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<mods:title id="EF5101B8E3803810F0C5DE233C8B35BE">Redescription of Rhyacophila clemens Tsuda 1940 (Trichoptera, Rhyacophilidae), with descriptions of five new, allied species from Japan</mods:title>
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<subSubSection id="F8E66537F7241177FF4CC059FD48A106" box="[151,730,1796,1823]" pageId="18" pageNumber="73" type="nomenclature">
<paragraph id="B04336BCF7241177FF4CC059FD48A106" blockId="18.[151,730,1796,1823]" box="[151,730,1796,1823]" pageId="18" pageNumber="73">
<heading id="EB0B81D0F7241177FF4CC059FD48A106" bold="true" box="[151,730,1796,1823]" fontSize="11" level="1" pageId="18" pageNumber="73" reason="1">
<emphasis id="8288EAAEF7241177FF4CC059FD48A106" bold="true" box="[151,730,1796,1823]" pageId="18" pageNumber="73">
<taxonomicName id="77FC4D3FF7241177FF4CC059FE08A106" ID-CoL="4SNQ4" authority="Tsuda, 1940" authorityName="Tsuda" authorityYear="1940" box="[151,410,1796,1823]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="18" pageNumber="73" phylum="Arthropoda" rank="species" species="clemens">
<emphasis id="8288EAAEF7241177FF4CC059FE08A106" bold="true" box="[151,410,1796,1823]" italics="true" pageId="18" pageNumber="73">Rhyacophila clemens</emphasis>
</taxonomicName>
Species Group
<bibRefCitation id="D46D4B4DF7241177FD86C058FD48A106" author="Ross, H. H." box="[605,730,1797,1823]" pageId="18" pageNumber="73" refId="ref18378" refString="Ross, H. H. (1956) Evolution and Classification of the Mountain Caddisflies. The University of Illinois Press, Urbana, Illinois, 213 pp." type="book" year="1956">Ross 1956</bibRefCitation>
</emphasis>
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<subSubSection id="F8E66537F7241171FF4CC010FBFBA704" lastPageId="20" lastPageNumber="75" pageId="18" pageNumber="73" type="discussion">
<paragraph id="B04336BCF7241171FF4CC010FBFBA704" blockId="18.[151,1437,1869,2039]" lastBlockId="20.[151,1437,151,285]" lastPageId="20" lastPageNumber="75" pageId="18" pageNumber="73">
In this study, I have recognized six Japanese species and one Korean species belonging to the
<taxonomicName id="77FC4D3FF7241177FB1FC013FAACA17E" authorityName="Tsuda" authorityYear="1940" box="[1220,1342,1869,1895]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="18" pageNumber="73" phylum="Arthropoda" rank="species" species="clemens">
<emphasis id="8288EAAEF7241177FB1FC013FAACA17E" box="[1220,1342,1869,1895]" italics="true" pageId="18" pageNumber="73">R. clemens</emphasis>
</taxonomicName>
Species Group. As mentioned above in the introduction, the males of this species group are similar to those of the
<taxonomicName id="77FC4D3FF7241177FAFDC02FFA0FA192" authorityName="Milne" authorityYear="1936" box="[1318,1437,1906,1931]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="18" pageNumber="73" phylum="Arthropoda" rank="species" species="viquaea">
<emphasis id="8288EAAEF7241177FAFDC02FFA0FA192" box="[1318,1437,1906,1931]" italics="true" pageId="18" pageNumber="73">R. viquaea</emphasis>
</taxonomicName>
Species Group in having two pairs of anal sclerites, but the males of the seven species of the
<taxonomicName id="77FC4D3FF7241177FB18C0CBFAACA1B6" authorityName="Tsuda" authorityYear="1940" box="[1219,1342,1941,1967]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="18" pageNumber="73" phylum="Arthropoda" rank="species" species="clemens">
