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<document ID-CLB-Dataset="55433" ID-DOI="http://dx.doi.org/10.3897/zookeys.1093.72339" ID-GBIF-Dataset="8bb856a5-26cb-4abc-8e4a-04d3716144b1" ID-PMC="PMC9010403" ID-Pensoft-Pub="1313-2970-1093-1" ID-Pensoft-UUID="D3A8D50FF61A5B61B8776D63EB0D3F4C" ID-PubMed="35586542" ID-ZooBank="23B7070849A94681AC20494D06F98CCE" ModsDocID="1313-2970-1093-1" checkinTime="1649449468463" checkinUser="pensoft" docAuthor="Benda, Daniel, Pohl, Hans, Nakase, Yuta, Beutel, Rolf &amp; Straka, Jakub" docDate="2022" docId="CE4A08BBFB945E69BC46E97B9D430985" docLanguage="en" docName="ZooKeys 1093: 1-134" docOrigin="ZooKeys 1093" docPubDate="2022-04-07" docSource="http://dx.doi.org/10.3897/zookeys.1093.72339" docTitle="Macroxenos Schultze 1925" docType="treatment" docVersion="7" id="D3A8D50FF61A5B61B8776D63EB0D3F4C" lastPageNumber="1" masterDocId="D3A8D50FF61A5B61B8776D63EB0D3F4C" masterDocTitle="A generic classification of Xenidae (Strepsiptera) based on the morphology of the female cephalothorax and male cephalotheca with a preliminary checklist of species" masterLastPageNumber="134" masterPageNumber="1" pageNumber="1" updateTime="1732615197594" updateUser="ExternalLinkService">
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<mods:title id="F8EDD30C2AD63AC514BD1E85D9AA0D88">A generic classification of Xenidae (Strepsiptera) based on the morphology of the female cephalothorax and male cephalotheca with a preliminary checklist of species</mods:title>
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<mods:namePart id="38175550CA0A403FD3C5E1ECFF2AC766">Benda, Daniel</mods:namePart>
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<mods:namePart id="3057282DA8523D934EEAC46DBAFE4F56">Pohl, Hans</mods:namePart>
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<mods:affiliation id="0F36B5EB6469E62940727A5B34DC7AFD">Institut fuer Zoologie und Evolutionsforschung, Friedrich-Schiller-Universitaet, Jena, Germany</mods:affiliation>
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<mods:namePart id="37F1497CB7593E7F13C0FA6A1B068D4E">Nakase, Yuta</mods:namePart>
<mods:affiliation id="01AA12CF9A984853D54C1CF2D43438F4">Department of Biology, Faculty of Science, Shinshu University, Matsumoto, Japan</mods:affiliation>
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<treatment id="CE4A08BBFB945E69BC46E97B9D430985" ID-GBIF-Taxon="194520406" LSID="urn:lsid:plazi:treatment:CE4A08BBFB945E69BC46E97B9D430985" httpUri="http://treatment.plazi.org/id/CE4A08BBFB945E69BC46E97B9D430985" lastPageNumber="1" pageId="0" pageNumber="1" scope_family="Xenidae" scope_order="Strepsiptera">
<subSubSection id="91F1B57BA7DEE7D442016AC9F0B681A8" pageId="0" pageNumber="1" type="nomenclature">
<paragraph id="535DAF12ED43F21E2168D085674DEC06" pageId="0" pageNumber="1">
<taxonomicName id="FDEBF4BBEAB51F45942F71C3FBBF733C" ID-CoL="BGJFR" LSID="CE4A08BB-FB94-5E69-BC46-E97B9D430985" authority="Schultze, 1925, stat. res." authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Macroxenos Schultze, 1925, stat. res.</taxonomicName>
</paragraph>
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<paragraph id="A16853A942A9C5AC6CF26D383CF1BF9E" pageId="0" pageNumber="1">
<taxonomicName id="3940F73F5F55C2C83D026E3C3B9F695A" authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Macroxenos</taxonomicName>
Schultze, 1925: 238. Type species:
<taxonomicName id="588A03210B61CB732A26CFD3F76BC7B9" authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos piercei" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="piercei">Macroxenos piercei</taxonomicName>
Schultze, 1925, by original designation.
