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<mods:title id="6370BE3328229FCA3E203FCDCA272B64">Additional records and descriptions of the ant-mimetic plant bug genus Pilophorus from Thailand (Hemiptera: Heteroptera: Miridae: Phylinae: Pilophorini)</mods:title>
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<mods:namePart id="7173BF1410C2A948AEA87D9ED738B0F0">Yamada, Kazutaka</mods:namePart>
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<subSubSection id="B3D3650CFFE2FFC5BBC4FE6D85B1FE37" box="[151,436,475,501]" pageId="2" pageNumber="3" type="nomenclature">
<paragraph id="FB763687FFE2FFC5BBC4FE6D85B1FE37" blockId="2.[151,436,475,501]" box="[151,436,475,501]" pageId="2" pageNumber="3">
<heading id="A03E81EBFFE2FFC5BBC4FE6D85B1FE37" bold="true" box="[151,436,475,501]" fontSize="10" level="2" pageId="2" pageNumber="3" reason="2">
<emphasis id="C9BDEA95FFE2FFC5BBC4FE6D85B1FE37" bold="true" box="[151,436,475,501]" pageId="2" pageNumber="3">
Genus
<taxonomicName id="3CC94D04FFE2FFC5BBB9FE6A85B1FE37" ID-CoL="9CLPJ" authority="Hahn" authorityName="Hahn" box="[234,436,475,501]" class="Insecta" family="Miridae" genus="Pilophorus" kingdom="Animalia" order="Hemiptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="C9BDEA95FFE2FFC5BBB9FE6A856FFE36" bold="true" box="[234,362,475,500]" italics="true" pageId="2" pageNumber="3">Pilophorus</emphasis>
Hahn
</taxonomicName>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection id="B3D3650CFFE2FFC5BBC4FD958733FCFA" pageId="2" pageNumber="3" type="diagnosis">
<paragraph id="FB763687FFE2FFC5BBC4FD958733FCFA" blockId="2.[151,1437,548,1509]" pageId="2" pageNumber="3">
<emphasis id="C9BDEA95FFE2FFC5BBC4FD958514FDFF" bold="true" box="[151,273,548,573]" pageId="2" pageNumber="3">Diagnosis.</emphasis>
Primarily recognized by more or less myrmecomorphic, elongate and always macropterous body form; partly aggregated scalelike setae on scutellum; presence of anterior band of scalelike setae on corium (a posterior band variable, sometimes interrupted or scattered); weakly or moderately flattened metatibia; fleshy, recurved, apically convergent parempodia; usually strongly sprayed-out left paramere; ovoid to elongate oval right paramere; and C- to J-shaped (but rarely nearly straight) endosoma often furnished with a median spine, flagellum or barb. Some species strikingly ant-like because of strong modifications of the pronotum that is conspicuously tightened, constricted, or horned (
<figureCitation id="63F22A02FFE2FFC5BAF5FD4D85EBFCD6" box="[422,494,764,789]" captionStart="FIGURE 3" captionStartId="6.[151,250,1851,1873]" captionTargetBox="[151,1436,301,1829]" captionTargetId="figure@6.[151,1436,301,1830]" captionTargetPageId="6" captionText="FIGURE 3. Habitus images of strikingly antlike (A D) or otherwise morphologically novel (E) members of Pilophorus, living adult individuals named on the figure." httpUri="https://zenodo.org/record/286017/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
).
<bibRefCitation id="9F584B76FFE2FFC5B954FD4D8699FCD6" author="Schuh" box="[519,668,764,789]" pageId="2" pageNumber="3" refString="Schuh, R. T. (1984) Revision of the Phylinae (Hemiptera, Miridae) of the Indo - Pacific. Bulletin of the American Museum of Natural History, 177, 1 - 476." type="journal article" year="1984">Schuh (1984</bibRefCitation>
,
<bibRefCitation id="9F584B76FFE2FFC5B9F8FD4D86E3FCD6" author="Schuh" box="[683,742,764,788]" pageId="2" pageNumber="3" refString="Schuh, R. T. (1991) Phylogenetic, host and biogeographic analysis of the Pilophorini (Heteroptera: Miridae: Phylinae). Cladistics, 7, 157 - 189. http: // dx. doi. org / 10.1111 / j. 1096 - 0031.1991. tb 00030. x" type="journal article" year="1991">1991</bibRefCitation>
), Schuh &amp; Schwartz (1988) and
<bibRefCitation id="9F584B76FFE2FFC5BF39FD4C8199FCD6" author="Yasunaga" box="[1130,1436,764,789]" pageId="2" pageNumber="3" refString="Yasunaga, T. &amp; Schuh, R. T. (2013) Morphologically novel members of the ant-mimetic plant bug genus Pilophorus Hahn found in Thailand, with descriptions of three new species (Heteroptera: Miridae: Phylinae: Pilophorini). American Museum Novitates, 3768, 1 - 18. http: // dx. doi. org / 10.1206 / 3768.2" type="journal article" year="2013">Yasunaga &amp; Schuh (2013)</bibRefCitation>
provided further diagnostic characters and synapomorphies.