<emphasis id="8288EAAEF7241177FB18C0CBFAACA1B6" box="[1219,1342,1941,1967]" italics="true" pageId="18" pageNumber="73">R. clemens</emphasis>
</taxonomicName>
Species Group in this study were distinguishable from those of the
<taxonomicName id="77FC4D3FF7241177FC9AC0E7FC29A1CA" authorityName="Milne" authorityYear="1936" box="[833,955,1978,2003]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="18" pageNumber="73" phylum="Arthropoda" rank="species" species="viquaea">
<emphasis id="8288EAAEF7241177FC9AC0E7FC29A1CA" box="[833,955,1978,2003]" italics="true" pageId="18" pageNumber="73">R. viquaea</emphasis>
</taxonomicName>
Species Group by the shape of the apical band: The apical band is large and oval in lateral view and independent of the root of the anal sclerites in the
<taxonomicName id="77FC4D3FF7241176FA5FC083FF63A132" authorityName="Tsuda" authorityYear="1940" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" lastPageId="19" lastPageNumber="74" order="Trichoptera" pageId="18" pageNumber="73" phylum="Arthropoda" rank="species" species="clemens">
<emphasis id="8288EAAEF7241176FA5FC083FF63A132" italics="true" lastPageId="19" lastPageNumber="74" pageId="18" pageNumber="73">R. clemens</emphasis>
</taxonomicName>
Species Group (
<figureCitation id="28C72A39F7251176FE69C04CFD83A132" box="[434,529,1809,1835]" captionStart="FIGURE 1" captionStartId="2.[151,250,1690,1714]" captionTargetBox="[279,1307,246,1655]" captionTargetId="figure-51@2.[271,1316,234,1667]" captionTargetPageId="2" captionText="FIGURE 1. Rhyacophila clemens Tsuda 1940. 1A1J: 1A, male habitus, left dorsolateral (photographed by K. Inazu); 1B, male head and pro-, meso- and metanota, dorsal; 1C, male head, left lateral; 1D, male right wings, dorsal; 1E, lectotype (in glass tube) and labels preserved in INHS; 1F, male right wings, paralectotype (balsam mounted slide) preserved in OMNH; 1G1J, male genitalia: 1G, left lateral; 1H, segment X and phallic apparatus, left lateral; 1I, same, posterodorsal; 1J, phallic apparatus, left dorsolateral. Abbreviations: I, II, III, IV, V = apical forks I, II, III, IV, V; IX, X = abdominal segments IX, X; A1, A2, A3 = anal veins 13; ab = apical band; aed = aedeagus; adl = apicodorsal lobe of segment IX; ahp = apical hinged process of segment X; as = anal sclerite; asia = apical segment of inferior appendage (paired); bsia = basal segment of inferior appendage (paired); C = costa; Cu1, Cu1a, Cu1b = cubital veins 1, 1a, 1b; hp = horizontal part of segment X; M1, M2, M3, M4 = median veins 14; m-cu = crossvein between median and cubital veins; phb = phallobase; R1, R2, R3, R4, R5 = radial veins 15; r-m = crossvein between radial and median veins; Sc = subcosta; tb = tergal band; TC = thyridial cell; vp = vertical part of segment X; vpl = ventral plate of aedeagus." figureDoi="http://doi.org/10.5281/zenodo.11119211" httpUri="https://zenodo.org/record/11119211/files/figure.png" pageId="19" pageNumber="74">Figs 1H</figureCitation>
,