</paragraph>
<paragraph id="7AA691F3AA4E579E74FDCE60E1211087" pageId="0" pageNumber="1">
<taxonomicName id="DE8D8F91DCE03834E33BAE6C93EE6C31" authorityName="Saunders" authorityYear="1872" class="Insecta" family="Xenidae" genus="Pseudoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">Pseudoxenos</taxonomicName>
Saunders, 1872 (partim!) (synonymy proposed by
<bibRefCitation id="ED89B9A6F0C46642815756C4B598EE83" DOI="https://doi.org/10.1155/1937/28072" author="Bohart, RM" journalOrPublisher="Psyche: A Journal of Entomology" pageId="0" pageNumber="1" pagination="132 - 137" refId="B7" refString="Bohart, RM, 1937. Synonymy of the Genus Pseudoxenos Saunders (Strepsiptera, Xenidae) and records of stylopized Hymenoptera from North Carolina. Psyche: A Journal of Entomology 44: 132 - 137, DOI: https://doi.org/10.1155/1937/28072" title="Synonymy of the Genus Pseudoxenos Saunders (Strepsiptera, Xenidae) and records of stylopized Hymenoptera from North Carolina." url="https://doi.org/10.1155/1937/28072" volume="44" year="1937">Bohart 1937</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="32EE18CADBBA16A5E52C433E685E19D8" pageId="0" pageNumber="1" type="diagnosis">
<paragraph id="87015C8B50AC531000814F1890714BAC" pageId="0" pageNumber="1">Diagnosis of female cephalothorax.</paragraph>
<paragraph id="49C7C8A9FCB266AA195C15ED264F34E2" pageId="0" pageNumber="1">
Maxilla reduced, not distinctly prominent (Fig.
<figureCitation id="A11BFBF080957BF3788938D95868B0E8" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34E</figureCitation>
). Two distinct dark spots present mesally on border between head and prothorax (Fig.
<figureCitation id="E0584A0FCB2B440FE6EE14E16C833C60" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Macroxenos cf. piercei, host, female, cephalothorax, photomicrographs A Anterhynchium flavomarginatum stylopized by female of M. cf. piercei, lateral view B detail of host abdomen with adult female inside C ventral side of cephalothorax D dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax." figureDoi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1">32D</figureCitation>
). Thoracic segments conspicuously sclerotized laterally from dorsal side (Fig.
<figureCitation id="EEF0E9894064999AA209B3BCF7360B98" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Macroxenos cf. piercei, host, female, cephalothorax, photomicrographs A Anterhynchium flavomarginatum stylopized by female of M. cf. piercei, lateral view B detail of host abdomen with adult female inside C ventral side of cephalothorax D dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax." figureDoi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1">32D</figureCitation>
). Lateral parts of abdomen posterior to spiracles always pale (Fig.
<figureCitation id="48EFC19454C13F6AD0AB66777C573DD7" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Macroxenos cf. piercei, host, female, cephalothorax, photomicrographs A Anterhynchium flavomarginatum stylopized by female of M. cf. piercei, lateral view B detail of host abdomen with adult female inside C ventral side of cephalothorax D dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax." figureDoi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1">32D</figureCitation>
). Clypeal region bulging, very distinctly separated from labral area (Fig.
<figureCitation id="D2B5850EAD67BEDDC3672EAB379581D3" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34D</figureCitation>
). Mandible not protruding from capsule. In contrast to
<taxonomicName id="7790B50D0E200E831BB32ADD7B057D39" authorityName="Ogloblin" authorityYear="1923" class="Insecta" family="Xenidae" genus="Paragioxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paragioxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="DF5A90D1050374B29647383BC0B0FED2" italics="true" pageId="0" pageNumber="1">Paragioxenos</emphasis>
</taxonomicName>
, head and prothorax ventrally delimited by birth opening medially and by suture laterally.
</paragraph>
</subSubSection>
<subSubSection id="440F758C614EACF90C719DEEA3123CE8" pageId="0" pageNumber="1" type="description">
<paragraph id="25B962D6AB1E3EE0D14459C6C86ACE01" pageId="0" pageNumber="1">Description of female cephalothorax.</paragraph>
<paragraph id="164099820942D3048347601B171D23B4" pageId="0" pageNumber="1">
<emphasis id="B7B5796CA6588DE0BC8C6A2483BAEEE4" bold="true" pageId="0" pageNumber="1">Shape and coloration.</emphasis>
Nearly as long as wide, or as long as or distinctly longer than wide. Very variable in size, length 0.8-1.82 mm, width 0.64-1.9 mm in midline. Anterior head margin evenly rounded or protruding. Thorax slightly or distinctly widening posteriorly. Cephalothorax with multiple brown shades forming distinct pattern.
