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<mods:title id="67D675D43CC74FE1E0924F86EF50D178">Taxonomic study of the subgenus Tropidocheila of the lace bug genus Tingis (Hemiptera: Heteroptera: Tingidae) from Japan, with description of Tingis (Tropidocheila) yamaboushi sp. nov. collected from Cornus kousa (Cornaceae)</mods:title>
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<mods:affiliation id="EE70D1E91566B372EECC32C05C983911">Entomological Laboratory, Graduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, Fukuoka, Japan. kodokusignal @ gmail. com; https: // orcid. org / 0000 - 0002 - 2238 - 5015 &amp; Research Fellowship for Young Scientists (DC 1), Japan Society for the Promotion of Science, Tokyo, Japan</mods:affiliation>
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<bibRefCitation id="E228A6B1DF72FFDDFE9EFE8D89B9326E" author="Takeya, C." box="[278,460,347,372]" pageId="3" pageNumber="287" pagination="41 - 75" refId="ref7934" refString="Takeya, C. (1962) Taxonomic revision of the Tingidae of Japan, Korea, the Ryukyus and Formosa Part 1 (Hemiptera). Mushi, 36 (5), 41 - 75." type="journal article" year="1962">Takeya (1962: 62)</bibRefCitation>
(monograph); 
<bibRefCitation id="E228A6B1DF72FFDDFDEAFE8D8BF53269" author="Drake, C. J. &amp; Ruhoff, F. A." box="[610,896,347,372]" pageId="3" pageNumber="287" refId="ref6776" refString="Drake, C. J. &amp; Ruhoff, F. A. (1965) Lacebugs of the World. A Catalog (Hemiptera: Tingidae). United States National Museum Bulletin 243. Smithsonian Institution, Washington, VIII + 634 pp., 56 pls., frontispiece. https: // doi. org / 10.5479 / si. 03629236.243.1" type="book" year="1965">Drake &amp; Ruhoff (1965: 408)</bibRefCitation>
(catalogue); 
<bibRefCitation id="E228A6B1DF72FFDDFB8BFE8D8CDD3269" author="Jing, H. - L." box="[1027,1192,348,372]" pageId="3" pageNumber="287" pagination="42 - 50" refId="ref7240" refString="Jing, H. - L. (1981) Tingidae. In: Hsiao, T. - Y., Ren, S. - Z., Zheng, L. - Y., Jing, H. - L., Zou, H. - G. &amp; Liu, S. - L. (Eds.), A handbook for the determination of the Chinese Hemiptera-Heteroptera. Vol. 2. Science Press, Beijing, pp. 271 - 368, pls. 42 - 50. [in Chinese, English summary]" type="book chapter" year="1981">Jing (1981: 304)</bibRefCitation>
(monograph); 
<bibRefCitation id="E228A6B1DF72FFDDFAB6FE8D89AE328F" author="Miyamoto, S. &amp; Yasunaga, T." pageId="3" pageNumber="287" pagination="151 - 188" refId="ref7504" refString="Miyamoto, S. &amp; Yasunaga, T. (1989) Heteroptera. In: Hirashima, Y. (Ed.), A Check List of Japanese Insects. I. Laboratory of Entomology, Faculty of Agriculture, Kyushu University, Fukuoka, pp. 151 - 188. [in Japanese]" type="book chapter" year="1989">Miyamoto &amp; Yasunaga (1989: 168)</bibRefCitation>
