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<document id="EAAA0454D25EF13DC6E94F0D72130CE8" ID-DOI="10.1111/j.1096-3642.2007.00314.x" ID-ISSN="0024-4082" ID-Zenodo-Dep="5432328" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630429767966" checkinUser="felipe" docAuthor="Sinclair, Bradley J., Borkent, Art &amp; Wood, D. Monty" docDate="2007" docId="0E458F02FF9BB246FC04FD99FEACFA00" docLanguage="en" docName="j.1096-3642.2007.00314.x.pdf" docOrigin="Zoological Journal of the Linnean Society 150 (4)" docSource="https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.2007.00314.x" docStyle="DocumentStyle:0DD8C314D74634CE09062A86991413F8.2:ZoolJLinnSoc.2002-2009.journal_article" docStyleId="0DD8C314D74634CE09062A86991413F8" docStyleName="ZoolJLinnSoc.2002-2009.journal_article" docStyleVersion="2" docTitle="Ceratopogonidae" docType="treatment" docVersion="3" lastPageNumber="730" masterDocId="F27CF77AFF89B255FFEFFF95FFA1FF85" masterDocTitle="The male genital tract and aedeagal components of the Diptera with a discussion of their phylogenetic significance" masterLastPageNumber="742" masterPageNumber="711" pageNumber="729" updateTime="1699266673815" updateUser="plazi" zenodo-license-document="CC-BY-4.0" zenodo-license-figures="CC-BY-4.0">
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<mods:title id="1305E91B069524474E61F508B25BE62D">The male genital tract and aedeagal components of the Diptera with a discussion of their phylogenetic significance</mods:title>
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<mods:namePart id="AFA915D160C7E49206F3BD58CC64D13B">Sinclair, Bradley J.</mods:namePart>
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<mods:namePart id="495C21D917568F3D2093A52C723C43D8">Borkent, Art</mods:namePart>
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<mods:namePart id="59B70A63CF3A9BA37DAF7A79D9DA946E">Wood, D. Monty</mods:namePart>
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<mods:title id="1965507EC912F80FE39581B669533750">Zoological Journal of the Linnean Society</mods:title>
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<treatment id="0E458F02FF9BB246FC04FD99FEACFA00" LSID="urn:lsid:plazi:treatment:0E458F02FF9BB246FC04FD99FEACFA00" httpUri="http://treatment.plazi.org/id/0E458F02FF9BB246FC04FD99FEACFA00" lastPageId="19" lastPageNumber="730" pageId="18" pageNumber="729">
<subSubSection id="CEF66D9FFF9BB247FC04FD99FB68FDA6" box="[1003,1225,524,548]" pageId="18" pageNumber="729" type="nomenclature">
<paragraph id="86533E14FF9BB247FC04FD99FB68FDA6" blockId="18.[1003,1225,524,548]" box="[1003,1225,524,548]" pageId="18" pageNumber="729">
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<taxonomicName id="41EC4597FF9BB247FC04FD99FB68FDA6" ID-CoL="7WQ" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[1003,1225,524,548]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="729" phylum="Arthropoda" rank="family">CERATOPOGONIDAE</taxonomicName>
</heading>
</paragraph>
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<subSubSection id="CEF66D9FFF9BB246FCC9FDA0FEACFA00" lastPageId="19" lastPageNumber="730" pageId="18" pageNumber="729" type="description">
<paragraph id="86533E14FF9BB247FCC9FDA0FBAAFDCF" blockId="18.[806,1035,565,586]" box="[806,1035,565,586]" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9FDA0FBAAFDCF" box="[806,1035,565,586]" italics="true" pageId="18" pageNumber="729">
Description (
<figureCitation id="1ED72291FF9BB247FC5BFDA0FBA2FDCF" box="[948,1027,565,586]" captionStart="Figure 7" captionStartId="16.[142,220,1586,1605]" captionTargetBox="[303,1261,194,1553]" captionTargetId="figure-176@16.[302,1262,193,1554]" captionTargetPageId="16" captionText="Figure 7. Male internal reproductive organs (dorsal view). A, Androprosopa japonica (Thaumaleidae). B, Austrothaumalea zentae (Thaumaleidae) (based on the specimen photographed in Fig. 5D). C, Culicoides nubeculosus (Meigen) (Ceratopogonidae) [modified from Pomerantzev (1932)]. D, Chironomus plumosus (L.) (Chironomidae) [modified from Wensler &amp; Rempel (1962)]. E, Austrosimulium furiosum species group (Simuliidae) (based on the specimen photographed in Fig. 5E). acc gl cplx, accessory gland complex; ej dt, ejaculatory duct; tes, testis; vas df, vas deferens. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5432342" httpUri="https://zenodo.org/record/5432342/files/figure.png" pageId="18" pageNumber="729">Fig. 7C</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="86533E14FF9BB247FCC9FDF6FB6BFD12" blockId="18.[806,1421,611,663]" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9FDF6FCCCFDFD" box="[806,877,611,632]" italics="true" pageId="18" pageNumber="729">Testis:</emphasis>
Each testis is narrow, spindle-shaped, tapered anteriorly, and organized into zones.