</paragraph>
<paragraph id="FA701C262C71265C6486D38B0FA6378E" pageId="0" pageNumber="1">
<emphasis id="655159324A6B558684E1A5B5D44960B4" bold="true" pageId="0" pageNumber="1">Head capsule.</emphasis>
Between ⅓ and&gt;
<normalizedToken id="4A6A2FFB09ADE808B5E90C94DF2A9E12" originalValue="½">1/2</normalizedToken>
<normalizedToken id="4201C9199268F85850491E47DA60F8E1" originalValue="×">x</normalizedToken>
as long as entire cephalothorax including the lateral cephalic extensions. Coloration forming specific pattern with pale and dark shades. Clypeal region very distinctly delimited from labral area (Fig.
<figureCitation id="4D8C9F0EA44CBD48F7A031A990258718" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34D</figureCitation>
), arcuate, or protruding and forming clypeal lobe. Surface smooth or distinctly wrinkled. Sensilla mainly concentrated on clypeal lobe. Border between clypeal area and frontal region clearly indicated by change in cuticular surface. Cuticle of frontal area variable, distinctly wrinkled or covered with papillae. Border between head and prothorax usually distinct on dorsal side, delimited by transverse stripe of distinctive coloration and two distinct dark spots on mesal region (Fig.
<figureCitation id="13F8CBEA49635D7F228FF6FF6BB41EBA" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Macroxenos cf. piercei, host, female, cephalothorax, photomicrographs A Anterhynchium flavomarginatum stylopized by female of M. cf. piercei, lateral view B detail of host abdomen with adult female inside C ventral side of cephalothorax D dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax." figureDoi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1">32D</figureCitation>
).
</paragraph>
<paragraph id="7C29F224935D722296812B9CB8576A7C" pageId="0" pageNumber="1">
<emphasis id="B7D09DE4E17D805F12EB3534BFAEC869" bold="true" pageId="0" pageNumber="1">Supra-antennal sensillary field.</emphasis>
Smooth, with dispersed sensilla. Furrow between supra-antennal sensillary field and frontal region absent, or very indistinct and only indicated by change in cuticular sculpture (Fig.
<figureCitation id="CF2DC270FBAA6BD920C7CAC75620D467" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34B</figureCitation>
).
</paragraph>
<paragraph id="A58FEF13090BA87531E0419DC77B1263" pageId="0" pageNumber="1">
<emphasis id="F71DCCB4EE56005BF3E3BE68E0E12843" bold="true" pageId="0" pageNumber="1">Antenna.</emphasis>
Preserved as poorly defined area, with several small, rounded plates, antennal sensilla, or cavity, in some cases all three combined. Periantennal area smooth or slightly wrinkled, sometimes indistinct.
</paragraph>
<paragraph id="537F2E709818F11850E0A0A04F71AF21" pageId="0" pageNumber="1">
<emphasis id="470F30DC2D8D56056919833F47604AA1" bold="true" pageId="0" pageNumber="1">Labrum.</emphasis>
Ventral field wider than long, elliptic to nearly circular. Dorsal field arcuate, distinctly raised (Fig.
<figureCitation id="9AA5FCB868D7F168AC3A9B596B77045F" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34D</figureCitation>
), sometimes very wide and narrow, ~ 5-8
<normalizedToken id="8DA8AFB74F8D07A603F2EDC2DDB818FD" originalValue="×">x</normalizedToken>
wider than long in midline. Dorsal field with 14-41 (or more) setae or sensilla inserted in cavities.
</paragraph>
<paragraph id="D03FC34359AC68F11A6C12583926E0D6" pageId="0" pageNumber="1">
<emphasis id="C9017377A7CA757D9FD236C3D194422D" bold="true" pageId="0" pageNumber="1">Mandible.</emphasis>
Anteromedially directed at angle of 30-35° and enclosed in mandibular capsule. Mandibular bulge distinctly raised, with several sensilla. Cuticle smooth or slightly sculptured, sometimes with longitudinal grooves (Fig.
<figureCitation id="CF4D78CF02B8594D6F6603DAD472F4CE" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34E</figureCitation>
). Tooth narrow or slightly widened, pointed apically or ventrally, more or less distinctly armed with spines.
</paragraph>
<paragraph id="16028D71CCBD2B143C408D792573B0C0" pageId="0" pageNumber="1">
<emphasis id="1C164AD8370B279432C652F6E1A07214" bold="true" pageId="0" pageNumber="1">Maxilla.</emphasis>
Almost completely fused with labial area, or slightly raised (Fig.
<figureCitation id="4A13AD52F57606A39D4C590A7DF849F2" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34E</figureCitation>
), not projecting beyond mandible. Cuticle smooth or wrinkled. Vestige of palp present as cavity or poorly defined area; usually located medially on ventral side of maxilla (Fig.
