<documentID-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 & Straka, Jakub"docDate="2022"docId="AF90B145A30D532BAB557499256C1D5B"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="Pseudoxenos Saunders 1872"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">
<mods:titleid="7CFC0B6658093B241F7BCF2FCDDC227A">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:affiliationid="014CDE23BB59C1174934ED157B858D76">Department of Zoology, Faculty of Science, Charles University, Prague, Czech Republic & Department of Entomology, National Museum, Prague, Czech Republic</mods:affiliation>
<mods:affiliationid="A1F24AAD1DD064326E5505EB723DA76B">Department of Biology, Faculty of Science, Shinshu University, Matsumoto, Japan</mods:affiliation>
<mods:affiliationid="DA1AAD7ADE7ED63B03591F029786A982">Department of Zoology, Faculty of Science, Charles University, Prague, Czech Republic</mods:affiliation>
<taxonomicNameid="C644B6688E28138CC3FA5989013E9021"authorityName="Benda & Pohl & Nakase & Beutel & 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">
<figureCitationid="6C06F707997A57932D9D32055D25C638"captionStart="Figure 41"captionStartId="F41"captionText="Figure 41. Pseudoxenos sp., 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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field."figureDoi="10.3897/zookeys.1093.72339.figure41"httpUri="https://binary.pensoft.net/fig/669363"pageId="0"pageNumber="1">41C</figureCitation>
<figureCitationid="8C65FD0F7543AC7A07EFE710FF2D6643"captionStart="Figure 39"captionStartId="F39"captionText="Figure 39. Pseudoxenos sp., host, male, female, cephalothorax, photomicrographs A Paradontodynerus sp. stylopized by male of Pseudoxenos sp., lateral view B detail of host abdomen with male puparium inside C ventral side of female cephalothorax D dorsal side of female cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax."figureDoi="10.3897/zookeys.1093.72339.figure39"httpUri="https://binary.pensoft.net/fig/669361"pageId="0"pageNumber="1">39C</figureCitation>
<taxonomicNameid="5B00EF2F054D166B9EFF35B3E8B11326"authorityName="Benda & Pohl & Nakase & Beutel & 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">
<figureCitationid="845442499F7E9298F74A164A5D9CC3D9"captionStart="Figure 41"captionStartId="F41"captionText="Figure 41. Pseudoxenos sp., 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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field."figureDoi="10.3897/zookeys.1093.72339.figure41"httpUri="https://binary.pensoft.net/fig/669363"pageId="0"pageNumber="1">41C, D</figureCitation>
<figureCitationid="1790B8592552286BD38258461D16C955"captionStart="Figure 39"captionStartId="F39"captionText="Figure 39. Pseudoxenos sp., host, male, female, cephalothorax, photomicrographs A Paradontodynerus sp. stylopized by male of Pseudoxenos sp., lateral view B detail of host abdomen with male puparium inside C ventral side of female cephalothorax D dorsal side of female cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax."figureDoi="10.3897/zookeys.1093.72339.figure39"httpUri="https://binary.pensoft.net/fig/669361"pageId="0"pageNumber="1">39D</figureCitation>
Compact, longer than wide, elliptic in cross-section. Meso-metathoracic segmental border not constricted laterally. Size fairly constant, length 1.08-1.44 mm, maximum width 1.02-1.4 mm. Anterior head margin rounded or protruding. Thorax slightly widening posteriorly. Coloration with multiple brown shades forming pattern.
Ca. ⅖ as long as entire cephalothorax including lateral extensions. Coloration mostly dark brown, often with specific patterns. Clypeal region delimited from labral area (Fig.
<figureCitationid="DFD69432724B0DE5C7B6F751CB99D4FA"captionStart="Figure 41"captionStartId="F41"captionText="Figure 41. Pseudoxenos sp., 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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field."figureDoi="10.3897/zookeys.1093.72339.figure41"httpUri="https://binary.pensoft.net/fig/669363"pageId="0"pageNumber="1">41D</figureCitation>
), arcuate, or protruding and forming clypeal lobe. Surface smooth or slightly wrinkled. Approximately 35-56 sensilla mainly concentrated anteriorly but dispersed over entire clypeal area. Border between clypeal area and frontal region hardly distinct but still recognizable. Frontal surface smooth (Fig.
