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<document id="5F1BED36A93C5060AF9EA7747AE1AF9F" ID-DOI="10.37520/aemnp.2023.020" ID-ISSN="1804-6487" ID-Zenodo-Dep="10621601" ID-ZooBank="996CCDF0-04C6-47A5-AAAA-CCA93ECD3BC2" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1707167483656" checkinUser="felipe" docAuthor="Sugaya, Kazuki, Kakizoe, Showtaro, Ooka, Sohei, Tamura, Hisao &amp; Sone, Shinzaburo" docDate="2023" docId="03DEBE061203FF8AFED21FDCFD0DFD22" docLanguage="en" docName="ActaEntMusNatPra.63.2.323-340.pdf" docOrigin="Acta Entomologica Musei Nationalis Pragae 63 (2)" docSource="http://dx.doi.org/10.37520/aemnp.2023.020" docStyle="DocumentStyle:BA2C5C8540539042E893A65A182CEA85.8:ActaEntMusNatPra.2018-.journal_article" docStyleId="BA2C5C8540539042E893A65A182CEA85" docStyleName="ActaEntMusNatPra.2018-.journal_article" docStyleVersion="8" docTitle="Ryukyuaphaenops Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone 2023, gen. nov." docType="treatment" docVersion="2" lastPageNumber="329" masterDocId="FFE7C67E1200FF8CFFFF1C40FFD4FFD7" masterDocTitle="Discovery of a troglomorphic trechine beetle from the Ryukyu Archipelago, Southwestern Japan (Coleoptera: Carabidae: Trechinae)" masterLastPageNumber="340" masterPageNumber="323" pageNumber="326" updateTime="1707167942509" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-3.0">
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<mods:title id="7E1335B527CCDE37684E56F02E5C2044">Discovery of a troglomorphic trechine beetle from the Ryukyu Archipelago, Southwestern Japan (Coleoptera: Carabidae: Trechinae)</mods:title>
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<mods:namePart id="DC3F0068CCCCD3469B79DCCC15AD708B">Sugaya, Kazuki</mods:namePart>
<mods:affiliation id="8CE7330ADCA316ECD2EBEB2336EDF934">Bioindicator Co., Ltd., Nikkô-Kagurazaka Building, 18 Iwato-chô, Shinjuku, Tokyo, 162 - 0832 Japan</mods:affiliation>
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<mods:namePart id="EE5754DAF282DADFFE1E9CEEFA76425A">Kakizoe, Showtaro</mods:namePart>
<mods:affiliation id="E52AF243CFF8556485F1A060359E1FB2">Center for Collections, National Museum of Nature and Science, Amakubo</mods:affiliation>
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<mods:namePart id="5FF39DFC53D9BD3CB247D2DDE60CF6B9">Ooka, Sohei</mods:namePart>
<mods:affiliation id="E3C2D93A04F8F3CAF37174404C460824">Nanto Co., Ltd., Maekawa 1336, Tamagusuku, Nanjo, Okinawa, 901 - 0616 Japan</mods:affiliation>
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<mods:namePart id="159B756E9ED4129DE54F379F03D6F831">Tamura, Hisao</mods:namePart>
<mods:affiliation id="E1010B12B6446480864DE6B38ED00E4B">-</mods:affiliation>
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<mods:namePart id="D1DDAE7C2F0F775D47FF5B525813A252">Sone, Shinzaburo</mods:namePart>
<mods:affiliation id="E5BFFF3DF34C49A8981353E420DEC5ED">Japan Plant Protection Association, Nakazato</mods:affiliation>
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<mods:date id="544125F9223422482184ECAD1AD76CE9">2023</mods:date>
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<treatment id="03DEBE061203FF8AFED21FDCFD0DFD22" LSID="urn:lsid:plazi:treatment:03DEBE061203FF8AFED21FDCFD0DFD22" httpUri="http://treatment.plazi.org/id/03DEBE061203FF8AFED21FDCFD0DFD22" lastPageId="6" lastPageNumber="329" pageId="3" pageNumber="326">
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<paragraph id="8BC80F101203FF8FFED21FDCFDB6FC63" blockId="3.[301,610,924,948]" box="[301,610,924,948]" pageId="3" pageNumber="326">
<heading id="D080B87C1203FF8FFED21FDCFDB6FC63" box="[301,610,924,948]" centered="true" fontSize="10" level="2" pageId="3" pageNumber="326" reason="3">
<emphasis id="B903D3021203FF8FFED21FDCFDB6FC63" bold="true" box="[301,610,924,948]" pageId="3" pageNumber="326">
<taxonomicName id="4C7774931203FF8FFED21FDCFE2CFC63" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[301,504,924,948]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021203FF8FFED21FDCFE2CFC63" bold="true" box="[301,504,924,948]" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops</emphasis>
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<taxonomicNameLabel id="A2306E791203FF8FFE011FDDFDB6FC63" box="[510,610,925,948]" pageId="3" pageNumber="326" rank="genus">gen. nov.</taxonomicNameLabel>
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Japanese name:
<emphasis id="B903D3021203FF8FFEC51FF8FD31FC1A" box="[314,741,952,973]" italics="true" pageId="3" pageNumber="326">リπ̚キπ̚7̐±fl̽yṽŧõiì^̐ȇ</emphasis>
</paragraph>
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<paragraph id="8BC80F101203FF8FFF681FA7FE91FBC9" blockId="3.[151,760,999,2078]" pageId="3" pageNumber="326">
<emphasis id="B903D3021203FF8FFF681FA7FEFAFC29" bold="true" box="[151,302,999,1022]" pageId="3" pageNumber="326">
<typeStatus id="54CCB1B21203FF8FFF681FA7FF1BFC29" box="[151,207,999,1022]" pageId="3" pageNumber="326">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="4C7774931203FF8FFEC71FA7FD46FC29" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[312,658,999,1022]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="species" species="pulcherrimus" status="sp. nov.">
<emphasis id="B903D3021203FF8FFEC71FA7FD46FC29" box="[312,658,999,1022]" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops pulcherrimus</emphasis>
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<taxonomicNameLabel id="A2306E791203FF8FFD631FA7FD20FC29" box="[668,756,999,1022]" pageId="3" pageNumber="326" rank="species">sp. nov.</taxonomicNameLabel>
, here designated.
