32DAC19FF914FFDFD851500FBD8" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
Mouthparts (
<figureCitation id="1359AFA8AC19FF914E0BFD851695FD58" box="[323,458,638,661]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Figs 11AE</figureCitation>
) with distribution of labral setae as in
<figureCitation id="1359AFA8AC19FF914E4FFD6516DEFD78" box="[263,385,670,693]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Figure 11A</figureCitation>
, with large and thick spiniform seta at antero-laterally; mandibles (
<figureCitation id="1359AFA8AC19FF914E90FD45156BFD18" box="[472,564,702,725]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11B</figureCitation>
) elongate triangle, asymmetrical, right mandible with some median tooth and irregular; median tooth longer than 1/4 of mandibles; left mandible without tooth, blunt at apex; maxilla with apical margin of lacinia (
<figureCitation id="1359AFA8AC19FF914EECFCC5155BFC98" box="[420,516,830,853]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11C</figureCitation>
) at obtuse angle; at tip with two spines very small and not closely interdigitating; 23 setae on inner margin of lacinia; labial palpus (
<figureCitation id="1359AFA8AC19FF914FD7FC65165FFC78" box="[159,256,926,949]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11D</figureCitation>
) with most setae located basally, small ligula; mentum (
<figureCitation id="1359AFA8AC19FF914FB7FC451601FC18" box="[255,350,958,981]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11E</figureCitation>
) with antero-lateral angles prolonged into spinose processes; anterior margin deeply truncate; small “v seta” far away from apical setae.
</paragraph>
<paragraph id="8BDDB32DAC19FF914FFDFBE515EDFBF8" blockId="19.[151,761,190,1845]" box="[181,690,1054,1077]" pageId="19" pageNumber="182">Pronotum shorter than elytra, wider than head.</paragraph>
<paragraph id="8BDDB32DAC19FF914FFDFBC51624FB58" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">Elytra subquadrate but wider than long, slightly narrower than pronotum; hind wings present but very small, apparently flightless.</paragraph>
<paragraph id="8BDDB32DAC19FF914FFDFB6515DBFB38" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
Male abdominal sternite VIII (
<figureCitation id="1359AFA8AC19FF914EB2FB65150AFB78" box="[506,597,1182,1205]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11F</figureCitation>
) with posterior margin slightly serrate. Median lobe of aedeagus as in
<figureCitation id="1359AFA8AC19FF914FDFFB251668FB38" box="[151,311,1246,1269]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Figures 11H, I</figureCitation>
; paramere (
<figureCitation id="1359AFA8AC19FF914EF0FB25154FFB38" box="[440,528,1246,1269]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11J</figureCitation>
) elongate.
</paragraph>
<paragraph id="8BDDB32DAC19FF914FFDFB051790FAB8" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
Female abdominal sternite VIII (
<figureCitation id="1359AFA8AC19FF914D64FB0515CDFAD8" box="[556,658,1278,1301]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11G</figureCitation>
) slightly serrate similar male on posterior margin. Spermatheca (
<figureCitation id="1359AFA8AC19FF914FD7FAC5165DFA98" box="[159,258,1342,1365]" captionStart="Fig" captionStartId="18.[151,182,2033,2052]" captionTargetBox="[155,1436,191,1952]" captionTargetId="figure-13@18.[151,1436,189,1973]" captionTargetPageId="18" captionText="Fig. 11. Bryothinusa hokkaidensis sp. nov. A labrum; B mandible, dorsal view; C lacinia, dorsal view; D labium; E mentum; F male sternite VIII; G female sternite VIII; H median lobe, ventral view; I median lobe, lateral view; J paramere; K spermatheca. Scale bars: 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5037354" httpUri="https://zenodo.org/record/5037354/files/figure.png" pageId="19" pageNumber="182">Fig. 11K</figureCitation>
) simplified in shape, about 1 time coiled near base.
</paragraph>
<paragraph id="8BDDB32DAC19FF914FDFFA851592FA38" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
<emphasis id="B9166F3FAC19FF914FDFFA851619FA58" bold="true" box="[151,326,1406,1429]" pageId="19" pageNumber="182">Measurements.</emphasis>
BL 1.802.20, FBL 0.901.00, HL 0.25 0.30, HW 0.380.40, PL 0.330.35, PW 0.400.43, EL 0.350.38, EW 0.350.40, HTL 0.250.35, AL 0.600.68, AS-I 0.100.11, AS-II 0.090.10, AS-III 0.040.05.
</paragraph>
</subSubSection>
<subSubSection id="C378E0A6AC19FF914FDFFA0515A7F958" pageId="19" pageNumber="182" type="diagnosis">
<paragraph id="8BDDB32DAC19FF914FDFFA0515A7F958" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
<emphasis id="B9166F3FAC19FF914FDFFA0516D1F9D8" bold="true" box="[151,398,1534,1557]" pageId="19" pageNumber="182">Differential diagnosis.</emphasis>
In general appearance, the species is similar to
<taxonomicName id="4C62C8AEAC19FF914E54F9E516EEF9F8" authority="Liu &amp; Ono &amp; Maruyama, 2021" authorityName="Liu &amp; Ono &amp; Maruyama" authorityYear="2021" box="[284,433,1565,1589]" class="Insecta" family="Staphylinidae" genus="Bryothinusa" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="182" phylum="Arthropoda" rank="species" species="fulvipennis" status="sp. nov.">
<emphasis id="B9166F3FAC19FF914E54F9E516EEF9F8" box="[284,433,1565,1589]" italics="true" pageId="19" pageNumber="182">B. fulvipennis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A225D244AC19FF914EFEF9E51555F9F8" box="[438,522,1566,1589]" pageId="19" pageNumber="182" rank="species">sp. nov.</taxonomicNameLabel>
and
<taxonomicName id="4C62C8AEAC19FF914D74F9E515C0F9F8" authority="Liu &amp; Ono &amp; Maruyama, 2021" authorityName="Liu &amp; Ono &amp; Maruyama" authorityYear="2021" box="[572,671,1566,1589]" class="Insecta" family="Staphylinidae" genus="Bryothinusa" kingdom="Animalia" order="Coleoptera" pageId="19" pageNumber="182" phylum="Arthropoda" rank="species" species="itsuroi" status="sp. nov.">
<emphasis id="B9166F3FAC19FF914D74F9E515C0F9F8" box="[572,671,1566,1589]" italics="true" pageId="19" pageNumber="182">B. itsuroi</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A225D244AC19FF914DEDF9E515A7F9F8" box="[677,760,1566,1589]" pageId="19" pageNumber="182" rank="species">sp. nov.</taxonomicNameLabel>
but distinguished from them by the combination of the following character states: the body colour (especially head and pronotum colour) and the spermathecal shape.