</paragraph>
</subSubSection>
<subSubSection id="B3D3650CFFE2FFC5BB94FCF5811DFC9F" box="[199,1304,836,861]" pageId="2" pageNumber="3" type="distribution">
<paragraph id="FB763687FFE2FFC5BB94FCF5811DFC9F" blockId="2.[151,1437,548,1509]" box="[199,1304,836,861]" pageId="2" pageNumber="3">
<emphasis id="C9BDEA95FFE2FFC5BB94FCF58565FC9F" bold="true" box="[199,352,836,861]" pageId="2" pageNumber="3">Distribution.</emphasis>
Holarctic, Oriental and Afrotropical regions, New
<collectingCountry id="83DE7617FFE2FFC5B8CEFCF587F4FC9F" box="[925,1009,836,861]" name="Guinea-Bissau" pageId="2" pageNumber="3">Guinea</collectingCountry>
, and some Pacific islands.
</paragraph>
</subSubSection>
<subSubSection id="B3D3650CFFE2FFC5BB94FCD68603FBF6" pageId="2" pageNumber="3" type="biology_ecology">
<paragraph id="FB763687FFE2FFC5BB94FCD68603FBF6" blockId="2.[151,1437,548,1509]" pageId="2" pageNumber="3">
<emphasis id="C9BDEA95FFE2FFC5BB94FCD68522FC42" bold="true" box="[199,295,871,896]" pageId="2" pageNumber="3">Biology.</emphasis>
Most members of
<taxonomicName id="3CC94D04FFE2FFC5B953FCD68678FC42" box="[512,637,871,896]" class="Insecta" family="Miridae" genus="Pilophorus" kingdom="Animalia" order="Hemiptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="C9BDEA95FFE2FFC5B953FCD68678FC42" box="[512,637,871,896]" italics="true" pageId="2" pageNumber="3">Pilophorus</emphasis>
</taxonomicName>
appear to be host plant specific, and the host plants range from forbs, broadleaf woody angiosperms to conifers, but the genus is also assumed to comprise predators, preying on other arthropods (
<bibRefCitation id="9F584B76FFE2FFC5BA71FC1E85CAFC0A" author="Wheeler" box="[290,463,943,968]" pageId="2" pageNumber="3" refString="Wheeler, A. G. Jr. (2001) Biology of the Plant Bugs (Hemiptera: Miridae), Pests, Predators, Opportunists. Cornell University Press, Ithaca and London, xv + 507 pp." type="book" year="2001">Wheeler, 2001</bibRefCitation>
;
<bibRefCitation id="9F584B76FFE2FFC5BA8CFC01869CFC0A" author="Yasunaga" box="[479,665,944,968]" pageId="2" pageNumber="3" refString="Yasunaga, T. (2001) Family Miridae Hahn, plant bugs. In: Yasunaga, T., Takai, M. &amp; Kawasawa, T. (Eds.), A Field Guide to Japanese Bugs II. Zenkoku Noson Kyoiku Kyokai, Publishing Co. Ltd., Tokyo, Japan, pp. 1 - 96, 111 - 351. [in Japanese]" type="book chapter" year="2001">Yasunaga, 2001</bibRefCitation>
;
<bibRefCitation id="9F584B76FFE2FFC5B9FAFC0187D2FC0A" author="Yasunaga" box="[681,983,943,968]" pageId="2" pageNumber="3" refString="Yasunaga, T. &amp; Schuh, R. T. (2013) Morphologically novel members of the ant-mimetic plant bug genus Pilophorus Hahn found in Thailand, with descriptions of three new species (Heteroptera: Miridae: Phylinae: Pilophorini). American Museum Novitates, 3768, 1 - 18. http: // dx. doi. org / 10.1206 / 3768.2" type="journal article" year="2013">Yasunaga &amp; Schuh, 2013</bibRefCitation>
). In
<collectingCountry id="83DE7617FFE2FFC5BF42FC018051FC05" box="[1041,1108,944,967]" name="Japan" pageId="2" pageNumber="3">Japan</collectingCountry>
,
<emphasis id="C9BDEA95FFE2FFC5BF30FC0080C1FC0A" box="[1123,1220,945,968]" italics="true" pageId="2" pageNumber="3">P. t y pi cu</emphasis>
s (Distant) and its closely related congeners are regarded to be potential candidates as indigenous natural enemies for biological control programs (
