Editing Strepsiptera (section)

==Taxonomy==
[[File:Mengenilla moldrzyki.jpg|left|thumb|''Mengenilla moldrzyki'' (Mengenilidae)]]
The order, named by [[William Kirby (entomologist)|William Kirby]] in 1813,<ref>{{Cite book|url=https://archive.org/details/biostor-235159|title=A monographic revision of the twisted winged insects comprising the order Strepsiptera Kirby|publisher=Government Printing Office|last=Pierce|first=William Dwight|year=1909|pages=[https://archive.org/details/biostor-235159/page/n212 209]}}</ref> is named for the [[hindwing]]s, which are held at a twisted angle when at rest (from [[Greek language|Greek]] {{wikt-lang|grc|στρέφω|στρέϕειν}} ({{transliteration|grc|strephein}}), to twist; and {{wikt-lang|grc|πτερόν}} ({{transliteration|grc|pteron}}), wing).<ref>{{Cite web|url=http://tolweb.org/Strepsiptera/8222|title=Strepsiptera|last=Kathirithamby|first=Jeyaraney|date=2002|website=The Tree of Life Web Project|archive-url=https://web.archive.org/web/20170715064446/http://tolweb.org/Strepsiptera/8222|archive-date=July 15, 2017}}</ref> The forewings are reduced to [[halteres]].

[[File:Odynerus spinipes^ Vespidae. See parasite note - Flickr - gailhampshire.jpg|thumb|A wasp (''[[Odynerus spinipes]]'') with a small portion of a strepsipteran's body protruding from its abdomen]]

Strepsiptera were once believed to be the sister group to the beetle families [[Meloidae]] and [[Ripiphoridae]], which have similar parasitic development and forewing reduction. Early molecular research suggested their inclusion as a sister group to the [[flies]],<ref name=insencyc/> in a clade called Halteria,<ref>{{cite journal | last1 = Whiting | first1 = Michael F. | year = 1998 | title = Long-Branch Distraction and the Strepsiptera | journal = Systematic Biology | volume = 47 | issue = 1 | pages = 134–138 | doi = 10.1080/106351598261076 | pmid = 12064233 | doi-access = free }}</ref> which have one pair of the wings modified into halteres,<ref name=strepprob>{{cite journal | last1 = Whiting | first1 = Michael F. | last2 = Carpenter | first2 = James C. | last3 = Wheeler | first3 = Quentin D. | last4 = Wheeler | first4 = Ward C. | year = 1997 | title = The Stresiptera Problem: Phylogeny of the Holometabolous Insect Orders Inferred from 18S and 28S Ribosomal DNA Sequences and Morphology | journal = Systematic Biology | volume = 46 | issue = 1| pages = 1–68 | doi=10.2307/2413635| jstor = 2413635 | pmid = 11975347 }}</ref> and failed to support their relationship to the beetles.<ref name=strepprob/> Further molecular studies, however, suggested they are outside the clade Mecopterida (containing the Diptera and Lepidoptera), but found no strong evidence for affinity with any other extant group.<ref>{{cite journal | last1 = Bonneton | first1 = F. | last2 = Brunet | first2 = F. G. | last3 = Kathirithamby | first3 = J. | last4 = Laudet | first4 = V. | year = 2006 | title = The rapid divergence of the ecdysone receptor is a synapomorphy for Mecopterida that clarifies the Strepsiptera problem | journal = Insect Molecular Biology | volume = 15 | issue = 3| pages = 351–362 | doi=10.1111/j.1365-2583.2006.00654.x | pmid=16756554| s2cid = 25178911 }}</ref> Study of their evolutionary position has been problematic due to difficulties in phylogenetic analysis arising from [[long branch attraction]].<ref>{{cite journal | last1 = Huelsenbeck | first1 = John P | year = 1998 | title = Systematic Bias in Phylogenetic Analysis: Is the Strepsiptera Problem Solved? | pmid = 12066692 | journal = Systematic Biology | volume = 47 | issue = 3| pages = 519–537 | jstor = 2585257}}</ref> Most modern molecular studies find strepsipterans as the [[sister group]] of beetles (Coleoptera), with both groups together forming the clade [[Coleopterida]].<ref>{{Cite journal |last1=Beutel |first1=Rolf G. |last2=Pohl |first2=Hans |last3=Yan |first3=Evgeny V. |last4=Anton |first4=Eric |last5=Liu |first5=Si-Pei |last6=Ślipiński |first6=Adam |last7=McKenna |first7=Duane |last8=Friedrich |first8=Frank |date=January 2019 |title=The phylogeny of Coleopterida (Hexapoda) – morphological characters and molecular phylogenies |journal=Systematic Entomology |language=en |volume=44 |issue=1 |pages=75–102 |doi=10.1111/syen.12316|s2cid=92390950 |doi-access=free }}</ref> The most [[Basal (phylogenetics)|basal]] strepsipteran is the fossil ''[[Protoxenos janzeni]]'' discovered in [[Eocene]] aged [[Baltic amber]],<ref>{{cite journal | last1 = Pohl | first1 = H. | last2 = Beutel | first2 = R.G. | last3 = Kinzelbach | first3 = R. | year = 2005 | title = Protoxenidae fam. nov. (Insecta, Strepsiptera) from Baltic amber—a 'missing link' in strepsipteran phylogeny | journal = Zoologica Scripta | volume = 34 | pages = 57–69 | doi=10.1111/j.1463-6409.2005.00173.x| s2cid = 85232914 }}</ref> whereas the most basal living strepsipteran is ''[[Bahiaxenos relictus]]'', the sole member of the family [[Bahiaxenidae]].<ref name = Bravoetal>{{cite journal | last1 = Bravo | first1 = Pohl | last2 = Silva-Neto | last3 = Beutel | year = 2009 | title = Bahiaxenidae, a "living fossil" and a new family of Strepsiptera (Hexapoda) discovered in Brazil | journal = Cladistics | volume = 25 | issue = 6| pages = 614–623 | doi = 10.1111/j.1096-0031.2009.00264.x | pmid = 34879590 | s2cid = 83936131 }}</ref> The earliest known strepsipteran fossils are those of ''[[Cretostylops engeli]]'' (Cretostylopdiae) and ''[[Kinzelbachilla ellenbergeri]]'', ''[[Phthanoxenos nervosus]]'' and ''[[Heterobathmilla kakopoios]]'' (Phthanoxenidae), discovered in middle Cretaceous [[Burmese amber]] from [[Myanmar]], around 99&nbsp;million years old, which all lie outside the [[crown group]], but are all more closely related to modern strepsiperans than ''Protoxenos'' is. The finding of a parasitic first [[instar]] in the same deposit indicates that the parasitic lifestyle of the group has likely existed nearly unchanged for 100&nbsp;million years, though their evolutionary history prior to this remains a mystery.<ref name=":0" /> The idea that mengellinids' targeting of zygentomans represents the ancestral ecology of the group as a whole has been considered questionable.<ref name=":5" />[[File:Stylops melittae m3.JPG|left|thumb|''[[Stylops melittae]]'' male]]
[[File:Heterobathmilla kakopoios dorsal.jpg|thumb|Stem-group strepsipteran ''[[Heterobathmilla|Heterobathmilla kakopoios]]'' (†[[Phthanoxenidae]]) in [[Burmese amber]], around 100&nbsp;million years old]]