</paragraph>
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<subSubSection id="C36D5C9B1203FF8FFF681867FB18F8E2" pageId="3" pageNumber="326" type="diagnosis">
<paragraph id="8BC80F101203FF8FFF681867FB3DFEE2" blockId="3.[151,760,999,2078]" lastBlockId="3.[828,1437,190,2069]" pageId="3" pageNumber="326">
<emphasis id="B903D3021203FF8FFF681867FED8FBE9" bold="true" box="[151,268,1063,1086]" pageId="3" pageNumber="326">Diagnosis.</emphasis>
<taxonomicName id="4C7774931203FF8FFEEA1867FE03FBE9" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[277,471,1063,1086]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021203FF8FFEEA1867FE03FBE9" box="[277,471,1063,1086]" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops</emphasis>
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<taxonomicNameLabel id="A2306E791203FF8FFE1F1867FD91FBE9" box="[480,581,1063,1086]" pageId="3" pageNumber="326" rank="genus">gen. nov.</taxonomicNameLabel>
is distinguished from all known genera of
<taxonomicName id="4C7774931203FF8FFE441807FDCDFB89" box="[443,537,1095,1118]" class="Insecta" family="Carabidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="subTribe" subTribe="Trechina">Trechina</taxonomicName>
in the world by the following combination of characteristics: 1) aphaenopsian facies; 2) both dorsal and ventral body surfaces sparsely covered with microscopic hairs (
<figureCitation id="134C13951203FF8FFE0C18E7FD97FB69" box="[499,579,1191,1214]" captionStart="Figs 2024" captionStartId="10.[151,188,2006,2025]" captionTargetBox="[284,1303,652,1994]" captionTargetId="figure-13@10.[284,1304,652,1994]" captionTargetPageId="10" captionText="Figs 2024. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 20 basal part of right elytron, left latero-dorsal (a) and right latero-dorsal (b) views; 21 apical part of right elytron, right latero-dorsal view; 22 sternites 3 and 4; 23 sternites 57; 24 microscopic hairs and microsculpture in head (a), pronotum (b) and elytra (c).Abbreviations: ac apical carina; as apical striole; pb prehumeral border." figureDoi="http://doi.org/10.5281/zenodo.10621621" httpUri="https://zenodo.org/record/10621621/files/figure.png" pageId="3" pageNumber="326">Fig. 24</figureCitation>
); 3) right mandible seemingly bidentate though actually tridentate, due to middle tooth frequently absent (
<figureCitation id="134C13951203FF8FFE0818A7FDEAFB29" box="[503,574,1255,1278]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="3" pageNumber="326">Figs 4</figureCitation>
,
<figureCitation id="134C13951203FF8FFDB218A7FDBEFB29" box="[589,618,1255,1278]" captionStart="Figs 510" captionStartId="7.[151,188,2006,2025]" captionTargetBox="[196,1394,202,1955]" captionTargetId="figure-17@7.[155,1432,166,1969]" captionTargetPageId="7" captionText="Figs 510. Body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., paratypes (5 KSES00018; 610 KSES00033). 5 left (a) and right (b) mandibles, dorsal view; 6 labrum, dorsal view; 7 labium, ventral view; 8 ligula and labial palpus, ventral view; 9 left maxilla, ventral view; 10 left protarsi of male, dorsal (a) and internal (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621615" httpUri="https://zenodo.org/record/10621615/files/figure.png" pageId="3" pageNumber="326">5b</figureCitation>
); 4) mentum fused with submentum; 5) submentum multisetose; 6) labial palpomere 2 quadrisetose; 7) maxillary palpi glabrous though palpomere 2 unisetose subapically, palpomere 3 longer than palpomere 4; 8) prothorax barrel-shaped, with notopleural suture completely invisible from above (
<figureCitation id="134C13951203FF8FFF6019E7FED4FA69" box="[159,256,1447,1470]" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="3" pageNumber="326">Fig. 19b</figureCitation>
), front and hind angles effaced; 9) scutellum distinct; 10) elytral basal transverse furrow absent; 11) elytral lateral margins ciliated (
<figureCitation id="134C13951203FF8FFE2019A7FDEEFA29" box="[479,570,1511,1534]" captionStart="Figs 2024" captionStartId="10.[151,188,2006,2025]" captionTargetBox="[284,1303,652,1994]" captionTargetId="figure-13@10.[284,1304,652,1994]" captionTargetPageId="10" captionText="Figs 2024. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 20 basal part of right elytron, left latero-dorsal (a) and right latero-dorsal (b) views; 21 apical part of right elytron, right latero-dorsal view; 22 sternites 3 and 4; 23 sternites 57; 24 microscopic hairs and microsculpture in head (a), pronotum (b) and elytra (c).Abbreviations: ac apical carina; as apical striole; pb prehumeral border." figureDoi="http://doi.org/10.5281/zenodo.10621621" httpUri="https://zenodo.org/record/10621621/files/figure.png" pageId="3" pageNumber="326">Fig. 20a</figureCitation>