<figureCitationid="0C924841546312BD8BBE205FF6DA79C8"captionStart="Figure 40"captionStartId="F40"captionText="Figure 40. Pseudoxenos sp., 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 detail of anterior border of cephalothorax, dorsal side. Abbreviations: a - vestigial antenna, fr - frontal region."figureDoi="10.3897/zookeys.1093.72339.figure40"httpUri="https://binary.pensoft.net/fig/669362"pageId="0"pageNumber="1">40F</figureCitation>
). Segmental border between head and prothorax clearly recognizable or indistinct on dorsal side, often indicated by dark brown stripes, and in some cases with two distinct dark spots on mesal region (Fig.
<figureCitationid="1F988210EA240198DACBF854E1FC6C85"captionStart="Figure 39"captionStartId="F39"captionText="Figure 39. Pseudoxenos sp., host, male, female, cephalothorax, photomicrographs A Paradontodynerus sp. stylopized by male of Pseudoxenos sp., lateral view B detail of host abdomen with male puparium inside C ventral side of female cephalothorax D dorsal side of female cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax."figureDoi="10.3897/zookeys.1093.72339.figure39"httpUri="https://binary.pensoft.net/fig/669361"pageId="0"pageNumber="1">39D</figureCitation>
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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field.
<figureCitationid="138A49D8FA7B1A4CF40B9203E730202F"captionStart="Figure 41"captionStartId="F41"captionText="Figure 41. Pseudoxenos sp., 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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field."figureDoi="10.3897/zookeys.1093.72339.figure41"httpUri="https://binary.pensoft.net/fig/669363"pageId="0"pageNumber="1">41B</figureCitation>
<figureCitationid="4B43C360CCCE0986753557CDC2537FC9"captionStart="Figure 40"captionStartId="F40"captionText="Figure 40. Pseudoxenos sp., 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 detail of anterior border of cephalothorax, dorsal side. Abbreviations: a - vestigial antenna, fr - frontal region."figureDoi="10.3897/zookeys.1093.72339.figure40"httpUri="https://binary.pensoft.net/fig/669362"pageId="0"pageNumber="1">40C</figureCitation>
<figureCitationid="3505E7A312870A6328040052DC74847E"captionStart="Figure 41"captionStartId="F41"captionText="Figure 41. Pseudoxenos sp., 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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field."figureDoi="10.3897/zookeys.1093.72339.figure41"httpUri="https://binary.pensoft.net/fig/669363"pageId="0"pageNumber="1">41C, D</figureCitation>
Mandibles anteromedially directed at an angle of 35-45° and nested in mandibular capsule. Mandibular bulge not or slightly raised, bears several sensilla. Cuticle of mandible sculptured to nearly smooth. Mandibular tooth narrow, pointed, straight or hook-shaped, armed with spines.
<figureCitationid="EB2834FD6D7B41F1C25A64E4241B43FC"captionStart="Figure 41"captionStartId="F41"captionText="Figure 41. Pseudoxenos sp., 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, sbhp - segmental border between head and prothorax, ssf - supra-antennal sensillary field."figureDoi="10.3897/zookeys.1093.72339.figure41"httpUri="https://binary.pensoft.net/fig/669363"pageId="0"pageNumber="1">41F</figureCitation>
), but not projecting beyond it anteriorly. Cuticle usually smooth, rarely wrinkled. Vestige of palp very distinct, with more or less distinct plates or cavity, located medially on ventral side of maxilla. Submaxillary groove more or less distinctly produced posterolaterally to maxillary base.
Labial area between maxillae flat but distinct, relatively large, delimited anteriorly by mouth opening and posteriorly by birth opening. As long as wide or longer than wide. Cuticular surface in most cases largely smooth and shiny, or faintly and uniformly sculptured.
Pro-mesothoracic and meso-metathoracic borders more or less distinct, separated by mesal furrows. Border between metathorax and abdomen formed by ridge. Cuticle of thoracic segments on ventral side reticulate with scattered small and pigmented papillae. Cuticle of dorsal side of thorax smooth or slightly wrinkled. Prosternal extension undifferentiated, anterior margin evenly arched. Meso- and metathorax transverse. Lateral parts of abdomen posterior to spiracle dark (Fig.
