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<document 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="15356098BBD55C09991750F497D92C96" 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="Tuberoxenos Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka 2022, gen. nov." docType="treatment" docUuid="99152C5A-B0FE-47A3-85B7-2A3F5ED548DA" docUuidSource="ZooBank" docVersion="5" 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="1668151620981" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>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>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Benda, Daniel</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-5729-0411</mods:nameIdentifier>
<mods:affiliation>Department of Zoology, Faculty of Science, Charles University, Prague, Czech Republic &amp; Department of Entomology, National Museum, Prague, Czech Republic</mods:affiliation>
<mods:nameIdentifier type="email">benda.daniel@email.cz</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Pohl, Hans</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-7090-6612</mods:nameIdentifier>
<mods:affiliation>Institut fuer Zoologie und Evolutionsforschung, Friedrich-Schiller-Universitaet, Jena, Germany</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Nakase, Yuta</mods:namePart>
<mods:affiliation>Department of Biology, Faculty of Science, Shinshu University, Matsumoto, Japan</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Beutel, Rolf</mods:namePart>
<mods:affiliation>Institut fuer Zoologie und Evolutionsforschung, Friedrich-Schiller-Universitaet, Jena, Germany</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Straka, Jakub</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-8987-1245</mods:nameIdentifier>
<mods:affiliation>Department of Zoology, Faculty of Science, Charles University, Prague, Czech Republic</mods:affiliation>
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<mods:title>ZooKeys</mods:title>
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<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-04-07</mods:number>
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<mods:detail type="volume">
<mods:number>1093</mods:number>
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<mods:start>1</mods:start>
<mods:end>134</mods:end>
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<mods:url>http://dx.doi.org/10.3897/zookeys.1093.72339</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.1093.72339</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-1093-1</mods:identifier>
<mods:identifier type="ZooBank">23B7070849A94681AC20494D06F98CCE</mods:identifier>
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<treatment ID-GBIF-Taxon="194520435" LSID="urn:lsid:zoobank.org:act:99152C5A-B0FE-47A3-85B7-2A3F5ED548DA" httpUri="http://treatment.plazi.org/id/15356098BBD55C09991750F497D92C96" lastPageNumber="1" pageId="0" pageNumber="1">
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
<paragraph pageId="0" pageNumber="1">
<taxonomicName LSID="http://zoobank.org/99152C5A-B0FE-47A3-85B7-2A3F5ED548DA" authority="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka, 2022" authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tuberoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus" status="gen. nov.">Tuberoxenos</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="1">gen. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="type species">
<paragraph pageId="0" pageNumber="1">Type species.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" baseAuthorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" baseAuthorityYear="2022" class="Insecta" family="Xenidae" genus="Xenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Xenos sphecidarum" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="sphecidarum">
<emphasis italics="true" pageId="0" pageNumber="1">Xenos sphecidarum</emphasis>
</taxonomicName>
Siebold, 1839, here designated.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="diagnosis">
<paragraph pageId="0" pageNumber="1">Diagnosis of female cephalothorax.</paragraph>
<paragraph pageId="0" pageNumber="1">
Distinguished from
<taxonomicName 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 italics="true" pageId="0" pageNumber="1">Pseudoxenos</emphasis>
</taxonomicName>
by conspicuously convex, round cephalothorax (Fig.
<figureCitation captionStart="Figure 43" captionStartId="F43" captionText="Figure 43. Tuberoxenos sphecidarum, host, male, female, cephalothorax, photomicrographs A Podalonia tydei stylopized by females of T. sphecidarum, lateral view B detail of host abdomen with three adult females inside C ventral side of female cephalothorax D dorsal side of female cephalothorax." figureDoi="10.3897/zookeys.1093.72339.figure43" httpUri="https://binary.pensoft.net/fig/669365" pageId="0" pageNumber="1">43C</figureCitation>
), and distinctly raised, anteriorly protruding dorsal labral field (Fig.