); 12) elytral bases distinctly depressed; 13) four pores of humeral set not ranged equidistantly and adjoining marginal gutter except for 3rd and 4th pores, 1st pore isolated anteriad from posterior pores, 2nd and 3rd pores close to each other, and 4th pore widely isolated posteriad from first three pores (
<figureCitation id="134C13951203FF8FFD541AC7FD27F949" box="[683,755,1671,1694]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="3" pageNumber="326">Figs 4</figureCitation>
,
<figureCitation id="134C13951203FF8FFF681AE7FF21F969" box="[151,245,1703,1726]" captionStart="Figs 2024" captionStartId="10.[151,188,2006,2025]" captionTargetBox="[284,1303,652,1994]" captionTargetId="figure-13@10.[284,1304,652,1994]" captionTargetPageId="10" captionText="Figs 2024. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 20 basal part of right elytron, left latero-dorsal (a) and right latero-dorsal (b) views; 21 apical part of right elytron, right latero-dorsal view; 22 sternites 3 and 4; 23 sternites 57; 24 microscopic hairs and microsculpture in head (a), pronotum (b) and elytra (c).Abbreviations: ac apical carina; as apical striole; pb prehumeral border." figureDoi="http://doi.org/10.5281/zenodo.10621621" httpUri="https://zenodo.org/record/10621621/files/figure.png" pageId="3" pageNumber="326">20a, 20b</figureCitation>
); each two pores of middle and apical sets distant from marginal gutter and isolated from other sets, close to each other in the former, while well distant from each other in the latter (
<figureCitation id="134C13951203FF8FFEDE1B47FEB2F8C9" box="[289,358,1799,1822]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="3" pageNumber="326">Figs 4</figureCitation>
,
<figureCitation id="134C13951203FF8FFE8C1B47FE44F8C9" box="[371,400,1799,1822]" captionStart="Figs 2024" captionStartId="10.[151,188,2006,2025]" captionTargetBox="[284,1303,652,1994]" captionTargetId="figure-13@10.[284,1304,652,1994]" captionTargetPageId="10" captionText="Figs 2024. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 20 basal part of right elytron, left latero-dorsal (a) and right latero-dorsal (b) views; 21 apical part of right elytron, right latero-dorsal view; 22 sternites 3 and 4; 23 sternites 57; 24 microscopic hairs and microsculpture in head (a), pronotum (b) and elytra (c).Abbreviations: ac apical carina; as apical striole; pb prehumeral border." figureDoi="http://doi.org/10.5281/zenodo.10621621" httpUri="https://zenodo.org/record/10621621/files/figure.png" pageId="3" pageNumber="326">21</figureCitation>
); 14) abdominal sternites 3 and 4 completely fused together; 15) protibiae wholly pubescent and not externally grooved; 16) protarsomere 1 not longer than protarsomeres 24 combined; 17) protarsomeres 1 and 2 of male dilated, inwardly denticulated at apices, and furnished beneath with adhesive appendages (
<figureCitation id="134C13951203FF8FFD331BE7FF60F809" captionStart="Figs 510" captionStartId="7.[151,188,2006,2025]" captionTargetBox="[196,1394,202,1955]" captionTargetId="figure-17@7.[155,1432,166,1969]" captionTargetPageId="7" captionText="Figs 510. Body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., paratypes (5 KSES00018; 610 KSES00033). 5 left (a) and right (b) mandibles, dorsal view; 6 labrum, dorsal view; 7 labium, ventral view; 8 ligula and labial palpus, ventral view; 9 left maxilla, ventral view; 10 left protarsi of male, dorsal (a) and internal (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621615" httpUri="https://zenodo.org/record/10621615/files/figure.png" pageId="3" pageNumber="326">Fig. 10</figureCitation>
); 18) median lobe of male genitalia with simple apical lobe neither reflexed dorsad nor bent ventrad; 19) internal sac without sclerotized teeth patches but with anisotopic copulatory piece; 20) styles elongate, with long setae at apices; and 21) female genitalia with distinct spermathecal complex not on common oviduct but on bursa copulatrix (
<figureCitation id="134C13951203FF8FFCBB1D5EFC47FEE2" box="[836,915,286,309]" captionStart="Figs 1115" captionStartId="8.[151,188,1977,1996]" captionTargetBox="[170,1420,218,1883]" captionTargetId="figure-13@8.[151,1436,195,1896]" captionTargetPageId="8" captionText="Figs 1115. Male (1113) and female (14, 15) genital organs of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov. 11, 13 holotype (KSES00004); 12, 14, 15 paratypes (12 KSES00005; 14 KSES00001; 15 KSES00037). 11 median lobe, left lateral (a), right lateral (b) and dorsal (c) views; 10 internal sac, left lateral (a), right lateral (b), front (c) and behind (d) views; 13 genital segment, reversed side view; 14 left gonocoxite, ventral view; 15 reproductive tract, left lateral view.Abbreviations: bc bursa copulatrix; co common oviduct; sp spermathecal complex; vg vagina." figureDoi="http://doi.org/10.5281/zenodo.10621617" httpUri="https://zenodo.org/record/10621617/files/figure.png" pageId="3" pageNumber="326">Fig. 15</figureCitation>
), gonocoxite 1 almost glabrous.