</paragraph>
<paragraph id="86533E14FF9BB247FCC9FD3AFB31FD40" blockId="18.[806,1168,687,709]" box="[806,1168,687,709]" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9FD3AFC11FD41" box="[806,944,687,708]" italics="true" pageId="18" pageNumber="729">Epididymis:</emphasis>
Not differentiated.
</paragraph>
<paragraph id="86533E14FF9BB247FCC9FD4BFB9CFC4F" blockId="18.[806,1422,733,970]" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9FD4BFC1AFD77" box="[806,955,733,755]" italics="true" pageId="18" pageNumber="729">Vas deferens:</emphasis>
The vasa deferentia are packed with mature spermatozoa, even at emergence (
<bibRefCitation id="E27D43E5FF9BB247FAAFFD69FCFEFCB4" author="Linley JR" pageId="18" pageNumber="729" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley, 1981</bibRefCitation>
). The ducts extend posteriorly to near segment 9, where they bend abruptly back, become fused to the midline of the accessory gland complex, and run juxtaposed to the anterior apex of the gland. The ducts enter their respective half of the gland through a sphincter (
<bibRefCitation id="E27D43E5FF9BB247FC4EFC21FB91FC4F" author="Linley JR" box="[929,1072,948,970]" pageId="18" pageNumber="729" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley, 1981</bibRefCitation>
).
</paragraph>
<paragraph id="86533E14FF9BB247FCC9FC77FA2EFA02" blockId="18.[806,1423,994,1415]" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9FC77FB7AFC72" box="[806,1243,994,1015]" italics="true" pageId="18" pageNumber="729">Accessory gland and seminal vesicle:</emphasis>
The accessory gland complex comprises three pairs of chambers in
<taxonomicName id="41EC4597FF9BB247FCC9FB8AFC49FBB0" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[806,1000,1055,1077]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="729" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
, united along the midline. The most anterior chamber was referred to as the seminal vesicle by
<bibRefCitation id="E27D43E5FF9BB247FC97FBC8FBB3FBF6" author="Linley JR" box="[888,1042,1117,1139]" pageId="18" pageNumber="729" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley (1981)</bibRefCitation>
, and is lined with an inner layer of narrow secretory cells with spherical vacuoles containing secretions. A narrow aperture separates the seminal vesicle from the middle chamber, referred to as the glutinous gland (
<bibRefCitation id="E27D43E5FF9BB247FBAAFB42FB76FB6B" author="Linley JR" box="[1093,1239,1239,1262]" pageId="18" pageNumber="729" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley, 1981</bibRefCitation>
). The secretory cells of this chamber produce a viscid material, which forms the envelope of the spermatophore. The third chamber is not easily observed in whole mounts and is not lined with glandular cells. The entire accessory gland complex is covered in a layer of circular muscle.
</paragraph>
<paragraph id="86533E14FF9BB247FCC9FA0AFBDCF9CB" blockId="18.[806,1422,1439,1614]" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9FA0AFC4AFA31" box="[806,1003,1439,1460]" italics="true" pageId="18" pageNumber="729">Ejaculatory duct:</emphasis>
This duct comprises two sections. In the anterior half, the ducts remain separate, whereas the ducts unite near the base of the genitalia and in this section the duct is lined with columnar glandular cells (
<bibRefCitation id="E27D43E5FF9BB247FC86F98CFC58F9B5" author="Linley JR" box="[873,1017,1561,1584]" pageId="18" pageNumber="729" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley, 1981</bibRefCitation>
). The ejaculatory duct is sheathed in a layer of circular muscles.