<figureCitation id="8D3A2FEF685243CB32152FE0051A310A" captionStart="Figure 34" captionStartId="F34" captionText="Figure 34. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A anterior part of cephalothorax, ventral side B anterior part of cephalothorax, dorsal side C mouthparts, ventral side D detail of anterior border of cephalothorax, ventral side E right mandible and maxilla, ventral side F left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1">34E</figureCitation>
). Submaxillary groove more or less distinctly produced anterolaterally to maxillary base.
</paragraph>
<paragraph id="AA4C7FBE9B3565F022D9B7E1ADD1F375" pageId="0" pageNumber="1">
<emphasis id="803E066EBFCA94A4281F8CE490D28556" bold="true" pageId="0" pageNumber="1">Labium.</emphasis>
Labial area between maxillae usually more or less distinct, delimited anteriorly by mouth opening and posteriorly by birth opening. Labial area wider than long in midline, flat or convex. Cuticular surface smooth or reticulated.
</paragraph>
<paragraph id="7950B5ADDBED350FED2C7CC2180CEEBA" pageId="0" pageNumber="1">
<emphasis id="F75C8DE33ABBDB9F7349538BDE3A8489" bold="true" pageId="0" pageNumber="1">Mouth opening.</emphasis>
Widely arcuate, sclerotized marginally.
</paragraph>
<paragraph id="9E46541ED04B9ADC3D9894D519B85391" pageId="0" pageNumber="1">
<emphasis id="FDEACC4D28439B1A58D26ED659BF79F2" bold="true" pageId="0" pageNumber="1">Thorax and abdominal segment I.</emphasis>
Pro-mesothoracic and meso-metathoracic borders more or less distinct, usually separated by mesal furrows, rarely combined with pigmented stripes or spots on dorsal and ventral side. Border between metathorax and abdomen usually formed by ridge or indicated by change in cuticular sculpture. Cuticle of thoracic segments on ventral side reticulate, with scattered inconspicuous or more distinct pigmented papillae. Dorsal surface of thorax smooth or slightly reticulated. Prosternal extension undifferentiated or distinct, in some cases extremely elongated. Thoracic segments conspicuously sclerotized laterally from dorsal side (Fig.
<figureCitation id="42F6961FA77BF1E49C8F5152517FAA78" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Macroxenos cf. piercei, host, female, cephalothorax, photomicrographs A Anterhynchium flavomarginatum stylopized by female of M. cf. piercei, lateral view B detail of host abdomen with adult female inside C ventral side of cephalothorax D dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax." figureDoi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1">32D</figureCitation>
). Shape of meso- and metathorax unmodified, transverse, or narrowed laterally in species with elongated head. Lateral parts of abdomen posterior to spiracles always pale (Fig.
<figureCitation id="44737C2BE15AE357D5FA1789A8CC3CA9" captionStart="Figure 32" captionStartId="F32" captionText="Figure 32. Macroxenos cf. piercei, host, female, cephalothorax, photomicrographs A Anterhynchium flavomarginatum stylopized by female of M. cf. piercei, lateral view B detail of host abdomen with adult female inside C ventral side of cephalothorax D dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax." figureDoi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1">32D</figureCitation>
). Setae present on lateral region of abdominal segment I (Fig.
<figureCitation id="7A139E2781B012C8E619A1AE4E4CFB9B" captionStart="Figure 33" captionStartId="F33" captionText="Figure 33. Macroxenos cf. piercei, female, cephalothorax, SEM micrographs A ventral side B dorsal side C left vestigial antenna, dorsal side D right vestigial antenna, dorsal side E left lateral border of abdominal segment I below spiracle, dorsal side F right lateral border of abdominal segment I below spiracle, dorsal side." figureDoi="10.3897/zookeys.1093.72339.figure33" httpUri="https://binary.pensoft.net/fig/669355" pageId="0" pageNumber="1">33E, F</figureCitation>
).