</paragraph>
<paragraph id="8BC80F101203FF8FFCA51D7EFB18F8E2" blockId="3.[828,1437,190,2069]" pageId="3" pageNumber="326">
Among all known genera of
<taxonomicName id="4C7774931203FF8FFB681D7EFB2CFE82" box="[1175,1272,318,341]" class="Insecta" family="Carabidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="subTribe" subTribe="Trechina">Trechina</taxonomicName>
,
<taxonomicName id="4C7774931203FF8FFAFB1D7EFCBAFEA2" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021203FF8FFAFB1D7EFCBAFEA2" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2306E791203FF8FFC881D1EFC0EFEA2" box="[887,986,350,373]" pageId="3" pageNumber="326" rank="genus">gen. nov.</taxonomicNameLabel>
shows a close resemblance to some Chinese aphaenopsian genera, in particularly
<taxonomicName id="4C7774931203FF8FFB091D3EFC6EFE62" authority="Ueno, 2002" authorityName="Ueno" authorityYear="2002" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFB091D3EFA48FE42" box="[1270,1436,382,405]" italics="true" pageId="3" pageNumber="326">Boreaphaenops</emphasis>
<bibRefCitation id="EFE672E11203FF8FFCC31DDEFC6EFE62" author="UENO S. - I." box="[828,954,414,437]" pageId="3" pageNumber="326" pagination="411 - 419" refId="ref14583" refString="UENO S. - I. 2002: A new genus and species of aphaenopsoid trechine beetle from the northern side of the Yangtze River. Elytra 30 (2): 411 - 419." type="journal article" year="2002">Uéno, 2002</bibRefCitation>
</taxonomicName>
and
<taxonomicName id="4C7774931203FF8FFC0B1DDEFAF4FE62" authority="Ueno, 2008" authorityName="Ueno" authorityYear="2008" box="[1012,1312,414,437]" class="Insecta" family="Carabidae" genus="Yanzaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFC0B1DDEFB4DFE62" box="[1012,1177,414,437]" italics="true" pageId="3" pageNumber="326">Yanzaphaenops</emphasis>
Uéno, 2008
</taxonomicName>
by sharing the following characteristics: 1) body surface sparsely covered with microscopic hairs; 2) elongated head with wide neck; 3) mentum fused with submentum; 4) pronotum barrel-shaped with incomplete lateral borders, basal area pre-pedunculated, front and hind angles degenerated; 5) elytral lateral margins ciliated; 6) humeral set of marginal umbilicate pores not aggregated; 7) protibiae wholly pubescent and not externally grooved; 8) protarsomere 1 not longer than protarsomeres 24 combined; 9) protarsomeres 1 and 2 modified in male; and 10) male genitalia with elongated aedeagus and styles, and anisotopic copulatory piece. However,
<taxonomicName id="4C7774931203FF8FFC551F5EFBB8FCE2" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[938,1132,798,821]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021203FF8FFC551F5EFBB8FCE2" box="[938,1132,798,821]" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2306E791203FF8FFB8D1F5EFB00FCE2" box="[1138,1236,798,821]" pageId="3" pageNumber="326" rank="genus">gen. nov.</taxonomicNameLabel>
can be clearly distinguished from the above two genera as follows: 1) labial palpomere 2 quadrisetose [bisetose in the two genera]; 2) maxillary palpomere 2 unisetose, palpomere 3 longer than palpomere 4 [palpomere 2 completely glabrous in the two genera; palpomere 3 shorter than palpomere
<quantity id="4C8FA2F51203FF8FFAFC1FFEFAFCFC02" box="[1283,1320,958,981]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" pageId="3" pageNumber="326" unit="in" value="4.0">4 in</quantity>
<taxonomicName id="4C7774931203FF8FFAD21FFEFCAEFC22" authorityName="Ueno" authorityYear="2002" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFAD21FFEFCAEFC22" italics="true" pageId="3" pageNumber="326">Boreaphaenops</emphasis>
</taxonomicName>
, almost as long as palpomere
<quantity id="4C8FA2F51203FF8FFB461F9EFB35FC22" box="[1209,1249,990,1013]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" pageId="3" pageNumber="326" unit="in" value="4.0">4 in</quantity>
<taxonomicName id="4C7774931203FF8FFB181F9EFA58FC22" authorityName="Ueno" authorityYear="2008" box="[1255,1420,990,1013]" class="Insecta" family="Carabidae" genus="Yanzaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFB181F9EFA58FC22" box="[1255,1420,990,1013]" italics="true" pageId="3" pageNumber="326">Yanzaphaenops</emphasis>
</taxonomicName>
]; 3) elytral setiferous dorsal pores on stria 3 composed of two pores [three pores in
<taxonomicName id="4C7774931203FF8FFBA7185EFB2AFBE2" authorityName="Ueno" authorityYear="2002" box="[1112,1278,1054,1077]" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFBA7185EFB2AFBE2" box="[1112,1278,1054,1077]" italics="true" pageId="3" pageNumber="326">Boreaphaenops</emphasis>
</taxonomicName>
, four pores in
<taxonomicName id="4C7774931203FF8FFCC3187EFC09FB82" authorityName="Ueno" authorityYear="2008" box="[828,989,1086,1109]" class="Insecta" family="Carabidae" genus="Yanzaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFCC3187EFC09FB82" box="[828,989,1086,1109]" italics="true" pageId="3" pageNumber="326">Yanzaphaenops</emphasis>
</taxonomicName>
]; 4) abdominal sternites 3 and 4 completely fused together [not fused in the two genera]; and 5) female genitalia with distinct spermathecal complex on bursa copulatrix [spermathecal complex unrecognizable in the two genera]. In addition to the above,
<taxonomicName id="4C7774931203FF8FFB2418FEFA49FB02" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[1243,1437,1214,1237]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021203FF8FFB2418FEFA49FB02" box="[1243,1437,1214,1237]" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2306E791203FF8FFCC3189EFC4FFB22" box="[828,923,1246,1269]" pageId="3" pageNumber="326" rank="genus">gen. nov.</taxonomicNameLabel>
also differs from
<taxonomicName id="4C7774931203FF8FFBAF189EFB22FB22" authorityName="Ueno" authorityYear="2002" box="[1104,1270,1246,1269]" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFBAF189EFB22FB22" box="[1104,1270,1246,1269]" italics="true" pageId="3" pageNumber="326">Boreaphaenops</emphasis>