<bibRefCitation id="9F584B76FFE2FFC5BA37FC49860FFBD2" author="Ito" box="[356,522,1015,1040]" pageId="2" pageNumber="3" refString="Ito, K., Nishikawa, H., Shimada, T., Ogawa, K., Minamiya, T., Tomoda, M., Nakahira, K., Kodama, R., Fukuda, T. &amp; Arakawa, R. (2011) Analysis of genetic variation and phylogeny of the predatory bug, Pilophorus typicus, in Japan using mitochondrial gene sequences. Journal of Insect Science, 11 (18), 1 - 13. http: // dx. doi. org / 10.1673 / 031.011.0118" type="journal article" year="2011">
Ito
<emphasis id="C9BDEA95FFE2FFC5BAD8FC4885C6FBD2" box="[395,451,1015,1040]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
, 2011
</bibRefCitation>
;
<bibRefCitation id="9F584B76FFE2FFC5B946FC4986CFFBD2" author="Yasunaga" box="[533,714,1016,1040]" pageId="2" pageNumber="3" refString="Yasunaga, T. (2001) Family Miridae Hahn, plant bugs. In: Yasunaga, T., Takai, M. &amp; Kawasawa, T. (Eds.), A Field Guide to Japanese Bugs II. Zenkoku Noson Kyoiku Kyokai, Publishing Co. Ltd., Tokyo, Japan, pp. 1 - 96, 111 - 351. [in Japanese]" type="book chapter" year="2001">Yasunaga, 2001</bibRefCitation>
). Both adults and immature forms are often observed to co-occur with the workers of various ants.
</paragraph>
</subSubSection>
<subSubSection id="B3D3650CFFE2FFC5BB94FB8E8528FB2A" pageId="2" pageNumber="3" type="discussion">
<paragraph id="FB763687FFE2FFC5BB94FB8E8528FB2A" blockId="2.[151,1437,548,1509]" pageId="2" pageNumber="3">
<emphasis id="C9BDEA95FFE2FFC5BB94FB8E8549FB9A" bold="true" box="[199,332,1087,1112]" pageId="2" pageNumber="3">Discussion.</emphasis>
Most members of
<taxonomicName id="3CC94D04FFE2FFC5B961FB8E86ABFB9A" box="[562,686,1087,1112]" class="Insecta" family="Miridae" genus="Pilophorus" kingdom="Animalia" order="Hemiptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="C9BDEA95FFE2FFC5B961FB8E86ABFB9A" box="[562,686,1087,1112]" italics="true" pageId="2" pageNumber="3">Pilophorus</emphasis>
</taxonomicName>
are assumed to have strong relationships to certain ant species (
<bibRefCitation id="9F584B76FFE2FFC5BBCDFBD58592FBBE" author="Nakatani" box="[158,407,1124,1149]" pageId="2" pageNumber="3" refString="Nakatani, Y., Komatsu, T., Itino, T., Shimizu-kaya, U., Itioka, R., Hashim, R. &amp; Ueda, S. (2013) New Pilophorus species associated with myrmecophilous Macaranga trees from the Malay Peninsula and Borneo (Heteroptera: Miridae: Phylinae). Tijdscrift voor Entomologie, 156, 113 - 126. http: // dx. doi. org / 10.1163 / 22119434 - 00002024" type="journal article" year="2013">
Nakatani
<emphasis id="C9BDEA95FFE2FFC5BA5CFBD4854EFBBF" box="[271,331,1124,1149]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
, 2013
</bibRefCitation>
;
<bibRefCitation id="9F584B76FFE2FFC5BAF6FBD486CBFBBE" author="Yasunaga" box="[421,718,1124,1149]" pageId="2" pageNumber="3" refString="Yasunaga, T. &amp; Schuh, R. T. (2013) Morphologically novel members of the ant-mimetic plant bug genus Pilophorus Hahn found in Thailand, with descriptions of three new species (Heteroptera: Miridae: Phylinae: Pilophorini). American Museum Novitates, 3768, 1 - 18. http: // dx. doi. org / 10.1206 / 3768.2" type="journal article" year="2013">Yasunaga &amp; Schuh, 2013</bibRefCitation>
), and quite a few undescribed species exist particularly in the subtropical and tropical Old World. There are at least three undetermined or undescribed species (now represented only by females, as in
