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<document ID-DOI="10.1016/j.ympev.2021.107115" ID-Zenodo-Dep="6643504" approvalRequired="21" approvalRequired_for_document="1" approvalRequired_for_taxonomicNames="20" checkinTime="1654106955118" checkinUser="diego" docAuthor="Bybee, Seth M., Kalkman, Vincent J., Erickson, Robert J., Frandsen, Paul B., Breinholt, Jesse W., Suvorov, Anton, Dijkstra, Klaas-Douwe B., Cordero-Rivera, Adolfo, Skevington, Jeffrey H., Abbott, John C., Herrera, Melissa Sanchez, Lemmon, Alan R., Lemmon, Emily Moriarty &amp; Ware, Jessica L." docDate="2021" docId="039687E7A861FFDFE433AA43FBE8EFED" docLanguage="en" docName="MolPhylogenetEvol.160.107115.pdf" docOrigin="Molecular Phylogenetics and Evolution 160" docTitle="Calopterygoidea Selys-Longchamps 1850" docType="treatment" docVersion="4" lastPageNumber="9" masterDocId="FFAFFF9FA869FFD7E457AC3AFFA9EC75" masterDocTitle="Phylogeny and classification of Odonata using targeted genomics" masterLastPageNumber="107115" masterPageNumber="107115" pageNumber="9" updateTime="1655236780830" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-4.0">
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<mods:title>Phylogeny and classification of Odonata using targeted genomics</mods:title>
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<mods:namePart>Kalkman, Vincent J.</mods:namePart>
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<mods:namePart>Frandsen, Paul B.</mods:namePart>
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<mods:namePart>Skevington, Jeffrey H.</mods:namePart>
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<mods:namePart>Abbott, John C.</mods:namePart>
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<mods:namePart>Herrera, Melissa Sanchez</mods:namePart>
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<mods:namePart>Lemmon, Alan R.</mods:namePart>
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<mods:namePart>Lemmon, Emily Moriarty</mods:namePart>
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<emphasis bold="true" box="[100,415,1656,1676]" italics="true" pageId="8" pageNumber="9">
4.1.5.4.
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1850" box="[189,333,1656,1675]" kingdom="Animalia" pageId="8" pageNumber="7" rank="superFamily" superFamily="Calopterygoidea">Calopterygoidea</taxonomicName>
group 4
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<paragraph blockId="8.[100,770,1656,1955]" pageId="8" pageNumber="9">
<emphasis bold="true" box="[421,455,1657,1676]" italics="true" pageId="8" pageNumber="9">(BS</emphasis>
=
<emphasis bold="true" box="[484,556,1657,1676]" italics="true" pageId="8" pageNumber="9">100, PP</emphasis>
=
<emphasis bold="true" box="[586,636,1657,1676]" italics="true" pageId="8" pageNumber="9">1, QS</emphasis>
=
<emphasis bold="true" italics="true" pageId="8" pageNumber="9">0.37/0.42/ 0.96).</emphasis>
</paragraph>
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<subSubSection pageId="8" pageNumber="9" type="discussion">
<paragraph blockId="8.[100,770,1656,1955]" lastBlockId="8.[818,1488,148,920]" pageId="8" pageNumber="9">
The fourth group contained within the Calopterygoidea consists of three pairs of families.