</taxonomicName>
in the first three pores of the humeral set of marginal umbilicate pores not ranged equidistantly and the aedeagus not reflexed dorsad, and from
<taxonomicName id="4C7774931203FF8FFC57197EFB99FA82" authorityName="Ueno" authorityYear="2008" box="[936,1101,1342,1365]" class="Insecta" family="Carabidae" genus="Yanzaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFC57197EFB99FA82" box="[936,1101,1342,1365]" italics="true" pageId="3" pageNumber="326">Yanzaphaenops</emphasis>
</taxonomicName>
in the 1st pore of the humeral set of marginal umbilicate pores adjoining marginal gutter. Incidentally, in
<taxonomicName id="4C7774931203FF8FFC1D193EFB53FA42" authorityName="Ueno" authorityYear="2008" box="[994,1159,1406,1429]" class="Insecta" family="Carabidae" genus="Yanzaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021203FF8FFC1D193EFB53FA42" box="[994,1159,1406,1429]" italics="true" pageId="3" pageNumber="326">Yanzaphaenops</emphasis>
</taxonomicName>
, it was found that the first three pores of the humeral umbilicate set are not ranged equidistantly, and the 1st pore is removed inward from marginal gutter, based on our re-examination of the
<typeStatus id="54CCB1B21203FF8FFA91199EFA48FA22" box="[1390,1436,1502,1525]" pageId="3" pageNumber="326">type</typeStatus>
series, although it was originally described with the first three pores ranged equidistantly and adjoining marginal gutter (UḖNO 2005, 2010). In addition,
<taxonomicName id="4C7774931203FF8FFB0A1A7EFB86F9A2" authority="Tian &amp; He, 2020" authorityName="Tian &amp; He" authorityYear="2020" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="species" species="liyuani">
<emphasis id="B903D3021203FF8FFB0A1A7EFC53F9A2" italics="true" pageId="3" pageNumber="326">Boreaphaenops liyuani</emphasis>
<bibRefCitation id="EFE672E11203FF8FFC6E1A1EFB86F9A2" author="TIAN M. &amp; HE L." box="[913,1106,1630,1653]" pageId="3" pageNumber="326" pagination="61 - 91" refId="ref14038" refString="TIAN M. &amp; HE L. 2020: A contribution to the knowledge of cavernicolous ground beetles from Sichuan Province, southwestern China (Coleoptera, Carabidae, Trechini, Platynini). ZooKeys 1008: 61 - 91." type="journal article" year="2020">Tian &amp; He, 2020</bibRefCitation>
</taxonomicName>
significantly differs from the
<typeStatus id="54CCB1B21203FF8FFCC31A3EFCBEF942" box="[828,874,1662,1685]" pageId="3" pageNumber="326">type</typeStatus>
species,
<taxonomicName id="4C7774931203FF8FFC311A3EFB00F942" authority="Ueno, 2002" authorityName="Ueno" authorityYear="2002" box="[974,1236,1662,1685]" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="species" species="angustus">
<emphasis id="B903D3021203FF8FFC311A3EFC0AF942" box="[974,990,1662,1685]" italics="true" pageId="3" pageNumber="326">B</emphasis>
.
<emphasis id="B903D3021203FF8FFC121A3EFB98F942" box="[1005,1100,1662,1685]" italics="true" pageId="3" pageNumber="326">angustus</emphasis>
<bibRefCitation id="EFE672E11203FF8FFBAB1A3EFB00F942" author="UENO S. - I." box="[1108,1236,1662,1685]" pageId="3" pageNumber="326" pagination="411 - 419" refId="ref14583" refString="UENO S. - I. 2002: A new genus and species of aphaenopsoid trechine beetle from the northern side of the Yangtze River. Elytra 30 (2): 411 - 419." type="journal article" year="2002">Uéno, 2002</bibRefCitation>
</taxonomicName>
, in the generically important characteristics, including the four pores of the humeral set of marginal umbilicate pores ranged equidistantly, and elytral lateral borders not ciliated (
<bibRefCitation id="EFE672E11203FF8FFADA1A9EFCA2F8C2" author="TIAN M. &amp; HE L." pageId="3" pageNumber="326" pagination="61 - 91" refId="ref14038" refString="TIAN M. &amp; HE L. 2020: A contribution to the knowledge of cavernicolous ground beetles from Sichuan Province, southwestern China (Coleoptera, Carabidae, Trechini, Platynini). ZooKeys 1008: 61 - 91." type="journal article" year="2020">TIAN &amp; HE 2020</bibRefCitation>
). Therefore,
<taxonomicName id="4C7774931203FF8FFBF91ABEFB1DF8C2" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[1030,1225,1790,1813]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021203FF8FFBF91ABEFB1DF8C2" box="[1030,1225,1790,1813]" italics="true" pageId="3" pageNumber="326">Ryukyuaphaenops</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2306E791203FF8FFB2D1ABEFAECF8C2" box="[1234,1336,1790,1813]" pageId="3" pageNumber="326" rank="genus">gen. nov.</taxonomicNameLabel>
is easily distinguished from
<taxonomicName id="4C7774931203FF8FFBF41B5EFBA7F8E2" authorityName="Tian &amp; He" authorityYear="2020" box="[1035,1139,1822,1845]" class="Insecta" family="Carabidae" genus="Boreaphaenops" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="species" species="liyuani">
<emphasis id="B903D3021203FF8FFBF41B5EFBCFF8E2" box="[1035,1051,1822,1845]" italics="true" pageId="3" pageNumber="326">B</emphasis>
.
<emphasis id="B903D3021203FF8FFBD61B5EFBA7F8E2" box="[1065,1139,1822,1845]" italics="true" pageId="3" pageNumber="326">liyuani</emphasis>
</taxonomicName>
as well.
</paragraph>
</subSubSection>
<subSubSection id="C36D5C9B1203FF8AFCC31B7EFED1FF22" lastPageId="6" lastPageNumber="329" pageId="3" pageNumber="326" type="description">
<paragraph id="8BC80F101203FF8FFCC31B7EFAFFF842" blockId="3.[828,1437,190,2069]" pageId="3" pageNumber="326">
<emphasis id="B903D3021203FF8FFCC31B7EFC13F882" bold="true" box="[828,967,1854,1877]" pageId="3" pageNumber="326">Description.</emphasis>
Medium-sized
<taxonomicName id="4C7774931203FF8FFB901B7EFB19F882" box="[1135,1229,1854,1877]" class="Insecta" family="Carabidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="3" pageNumber="326" phylum="Arthropoda" rank="subTribe" subTribe="Trechina">Trechina</taxonomicName>
with aphaenopsian facies; apterous and depigmented; body surface sparsely covered with very minute microscopical hairs.