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<bibRefCitation id="E228A6B1DF72FFDDFD2BFEAB8BB0328F" author="Pericart, J. &amp; Golub, V. B." box="[675,965,378,402]" pageId="3" pageNumber="287" pagination="3 - 78" refId="ref7692" refString="Pericart, J. &amp; Golub, V. B. (1996) Superfamily Tingoidea Laporte, 1832. In: Aukema, B. &amp; Rieger, C. (Eds.), Catalogue of the Heteroptera of the Palaearctic Region. Vol. 2, Cimicomorpha I. The Netherlands Entomological Society, Amsterdam, pp. 3 - 78." type="book chapter" year="1996">Péricart &amp; Golub (1996: 69)</bibRefCitation>
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<bibRefCitation id="E228A6B1DF72FFDDFB49FEAB894B32AC" author="Yamada, K. &amp; Tomokuni, M." pageId="3" pageNumber="287" pagination="2 - 13" refId="ref7972" refString="Yamada, K. &amp; Tomokuni, M. (2012) Family Tingidae Laporte, 1832. In: Ishikawa, T., Takai, M. &amp; Yasunaga, T. (Eds.), A field guide to Japanese bugs. Terrestrial heteropterans III. Zenkoku Noson Kyoiku Kyokai, Tokyo, pp. 180 - 213, pls. 2 - 13. [in Japanese]" type="book chapter" year="2012">Yamada &amp; Tomokuni (2012: 210)</bibRefCitation>
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Yano 
<emphasis id="B4CD0752DF72FFDDFD98FE4B8A4932AC" box="[528,572,409,433]" italics="true" pageId="3" pageNumber="287">et al</emphasis>
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; 
<materialsCitation id="36D1D11DDF72FFDDFE4FFDD28974315F" ID-GBIF-Occurrence="3015254321" accessionNumber="N 13416" collectingDate="2018-11-06" collectionCode="TPM" collectorName="Y. Obae" country="Japan" location="Syouzan" municipality="Tottori-shi" pageId="3" pageNumber="287" specimenCount="1" specimenCount-male="1" stateProvince="Tottori">
<collectingRegion id="447D15A2DF72FFDDFE4FFDD28A273100" box="[455,594,515,541]" country="Japan" name="Tottori" pageId="3" pageNumber="287">Tottori Pref.</collectingRegion>
, 
<collectingMunicipality id="6662413ADF72FFDDFDD2FDD28AA63100" box="[602,723,515,541]" pageId="3" pageNumber="287">Tottori-shi</collectingMunicipality>
, 
<location id="83668D9BDF72FFDDFD55FDD28B353100" LSID="urn:lsid:plazi:treatment:0E106A56DF72FFDDFF1FFF468BC33740:83668D9BDF72FFDDFD55FDD28B353100" box="[733,832,515,541]" country="Japan" municipality="Tottori-shi" name="Syouzan" pageId="3" pageNumber="287" stateProvince="Tottori">Syouzan</location>
, 3526'33.2&quot; 
<accessionNumber id="99EA46A3DF72FFDDFC4AFDD28C523100" box="[962,1063,515,542]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/N13416" pageId="3" pageNumber="287" type="EnaNcbi">N 13416</accessionNumber>
'03.8&quot;E, 
<date id="F207FD80DF72FFDDFB0BFDD28C9A3100" box="[1155,1263,515,541]" pageId="3" pageNumber="287" value="2018-11-06">
<collectingDate id="E2430468DF72FFDDFB0BFDD28C9A3100" box="[1155,1263,515,541]" pageId="3" pageNumber="287" value="2018-11-06">6.xi.2018</collectingDate>
</date>
, leg. 
<collectorName id="2B4CBE96DF72FFDDFAAFFDD58D0A3100" box="[1319,1407,515,542]" pageId="3" pageNumber="287">Y. Obae</collectorName>
(
<specimenCount id="90BF10C9DF72FFDDFA18FDD288DB315F" pageId="3" pageNumber="287" type="male">1 ♂</specimenCount>
, 
<collectionCode id="E0A84385DF72FFDDFF32FDF9888C315F" box="[186,249,552,578]" pageId="3" pageNumber="287">TPM</collectionCode>
)
</materialsCitation>
. 