<taxonomicName authorityName="Kennedy" authorityYear="1925" box="[336,451,1712,1731]" class="Insecta" family="Philosinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Philosinidae</taxonomicName>
(BS = 100, PP = 1, QS = 1/NA/1) includes the Asian genera
<taxonomicName authorityName="Ris" authorityYear="1917" box="[342,423,1740,1759]" class="Insecta" family="Megapodagrionidae" genus="Philosina" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[342,423,1740,1759]" italics="true" pageId="8" pageNumber="9">Philosina</emphasis>
</taxonomicName>
and
<emphasis bold="true" box="[467,570,1740,1759]" italics="true" pageId="8" pageNumber="9">
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[467,566,1740,1759]" class="Insecta" family="Megapodagrionidae" genus="Rhinagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">Rhinagrion</taxonomicName>
,
</emphasis>
which resemble each other strongly in adult morphology and have a clear apomorphy in the nymphal stage in the tube-shaped caudal gills, i.e. the outer gills are folded around the median gill (
<bibRefCitation author="Kalkman, V. J. &amp; Choong, C. Y. &amp; Orr, A. G. &amp; Schutte, K." box="[387,578,1824,1844]" journalOrPublisher="Int. J. Odonatol." pageId="8" pageNumber="9" pagination="119 - 135" part="13" refId="ref18012" refString="Kalkman, V. J., Choong, C. Y., Orr, A. G., Schutte, K., 2010. Remarks on the taxonomy of Megapodagrionidae with emphasis on the larval gills (Odonata). Int. J. Odonatol. 13, 119 - 135. https: // doi. org / 10.1080 / 13887890.2010.9748366." title="Remarks on the taxonomy of Megapodagrionidae with emphasis on the larval gills (Odonata)" type="journal article" year="2010">Kalkman et al., 2010</bibRefCitation>
). The
<taxonomicName box="[636,770,1824,1843]" class="Insecta" family="Philogangidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Philogangidae</taxonomicName>
with its single genus
<taxonomicName authorityName="Kirby" authorityYear="1890" box="[305,402,1852,1871]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[305,402,1852,1871]" italics="true" pageId="8" pageNumber="9">Philoganga</emphasis>
</taxonomicName>
is its sister group with strong support (BS = 100, PP = 1, QS = 0.92/0/1). Adult
<taxonomicName authorityName="Kirby" authorityYear="1890" box="[521,618,1880,1899]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[521,618,1880,1899]" italics="true" pageId="8" pageNumber="9">Philoganga</emphasis>
</taxonomicName>
resembles
<taxonomicName authorityName="Kennedy" authorityYear="1925" class="Insecta" family="Philosinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Philosinidae</taxonomicName>
in general appearance, being relatively large and robust, and resting with wings outstretched. The main difference is in the denser venation, with two antenodal crossveins in
<taxonomicName authorityName="Kennedy" authorityYear="1925" box="[1223,1341,148,167]" class="Insecta" family="Philosinidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Philosinidae</taxonomicName>
, but
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<taxonomicName box="[818,955,176,195]" class="Insecta" family="Philogangidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Philogangidae</taxonomicName>
. The nymphs of both families also resemble each other in general build, with long lateral outer caudal gills and a slightly shorter central caudal gill, although the lateral ones are not tube-shaped in
<taxonomicName authorityName="Kirby" authorityYear="1890" box="[818,915,259,278]" class="Insecta" family="Lestoideidae" genus="Philoganga" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[818,915,259,278]" italics="true" pageId="8" pageNumber="9">Philoganga</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="8.[818,1488,148,920]" pageId="8" pageNumber="9">
For about a century,
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[1042,1228,287,306]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
served as a dustbin family for damselfly genera with unclear relationships. Based on present and recent work (
<bibRefCitation author="Dijkstra, K. - D. - B. &amp; Kalkman, V. J. &amp; Dow, R. A. &amp; Stokvis, F. R. &amp; Van Tol, J. A. N." box="[948,1144,343,362]" journalOrPublisher="Syst. Entomol." pageId="8" pageNumber="9" pagination="68 - 96" part="39" refId="ref17175" refString="Dijkstra, K. - D. - B., Kalkman, V. J., Dow, R. A., Stokvis, F. R., Van Tol, J. A. N., 2014. Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata). Syst. Entomol. 39, 68 - 96." title="Redefining the damselfly families: a comprehensive molecular phylogeny of Z ygoptera (O donata)" type="journal article" year="2014">Dijkstra et al., 2014</bibRefCitation>
;