</paragraph>
<caption id="3EA073C64278350B9DAC8B736DF3E794" doi="10.3897/zookeys.1093.72339.figure32" httpUri="https://binary.pensoft.net/fig/669354" pageId="0" pageNumber="1" start="Figure 32" startId="F32">
<paragraph id="25686063D421AB7440030F8F93535CCC" pageId="0" pageNumber="1">
<emphasis id="837B9AFB3B729B5A3421B96F99D25CF5" bold="true" pageId="0" pageNumber="1">Figure 32.</emphasis>
<taxonomicName id="5D7C0AA4D3CBEB602A676C96F4DB9ED2" authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos piercei" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="piercei">Macroxenos cf. piercei</taxonomicName>
, host, female, cephalothorax, photomicrographs
<emphasis id="9F2C7AADDFD038B4DE5A073F7011CB7A" bold="true" pageId="0" pageNumber="1">A</emphasis>
<taxonomicName id="8CF01D6208E43FCD6DC76450D551B598" baseAuthorityName="Smith" baseAuthorityYear="1852" class="Insecta" family="Eumenidae" genus="Anterhynchium" kingdom="Animalia" lsidName="Anterhynchium flavomarginatum" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="flavomarginatum">
<emphasis id="9296F859223FA3D6773AB7765B86ACB8" italics="true" pageId="0" pageNumber="1">Anterhynchium flavomarginatum</emphasis>
</taxonomicName>
stylopized by female of
<taxonomicName id="51F19B883D51E8A5E2373031DCAA0120" lsidName="M. piercei" pageId="0" pageNumber="1" rank="species" species="piercei">M. cf. piercei</taxonomicName>
, lateral view
<emphasis id="F477B3416D1181BA5BE1F9C269695C87" bold="true" pageId="0" pageNumber="1">B</emphasis>
detail of host abdomen with adult female inside
<emphasis id="BE9FE5C487D3A9525FF8DD471D9D9A37" bold="true" pageId="0" pageNumber="1">C</emphasis>
ventral side of cephalothorax
<emphasis id="62F4EE1FDCA7EE2A96A81672D8627593" bold="true" pageId="0" pageNumber="1">D</emphasis>
dorsal side of cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax, tx - thorax.
</paragraph>
</caption>
<caption id="3592520065920DDAEAD87BCFBF07779D" doi="10.3897/zookeys.1093.72339.figure33" httpUri="https://binary.pensoft.net/fig/669355" pageId="0" pageNumber="1" start="Figure 33" startId="F33">
<paragraph id="4076409BEE8F04A39EBF9126CDF09A16" pageId="0" pageNumber="1">
<emphasis id="2E0DE79ED538354AAB962C6754750BA4" bold="true" pageId="0" pageNumber="1">Figure 33.</emphasis>
<taxonomicName id="8ADE4706CF64FB45C3736CFB93DCB93D" authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos piercei" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="piercei">Macroxenos cf. piercei</taxonomicName>
, female, cephalothorax, SEM micrographs
<emphasis id="29D134353213630C6FAB9ACD8B55647C" bold="true" pageId="0" pageNumber="1">A</emphasis>
ventral side
<emphasis id="8E68841550560C068A60EDD6FBB3F957" bold="true" pageId="0" pageNumber="1">B</emphasis>
dorsal side
<emphasis id="796AD3AB33AB303C6DB579868466DCFF" bold="true" pageId="0" pageNumber="1">C</emphasis>
left vestigial antenna, dorsal side
<emphasis id="D6F7F2B508CC2ED0CED1385AFADEE132" bold="true" pageId="0" pageNumber="1">D</emphasis>
right vestigial antenna, dorsal side
<emphasis id="4C9740A9A5EC026915E7C13B9A7BC342" bold="true" pageId="0" pageNumber="1">E</emphasis>
left lateral border of abdominal segment I below spiracle, dorsal side
<emphasis id="6272F3F7CDD9BEF5AC918495C212BFA1" bold="true" pageId="0" pageNumber="1">F</emphasis>
right lateral border of abdominal segment I below spiracle, dorsal side.