</paragraph>
<paragraph id="8BC80F101203FF89FCA51BDEFD9BFCE2" blockId="3.[828,1437,190,2069]" lastBlockId="5.[151,767,190,2069]" lastPageId="5" lastPageNumber="328" pageId="3" pageNumber="326">
Head elongated subquadrate, gradually narrowing posteriad with wide neck; dorsum feebly convex; frontal furrows absent posteriorly; two pairs of supraorbital pores present; ventral surface of genae (
<figureCitation id="134C13951203FF8FFBC41BBEFB13F7C2" box="[1083,1223,2046,2069]" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="3" pageNumber="326">Figs 18b, 18c</figureCitation>
) bearing pair of long hairs before neck constriction; eyes completely missing; clypeus quadrisetose. Labrum sextisetose. Mandibles long and slender, falcate. Right mandible visually bidentate to tridentate, composed of well-developed premolar and wide retinaculum; retinaculum bidentate, though proximal tooth frequently missing while distal one always developed. Left mandible bidentate, with two partially overlapping undeveloped teeth on dorsal and ventral sides.
<collectingRegion id="49B3C1F21205FF89FDB51DDEFD76FE62" box="[586,674,414,437]" country="Myanmar" name="Chin" pageId="5" pageNumber="328">Mentum</collectingRegion>
bisetose and fused with multisetose submentum; ligula trapezoidal in apical part, with two long setae in middle of truncated portion, and three lateral setae on each oblique part; paraglossae elongate, moderately arcuate inwardly, extending much beyond ligula, with minute hairs along inner sides; palps elongate and glabrous, but with labial palpomere 2 quadrisetose, two on inner and outer margins, respectively, and with maxillary palpomere 2 unisetose on inner margin subapically; penultimate palpomere longer than apical one in labium and maxillae, respectively.Antennae exceedingly elongated filiform, longer in male than female, wholly pubescent except on basal half of segment 1.
</paragraph>
<caption id="DF085F981204FF88FF681BBFFB36F7C5" ID-DOI="http://doi.org/10.5281/zenodo.10621611" ID-Zenodo-Dep="10621611" box="[151,1250,2047,2066]" httpUri="https://zenodo.org/record/10621611/files/figure.png" pageId="4" pageNumber="327" startId="4.[151,182,2047,2066]" targetBox="[151,1436,187,2001]" targetPageId="4" targetType="figure">
<paragraph id="8BC80F101204FF88FF681BBFFB36F7C5" blockId="4.[151,1250,2047,2066]" box="[151,1250,2047,2066]" pageId="4" pageNumber="327">
Fig. 3. Habitus of
<taxonomicName id="4C7774931204FF88FECD1BBFFD91F7C5" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[306,581,2047,2066]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="327" phylum="Arthropoda" rank="species" species="pulcherrimus">
<emphasis id="B903D3021204FF88FECD1BBFFD91F7C5" box="[306,581,2047,2066]" italics="true" pageId="4" pageNumber="327">Ryukyuaphaenops pulcherrimus</emphasis>
</taxonomicName>
gen. &amp; sp. nov., holotype male (KSES00004), dorsal (a) and lateral (b) views.
</paragraph>
</caption>
<paragraph id="8BC80F101205FF89FF4A1F7EFE3CFA82" blockId="5.[151,767,190,2069]" pageId="5" pageNumber="328">
Prothorax hardly tumid laterad. Pronotum barrel-shaped; dorsum moderately convex; median line distinct, not reaching apical nor basal margins; apical transverse impression distinct though superficial, with longitudinal fine wrinkles; basal transverse impression (
<figureCitation id="134C13951205FF89FD8C1FFEFD68FC02" box="[627,700,958,981]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="5" pageNumber="328">Figs 4</figureCitation>
,
<figureCitation id="134C13951205FF89FD341FFEFF6BFC22" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="5" pageNumber="328">19a, 19b</figureCitation>
) distinct, not continuous and merging on each side into basal fovea; sides (
<figureCitation id="134C13951205FF89FE9C1FBEFE22FBC2" box="[355,502,1022,1045]" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="5" pageNumber="328">Figs 19a, 19b</figureCitation>
) not ciliated, incompletely and very finely bordered, and deflexed ventrad before anterior latero-marginal pore; marginal gutters vestigial; both apical and basal margins not bordered entirely; basal margin (
<figureCitation id="134C13951205FF89FF0B183EFE9DFB42" box="[244,329,1150,1173]" captionStart="Fig" captionStartId="4.[151,182,2047,2066]" captionTargetBox="[151,1436,187,2001]" captionTargetId="figure-13@4.[151,1436,187,2001]" captionTargetPageId="4" captionText="Fig. 3. Habitus of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype male (KSES00004), dorsal (a) and lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621611" httpUri="https://zenodo.org/record/10621611/files/figure.png" pageId="5" pageNumber="328">Figs 3a</figureCitation>
,
<figureCitation id="134C13951205FF89FEA7183EFEB3FB42" box="[344,359,1150,1173]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="5" pageNumber="328">4</figureCitation>
,
<figureCitation id="134C13951205FF89FE89183EFE03FB42" box="[374,471,1150,1173]" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="5" pageNumber="328">19a, 19b</figureCitation>
) almost straight or feebly emarginated in middle, obliquely rounded at just inside of hind angles; front angles effaced; hind angles (
<figureCitation id="134C13951205FF89FD6C18FEFF14FB22" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="5" pageNumber="328">Figs 19a, 19b</figureCitation>
) never produced, subrectangular or entirely rounded. Propleura (
<figureCitation id="134C13951205FF89FEF218BEFE85FAC2" box="[269,337,1278,1301]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="5" pageNumber="328">Figs 4</figureCitation>
,
<figureCitation id="134C13951205FF89FEA318BEFE6CFAC2" box="[348,440,1278,1301]" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="5" pageNumber="328">19a, 19b</figureCitation>
) almost invisible from above; notopleural sutures (
<figureCitation id="134C13951205FF89FE92195EFE64FAE2" box="[365,432,1310,1333]" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="5" pageNumber="328">Figs 4</figureCitation>
,
<figureCitation id="134C13951205FF89FE45195EFDC0FAE2" box="[442,532,1310,1333]" captionStart="Figs 1619" captionStartId="9.[151,188,2008,2027]" captionTargetBox="[216,1369,529,1995]" captionTargetId="figure-17@9.[216,1369,529,1996]" captionTargetPageId="9" captionText="Figs 1619. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 16 labrum; 17 labium; 18 head; 19 prothorax. a dorsal view; b dorsolateral view; c ventral view. Abbreviations: ha hind angle; ld lateral declivity; nps notopleural suture; pp propleura; sb side border." figureDoi="http://doi.org/10.5281/zenodo.10621619" httpUri="https://zenodo.org/record/10621619/files/figure.png" pageId="5" pageNumber="328">19a, 19b</figureCitation>
) completely invisible from above. Scutellum distinct.