<figureCitation captionStart="Figure 45" captionStartId="F45" captionText="Figure 45. Tuberoxenos sphecidarum, 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, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1">45D</figureCitation>
). Differring from other genera by the following combination of characters. Maxilla well-developed and clearly separated from labial area, prominent and directed anteriorly (Fig.
<figureCitation captionStart="Figure 45" captionStartId="F45" captionText="Figure 45. Tuberoxenos sphecidarum, 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, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1">45E</figureCitation>
). Mandibular tooth narrow or slightly widened. Prosternal extension undifferentiated, evenly arched but in some cases protruding and overlapping with maxillolabial area and posterior part of mandibles. Differing from
<taxonomicName authorityName="Kifune &amp; Maeta" authorityYear="1975" class="Insecta" family="Xenidae" genus="Nipponoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Nipponoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Nipponoxenos</emphasis>
</taxonomicName>
by mandible nested in capsule. In contrast to
<taxonomicName 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 italics="true" pageId="0" pageNumber="1">Paragioxenos</emphasis>
</taxonomicName>
, head and prothorax ventrally delimited by birth opening medially and by suture laterally.
</paragraph>
<caption doi="10.3897/zookeys.1093.72339.figure43" httpUri="https://binary.pensoft.net/fig/669365" pageId="0" pageNumber="1" start="Figure 43" startId="F43">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 43.</emphasis>
<taxonomicName authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" baseAuthorityName="Siebold" baseAuthorityYear="1839" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tuberoxenos sphecidarum" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="sphecidarum">
<emphasis italics="true" pageId="0" pageNumber="1">Tuberoxenos sphecidarum</emphasis>
</taxonomicName>
, host, male, female, cephalothorax, photomicrographs
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
<taxonomicName baseAuthorityName="Le Guillou" baseAuthorityYear="1841" class="Insecta" family="Sphecidae" genus="Podalonia" kingdom="Animalia" lsidName="Podalonia tydei" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="tydei">
<emphasis italics="true" pageId="0" pageNumber="1">Podalonia tydei</emphasis>
</taxonomicName>
stylopized by females of
<taxonomicName lsidName="T. sphecidarum" pageId="0" pageNumber="1" rank="species" species="sphecidarum">
<emphasis italics="true" pageId="0" pageNumber="1">T. sphecidarum</emphasis>
</taxonomicName>
, lateral view
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
detail of host abdomen with three adult females inside
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
ventral side of female cephalothorax
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
dorsal side of female cephalothorax.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph pageId="0" pageNumber="1">Description of female cephalothorax.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Shape and coloration.</emphasis>
Compact, ca. as long as wide or longer than wide. In ventral view appearing conspicuously convex, rotund (Fig.
<figureCitation captionStart="Figure 43" captionStartId="F43" captionText="Figure 43. Tuberoxenos sphecidarum, host, male, female, cephalothorax, photomicrographs A Podalonia tydei stylopized by females of T. sphecidarum, lateral view B detail of host abdomen with three adult females inside C ventral side of female cephalothorax D dorsal side of female cephalothorax." figureDoi="10.3897/zookeys.1093.72339.figure43" httpUri="https://binary.pensoft.net/fig/669365" pageId="0" pageNumber="1">43C</figureCitation>
), high-elliptic in cross-section. Species rather constant in size, length 1.06-1.34 mm, maximum width 0.94-1.4 mm. Anterior head margin evenly rounded or very slightly protruding. Thorax slightly or distinctly widening posteriorly. Coloration with multiple brown shades forming distinct pattern, mostly dark.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Head capsule.</emphasis>
Ca. ⅓ - ⅖ as long as entire cephalothorax including lateral cephalic extension. Coloration of head dominantly pale or brown, forming specific color pattern. Clypeal region well delimited from labral area, arcuate, or very slightly protruding and forming clypeal lobe. Surface smooth or slightly wrinkled. Ca. 50-95 sensilla regularly dispersed on clypeal area. Border between clypeal area and frontal region clearly recognizable or indistinct. Frontal region smooth or slightly wrinkled. Segmental border between head and prothorax quite distinct on dorsal side, indicated by furrow, change in cuticular sculpture or coloration.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Supra-antennal sensillary field.</emphasis>
Slightly wrinkled or reticulated, delimited by more or less distinct furrow on medial side (Fig.