</paragraph>
<paragraph id="86533E14FF9BB246FCC9F9F3FD0AFF7D" blockId="18.[806,1422,1638,1905]" lastBlockId="19.[162,778,195,248]" lastPageId="19" lastPageNumber="730" pageId="18" pageNumber="729">
<emphasis id="B498E206FF9BB247FCC9F9F3FA2CF9FE" box="[806,1421,1638,1660]" italics="true" pageId="18" pageNumber="729">Ejaculatory apodeme, sperm pump, and aedeagus:</emphasis>
The ejaculatory apodeme is absent and the true aedeagus is membranous. The ventral plate with its recurved tip, present in most
<taxonomicName id="41EC4597FF9BB247FB4BF957FAC6F95D" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[1188,1383,1730,1752]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="18" pageNumber="729" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
, is pressed into the opening of the dorsal wall of the common female spermathecal duct and may aid in stretching the opening (
<bibRefCitation id="E27D43E5FF9BB247FBEDF88BFB39F8B1" author="Linley JR" box="[1026,1176,1822,1844]" pageId="18" pageNumber="729" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley, 1981</bibRefCitation>
). The ventral plate (termed aedeagus by authors in this family) is present throughout the family, including
<taxonomicName id="41EC4597FF9BB246FB45F8C9FF48FF5C" authority="Wirth &amp; Lee" authorityName="Wirth &amp; Lee" class="Insecta" family="Ceratopogonidae" genus="Austroconops" kingdom="Animalia" lastPageId="19" lastPageNumber="730" order="Diptera" pageId="18" pageNumber="729" phylum="Arthropoda" rank="genus">
<emphasis id="B498E206FF9BB247FB45F8C9FAE0F8F4" box="[1194,1345,1884,1905]" italics="true" pageId="18" pageNumber="729">Austroconops</emphasis>
Wirth &amp; Lee
</taxonomicName>
, an early lineage of
<taxonomicName id="41EC4597FF9AB246FE3BFF56FD35FF5C" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[468,660,195,217]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
(
<bibRefCitation id="E27D43E5FF9AB246FD4BFF56FE28FF72" author="Borkent A &amp; Wirth WW &amp; Dyce AL" pageId="19" pageNumber="730" pagination="587 - 606" refId="ref20630" refString="Borkent A, Wirth WW, Dyce AL. 1987. The newly discovered male of Austroconops (Ceratopogonidae: Diptera) with a discussion of the phylogeny of the basal lineages of the Ceratopogonidae. Proceedings of the Entomological Society of Washington 89: 587 - 606." type="journal article" year="1987">Borkent, Wirth &amp; Dyce, 1987</bibRefCitation>
;
<bibRefCitation id="E27D43E5FF9AB246FE7AFF77FD3EFF7D" author="Borkent A &amp; Craig DA" box="[405,671,226,248]" pageId="19" pageNumber="730" refId="ref20542" refString="Borkent A, Craig DA. 2004. Austroconops Wirth and Lee, a Lower Cretaceous genus of biting midges yet living in Western Australia: a new species, first description of the immatures and discussion of their biology and phylogeny (Diptera: Ceratopogonidae). American Museum Novitates no. 3449." type="book" year="2004">Borkent &amp; Craig, 2004</bibRefCitation>
).
</paragraph>
<paragraph id="86533E14FF9AB246FF4DFE85FECBFC94" blockId="19.[162,778,272,785]" pageId="19" pageNumber="730">
<emphasis id="B498E206FF9AB246FF4DFE85FEAFFEA0" box="[162,270,272,293]" italics="true" pageId="19" pageNumber="730">Remarks:</emphasis>
The male genital tract of
<taxonomicName id="41EC4597FF9AB246FDA6FE85FCA8FEA3" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[585,777,272,294]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
was first described by
<bibRefCitation id="E27D43E5FF9AB246FE58FEBAFD00FEC0" author="Pomerantzev BI" box="[439,673,303,325]" pageId="19" pageNumber="730" pagination="183 - 214" refId="ref23129" refString="Pomerantzev BI. 1932. Beitrage zur Morphologie und Anatomie der Genitalien von Culicoides (Diptera, Nematocera). Magazin de Parasitologie (Leningrad) 3: 183 - 214 [in Russian, with a German summary]." type="journal article" year="1932">Pomerantzev (1932)</bibRefCitation>
and the method of spermatophore formation in
<taxonomicName id="41EC4597FF9AB246FD7BFED8FEA4FE07" authority="Latreille" authorityName="Latreille" authorityYear="1809" class="Insecta" family="Ceratopogonidae" genus="Culicoides" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="genus">
<emphasis id="B498E206FF9AB246FD7BFED8FCABFEE7" box="[660,778,333,354]" italics="true" pageId="19" pageNumber="730">Culicoides</emphasis>