</paragraph>
</caption>
<caption id="E6EA85883708984CDC03F903376DBF86" doi="10.3897/zookeys.1093.72339.figure34" httpUri="https://binary.pensoft.net/fig/669356" pageId="0" pageNumber="1" start="Figure 34" startId="F34">
<paragraph id="8BBEFDC736E7D0B0DC54B5F3C57AAAB5" pageId="0" pageNumber="1">
<emphasis id="5C4648E50E91D3701CCD0E516C8F1C05" bold="true" pageId="0" pageNumber="1">Figure 34.</emphasis>
<taxonomicName id="182A65FEAB953BEF622849299C7751FF" authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos piercei" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="piercei">Macroxenos cf. piercei</taxonomicName>
, female, cephalothorax, SEM micrographs
<emphasis id="5AAA0552FB909E17D73FE3C0AB83C13F" bold="true" pageId="0" pageNumber="1">A</emphasis>
anterior part of cephalothorax, ventral side
<emphasis id="7DF2A1235CE7444B777C7BAE9D9D2CDE" bold="true" pageId="0" pageNumber="1">B</emphasis>
anterior part of cephalothorax, dorsal side
<emphasis id="BC5A92D5449AD52ED71AFF2E03393151" bold="true" pageId="0" pageNumber="1">C</emphasis>
mouthparts, ventral side
<emphasis id="9335166B8CCF6819C7313A25DB2ED908" bold="true" pageId="0" pageNumber="1">D</emphasis>
detail of anterior border of cephalothorax, ventral side
<emphasis id="FA465654B83C3EEF3F31C99C14FA86CF" bold="true" pageId="0" pageNumber="1">E</emphasis>
right mandible and maxilla, ventral side
<emphasis id="1DF2F78D7D7C4ED34E9630B713F7E9B5" bold="true" pageId="0" pageNumber="1">F</emphasis>
left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, md - mandible, mx - vestige of maxilla, mxp - vestige of maxillary palp, sbcl - segmental border between clypeus and labrum, smxg - submaxillary groove, ssf - supra-antennal sensillary field.
</paragraph>
</caption>
<paragraph id="BECCF7545AD882826C8F4E64EBC852B6" pageId="0" pageNumber="1">
<emphasis id="F826A5FA3F273C69016BDEE759B5EB0A" bold="true" pageId="0" pageNumber="1">Spiracles.</emphasis>
Spiracles on posterior ~ ⅓ of cephalothorax slightly elevated, with anterodorsal and anterolateral orientation.
</paragraph>
</subSubSection>
<subSubSection id="E5C7A3B394887A01DFFD21CE82EE9908" pageId="0" pageNumber="1" type="diagnosis">
<paragraph id="DEF4C5710211FCF7474E91FACBDD58FB" pageId="0" pageNumber="1">Diagnosis of male cephalotheca.</paragraph>
<paragraph id="C351B13C09C80A2BEF3EDABEB1026E6A" pageId="0" pageNumber="1">Male cephalotheca unknown.</paragraph>
</subSubSection>
<subSubSection id="2465D28E406EA1959506DA4D6607199E" pageId="0" pageNumber="1" type="phylogenetic relationships">
<paragraph id="ECB4008A3CF18D8B76A78EF123F76C70" pageId="0" pageNumber="1">Phylogenetic relationships.</paragraph>
<paragraph id="83C3D13F85A04B122EB541BF7BDC70AF" pageId="0" pageNumber="1">
The phylogenetic position is unstable.
<bibRefCitation id="54761E39B16798136C241ED0B725119B" DOI="https://doi.org/10.1016/j.ympev.2019.02.023" author="Benda, D" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="1" pagination="67 - 77" refId="B3" refString="Benda, D, Nakase, Y, Straka, J, 2019. Frozen Antarctic path for dispersal initiated parallel host-parasite evolution on different continents. Molecular Phylogenetics and Evolution 135: 67 - 77, DOI: https://doi.org/10.1016/j.ympev.2019.02.023" title="Frozen Antarctic path for dispersal initiated parallel host-parasite evolution on different continents." url="https://doi.org/10.1016/j.ympev.2019.02.023" volume="135" year="2019">Benda et al. (2019)</bibRefCitation>
revealed it as sister to a lineage including
<taxonomicName id="CA905F7A8999182A749A5DDEAC63620F" authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Sphecixenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphecixenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="099AB004ABAC67B213E6B02C4B89214E" italics="true" pageId="0" pageNumber="1">Sphecixenos</emphasis>
</taxonomicName>
,
<taxonomicName id="12C4C9DB403F357C5DC5DCEB39E30E5F" authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tuberoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="31C576E8D8BD1A32DCB264AC911B5DF9" italics="true" pageId="0" pageNumber="1">Tuberoxenos</emphasis>