<figureCitation captionStart="Figure 45" captionStartId="F45" captionText="Figure 45. Tuberoxenos sphecidarum, 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, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1">45B</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Antenna.</emphasis>
Preserved as poorly defined area, in some cases indistinct (Fig.
<figureCitation captionStart="Figure 44" captionStartId="F44" captionText="Figure 44. Tuberoxenos sphecidarum, 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. Abbreviations: a - vestigial antenna, paa - periantennal area." figureDoi="10.3897/zookeys.1093.72339.figure44" httpUri="https://binary.pensoft.net/fig/669366" pageId="0" pageNumber="1">44C</figureCitation>
). With cavities, several small, rounded plates, or sensilla, the latter combined in some cases. Periantennal area smooth or slightly wrinkled (Fig.
<figureCitation captionStart="Figure 44" captionStartId="F44" captionText="Figure 44. Tuberoxenos sphecidarum, 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. Abbreviations: a - vestigial antenna, paa - periantennal area." figureDoi="10.3897/zookeys.1093.72339.figure44" httpUri="https://binary.pensoft.net/fig/669366" pageId="0" pageNumber="1">44C</figureCitation>
).
</paragraph>
<caption doi="10.3897/zookeys.1093.72339.figure44" httpUri="https://binary.pensoft.net/fig/669366" pageId="0" pageNumber="1" start="Figure 44" startId="F44">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 44.</emphasis>
<taxonomicName authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" baseAuthorityName="Siebold" baseAuthorityYear="1839" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tuberoxenos sphecidarum" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="sphecidarum">
<emphasis italics="true" pageId="0" pageNumber="1">Tuberoxenos sphecidarum</emphasis>
</taxonomicName>
, female, cephalothorax, SEM micrographs
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
ventral side
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
dorsal side
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
left vestigial antenna, dorsal side
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
right vestigial antenna, dorsal side
<emphasis bold="true" pageId="0" pageNumber="1">E</emphasis>
left lateral border of abdominal segment I below spiracle, dorsal side
<emphasis bold="true" pageId="0" pageNumber="1">F</emphasis>
right lateral border of abdominal segment I below spiracle, dorsal side. Abbreviations: a - vestigial antenna, paa - periantennal area.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Labrum.</emphasis>
Ventral field at least slightly wider than long, elliptical or semicircular. Dorsal field widely arcuate, ~ 5
<normalizedToken originalValue="×">x</normalizedToken>
wider than long in midline, distinctly raised (Fig.
<figureCitation captionStart="Figure 45" captionStartId="F45" captionText="Figure 45. Tuberoxenos sphecidarum, 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, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1">45D</figureCitation>
). Dorsal field with ~ 17-28 pointed or blunt setae on its surface.
</paragraph>
<caption doi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1" start="Figure 45" startId="F45">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 45.</emphasis>
<taxonomicName authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" baseAuthorityName="Siebold" baseAuthorityYear="1839" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tuberoxenos sphecidarum" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="sphecidarum">
<emphasis italics="true" pageId="0" pageNumber="1">Tuberoxenos sphecidarum</emphasis>
</taxonomicName>
, female, cephalothorax, SEM micrographs
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
anterior part of cephalothorax, ventral side
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
anterior part of cephalothorax, dorsal side
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
mouthparts, ventral side
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
detail of anterior border of cephalothorax, ventral side
<emphasis bold="true" pageId="0" pageNumber="1">E</emphasis>
right mandible and maxilla, ventral side
<emphasis bold="true" pageId="0" pageNumber="1">F</emphasis>
left mandible and maxilla, ventral side. Abbreviations: dlf - dorsal field of labral area, fr - frontal region, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Mandible.</emphasis>
Anteromedially directed at angle of 20-40°, enclosed in mandibular capsule. Mandibular bulge slightly or distinctly raised, with several sensilla. Cuticle smooth, slightly sculptured or reticulated. Longitudinal grooves on articular area present. Tooth narrow, pointed, more or less armed with spines.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Maxilla.</emphasis>
Well developed and clearly separated from labial area, prominent and anteriorly directed. Protruding maxillary part usually slightly overlapping with proximal portion of mandible (Fig.