Latreille
</taxonomicName>
was thoroughly described and excellently illustrated by
<bibRefCitation id="E27D43E5FF9AB246FEA4FE1EFE49FE24" author="Linley JR" box="[331,488,395,417]" pageId="19" pageNumber="730" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley (1981)</bibRefCitation>
. Two packages of sperm are prepared in the long, narrow, tapering glutinous gland (middle chamber) and are pushed into paired, secreted sheaths upon exiting, which fuse medially further down the ejaculatory duct. The spermatophore is then passed to the female during copulation. As observed by
<bibRefCitation id="E27D43E5FF9AB246FEDFFDD6FE68FDDC" author="Linley JR" box="[304,457,579,601]" pageId="19" pageNumber="730" pagination="332 - 346" refId="ref22288" refString="Linley JR. 1981. Ejaculation and spermatophore formation in Culicoides melleus (Coq.) (Diptera: Ceratopogonidae). Canadian Journal of Zoology 59: 332 - 346." type="journal article" year="1981">Linley (1981)</bibRefCitation>
, spermatozoa remain in the vasa deferentia until ejaculation. In the present study, spermatozoa-packed vasa deferentia were observed in freshly collected, nonmated males of
<taxonomicName id="41EC4597FF9AB246FDB6FD0AFD72FD30" box="[601,723,671,693]" class="Insecta" family="Bibionidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Bibionidae</taxonomicName>
and
<taxonomicName id="41EC4597FF9AB246FF4DFD28FEE7FD56" baseAuthorityName="Wood &amp; Borkent" baseAuthorityYear="1989" box="[162,326,701,723]" class="Insecta" family="Thaumaleidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Thaumaleidae</taxonomicName>
(
<figureCitation id="1ED72291FF9AB246FEB7FD28FE75FD56" box="[344,468,701,723]" captionStart="Figure 3" captionStartId="8.[142,220,1579,1598]" captionTargetBox="[303,1259,197,1547]" captionTargetId="figure-161@8.[302,1262,196,1547]" captionTargetPageId="8" captionText="Figure 3. Male internal reproductive organs (dorsal view). A, Hesperinus rohdendorfi (Hesperinidae). B, Plecia membranifera (Bibionidae). C, Bibio flavihalter (Bibionidae), left vas deferens and testis removed (based on the specimen photographed in Fig. 4A). D, Pachyneura fasciata (Pachyneuridae). acc gl cplx, accessory gland complex; ej dt, ejaculatory duct; tes, testis; vas df, vas deferens. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5432334" httpUri="https://zenodo.org/record/5432334/files/figure.png" pageId="19" pageNumber="730">Figs 3B, D</figureCitation>
,
<figureCitation id="1ED72291FF9AB246FE0CFD2BFDA4FD51" box="[483,517,702,724]" captionStart="Figure 7" captionStartId="16.[142,220,1586,1605]" captionTargetBox="[303,1261,194,1553]" captionTargetId="figure-176@16.[302,1262,193,1554]" captionTargetPageId="16" captionText="Figure 7. Male internal reproductive organs (dorsal view). A, Androprosopa japonica (Thaumaleidae). B, Austrothaumalea zentae (Thaumaleidae) (based on the specimen photographed in Fig. 5D). C, Culicoides nubeculosus (Meigen) (Ceratopogonidae) [modified from Pomerantzev (1932)]. D, Chironomus plumosus (L.) (Chironomidae) [modified from Wensler &amp; Rempel (1962)]. E, Austrosimulium furiosum species group (Simuliidae) (based on the specimen photographed in Fig. 5E). acc gl cplx, accessory gland complex; ej dt, ejaculatory duct; tes, testis; vas df, vas deferens. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5432342" httpUri="https://zenodo.org/record/5432342/files/figure.png" pageId="19" pageNumber="730">7A</figureCitation>
), suggesting a similar process of spermatophore production as is present in
<taxonomicName id="41EC4597FF9AB246FF4DFD6EFEC7FC94" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[162,358,763,785]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
.
</paragraph>
<paragraph id="86533E14FF9AB246FF4DFCBCFE34FB93" blockId="19.[162,778,809,1413]" pageId="19" pageNumber="730">
<emphasis id="B498E206FF9AB246FF4DFCBCFDF8FCBB" box="[162,601,809,830]" italics="true" pageId="19" pageNumber="730">
Remarks – features of
<taxonomicName id="41EC4597FF9AB246FE4CFCBCFDF4FCBB" authorityName="Macquart" authorityYear="1838" box="[419,597,809,830]" class="Insecta" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="738" phylum="Arthropoda" rank="superFamily" superFamily="Chironomoidea">Chironomoidea</taxonomicName>
.