<bibRefCitation author="Kalkman, V. J. &amp; Theischinger, G." box="[1156,1473,343,362]" journalOrPublisher="Int. J. Odonatol." pageId="8" pageNumber="9" pagination="1 - 52" part="16" refId="ref18079" refString="Kalkman, V. J., Theischinger, G., 2013. Generic revision of Argiolestidae (Odonata), with four new genera. Int. J. Odonatol. 16, 1 - 52." title="Generic revision of Argiolestidae (Odonata), with four new genera" type="journal article" year="2013">Kalkman and Theischinger, 2013</bibRefCitation>
), these are now divided across no less than fifteen families. The true
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[818,1004,399,418]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
are limited to the genera
<emphasis bold="true" italics="true" pageId="8" pageNumber="9">
<taxonomicName authorityName="Selys-Longchamps" authorityYear="1885" box="[1271,1409,398,417]" class="Insecta" family="Megapodagrionidae" genus="Megapodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">Megapodagrion</taxonomicName>
,
<taxonomicName authorityName="Forster" authorityYear="1910" class="Insecta" family="Megapodagrionidae" genus="Allopodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">Allopodagrion</taxonomicName>
</emphasis>
and
<taxonomicName authorityName="De Marmels" authorityYear="2001" box="[937,1075,426,445]" class="Insecta" family="Megapodagrionidae" genus="Teinopodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[937,1075,426,445]" italics="true" pageId="8" pageNumber="9">Teinopodagrion</emphasis>
</taxonomicName>
with a total of only 29 species limited to tropical America. The only genus included in our study (
<taxonomicName authorityName="De Marmels" authorityYear="2001" box="[1341,1479,454,473]" class="Insecta" family="Megapodagrionidae" genus="Teinopodagrion" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="genus">
<emphasis bold="true" box="[1341,1479,454,473]" italics="true" pageId="8" pageNumber="9">Teinopodagrion</emphasis>
</taxonomicName>
) was found to be sister to
<taxonomicName box="[1049,1173,482,501]" class="Insecta" family="Argiolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Argiolestidae</taxonomicName>
(BS = 100, PP = 1, QS = 0.86/0/ 0.99), a group restricted to the Afrotropics and Australasia.
<taxonomicName class="Insecta" family="Argiolestidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Argiolestidae</taxonomicName>
was until recently considered as a subfamily of
<taxonomicName authorityName="Calvert" authorityYear="1913" box="[1295,1484,538,557]" class="Insecta" family="Megapodagrionidae" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Megapodagrionidae</taxonomicName>
, but raised to family level based on the morphology of caudal gills of the nymphs that are distinctively flat and fan-shaped and held in a horizontal plane (
<bibRefCitation author="Kalkman, V. J. &amp; Theischinger, G." box="[947,1253,622,641]" journalOrPublisher="Int. J. Odonatol." pageId="8" pageNumber="9" pagination="1 - 52" part="16" refId="ref18079" refString="Kalkman, V. J., Theischinger, G., 2013. Generic revision of Argiolestidae (Odonata), with four new genera. Int. J. Odonatol. 16, 1 - 52." title="Generic revision of Argiolestidae (Odonata), with four new genera" type="journal article" year="2013">Kalkman and Theischinger 2013</bibRefCitation>
). True megapodagrionid caudal gills lie in a vertical plane with the lateral pair triquetral and the median foliaceous (
<bibRefCitation author="De Marmels, J." box="[1003,1175,678,697]" journalOrPublisher="Boletin de Entomologia Venezolana" pageId="8" pageNumber="9" pagination="27 - 36" part="14" refId="ref17071" refString="De Marmels, J., 1999. A new species of Dimeragrion Calvert 1913 from Pantepui, Venezuela (Odonata: Megapodagrionidae). Boletin de Entomologia Venezolana 14, 27 - 36." title="A new species of Dimeragrion Calvert 1913 from Pantepui, Venezuela (Odonata: Megapodagrionidae)" type="journal article" year="1999">De Marmels, 1999</bibRefCitation>
).
</paragraph>
<paragraph blockId="8.[818,1488,148,920]" pageId="8" pageNumber="9">
Finally, the well supported monophyletic families
<taxonomicName authorityName="de Selys-Longchamps &amp; Hagen" authorityYear="1815" box="[1345,1487,705,724]" class="Insecta" family="Calopterygidae" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Calopterygidae</taxonomicName>
(BS = 100, PP = 1, QS = 1/NA/1) and
<taxonomicName authorityName="Cowley" authorityYear="1937" box="[1181,1329,733,752]" class="Insecta" family="Chlorocyphidae" kingdom="Animalia" order="Odonata" pageId="8" pageNumber="9" phylum="Arthropoda" rank="family">Chlorocyphidae</taxonomicName>
(BS = 100, PP = 1, QS = 1/NA/1) form the core of a group which is colloquially often addressed as Caloptera. However, Caloptera has poor QS values (BS = 100, PP = 1, QS = -0.25/0.23/1). Nonetheless, all species are restricted to running waters and the majority of males have brightly colored (both metallic and pigmented) bodies and often wings used in wonderfully elaborate courtship displays.
</paragraph>
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