<figureCitation id="28C72A39F7251176FDFBC04CFDD1A132" box="[544,579,1809,1835]" captionStart="FIGURE 3" captionStartId="7.[151,250,1726,1750]" captionTargetBox="[278,1310,194,1690]" captionTargetId="figure-16@7.[265,1322,181,1702]" captionTargetPageId="7" captionText="FIGURE 3. Rhyacophila vesperalis sp. nov. 3A3E, male genitalia: 3A, left lateral; 3B, segment X and phallic apparatus, left lateral; 3C, dorsal (left inferior appendage omitted); 3D, ventral (left inferior appendage and apical segment of right inferior appendage omitted); 3E1, apical hinged process of segment X, enlarged, ventral; 3E2, ventral plate of aedeagus, enlarged, ventral. 3F3K, female genitalia: 3F, left lateral; 3G, segment VIII, left lateral; 3H, same dorsal; 3I, same ventral; 3J, vaginal apparatus, left lateral; 3K, same, ventral. Arrows see text." figureDoi="http://doi.org/10.5281/zenodo.11119215" httpUri="https://zenodo.org/record/11119215/files/figure.png" pageId="19" pageNumber="74">3B</figureCitation>
,
<figureCitation id="28C72A39F7251176FD8AC04CFDE4A132" box="[593,630,1809,1835]" captionStart="FIGURE 4" captionStartId="10.[151,250,1810,1834]" captionTargetBox="[288,1300,430,1774]" captionTargetId="figure-170@10.[275,1313,417,1787]" captionTargetPageId="10" captionText="FIGURE 4. Rhyacophila inazui sp. nov. 4A, male habitus, left dorsolateral (photographed by K. Inazu); 4B, female right forewing (in alcohol), dorsal; 4C4H, male genitalia: 4C, left lateral; 4D, segment X and phallic apparatus, left lateral; 4E, dorsal (left inferior appendage omitted); 4F, ventral (left inferior appendage and apical segment of right inferior appendage omitted); 4G1, apical hinged process of segment X, enlarged, ventral; 4G2, ventral plate of aedeagus, enlarged, ventral; 4H, posterodorsal. 4I4N, female genitalia: 4I, left lateral; 4J, ventral; 4K, segment VIII, left lateral; 4L, same, ventral; 4M, vaginal apparatus, left lateral; 4N, same, ventral. Abbreviation: vr = ventral ridge. Arrows see text." figureDoi="http://doi.org/10.5281/zenodo.11119217" httpUri="https://zenodo.org/record/11119217/files/figure.png" pageId="19" pageNumber="74">4D</figureCitation>
,
<figureCitation id="28C72A39F7251176FD5FC04CFD34A135" box="[644,678,1809,1836]" captionStart="FIGURE 5" captionStartId="13.[151,250,1525,1549]" captionTargetBox="[284,1304,286,1489]" captionTargetId="figure-44@13.[271,1316,273,1501]" captionTargetPageId="13" captionText="FIGURE 5. Rhyacophila angulicauda sp. nov. 5A5D, male genitalia: 5A, left lateral; 5B, segment X and phallic apparatus, left lateral; 5C, dorsal (left inferior appendage omitted); 5D, ventral (left inferior appendage and apex of apical segment of right inferior appendage omitted). 5E5I, female genitalia: 5E, left lateral; 5F, segment VIII, left lateral; 5G, same, ventral; 5H, vaginal apparatus, left lateral; 5I, same, ventral. Arrows see text." figureDoi="http://doi.org/10.5281/zenodo.11119221" httpUri="https://zenodo.org/record/11119221/files/figure.png" pageId="19" pageNumber="74">5B</figureCitation>
,
<figureCitation id="28C72A39F7251176FD6EC04CFD4AA132" box="[693,728,1809,1835]" captionStart="FIGURE 6" captionStartId="15.[151,250,1368,1392]" captionTargetBox="[298,1289,250,1332]" captionTargetId="figure-43@15.[285,1301,237,1345]" captionTargetPageId="15" captionText="FIGURE 6. Rhyacophila parvicauda sp. nov. 6A6D, male genitalia: 6A, left lateral; 6B, segment X and phallic apparatus, left lateral; 6C, dorsal (left inferior appendage omitted); 6D, ventral (left inferior appendage omitted). 6E6H, female genitalia: 6E, segment VIII, left lateral; 6F, same, ventral; 6G, vaginal apparatus, left lateral; 6H, same, ventral. Abbreviation: vr = ventral ridge.Arrows see text." figureDoi="http://doi.org/10.5281/zenodo.11119223" httpUri="https://zenodo.org/record/11119223/files/figure.png" pageId="19" pageNumber="74">6B</figureCitation>