<materialsCitation id="36D1D11DDF72FFDDFE84FDF68BA1315F" ID-GBIF-Occurrence="3015254302" box="[268,980,551,578]" collectingDate="1951-08-15" collectionCode="ELKU" collectorName="T. Esaki" country="Japan" county="Shikoku" location="Omogo" municipality="Iyo" pageId="3" pageNumber="287" specimenCount="1" specimenCount-female="1" stateProvince="Tottori">
<collectingCounty id="6F67A3CCDF72FFDDFE84FDF6891B315C" box="[268,366,551,577]" pageId="3" pageNumber="287">Shikoku</collectingCounty>
: 
<collectingMunicipality id="6662413ADF72FFDDFEF1FDF989D4315F" box="[377,417,552,578]" pageId="3" pageNumber="287">Iyo</collectingMunicipality>
, 
<location id="83668D9BDF72FFDDFE25FDF68A71315C" LSID="urn:lsid:plazi:treatment:0E106A56DF72FFDDFF1FFF468BC33740:83668D9BDF72FFDDFE25FDF68A71315C" box="[429,516,551,577]" country="Japan" county="Shikoku" municipality="Iyo" name="Omogo" pageId="3" pageNumber="287" stateProvince="Tottori">Omogo</location>
, 
<date id="F207FD80DF72FFDDFD87FDF68AEC315F" box="[527,665,551,578]" pageId="3" pageNumber="287" value="1951-08-15">
<collectingDate id="E2430468DF72FFDDFD87FDF68AEC315F" box="[527,665,551,578]" pageId="3" pageNumber="287" value="1951-08-15">15.viii.1951</collectingDate>
</date>
, leg. 
<collectorName id="2B4CBE96DF72FFDDFD5EFDF98B47315C" box="[726,818,551,578]" pageId="3" pageNumber="287">T. Esaki</collectorName>
(
<specimenCount id="90BF10C9DF72FFDDFCCCFDF68B18315F" box="[836,877,551,578]" pageId="3" pageNumber="287" type="female">1 ♀</specimenCount>
, 
<collectionCode id="E0A84385DF72FFDDFCF4FDF98BB9315F" box="[892,972,552,578]" pageId="3" pageNumber="287">ELKU</collectionCode>
)
</materialsCitation>
. 
<materialsCitation id="36D1D11DDF72FFDDFC57FDF689C8317B" ID-GBIF-Occurrence="3015254330" collectingDate="1935-07-26" collectionCode="ELKU" collectorName="S. Oishi" country="Japan" county="Kyushu" location="Sobo-san" municipality="Bungo" pageId="3" pageNumber="287" specimenCount="1" specimenCount-male="1" stateProvince="Tottori">
<collectingCounty id="6F67A3CCDF72FFDDFC57FDF68C4E315C" box="[991,1083,551,577]" pageId="3" pageNumber="287">Kyushu</collectingCounty>
: 
<collectingMunicipality id="6662413ADF72FFDDFBCDFDF98CE7315F" box="[1093,1170,552,578]" pageId="3" pageNumber="287">Bungo</collectingMunicipality>
, 
<location id="83668D9BDF72FFDDFB17FDF68D7F315C" LSID="urn:lsid:plazi:treatment:0E106A56DF72FFDDFF1FFF468BC33740:83668D9BDF72FFDDFB17FDF68D7F315C" box="[1183,1290,551,577]" country="Japan" county="Kyushu" municipality="Bungo" name="Sobo-san" pageId="3" pageNumber="287" stateProvince="Tottori">Sobo-san</location>
, 
<date id="F207FD80DF72FFDDFA9EFDF68DED315F" box="[1302,1432,551,578]" pageId="3" pageNumber="287" value="1935-07-26">
<collectingDate id="E2430468DF72FFDDFA9EFDF68DED315F" box="[1302,1432,551,578]" pageId="3" pageNumber="287" value="1935-07-26">26.vii.1935</collectingDate>
</date>
, leg. 