<figureCitation captionStart="Figure 45" captionStartId="F45" captionText="Figure 45. Tuberoxenos sphecidarum, 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, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1">45E</figureCitation>
), but not projecting beyond mandible anteriorly. Cuticle smooth or very slightly wrinkled. Vestige of palp inconspicuous, preserved as small bulge with indistinct plates, located anteromedially on ventral side of maxilla (Fig.
<figureCitation captionStart="Figure 45" captionStartId="F45" captionText="Figure 45. Tuberoxenos sphecidarum, 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, fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla, mxp - vestige of maxillary palp, smxg - submaxillary groove, ssf - supra-antennal sensillary field." figureDoi="10.3897/zookeys.1093.72339.figure45" httpUri="https://binary.pensoft.net/fig/669367" pageId="0" pageNumber="1">45E</figureCitation>
). Maxillary base distinctly produced anterolaterally as submaxillary groove.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Labium.</emphasis>
Labial area distinct between maxillae, delimited anteriorly by mouth opening and posteriorly by birth opening. Labial area wider than long in midline, flat or slightly convex. Cuticular surface smooth or slightly reticulated.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Mouth opening.</emphasis>
Arcuate, nearly straight, or bi-arcuate, sclerotized marginally.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Thorax and abdominal segment I.</emphasis>
Pro-mesothoracic and meso-metathoracic borders distinct, usually separated by mesal furrows, often combined with color stripes or spots on dorsal and ventral sides. Border between metathorax and abdomen usually indicated by change of cuticular sculpture and very indistinct ridge. Cuticle of thoracic segments on ventral side reticulate with scattered small and pigmented papillae. Dorsal side of thorax smooth or slightly reticulated. Prosternal extension undifferentiated, prosternal margin evenly arched but in some cases protruding and overlapping with maxillolabial area and posterior part of mandibles. Meso- and metathorax of standard transverse shape. Setae present on lateral region of abdominal segment I (Fig.
<figureCitation captionStart="Figure 44" captionStartId="F44" captionText="Figure 44. Tuberoxenos sphecidarum, 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. Abbreviations: a - vestigial antenna, paa - periantennal area." figureDoi="10.3897/zookeys.1093.72339.figure44" httpUri="https://binary.pensoft.net/fig/669366" pageId="0" pageNumber="1">44 E, F</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Spiracles.</emphasis>
Spiracles on posterior half or third of cephalothorax slightly elevated, with lateral or anterolateral orientation.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="diagnosis">
<paragraph pageId="0" pageNumber="1">Diagnosis of male cephalotheca.</paragraph>
<paragraph pageId="0" pageNumber="1">
Differing from other genera by the following combination of characters. Diameter of genae between maxillary base and compound eye ~ 1.5
<normalizedToken originalValue="×">x</normalizedToken>
larger than diameter of vestigial antenna. Occipital bulge absent and frontal impression indistinct or missing. Distinct paired furrows of supra-antennal sensillary field present (Fig.
<figureCitation captionStart="Figure 46" captionStartId="F46" captionText="Figure 46. Tuberoxenos sphecidarum, male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla." figureDoi="10.3897/zookeys.1093.72339.figure46" httpUri="https://binary.pensoft.net/fig/669368" pageId="0" pageNumber="1">46A, D</figureCitation>
). Cephalotheca always appearing rotund.