<figureCitation id="63F22A02FFE2FFC5BAC6FB1D85D6FB06" box="[405,467,1196,1221]" captionStart="FIGURE 1" captionStartId="4.[151,250,1821,1843]" captionTargetBox="[151,1436,193,1799]" captionTargetId="figure@4.[151,1436,193,1799]" captionTargetPageId="4" captionText="FIGURE 1. Habitus images of Pilophorus spp., living individuals. A &amp; B. P. suwimonae, holotype male (AMNH _ PBI 00380149). C. P. subparallelus, male (00380162). D. P. meteorus, holotype male (00380150). E. Same, female (00380152). F. P. typicus, male (00380192). G. Same, 5 th instar immature (from SERS). H. Undetermined species, female (SERS)." httpUri="https://zenodo.org/record/286015/files/figure.png" pageId="2" pageNumber="3">Fig. 1</figureCitation>
H) present in
<collectingCountry id="83DE7617FFE2FFC5B922FB1D86DDFB07" box="[625,728,1196,1221]" name="Thailand" pageId="2" pageNumber="3">Thailand</collectingCountry>
. We refrain from describing such species until male specimens are available.
</paragraph>
</subSubSection>
<subSubSection id="B3D3650CFFE2FFC5BB94FB458141FA26" pageId="2" pageNumber="3" type="reference_group">
<paragraph id="FB763687FFE2FFC5BB94FB458141FA26" blockId="2.[151,1437,548,1509]" pageId="2" pageNumber="3">
<taxonomicName id="3CC94D04FFE2FFC5BB94FB458546FACF" ID-CoL="9CLPJ" box="[199,323,1268,1293]" class="Insecta" family="Miridae" genus="Pilophorus" kingdom="Animalia" order="Hemiptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">
<emphasis id="C9BDEA95FFE2FFC5BB94FB458546FACF" box="[199,323,1268,1293]" italics="true" pageId="2" pageNumber="3">Pilophorus</emphasis>
</taxonomicName>
comprises morphologically very diverse species; some members are strikingly antlike (
<figureCitation id="63F22A02FFE2FFC5BE1FFB458196FACE" box="[1356,1427,1268,1293]" captionStart="FIGURE 3" captionStartId="6.[151,250,1851,1873]" captionTargetBox="[151,1436,301,1829]" captionTargetId="figure@6.[151,1436,301,1830]" captionTargetPageId="6" captionText="FIGURE 3. Habitus images of strikingly antlike (A D) or otherwise morphologically novel (E) members of Pilophorus, living adult individuals named on the figure." httpUri="https://zenodo.org/record/286017/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
) because of conspicuous modification of the pronota, and
<emphasis id="C9BDEA95FFE2FFC5B872FAA887C2FAF2" box="[801,967,1305,1328]" italics="true" pageId="2" pageNumber="3">
<taxonomicName id="3CC94D04FFE2FFC5B872FAA887C6FAF2" box="[801,963,1305,1328]" class="Insecta" family="Miridae" genus="Pilophorus" kingdom="Animalia" order="Hemiptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="species" species="portentosus">P. portentosus</taxonomicName>
,
</emphasis>
which has the excessively long antennae and legs (
<figureCitation id="63F22A02FFE2FFC5BA56FA8D8542FA96" box="[261,327,1340,1365]" captionStart="FIGURE 3" captionStartId="6.[151,250,1851,1873]" captionTargetBox="[151,1436,301,1829]" captionTargetId="figure@6.[151,1436,301,1830]" captionTargetPageId="6" captionText="FIGURE 3. Habitus images of strikingly antlike (A D) or otherwise morphologically novel (E) members of Pilophorus, living adult individuals named on the figure." httpUri="https://zenodo.org/record/286017/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