<collectorName id="2B4CBE96DF72FFDDFF4FFD9A89573175" box="[199,290,587,616]" pageId="3" pageNumber="287">S. Ôishi</collectorName>
(
<specimenCount id="90BF10C9DF72FFDDFEBCFD9A892B317B" box="[308,350,587,614]" pageId="3" pageNumber="287" type="male">1 ♂</specimenCount>
, 
<collectionCode id="E0A84385DF72FFDDFEE2FD9D89C2317B" box="[362,439,588,614]" pageId="3" pageNumber="287">ELKU</collectionCode>
)
</materialsCitation>
(
<figureCitation id="1E82C7C5DF72FFDDFE45FD9A8A67317B" box="[461,530,587,614]" captionStart="FIGURE 7" captionStartId="8.[151,250,1898,1922]" captionTargetBox="[151,1436,993,1872]" captionTargetId="figure-359@8.[151,1436,993,1875]" captionTargetPageId="8" captionText="FIGURE 7. Collection sites of three species of Tingis (Tropidocheila) from Japan used in present study. Blue icons = T. (Tr.) matsumurai; green icons = T. (Tr.) shaowuana; red icons = T. (Tr.) yamaboushi sp. nov. Circle = holotype locality; triangles = other localities." figureDoi="http://doi.org/10.5281/zenodo.4417720" httpUri="https://zenodo.org/record/4417720/files/figure.png" pageId="3" pageNumber="287">Fig. 7</figureCitation>
).
</paragraph>
<paragraph id="8606DB40DF72FFDDFF4FFDBE898B30B7" blockId="3.[151,1437,479,1118]" pageId="3" pageNumber="287">
<emphasis id="B4CD0752DF72FFDDFF4FFDBE89A73194" bold="true" box="[199,466,623,649]" pageId="3" pageNumber="287">Diagnostic characters.</emphasis>
Recognized among other species of 
<taxonomicName id="41B9A0C3DF72FFDDFCFDFDBE8C643194" ID-CoL="8GPV" authorityName="Fieber" authorityYear="1844" box="[885,1041,623,649]" class="Insecta" family="Tingidae" genus="Tingis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="285" phylum="Arthropoda" rank="subGenus" subGenus="Tropidocheila">
<emphasis id="B4CD0752DF72FFDDFCFDFDBE8C643194" box="[885,1041,623,649]" italics="true" pageId="3" pageNumber="287">Tropidocheila</emphasis>
</taxonomicName>
by a combination of the following characters: general color brown (
<figureCitation id="1E82C7C5DF72FFDDFD8DFD428A2E31B0" box="[517,603,659,685]" captionStart="FIGURE 1" captionStartId="4.[151,250,1917,1941]" captionTargetBox="[164,1426,425,1887]" captionTargetId="figure-151@4.[151,1436,417,1894]" captionTargetPageId="4" captionText="FIGURE 1. Three species of Tingis (Tropidocheila) from Japan, dorsal and lateral views: T. (Tr.) matsumurai (A); T. (Tr.) shaowuana (B); T. (Tr.) yamaboushi sp. nov., male (C–E) and female (F, G). Arrow marks antenna with segmental oligomery. Scale bar: 1.0 mm." figureDoi="http://doi.org/10.5281/zenodo.4417708" httpUri="https://zenodo.org/record/4417708/files/figure.png" pageId="3" pageNumber="287">Fig. 1B</figureCitation>
); pubescence on dorsum longer than diameter of compound eye; head with five spines (
<figureCitation id="1E82C7C5DF72FFDDFE96FD66890331CC" box="[286,374,695,721]" captionStart="FIGURE 4" captionStartId="7.[151,250,895,919]" captionTargetBox="[170,1414,186,866]" captionTargetId="figure-20@7.[165,1422,181,872]" captionTargetPageId="7" captionText="FIGURE 4. Heads and pronota of three species of Tingis (Tropidocheila) from Japan: T. (Tr.) matsumurai (A); T. (Tr.) shaowuana (B); T. (Tr.) yamaboushi sp. nov. (C). Scale bars: 0.2 mm." figureDoi="http://doi.org/10.5281/zenodo.4417714" httpUri="https://zenodo.org/record/4417714/files/figure.png" pageId="3" pageNumber="287">Fig. 4B</figureCitation>
); a pair of frontal spines reaching behind tip of clypeus; median spine reaching level far remote from bases of frontal spines; a pair of occipital spines reaching posterior margin of compound eye; rostrum reaching posterior margin of mesosternum; anterior margin of hood curved outward throughout its length; lateral carinae of pronotum distinct on pronotal disc; paranotum laminate, subhorizontal, with 3 rows of areolae at widest part; costal area of hemelytron with 3 rows of areolae at widest part, as wide as subcostal area at widest part of each; subcostal area with 3 rows of areolae at widest part; discoidal area with 6 rows of areolae at widest part; and sutural area with 8 rows of areolae at widest part.