</emphasis>
The accessory gland complex of
<taxonomicName id="41EC4597FF9AB246FE97FCD2FD87FCD8" authorityName="Macquart" authorityYear="1838" box="[376,550,839,861]" class="Insecta" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="738" phylum="Arthropoda" rank="superFamily" superFamily="Chironomoidea">Chironomoidea</taxonomicName>
is similar in form among all four families. The elongate second chamber with its glandular cells is similar in
<taxonomicName id="41EC4597FF9AB246FDADFC10FCA7FC1E" baseAuthorityName="Linley" baseAuthorityYear="1981" box="[578,774,901,923]" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
,
<taxonomicName id="41EC4597FF9AB246FF4DFC36FEBFFC3C" authorityName="Newman" authorityYear="1834" box="[162,286,931,953]" class="Insecta" family="Simuliidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Simuliidae</taxonomicName>
(
<figureCitation id="1ED72291FF9AB246FEDBFC36FE29FC3C" box="[308,392,931,953]" captionStart="Figure 5" captionStartId="12.[142,220,1269,1288]" captionTargetBox="[305,1262,194,1237]" captionTargetId="figure-333@12.[302,1262,194,1237]" captionTargetPageId="12" captionText="Figure 5. Male internal reproductive organs (dorsal view). A, Trichocera sp. (Trichoceridae). B, Dixa yamatoma (Dixidae). C, Dixa yamatoma (Dixidae). D, Austrothaumalea zentae (Thaumaleidae). E, Austrosimulium furiosum group (Simuliidae). F, Chironomus plumosus (Chironomidae). acc gl cplx, accessory gland complex; ej dt, ejaculatory duct; tes, testis; vas df, vas deferens. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5432338" httpUri="https://zenodo.org/record/5432338/files/figure.png" pageId="19" pageNumber="730">Fig. 5C</figureCitation>
;
<bibRefCitation id="E27D43E5FF9AB246FE74FC36FDE2FC3C" author="Rubtsov IA" box="[411,579,931,953]" pageId="19" pageNumber="730" pagination="1 - 1042" refId="ref23292" refString="Rubtsov IA. 1989. Blackflies (Simuliidae). Fauna of the USSR 6 (6): 1 - 1042." type="journal article" year="1989">Rubtsov, 1989</bibRefCitation>
: fig. 10A, epsd), and
<taxonomicName id="41EC4597FF9AB246FF3BFC57FED4FC5D" box="[212,373,962,984]" class="Insecta" family="Chironomidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Chironomidae</taxonomicName>
(
<figureCitation id="1ED72291FF9AB246FE6BFC57FE77FC5D" box="[388,470,962,984]" captionStart="Figure 5" captionStartId="12.[142,220,1269,1288]" captionTargetBox="[305,1262,194,1237]" captionTargetId="figure-333@12.[302,1262,194,1237]" captionTargetPageId="12" captionText="Figure 5. Male internal reproductive organs (dorsal view). A, Trichocera sp. (Trichoceridae). B, Dixa yamatoma (Dixidae). C, Dixa yamatoma (Dixidae). D, Austrothaumalea zentae (Thaumaleidae). E, Austrosimulium furiosum group (Simuliidae). F, Chironomus plumosus (Chironomidae). acc gl cplx, accessory gland complex; ej dt, ejaculatory duct; tes, testis; vas df, vas deferens. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5432338" httpUri="https://zenodo.org/record/5432338/files/figure.png" pageId="19" pageNumber="730">Fig. 5F</figureCitation>
). Only in
<taxonomicName id="41EC4597FF9AB246FDABFC57FD49FC5D" baseAuthorityName="Wood &amp; Borkent" baseAuthorityYear="1989" box="[580,744,962,984]" class="Insecta" family="Thaumaleidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Thaumaleidae</taxonomicName>
do the vasa deferentia enter the accessory/seminal gland subapically (
<figureCitation id="1ED72291FF9AB246FEDBFC6AFE28FB93" box="[308,393,1023,1046]" captionStart="Figure 7" captionStartId="16.[142,220,1586,1605]" captionTargetBox="[303,1261,194,1553]" captionTargetId="figure-176@16.[302,1262,193,1554]" captionTargetPageId="16" captionText="Figure 7. Male internal reproductive organs (dorsal view). A, Androprosopa japonica (Thaumaleidae). B, Austrothaumalea zentae (Thaumaleidae) (based on the specimen photographed in Fig. 5D). C, Culicoides nubeculosus (Meigen) (Ceratopogonidae) [modified from Pomerantzev (1932)]. D, Chironomus plumosus (L.) (Chironomidae) [modified from Wensler &amp; Rempel (1962)]. E, Austrosimulium furiosum species group (Simuliidae) (based on the specimen photographed in Fig. 5E). acc gl cplx, accessory gland complex; ej dt, ejaculatory duct; tes, testis; vas df, vas deferens. Scale bars = 0.1 mm." figureDoi="http://doi.org/10.5281/zenodo.5432342" httpUri="https://zenodo.org/record/5432342/files/figure.png" pageId="19" pageNumber="730">Fig. 7A</figureCitation>
).