</paragraph>
<paragraph id="8BC80F101205FF89FF4A191EFEB7F7C2" blockId="5.[151,767,190,2069]" pageId="5" pageNumber="328">
Elytra fused together, elongated pyriform; basal peduncle without transverse furrow; dorsum convex though basal area longitudinally and obliquely depressed except for scutellar area; sides narrowly bordered throughout and very finely ciliated; marginal gutters distinct; stria 1 (
<figureCitation id="134C13951205FF89FD37199EFF72F9C2" captionStart="Fig" captionStartId="5.[828,859,2025,2044]" captionTargetBox="[828,1428,992,2005]" captionTargetId="figure-704@5.[825,1438,989,2019]" captionTargetPageId="5" captionText="Fig. 4. Chaetotaxy of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views." figureDoi="http://doi.org/10.5281/zenodo.10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="5" pageNumber="328">Figs 4</figureCitation>
,
<figureCitation id="134C13951205FF89FF4F19BEFF0EF9C2" box="[176,218,1534,1557]" captionStart="Figs 2024" captionStartId="10.[151,188,2006,2025]" captionTargetBox="[284,1303,652,1994]" captionTargetId="figure-13@10.[284,1304,652,1994]" captionTargetPageId="10" captionText="Figs 2024. SEM photos of body parts of Ryukyuaphaenops pulcherrimus gen. &amp; sp. nov., holotype (KSES00004). 20 basal part of right elytron, left latero-dorsal (a) and right latero-dorsal (b) views; 21 apical part of right elytron, right latero-dorsal view; 22 sternites 3 and 4; 23 sternites 57; 24 microscopic hairs and microsculpture in head (a), pronotum (b) and elytra (c).Abbreviations: ac apical carina; as apical striole; pb prehumeral border." figureDoi="http://doi.org/10.5281/zenodo.10621621" httpUri="https://zenodo.org/record/10621621/files/figure.png" pageId="5" pageNumber="328">20b</figureCitation>
) adjoining elytral suture except for basal area; basal pore present; two setiferous dorsal pores present on stria 3; preapical pore located at apical anastomosis of striae 2 and 3; two apical pores present; marginal umbilicate pores not aggregated; four pores of humeral set of marginal umbilicate pores not ranged equidistantly and adjoining marginal gutter except for 3rd and 4th pores, 1st pore isolated anteriad from posterior two pores, 2nd and 3rd pores close to each other, and 4th pore widely isolated posteriad from first three pores; two pores of middle set of marginal umbilicate pores close to each other, both isolated from humeral set and widely distant from marginal gutter, 1st pore more removed inward from marginal gutter than the 2nd pore; two pores of apical set of marginal umbilicate pores well distant from each other, isolated from middle set, 1st pore widely distant from marginal gutter while 2nd pore not so distant.
</paragraph>
<paragraph id="8BC80F101205FF89FCA51CFEFA82FEE2" blockId="5.[828,1437,190,949]" pageId="5" pageNumber="328">Abdominal sternites 3 and 4 completely fused together, suture between them completely absent; sternites 46 each bearing pair of paramedian setae near posterior margin; sternite 7 bisetose in male, quadrisetose in female.</paragraph>
<paragraph id="8BC80F101205FF89FCA51D7EFACBFE02" blockId="5.[828,1437,190,949]" pageId="5" pageNumber="328">Legs exceedingly long; protibiae wholly pubescent and not externally grooved; tarsomeres 4 with long hyaline ventral apophysis in pro- and mesotarsi; in male,protarsomeres 1 and 2 dilated, inwardly denticulate at apices, and furnished beneath with adhesive appendages.</paragraph>
<paragraph id="8BC80F101205FF89FCA51D9EFA83FD02" blockId="5.[828,1437,190,949]" pageId="5" pageNumber="328">Male genital organ small and slightly sclerotized; median lobe elongated tubular, gradually tapered toward apex; apical lobe almost straight, with simple extremity; sagittal aileron distinct. Internal sac without sclerotized teeth patches; copulatory piece anisotopic and spatulate. Styles elongate; left style obviously more elongate than right one, each usually bearing four long setae at apex. Genital segment devoid of distinct basal peduncle.</paragraph>
<paragraph id="8BC80F101205FF8AFCA51E9EFED1FF22" blockId="5.[828,1437,190,949]" lastBlockId="6.[151,760,190,757]" lastPageId="6" lastPageNumber="329" pageId="5" pageNumber="328">Female genital organ with distinct spermathecal complex on dorsal surface of bursa copulatrix near its junction with vagina. Gonocoxite 1 short, pick-shaped and almost glabrous, with a few short apical fringe setae (afs) along apical margin ventrally, two elongated apico-medial setae (ams) on apico-internal corner dorsally; gonocoxite 2 subconical, with two long dorsal ensiform setae (des) proximally, two apical nematiform setae (ans) preapico- -ventrally.</paragraph>
</subSubSection>
<caption id="DF085F981205FF89FCC31BA9FA85F7C1" ID-DOI="http://doi.org/10.5281/zenodo.10621613" ID-Zenodo-Dep="10621613" httpUri="https://zenodo.org/record/10621613/files/figure.png" pageId="5" pageNumber="328" startId="5.[828,859,2025,2044]" targetBox="[828,1428,992,2005]" targetPageId="5" targetType="figure">
<paragraph id="8BC80F101205FF89FCC31BA9FA85F7C1" blockId="5.[828,1437,2025,2070]" pageId="5" pageNumber="328">
Fig. 4. Chaetotaxy of
<taxonomicName id="4C7774931205FF89FC051BA9FAD9F82B" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[1018,1293,2025,2044]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="5" pageNumber="328" phylum="Arthropoda" rank="species" species="pulcherrimus">
<emphasis id="B903D3021205FF89FC051BA9FAD9F82B" box="[1018,1293,2025,2044]" italics="true" pageId="5" pageNumber="328">Ryukyuaphaenops pulcherrimus</emphasis>
</taxonomicName>
gen. &amp; sp. nov., holotype (KSES00004), dorsal (a) and dorso-lateral (b) views.