</paragraph>
</subSubSection>
<subSubSection id="CEA388CBDF72FFDDFF4FFC628B3730D0" box="[199,834,947,974]" pageId="3" pageNumber="287" type="distribution">
<paragraph id="8606DB40DF72FFDDFF4FFC628B3730D0" blockId="3.[151,1437,479,1118]" box="[199,834,947,974]" pageId="3" pageNumber="287">
<emphasis id="B4CD0752DF72FFDDFF4FFC62891430D0" bold="true" box="[199,353,947,973]" pageId="3" pageNumber="287">Distribution.</emphasis>
<collectingCountry id="FEAE9BD0DF72FFDDFEEFFC6589D230D3" box="[359,423,948,974]" name="Japan" pageId="3" pageNumber="287">Japan</collectingCountry>
(Honshu, Shikoku, Kyushu); 
<collectingCountry id="FEAE9BD0DF72FFDDFD7FFC628B4B30D0" box="[759,830,947,973]" name="China" pageId="3" pageNumber="287">China</collectingCountry>
.
</paragraph>
</subSubSection>
<subSubSection id="CEA388CBDF72FFDDFF4FFC068BC33740" pageId="3" pageNumber="287" type="biology_ecology">
<paragraph id="8606DB40DF72FFDDFF4FFC068D383708" blockId="3.[151,1437,479,1118]" pageId="3" pageNumber="287">
<emphasis id="B4CD0752DF72FFDDFF4FFC06893830EC" bold="true" box="[199,333,983,1009]" pageId="3" pageNumber="287">Host plant.</emphasis>
In 
<collectingCountry id="FEAE9BD0DF72FFDDFEFFFC0989CF30EF" box="[375,442,984,1010]" name="Japan" pageId="3" pageNumber="287">Japan</collectingCountry>
, the host plant of 
<taxonomicName id="41B9A0C3DF72FFDDFD02FC098C7A30EC" authorityName="Drake &amp; Maa" authorityYear="1953" box="[650,1039,983,1010]" class="Insecta" family="Tingidae" genus="Tingis" kingdom="Animalia" order="Hemiptera" pageId="3" pageNumber="287" phylum="Arthropoda" rank="species" species="shaowuana" subGenus="Tropidocheila">
<emphasis id="B4CD0752DF72FFDDFD02FC098ABB30EC" box="[650,718,984,1009]" italics="true" pageId="3" pageNumber="287">Tingis</emphasis>
(
<emphasis id="B4CD0752DF72FFDDFD69FC068B0830EC" box="[737,893,983,1009]" italics="true" pageId="3" pageNumber="287">Tropidocheila</emphasis>
) 
<emphasis id="B4CD0752DF72FFDDFC07FC068C7A30EC" box="[911,1039,983,1009]" italics="true" pageId="3" pageNumber="287">shaowuana</emphasis>
</taxonomicName>
remains unknown (
<bibRefCitation id="E228A6B1DF72FFDDFB7AFC0689443708" author="Yamada, K. &amp; Tomokuni, M." pageId="3" pageNumber="287" pagination="2 - 13" refId="ref7972" refString="Yamada, K. &amp; Tomokuni, M. (2012) Family Tingidae Laporte, 1832. In: Ishikawa, T., Takai, M. &amp; Yasunaga, T. (Eds.), A field guide to Japanese bugs. Terrestrial heteropterans III. Zenkoku Noson Kyoiku Kyokai, Tokyo, pp. 180 - 213, pls. 2 - 13. [in Japanese]" type="book chapter" year="2012">Yamada &amp; Tomokuni 2012</bibRefCitation>
). In 
<collectingCountry id="FEAE9BD0DF72FFDDFEE9FC2A89D23708" box="[353,423,1019,1045]" name="China" pageId="3" pageNumber="287">China</collectingCountry>
, this lace bug has been collected from 