</paragraph>
<caption doi="10.3897/zookeys.1093.72339.figure46" httpUri="https://binary.pensoft.net/fig/669368" pageId="0" pageNumber="1" start="Figure 46" startId="F46">
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Figure 46.</emphasis>
<taxonomicName authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" baseAuthorityName="Siebold" baseAuthorityYear="1839" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tuberoxenos sphecidarum" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="species" species="sphecidarum">
<emphasis italics="true" pageId="0" pageNumber="1">Tuberoxenos sphecidarum</emphasis>
</taxonomicName>
, male, cephalotheca, photomicrographs, SEM micrographs
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
frontal view
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
lateral view
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
vestigial antenna
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
frontal view
<emphasis bold="true" pageId="0" pageNumber="1">E</emphasis>
mouthparts. Abbreviations: fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph pageId="0" pageNumber="1">Description of male cephalotheca.</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Shape and coloration.</emphasis>
In frontal view rounded, almost circular (Fig.
<figureCitation captionStart="Figure 46" captionStartId="F46" captionText="Figure 46. Tuberoxenos sphecidarum, male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla." figureDoi="10.3897/zookeys.1093.72339.figure46" httpUri="https://binary.pensoft.net/fig/669368" pageId="0" pageNumber="1">46A</figureCitation>
), in lateral view pointed anteriorly. Coloration forming pattern of pale and dark shades.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Cephalothecal capsule.</emphasis>
Compound eyes with darker individual ommatidia well visible on pale ocular background. Conspicuous clypeal lobe arcuate in frontal view, prominent in lateral view. Sensilla dispersed over entire clypeal area. Paired furrows of supra-antennal sensillary field distinctly presented but impression lacking on frontal region. Occipital bulge absent. Diameter of genae between maxillary base and compound eye small, ~ 1.5
<normalizedToken originalValue="×">x</normalizedToken>
larger than diameter of vestigial antenna.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Supra-antennal sensillary field.</emphasis>
Kidney-shaped and bulging, medially delimited by distinct furrow (Fig.
<figureCitation captionStart="Figure 46" captionStartId="F46" captionText="Figure 46. Tuberoxenos sphecidarum, male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla." figureDoi="10.3897/zookeys.1093.72339.figure46" httpUri="https://binary.pensoft.net/fig/669368" pageId="0" pageNumber="1">46A, D</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Antenna.</emphasis>
Large, with complete torulus. Periantennal area not clearly delimited from supra-antennal sensillary field. Small plates, cavities and sensilla present (Fig.
<figureCitation captionStart="Figure 46" captionStartId="F46" captionText="Figure 46. Tuberoxenos sphecidarum, male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fssf - furrow of supra-antennal sensillary field, mx - vestige of maxilla." figureDoi="10.3897/zookeys.1093.72339.figure46" httpUri="https://binary.pensoft.net/fig/669368" pageId="0" pageNumber="1">46C</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Labrum.</emphasis>
Labral area distinct. Setae on dorsal field present.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Mandible.</emphasis>
Anteromedially directed. Tooth pointed, not reaching area of mandibular bulge basally. Bulge with sensilla.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Maxilla.</emphasis>
Distinct, prominent. Coloration completely dark or brighter around distinct vestige of maxillary palp.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Labium and hypopharynx.</emphasis>
Labium distinct between and below maxillae, dark. Praementum and postmentum separated by furrow. Hypopharyngeal protuberance indistinct or absent.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Mouth opening.</emphasis>
Well visible, not covered by ventral labral field, slightly arcuate.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="phylogenetic relationships">
<paragraph pageId="0" pageNumber="1">Phylogenetic relationships.</paragraph>
<paragraph pageId="0" pageNumber="1">
Deeply nested within
<taxonomicName class="Insecta" family="Xenidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Xenidae</taxonomicName>
(
<bibRefCitation 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 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>