</paragraph>
</subSubSection>
<subSubSection id="C378E0A6AC19FF914FDFF9651567F918" pageId="19" pageNumber="182" type="etymology">
<paragraph id="8BDDB32DAC19FF914FDFF9651567F918" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
<emphasis id="B9166F3FAC19FF914FDFF9651645F978" bold="true" box="[151,282,1694,1717]" pageId="19" pageNumber="182">Etymology.</emphasis>
Toponym. Named after
<collectingRegion id="49A67DCFAC19FF914D6EF96515CBF978" box="[550,660,1694,1717]" country="Japan" name="Hokkaido" pageId="19" pageNumber="182">Hokkaidô</collectingRegion>
, the
<typeStatus id="54D90D8FAC19FF914D81F96515A8F978" box="[713,759,1694,1717]" pageId="19" pageNumber="182">type</typeStatus>
locality of this new species; adjective.
</paragraph>
</subSubSection>
<subSubSection id="C378E0A6AC19FF914FDFF92515A6F938" box="[151,761,1758,1781]" pageId="19" pageNumber="182" type="distribution">
<paragraph id="8BDDB32DAC19FF914FDFF92515A6F938" blockId="19.[151,761,190,1845]" box="[151,761,1758,1781]" pageId="19" pageNumber="182">
<emphasis id="B9166F3FAC19FF914FDFF9251676F938" bold="true" box="[151,297,1758,1781]" pageId="19" pageNumber="182">Distribution.</emphasis>
<collectingCountry id="F375F3BDAC19FF914E66F9251634F938" box="[302,363,1758,1781]" name="Japan" pageId="19" pageNumber="182">Japan</collectingCountry>
(
<emphasis id="B9166F3FAC19FF914E30F92516B7F938" bold="true" box="[376,488,1758,1781]" pageId="19" pageNumber="182">
<collectingRegion id="49A67DCFAC19FF914E30F92516B7F938" box="[376,488,1758,1781]" country="Japan" name="Hokkaido" pageId="19" pageNumber="182">Hokkaidô</collectingRegion>
</emphasis>
;
<emphasis id="B9166F3FAC19FF914EBDF9251508F938" bold="true" box="[501,599,1758,1781]" pageId="19" pageNumber="182">Kyûshû:</emphasis>
Fukuoka-ken).
</paragraph>
</subSubSection>
<subSubSection id="C378E0A6AC19FF914FDFF90515C8F8F8" pageId="19" pageNumber="182" type="discussion">
<paragraph id="8BDDB32DAC19FF914FDFF90515C8F8F8" blockId="19.[151,761,190,1845]" pageId="19" pageNumber="182">
<emphasis id="B9166F3FAC19FF914FDFF9051657F8D8" bold="true" box="[151,264,1790,1813]" pageId="19" pageNumber="182">Remarks.</emphasis>
A distribution map of examined specimens of this species in
<collectingCountry id="F375F3BDAC19FF914E70F8E51629F8F8" box="[312,374,1822,1845]" name="Japan" pageId="19" pageNumber="182">Japan</collectingCountry>
is illustrated in
<figureCitation id="1359AFA8AC19FF914D6FF8E515CCF8F8" box="[551,659,1822,1845]" captionStart="Fig" captionStartId="35.[150,181,1388,1407]" captionTargetBox="[150,1434,191,1370]" captionTargetId="figure-236@35.[150,1435,191,1371]" captionTargetPageId="35" captionText="Fig. 22. Map of examined specimens of Bryothinusa spp. from Japan: white inverted triangle B. algarum Sawada, 1971; black inverted triangle B. minuta (Sawada, 1955); white star B. tsutsuii (Sawada, 1955); black star B. koreana Ahn &amp; Jeon, 2004; black square B. yoshigoui sp. nov.; white triangle B. japonica Liu, Ono &amp; Maruyama, 2020; black diamond B. sakishimana Sawada, 1991; white diamond B. constricta sp. nov.; white square B. hokkaidensis sp. nov.; black triangle B. fulvipennis sp. nov.; double circle B. itsuroi sp. nov.; arrow B. aikoae sp. nov.; black hart B. okinawana sp. nov.; circled dot B. nigra sp. nov.; white circle B. nakanei (Sawada, 1955); white hart B. moriguchii sp. nov.; black circle B. gangjinensis Ahn &amp; Jeon, 2004." figureDoi="http://doi.org/10.5281/zenodo.5037376" httpUri="https://zenodo.org/record/5037376/files/figure.png" pageId="19" pageNumber="182">Figure 22</figureCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>