</paragraph>
<paragraph id="86533E14FF9AB246FF54FB8BFEACFA00" blockId="19.[162,778,809,1413]" pageId="19" pageNumber="730">
The genital tract of the
<taxonomicName id="41EC4597FF9AB246FE3BFB8BFD23FBB1" authorityName="Macquart" authorityYear="1838" box="[468,642,1054,1076]" class="Insecta" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="738" phylum="Arthropoda" rank="superFamily" superFamily="Chironomoidea">Chironomoidea</taxonomicName>
is modified for the passage of a preformed spermatophore (
<bibRefCitation id="E27D43E5FF9AB246FD2CFBA8FF7AFBF4" author="Wood DM" pageId="19" pageNumber="730" pagination="1297 - 1337" refId="ref24053" refString="Wood DM. 1978. Taxonomy of the Nearctic species of Twinnia and Gymnopais (Diptera: Simuliidae) and a discussion of the ancestry of the Simuliidae. Canadian Entomologist 110: 1297 - 1337." type="journal article" year="1978">Wood, 1978</bibRefCitation>
;
<bibRefCitation id="E27D43E5FF9AB246FF07FBCEFE53FBF7" author="Wood DM &amp; Borkent A" box="[232,498,1115,1138]" pageId="19" pageNumber="730" pagination="1333 - 1370" refId="ref24188" refString="Wood DM, Borkent A. 1989. Phylogeny and classification of the Nematocera. Agriculture Canada Monograph 32: 1333 - 1370." type="journal article" year="1989">Wood &amp; Borkent, 1989</bibRefCitation>
), the existence of which has been confirmed in the families
<taxonomicName id="41EC4597FF9AB246FDC3FBEFFD0BFB15" authorityName="Newman" authorityYear="1834" box="[556,682,1146,1168]" class="Insecta" family="Simuliidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Simuliidae</taxonomicName>
,
<taxonomicName id="41EC4597FF9AB246FD5AFBEFFEB6FB2A" baseAuthorityName="Linley" baseAuthorityYear="1981" class="Insecta" family="Ceratopogonidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Ceratopogonidae</taxonomicName>
, and
<taxonomicName id="41EC4597FF9AB246FEB0FB0CFDA3FB2A" box="[351,514,1177,1199]" class="Insecta" family="Chironomidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Chironomidae</taxonomicName>
, and presumably also the
<taxonomicName id="41EC4597FF9AB246FF38FB22FEDCFB48" baseAuthorityName="Wood &amp; Borkent" baseAuthorityYear="1989" box="[215,381,1207,1229]" class="Insecta" family="Thaumaleidae" kingdom="Animalia" order="Diptera" pageId="19" pageNumber="730" phylum="Arthropoda" rank="family">Thaumaleidae</taxonomicName>
, given the configuration of the internal genital tract (
<bibRefCitation id="E27D43E5FF9AB246FE44FB43FDFAFB69" author="Sinclair BJ" box="[427,603,1238,1260]" pageId="19" pageNumber="730" pagination="491 - 499" refId="ref23433" refString="Sinclair BJ. 1992 a. A new species of Trichothaumalea (Diptera: Thaumaleidae) from eastern North America and a discussion of male genitalic homologies. Canadian Entomologist 124: 491 - 499." type="journal article" year="1992">Sinclair, 1992a</bibRefCitation>
). The enlarged accessory gland complex is quite stout and generally easily identified in whole mount dissections, whereas the vasa deferentia and testes are often more fragile, the latter often distinctively clothed in pigment/fat granules.
</paragraph>
</subSubSection>
</treatment>
</document>
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