), part of a clade of an Old Word origin, with
<taxonomicName 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 italics="true" pageId="0" pageNumber="1">Pseudoxenos</emphasis>
</taxonomicName>
Saunders as sister group.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="distribution">
<paragraph pageId="0" pageNumber="1">Diversity and distribution.</paragraph>
<paragraph pageId="0" pageNumber="1">
A lineage of Afrotropical-Palearctic origin, comprising 5 currently valid species, restricted to these regions. It is an example of connectivity between both biogeographic regions (
<bibRefCitation 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>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="host">
<paragraph pageId="0" pageNumber="1">Hosts.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName authorityName="W. Kirby" authorityYear="1798" class="Liliopsida" family="Poaceae" genus="Ammophila" kingdom="Plantae" lsidName="Ammophila" order="Poales" pageId="0" pageNumber="1" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Ammophila</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Fernald" authorityYear="1927" class="Insecta" family="Sphecidae" genus="Podalonia" kingdom="Animalia" lsidName="Podalonia" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Podalonia</emphasis>
</taxonomicName>
spp. (
<taxonomicName authorityName="Latreille" authorityYear="1802" class="Insecta" family="Sphecidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Sphecidae</taxonomicName>
:
<taxonomicName authorityName="Andre" authorityYear="1886" lsidName="" pageId="0" pageNumber="1" rank="subFamily" subFamily="Ammophilinae">Ammophilinae</taxonomicName>
), rarely
<taxonomicName authorityName="Vander Linden" authorityYear="1827" class="Insecta" family="Sphecidae" genus="Prionyx" kingdom="Animalia" lsidName="Prionyx" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Prionyx</emphasis>
</taxonomicName>
spp. (
<taxonomicName authorityName="Latreille" authorityYear="1802" class="Insecta" family="Sphecidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Sphecidae</taxonomicName>
:
<taxonomicName authorityName="Latreille" authorityYear="1802" lsidName="" pageId="0" pageNumber="1" rank="subFamily" subFamily="Sphecinae">Sphecinae</taxonomicName>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="etymology">
<paragraph pageId="0" pageNumber="1">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="1">
From the Latin substantive
<emphasis italics="true" pageId="0" pageNumber="1">tuber</emphasis>
, meaning a swelling. The name refers to conspicuous swellings on the host abdomen caused by protruded xenid specimens under tergites or sternites. Gender masculine.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="comments">
<paragraph pageId="0" pageNumber="1">Comments.</paragraph>
<paragraph pageId="0" pageNumber="1">
All described species of
<taxonomicName authorityName="Benda &amp; Pohl &amp; Nakase &amp; Beutel &amp; Straka" authorityYear="2022" class="Insecta" family="Xenidae" genus="Tuberoxenos" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Tuberoxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Tuberoxenos</emphasis>
</taxonomicName>
gen. nov. were previously placed in
<taxonomicName authorityName="Saunders" authorityYear="1872" class="Insecta" family="Xenidae" genus="Paraxenos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paraxenos" order="Strepsiptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Paraxenos</emphasis>
</taxonomicName>
based on parasitising
<taxonomicName authorityName="Latreille" authorityYear="1802" class="Insecta" family="Sphecidae" kingdom="Animalia" lsidName="" order="Hymenoptera" pageId="0" pageNumber="1" phylum="Arthropoda" rank="family">Sphecidae</taxonomicName>
(
<bibRefCitation author="Kinzelbach, R" journalOrPublisher="Schweizerbart'sche Verlagsbuchhandlung, Stuttgart" pageId="0" pageNumber="1" refId="B54" refString="Kinzelbach, R, 1971b. Morphologische Befunde an Faecherflueglern und ihre phylogenetische Bedeutung (Insecta: Strepsiptera). Schweizerbart'sche Verlagsbuchhandlung, Stuttgart" title="Morphologische Befunde an Faecherflueglern und ihre phylogenetische Bedeutung (Insecta: Strepsiptera)." year="1971 b">Kinzelbach 1971b</bibRefCitation>
). Despite this concept, this group is morphologically well defined. We classify it as a separate genus, based on molecular phylogenies (
<bibRefCitation 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 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 in this paper.
</paragraph>
</subSubSection>
</treatment>
</document>