<figureCitationid="DA003C1DD81F8C2DFB62D5CC08FB8945"captionStart="Figure 39"captionStartId="F39"captionText="Figure 39. Pseudoxenos sp., host, male, female, cephalothorax, photomicrographs A Paradontodynerus sp. stylopized by male of Pseudoxenos sp., lateral view B detail of host abdomen with male puparium inside C ventral side of female cephalothorax D dorsal side of female cephalothorax. Abbreviations: asI - abdominal segment I, sbhp - segmental border between head and prothorax."figureDoi="10.3897/zookeys.1093.72339.figure39"httpUri="https://binary.pensoft.net/fig/669361"pageId="0"pageNumber="1">39D</figureCitation>
<figureCitationid="38495FE561074894A986834C3A8DC2FD"captionStart="Figure 42"captionStartId="F42"captionText="Figure 42. Pseudoxenos sp., male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fi - frontal impression, ob - occipital bulge."figureDoi="10.3897/zookeys.1093.72339.figure42"httpUri="https://binary.pensoft.net/fig/669364"pageId="0"pageNumber="1">42D</figureCitation>
<figureCitationid="969A16E8CA0277FE75E2505C9DE363AB"captionStart="Figure 42"captionStartId="F42"captionText="Figure 42. Pseudoxenos sp., male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fi - frontal impression, ob - occipital bulge."figureDoi="10.3897/zookeys.1093.72339.figure42"httpUri="https://binary.pensoft.net/fig/669364"pageId="0"pageNumber="1">42D</figureCitation>
Compound eyes with darker individual ommatidia well visible on pale ocular background. Clypeal lobe straight or slightly arcuate in frontal view, prominent in lateral view. Sensilla mainly concentrated medially. Frontal impression distinctly present (Fig.
<figureCitationid="D43AA9367DE1EBF6876A670DCD4A12F2"captionStart="Figure 42"captionStartId="F42"captionText="Figure 42. Pseudoxenos sp., male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fi - frontal impression, ob - occipital bulge."figureDoi="10.3897/zookeys.1093.72339.figure42"httpUri="https://binary.pensoft.net/fig/669364"pageId="0"pageNumber="1">42D</figureCitation>
<figureCitationid="5A0978A2F82EBF366966950D50967EA8"captionStart="Figure 42"captionStartId="F42"captionText="Figure 42. Pseudoxenos sp., male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fi - frontal impression, ob - occipital bulge."figureDoi="10.3897/zookeys.1093.72339.figure42"httpUri="https://binary.pensoft.net/fig/669364"pageId="0"pageNumber="1">42D</figureCitation>
Vestiges large, with complete torulus. Periantennal area not clearly delimited from supra-antennal sensillary field. Small plates or sensilla present (Fig.
<figureCitationid="ED831EC3D3BF779BA367C2F6A0124C39"captionStart="Figure 27"captionStartId="F27"captionText="Figure 27. Leionotoxenos sp., male, cephalotheca, photomicrographs, SEM micrographs A frontal view B lateral view C vestigial antenna D frontal view E mouthparts. Abbreviations: fi - frontal impression, mx - vestige of maxilla, ob - occipital bulge."figureDoi="10.3897/zookeys.1093.72339.figure27"httpUri="https://binary.pensoft.net/fig/669349"pageId="0"pageNumber="1">27C</figureCitation>
<bibRefCitationid="EEF634216FB4A9D6AFFE6567B52B3EAB"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>
<bibRefCitationid="6E43CB392C6C9D7DDEB7EC257AAF6CEE"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>
<taxonomicNameid="38C8545B908C49A9E5A653B651BF45BD"authorityName="Benda & Pohl & Nakase & Beutel & 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">
<bibRefCitationid="058FF9A3721A0DE2FFC4668630ABB0E7"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>
<bibRefCitationid="0C04AF331E192CD2E6240FE81BB15CE7"DOI="https://doi.org/10.1111/j.1365-2311.1872.tb01884.x"author="Saunders, SS"journalOrPublisher="Transactions of the Entomological Society of London"pageId="0"pageNumber="1"pagination="1 - 48"refId="B103"refString="Saunders, SS, 1872. Stylopidarum, ordinem Strepsipterorum Kirbii constituentium, mihi tamen potius Coleopterorum Familiae, Rhipiphoridis Meloidisque propinquae, Monographia. Transactions of the Entomological Society of London 20: 1 - 48, DOI: https://doi.org/10.1111/j.1365-2311.1872.tb01884.x"title="Stylopidarum, ordinem Strepsipterorum Kirbii constituentium, mihi tamen potius Coleopterorum Familiae, Rhipiphoridis Meloidisque propinquae, Monographia."url="https://doi.org/10.1111/j.1365-2311.1872.tb01884.x"volume="20"year="1872">Saunders (1872)</bibRefCitation>
<bibRefCitationid="72E87CFE0AEE676B1587E75E3278FA84"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>
<bibRefCitationid="BD337CCBE33DB96E9B77A9AB88F07102"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>
<bibRefCitationid="27A4052CE1DCCF8D0EC26FF571776365"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>
<bibRefCitationid="F491628542731670E042D22759D0841D"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>