,
<figureCitation id="28C72A39F7251176FD3DC04CFC9BA132" box="[742,777,1809,1835]" captionStart="FIGURE 7" captionStartId="16.[151,250,1907,1931]" captionTargetBox="[309,1278,737,1871]" captionTargetId="figure-194@16.[296,1291,724,1884]" captionTargetPageId="16" captionText="FIGURE 7. Rhyacophila furcicauda sp. nov. 7A7F, male genitalia: 7A, left lateral; 7B, segment X and phallic apparatus, left lateral; 7C, same, posterodorsal; 7D, dorsal (left inferior appendage omitted); 7E, apical segment of right inferior appendage, mesal; 7F, segment X and phallic apparatus, ventral. 7G7J, female genitalia: 7G, segment VIII, left lateral; 7H, same, ventral; 7I, vaginal apparatus, left lateral; 7J, same, ventral. Abbreviation: vr = ventral ridge. Arrows see text." figureDoi="http://doi.org/10.5281/zenodo.11119225" httpUri="https://zenodo.org/record/11119225/files/figure.png" pageId="19" pageNumber="74">7B</figureCitation>
), but the ends of the apical band are joined with the root of the ventral pair of the anal sclerites in the
<taxonomicName id="77FC4D3FF7251176FD78C06BFC8CA156" authorityName="Milne" authorityYear="1936" box="[675,798,1846,1871]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="19" pageNumber="74" phylum="Arthropoda" rank="species" species="viquaea">
<emphasis id="8288EAAEF7251176FD78C06BFC8CA156" box="[675,798,1846,1871]" italics="true" pageId="19" pageNumber="74">R. viquaea</emphasis>
</taxonomicName>
Species Group (
<bibRefCitation id="D46D4B4DF7251176FC05C068FBCEA149" author="Ross, H. H." box="[990,1116,1845,1872]" pageId="19" pageNumber="74" refId="ref18378" refString="Ross, H. H. (1956) Evolution and Classification of the Mountain Caddisflies. The University of Illinois Press, Urbana, Illinois, 213 pp." type="book" year="1956">Ross 1956</bibRefCitation>
).
<bibRefCitation id="D46D4B4DF7251176FBA9C068FA89A156" author="Schmid, F." box="[1138,1307,1845,1871]" pageId="19" pageNumber="74" pagination="5 - 230" refId="ref18408" refString="Schmid, F. (1970) Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Memoires de la Societe Entomologique du Canada, 66, 5 - 230, pls. 1 - 52. https: // doi. org / 10.4039 / entm 10266 fv" type="journal article" year="1970">Schmid (1970)</bibRefCitation>
treated the ventral pair of the anal sclerites in the
<taxonomicName id="77FC4D3FF7251176FD9FC007FD2FA16A" authorityName="Milne" authorityYear="1936" box="[580,701,1882,1907]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="19" pageNumber="74" phylum="Arthropoda" rank="species" species="viquaea">
<emphasis id="8288EAAEF7251176FD9FC007FD2FA16A" box="[580,701,1882,1907]" italics="true" pageId="19" pageNumber="74">R. viquaea</emphasis>
</taxonomicName>
Species Group as the apical band of the tergal strap, whereas
<bibRefCitation id="D46D4B4DF7251176FAA9C007FED8A18E" author="Lee, J. J. &amp; Ruiter, D. E." pageId="19" pageNumber="74" pagination="126 - 130" refId="ref17936" refString="Lee, J. J. &amp; Ruiter, D. E. (2011) Rhyacophila weitchpec sp. nov. from northern California, with discussion of the Rhyacophila viquaea Milne 1936 species group (Trichoptera: Rhyacophilidae). The Pan-Pacific Entomologist, 86, 126 - 130. https: // doi. org / 10.3956 / 2010 - 22.1" type="journal article" year="2011">Lee &amp; Ruiter (2011)</bibRefCitation>