</taxonomicName>
, and
<taxonomicName id="B887B56FB96DD9BE19B83C1E732FCB7F" authorityName="Saunders" authorityYear="1872" class="Insecta" family="Xenidae" genus="Pseudoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="148BDB9A40F55F12D5C7927ABEA8E85C" italics="true" pageId="0" pageNumber="1">Pseudoxenos</emphasis>
</taxonomicName>
in our concept. In contrast,
<bibRefCitation id="D79C695B0DB82BDC51538331EF50672F" DOI="https://doi.org/10.1111/syen.12460" author="Benda, D" journalOrPublisher="Systematic Entomology" pageId="0" pageNumber="1" pagination="252 - 265" refId="B4" refString="Benda, D, Votypkova, K, Nakase, Y, Straka, J, 2021. Unexpected cryptic species diversity of parasites of the family Xenidae (Strepsiptera) with a constant diversification rate over time. Systematic Entomology 46: 252 - 265, DOI: https://doi.org/10.1111/syen.12460" title="Unexpected cryptic species diversity of parasites of the family Xenidae (Strepsiptera) with a constant diversification rate over time." url="https://doi.org/10.1111/syen.12460" volume="46" year="2021">Benda et al. (2021)</bibRefCitation>
resolved its position as sister to a clade including
<taxonomicName id="D588D120FC4BF153E73D95E6E6C1973C" authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Sphecixenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Sphecixenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="8F313B50A4FF92E5C2642C82832CE7EE" italics="true" pageId="0" pageNumber="1">Sphecixenos</emphasis>
</taxonomicName>
,
<taxonomicName id="7B03E4FA7A31855CE3DBB821331EEB36" authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Tuberoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="7D154D10C039D4F29F65F123C7BCADC4" italics="true" pageId="0" pageNumber="1">Tuberoxenos</emphasis>
</taxonomicName>
,
<taxonomicName id="FB90659B08307B759D1B106FCCC05C49" authorityName="Saunders" authorityYear="1872" class="Insecta" family="Xenidae" genus="Pseudoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="5A1C06F2E0043B868D1DC558BF633D66" italics="true" pageId="0" pageNumber="1">Pseudoxenos</emphasis>
</taxonomicName>
,
<taxonomicName id="D60591A30D263BA8ADD2F81EDB0ABAC1" authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Deltoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Deltoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="BC8F27903A2E7A03D2669E8C4E5F6AC9" italics="true" pageId="0" pageNumber="1">Deltoxenos</emphasis>
</taxonomicName>
, and
<taxonomicName id="2B86011601E374CFD93B0EB5DDC9213E" authorityName="Rossi" authorityYear="1794" class="Insecta" family="Xenidae" genus="Xenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="65980DD4A04B6155D9D91A58F007A704" italics="true" pageId="0" pageNumber="1">Xenos</emphasis>
</taxonomicName>
. In both cases, the support was very weak. Further phylogenomic investigations with robust data are needed to resolve the intergeneric relationships.
</paragraph>
</subSubSection>
<subSubSection id="9D5F7F4FC6E445A785D61EE31599D787" pageId="0" pageNumber="1" type="distribution">
<paragraph id="F2F4A40B4428EE819AD15D099C486F1C" pageId="0" pageNumber="1">Diversity and distribution.</paragraph>
<paragraph id="1D28BF85954AD25A09B83D3963CE2BCC" pageId="0" pageNumber="1">
A lineage of Australasian origin, with dispersion into the Indomalayan region (
<bibRefCitation id="04C3126A740F699830C5A0C877ECCB4E" DOI="https://doi.org/10.1016/j.ympev.2019.02.023" author="Benda, D" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="1" pagination="67 - 77" refId="B3" refString="Benda, D, Nakase, Y, Straka, J, 2019. Frozen Antarctic path for dispersal initiated parallel host-parasite evolution on different continents. Molecular Phylogenetics and Evolution 135: 67 - 77, DOI: https://doi.org/10.1016/j.ympev.2019.02.023" title="Frozen Antarctic path for dispersal initiated parallel host-parasite evolution on different continents." url="https://doi.org/10.1016/j.ympev.2019.02.023" volume="135" year="2019">Benda et al. 2019</bibRefCitation>
). The two currently known species are restricted to these two biogeographic regions.