H), resembles wasp rather than ant (
<bibRefCitation id="9F584B76FFE2FFC5B9B2FA8D8002FA96" author="Yasunaga" box="[737,1031,1340,1365]" pageId="2" pageNumber="3" refString="Yasunaga, T. &amp; Schuh, R. T. (2013) Morphologically novel members of the ant-mimetic plant bug genus Pilophorus Hahn found in Thailand, with descriptions of three new species (Heteroptera: Miridae: Phylinae: Pilophorini). American Museum Novitates, 3768, 1 - 18. http: // dx. doi. org / 10.1206 / 3768.2" type="journal article" year="2013">Yasunaga &amp; Schuh, 2013</bibRefCitation>
). On the other hand,
<bibRefCitation id="9F584B76FFE2FFC5BFA5FA8D84E4FABA" pageId="2" pageNumber="3" refString="Nakatani, Y., Komatsu, T., Itino, T., Shimizu-kaya, U., Itioka, R., Hashim, R. &amp; Ueda, S. (2013) New Pilophorus species associated with myrmecophilous Macaranga trees from the Malay Peninsula and Borneo (Heteroptera: Miridae: Phylinae). Tijdscrift voor Entomologie, 156, 113 - 126. http: // dx. doi. org / 10.1163 / 22119434 - 00002024" type="journal article">
Nakatani
<emphasis id="C9BDEA95FFE2FFC5BE30FA8C8199FA97" box="[1379,1436,1340,1365]" italics="true" pageId="2" pageNumber="3">et al.</emphasis>
(2013)
</bibRefCitation>
reported seven unique species that are predominantly associated with
<taxonomicName id="3CC94D04FFE2FFC5B8ADFAD08084FABA" ID-CoL="8VSBG" box="[1022,1153,1377,1400]" class="Magnoliopsida" family="Euphorbiaceae" genus="Macaranga" kingdom="Plantae" order="Malpighiales" pageId="2" pageNumber="3" phylum="Tracheophyta" rank="genus">
<emphasis id="C9BDEA95FFE2FFC5B8ADFAD08084FABA" box="[1022,1153,1377,1400]" italics="true" pageId="2" pageNumber="3">Macaranga</emphasis>
</taxonomicName>
trees (
<taxonomicName id="3CC94D04FFE2FFC5BF9EFAEE8172FABA" ID-CoL="9Y7" box="[1229,1399,1375,1400]" class="Magnoliopsida" family="Euphorbiaceae" kingdom="Plantae" order="Malpighiales" pageId="2" pageNumber="3" phylum="Tracheophyta" rank="family">Euphorbiaceae</taxonomicName>
) in
<collectingCountry id="83DE7617FFE2FFC5BBC4FA358504FA5F" box="[151,257,1412,1437]" name="Malaysia" pageId="2" pageNumber="3">Malaysia</collectingCountry>
; these members uniquely have the strongly protruded scutella. In spite of such great interspecific variation exhibited in some special members, the monophyly of the genus may be supported by the recent phylogenetic analyses based on both morphology and DNA sequence data (
<bibRefCitation id="9F584B76FFE2FFC5B8B6FA7D80EAFA26" author="Schuh" box="[997,1263,1484,1509]" pageId="2" pageNumber="3" refString="Schuh, R. T. &amp; Menard, K. (2011) Santatalean - feeding plant bugs: ten new species in the genus Hypseloecus Reuter from Australia and South Africa (Heteroptera: Miridae: Phylinae): their hosts and placement in the Pilophorini. Australian Journal of Entomology, 50, 365 - 392." type="journal article" year="2011">Schuh &amp; Menard, 2011</bibRefCitation>
;
<bibRefCitation id="9F584B76FFE2FFC5BFA8FA7D813DFA26" author="Menard" box="[1275,1336,1484,1508]" pageId="2" pageNumber="3" refString="Schuh, R. T. &amp; Menard, K. (2013) A revised classification of the Phylinae (Insecta: Heteroptera: Miridae): Arguments for the placement of genera. American Museum Novitates, 3785, 1 - 72. http: // dx. doi. org / 10.1206 / 3785.2" type="journal article" year="2013">2013</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>