agreed with Rosss interpretation. Furthermore, the male of each species in the
<taxonomicName id="77FC4D3FF7251176FB1CC023FAD2A18E" authorityName="Tsuda" authorityYear="1940" box="[1223,1344,1917,1943]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="19" pageNumber="74" phylum="Arthropoda" rank="species" species="clemens">
<emphasis id="8288EAAEF7251176FB1CC023FAD2A18E" box="[1223,1344,1917,1943]" italics="true" pageId="19" pageNumber="74">R. clemens</emphasis>
</taxonomicName>
Species Group bears the apical hinged process on segment X posteroventrally and the ventral plate of the aedeagus, but those of the
<taxonomicName id="77FC4D3FF7251176FEC4C09BFE05A1C6" authorityName="Milne" authorityYear="1936" box="[287,407,1990,2015]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="19" pageNumber="74" phylum="Arthropoda" rank="species" species="viquaea">
<emphasis id="8288EAAEF7251176FEC4C09BFE05A1C6" box="[287,407,1990,2015]" italics="true" pageId="19" pageNumber="74">R. viquaea</emphasis>
</taxonomicName>
Species Group lack such a process and plate (
<bibRefCitation id="D46D4B4DF7251176FC4DC098FB82A1F9" author="Ross, H. H." box="[918,1040,1989,2016]" pageId="19" pageNumber="74" refId="ref18378" refString="Ross, H. H. (1956) Evolution and Classification of the Mountain Caddisflies. The University of Illinois Press, Urbana, Illinois, 213 pp." type="book" year="1956">Ross 1956</bibRefCitation>
,
<bibRefCitation id="D46D4B4DF7251176FBC1C098FB23A1C6" author="Schmid, F." box="[1050,1201,1989,2015]" pageId="19" pageNumber="74" pagination="5 - 230" refId="ref18408" refString="Schmid, F. (1970) Le genre Rhyacophila et la famille des Rhyacophilidae (Trichoptera). Memoires de la Societe Entomologique du Canada, 66, 5 - 230, pls. 1 - 52. https: // doi. org / 10.4039 / entm 10266 fv" type="journal article" year="1970">Schmid 1970</bibRefCitation>
,
<bibRefCitation id="D46D4B4DF7251176FB67C09BFA03A1C6" author="Lee, J. J. &amp; Ruiter, D. E." box="[1212,1425,1989,2016]" pageId="19" pageNumber="74" pagination="126 - 130" refId="ref17936" refString="Lee, J. J. &amp; Ruiter, D. E. (2011) Rhyacophila weitchpec sp. nov. from northern California, with discussion of the Rhyacophila viquaea Milne 1936 species group (Trichoptera: Rhyacophilidae). The Pan-Pacific Entomologist, 86, 126 - 130. https: // doi. org / 10.3956 / 2010 - 22.1" type="journal article" year="2011">Lee &amp; Ruiter 2011</bibRefCitation>
). Consequently, the male diagnostic characters for the
<taxonomicName id="77FC4D3FF7221171FD31C7CAFCF1A6A8" authorityName="Tsuda" authorityYear="1940" box="[746,867,151,177]" class="Insecta" family="Rhyacophilidae" genus="Rhyacophila" kingdom="Animalia" order="Trichoptera" pageId="20" pageNumber="75" phylum="Arthropoda" rank="species" species="clemens">
<emphasis id="8288EAAEF7221171FD31C7CAFCF1A6A8" box="[746,867,151,177]" italics="true" pageId="20" pageNumber="75">R. clemens</emphasis>
</taxonomicName>
Species Group are as follows: Segment IX is long dorsally; the preanal appendages are absent; segment X bears an apical hinged process posteroventrally; two pairs of anal sclerites have a common internal root; a pair of large apical bands surrounds the anus and the root of the anal sclerites; the aedeagus bears a large sclerotized ventral plate; and parameres are absent.
</paragraph>
</subSubSection>
</treatment>
</document>