</paragraph>
</caption>
<subSubSection id="C36D5C9B1206FF8AFF681CBEFD44FE02" pageId="6" pageNumber="329" type="etymology">
<paragraph id="8BC80F101206FF8AFF681CBEFD44FE02" blockId="6.[151,760,190,757]" pageId="6" pageNumber="329">
<emphasis id="B903D3021206FF8AFF681CBEFEC0FEC2" bold="true" box="[151,276,254,277]" pageId="6" pageNumber="329">Etymology.</emphasis>
A combination of “Ryukyu” and “
<taxonomicName id="4C7774931206FF8AFD8D1CBEFD32FEC2" authorityName="Bonvouioir" authorityYear="1862" box="[626,742,254,277]" class="Insecta" family="Carabidae" genus="Aphaenops" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="329" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021206FF8AFD8D1CBEFD32FEC2" box="[626,742,254,277]" italics="true" pageId="6" pageNumber="329">Aphaenops</emphasis>
</taxonomicName>
”. “Ryukyu [
<emphasis id="B903D3021206FF8AFEF81D5CFEE9FEE2" box="[263,317,284,309]" italics="true" pageId="6" pageNumber="329">ñṉ</emphasis>
]” refers to the range of the new trechine beetle.
<taxonomicName id="4C7774931206FF8AFF1B1D7EFDCFFE82" authority="Bonvouioir, 1862" authorityName="Bonvouioir" authorityYear="1862" box="[228,539,318,341]" class="Insecta" family="Carabidae" genus="Aphaenops" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="329" phylum="Arthropoda" rank="genus">
<emphasis id="B903D3021206FF8AFF1B1D7EFE8FFE82" box="[228,347,318,341]" italics="true" pageId="6" pageNumber="329">Aphaenops</emphasis>
Bonvouioir, 1862
</taxonomicName>
is a generic name of the famous Pyrenean trechine beetle with a strongly developed troglomorphism and is also traditionally used as root of the generic name of other strongly troglomorphic (aphaenopsian) trechines. Gender is masculine.
</paragraph>
</subSubSection>
<subSubSection id="C36D5C9B1206FF8AFF681D9EFD0DFD22" pageId="6" pageNumber="329" type="discussion">
<paragraph id="8BC80F101206FF8AFF681D9EFD0DFD22" blockId="6.[151,760,190,757]" pageId="6" pageNumber="329">
<emphasis id="B903D3021206FF8AFF681D9EFED2FE22" bold="true" box="[151,262,478,501]" pageId="6" pageNumber="329">Remarks.</emphasis>
Though the direction of apical striole has been traditionally considered important to the trechine taxonomy, we do not use this character in generic diagnosis of
<taxonomicName id="4C7774931206FF8AFF681E7EFE80FD82" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[151,340,574,597]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="329" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021206FF8AFF681E7EFE80FD82" box="[151,340,574,597]" italics="true" pageId="6" pageNumber="329">Ryukyuaphaenops</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2306E791206FF8AFEA61E7EFE6CFD82" box="[345,440,574,597]" pageId="6" pageNumber="329" rank="genus">gen. nov.</taxonomicNameLabel>
, because the character has been recently reported to be unstable in several genera and/or species (UḖNO 1999,
<bibRefCitation id="EFE672E11206FF8AFE831E3EFD4EFD42" author="BELOUSOV I. A. &amp; KABAK I. I." box="[380,666,638,661]" pageId="6" pageNumber="329" pagination="15 - 86" refId="ref12221" refString="BELOUSOV I. A. &amp; KABAK I. I. 2003: New Trechini from China (Coleoptera, Carabidae). Tethys Entomological Research 8: 15 - 86." type="journal article" year="2003">BELOUSOV &amp; KABAK 2003</bibRefCitation>
). In fact, the character state of the
<typeStatus id="54CCB1B21206FF8AFE671EDEFE12FD62" box="[408,454,670,693]" pageId="6" pageNumber="329">type</typeStatus>
species of
<taxonomicName id="4C7774931206FF8AFDC81EDEFD2CFD62" authority="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone, 2023" authorityName="Sugaya &amp; Kakizoe &amp; Ooka &amp; Tamura &amp; Sone" authorityYear="2023" box="[567,760,670,693]" class="Insecta" family="Carabidae" genus="Ryukyuaphaenops" kingdom="Animalia" order="Coleoptera" pageId="6" pageNumber="329" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B903D3021206FF8AFDC81EDEFD2CFD62" box="[567,760,670,693]" italics="true" pageId="6" pageNumber="329">Ryukyuaphaenops</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A2306E791206FF8AFF681EFEFF2DFD02" box="[151,249,702,725]" pageId="6" pageNumber="329" rank="genus">gen. nov.</taxonomicNameLabel>
is unstable, and in many cases, the direction of apical striole is undetermined, due to its degeneration.
</paragraph>
</subSubSection>
</treatment>
</document>