<taxonomicName id="41B9A0C3DF72FFDDFCD0FC2A8BC83708" box="[856,957,1019,1045]" class="Magnoliopsida" family="Scrophulariaceae" genus="Buddleja" kingdom="Plantae" order="Lamiales" pageId="3" pageNumber="287" phylum="Tracheophyta" rank="genus">
<emphasis id="B4CD0752DF72FFDDFCD0FC2A8BC83708" box="[856,957,1019,1045]" italics="true" pageId="3" pageNumber="287">Buddleja</emphasis>
</taxonomicName>
sp. (
<taxonomicName id="41B9A0C3DF72FFDDFC7BFC2A8CCD370B" authorityName="de Jussieu" authorityYear="1789" box="[1011,1208,1019,1046]" class="Magnoliopsida" family="Scrophulariaceae" kingdom="Plantae" order="Lamiales" pageId="3" pageNumber="287" phylum="Tracheophyta" rank="family">Scrophulariaceae</taxonomicName>
) (
<bibRefCitation id="E228A6B1DF72FFDDFB46FC2A8D343708" author="Jing, H. - L." box="[1230,1345,1019,1045]" pageId="3" pageNumber="287" pagination="42 - 50" refId="ref7240" refString="Jing, H. - L. (1981) Tingidae. In: Hsiao, T. - Y., Ren, S. - Z., Zheng, L. - Y., Jing, H. - L., Zou, H. - G. &amp; Liu, S. - L. (Eds.), A handbook for the determination of the Chinese Hemiptera-Heteroptera. Vol. 2. Science Press, Beijing, pp. 271 - 368, pls. 42 - 50. [in Chinese, English summary]" type="book chapter" year="1981">Jing 1981</bibRefCitation>
).
</paragraph>
<paragraph id="8606DB40DF72FFDDFF4FFBCE8BC33740" blockId="3.[151,1437,479,1118]" pageId="3" pageNumber="287">
<emphasis id="B4CD0752DF72FFDDFF4FFBCE89523724" bold="true" box="[199,295,1055,1081]" pageId="3" pageNumber="287">Biology.</emphasis>
In 
<collectingCountry id="FEAE9BD0DF72FFDDFEC3FBF189FB3727" box="[331,398,1056,1082]" name="Japan" pageId="3" pageNumber="287">Japan</collectingCountry>
, adults were collected in April, from June to August, and in November (
<bibRefCitation id="E228A6B1DF72FFDDFB22FBCE8D4B3724" author="Takeya, C." box="[1194,1342,1055,1081]" pageId="3" pageNumber="287" pagination="41 - 75" refId="ref7934" refString="Takeya, C. (1962) Taxonomic revision of the Tingidae of Japan, Korea, the Ryukyus and Formosa Part 1 (Hemiptera). Mushi, 36 (5), 41 - 75." type="journal article" year="1962">Takeya 1962</bibRefCitation>
; 
<bibRefCitation id="E228A6B1DF72FFDDFAC0FBF188833740" author="Yano, S. &amp; Takechi, L. &amp; Kikuhara, Y." pageId="3" pageNumber="287" pagination="23 - 52" refId="ref8045" refString="Yano, S., Takechi, L. &amp; Kikuhara, Y. (2013) Heteroptera of Ehime Prefecture, Shikoku, Japan (Insecta, Hemiptera) I. Bulletin of Omogo Mountain Museum, 5, 23 - 52. [in Japanese]" type="journal article" year="2013">
Yano 
<emphasis id="B4CD0752DF72FFDDFA00FBF188D83740" italics="true" pageId="3" pageNumber="287">et al</emphasis>
. 2013
</bibRefCitation>
; present study). Nymph and overwintering stage are unknown.
</paragraph>
</subSubSection>
</treatment>
</document>