</paragraph>
</subSubSection>
<subSubSection id="4731A776868874D2E8AF7ABC52D420C6" pageId="0" pageNumber="1" type="host">
<paragraph id="C6B0A3F7A0BBAC7B1DC742730A6EDE1F" pageId="0" pageNumber="1">Hosts.</paragraph>
<paragraph id="312668F088ABE7B6DD5026FD7D98C5C4" pageId="0" pageNumber="1">
Various genera of
<taxonomicName id="1958225FB33162832CFBA0758F330582" lsidName="" pageId="0" pageNumber="1" rank="tribe" tribe="Odynerini">Odynerini</taxonomicName>
(
<taxonomicName id="CCC170E147C4DE56A47902850B8E5643" class="Insecta" family="Vespidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Vespidae</taxonomicName>
:
<taxonomicName id="E4E685752F4EE3C8CE6A0A91997DE461" authorityName="Leach" authorityYear="1815" lsidName="" pageId="0" pageNumber="1" rank="subFamily" subFamily="Eumeninae">Eumeninae</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection id="54E43DCD01B7E3AB3F016AB013E528AE" pageId="0" pageNumber="1" type="comments">
<paragraph id="430EEE6A5217A4354C2D80A7BD11891A" pageId="0" pageNumber="1">Comments.</paragraph>
<paragraph id="21CEAD229ABE0E90D8654D3A8D57388F" pageId="0" pageNumber="1">
The genus
<taxonomicName id="1E1FBFDA9A05E388CD731BE23A51C503" authorityName="Schultze" authorityYear="1925" class="Insecta" family="Xenidae" genus="Macroxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Macroxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="215D8D03B709BDB971EE155E01EABBF5" italics="true" pageId="0" pageNumber="1">Macroxenos</emphasis>
</taxonomicName>
was described by
<bibRefCitation id="AEDB52013F641FCA246B92ECB80B15CE" author="Schultze, W" journalOrPublisher="Phillippine Journal of Science" pageId="0" pageNumber="1" pagination="235 - 241" refId="B104" refString="Schultze, W, 1925. Macroxenos piercei (Order Strepsiptera), a new genus and species of wasp parasites of the Philippine Islands. Phillippine Journal of Science 27: 235 - 241" title="Macroxenos piercei (Order Strepsiptera), a new genus and species of wasp parasites of the Philippine Islands." volume="27" year="1925">Schultze (1925)</bibRefCitation>
but the descriptions of male and female was superficial. Later,
<bibRefCitation id="8FD1F2B35C7BFF0E2FF69B09C0153EBA" DOI="https://doi.org/10.1155/1937/28072" author="Bohart, RM" journalOrPublisher="Psyche: A Journal of Entomology" pageId="0" pageNumber="1" pagination="132 - 137" refId="B7" refString="Bohart, RM, 1937. Synonymy of the Genus Pseudoxenos Saunders (Strepsiptera, Xenidae) and records of stylopized Hymenoptera from North Carolina. Psyche: A Journal of Entomology 44: 132 - 137, DOI: https://doi.org/10.1155/1937/28072" title="Synonymy of the Genus Pseudoxenos Saunders (Strepsiptera, Xenidae) and records of stylopized Hymenoptera from North Carolina." url="https://doi.org/10.1155/1937/28072" volume="44" year="1937">Bohart (1937)</bibRefCitation>
synonymized it with
<taxonomicName id="5892C71CAFCA46763AFB3027F1F29961" authorityName="Saunders" authorityYear="1872" class="Insecta" family="Xenidae" genus="Pseudoxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis id="4B4515973C7834597A814D7DD8C075E7" italics="true" pageId="0" pageNumber="1">Pseudoxenos</emphasis>
</taxonomicName>
. We classify this lineage as a separate genus, based on molecular phylogenies (
<bibRefCitation id="C53BA685497B51DBF20DF6789EBC801B" DOI="https://doi.org/10.1016/j.ympev.2019.02.023" author="Benda, D" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="1" pagination="67 - 77" refId="B3" refString="Benda, D, Nakase, Y, Straka, J, 2019. Frozen Antarctic path for dispersal initiated parallel host-parasite evolution on different continents. Molecular Phylogenetics and Evolution 135: 67 - 77, DOI: https://doi.org/10.1016/j.ympev.2019.02.023" title="Frozen Antarctic path for dispersal initiated parallel host-parasite evolution on different continents." url="https://doi.org/10.1016/j.ympev.2019.02.023" volume="135" year="2019">Benda et al. 2019</bibRefCitation>
,
<bibRefCitation id="C074635DE37B00D22F1C95594DA12A92" DOI="https://doi.org/10.1111/syen.12460" author="Benda, D" journalOrPublisher="Systematic Entomology" pageId="0" pageNumber="1" pagination="252 - 265" refId="B4" refString="Benda, D, Votypkova, K, Nakase, Y, Straka, J, 2021. Unexpected cryptic species diversity of parasites of the family Xenidae (Strepsiptera) with a constant diversification rate over time. Systematic Entomology 46: 252 - 265, DOI: https://doi.org/10.1111/syen.12460" title="Unexpected cryptic species diversity of parasites of the family Xenidae (Strepsiptera) with a constant diversification rate over time." url="https://doi.org/10.1111/syen.12460" volume="46" year="2021">2021</bibRefCitation>
) and morphological characters newly reported here. However, this genus is quite complicated to diagnose because of a high morphological variability of species. More samples are still needed for a better characterization and recognition of this formerly overlooked group.
</paragraph>
</subSubSection>
</treatment>
</document>