===Families===
[[File:Andrena vaga m Stylops melittae fm3.JPG|thumb|''[[Andrena vaga]]'' male bee, with ''[[Stylops melittae]]'' mating on its abdomen]]
The vast majority of living strepispterans are placed within the grouping Stylopidia, which includes the families [[Corioxenidae]], [[Halictophagidae]], [[Callipharixenidae]], [[Bohartillidae]], [[Elenchidae]], [[Myrmecolacidae]], [[Stylopidae]], [[Protelencholacidae]] (extinct) and [[Xenidae]].<ref name=":0" /> All Stylopidia have endoparasitic females that have multiple genital openings.<ref name="insencyc" /> Two living families, [[Mengenillidae]] and [[Bahiaxenidae]], are placed outside of this group, along with several extinct families.<ref name=":0" />

The Stylopidae have four-segmented tarsi and four- to six-segmented antennae, with the third segment having a lateral process. The family Stylopidae may be [[paraphyletic]].<ref name="insencyc" /> The Elenchidae have two-segmented tarsi and four-segmented antennae, with the third segment having a lateral process. The Halictophagidae have three-segmented tarsi and seven-segmented antennae, with lateral processes from the third and fourth segments.<ref name="borror" />
The Stylopidae mostly parasitize wasps and bees, the Elenchidae are known to parasitize [[Fulgoroidea]], whereas the Halictophagidae are found on leafhoppers, treehoppers, and mole cricket hosts.<ref name="borror" />

Strepsipteran insects in the genus [[Xenos (insect)|''Xenos'']] parasitize ''[[Polistes carnifex]]'', a species of social wasps.<ref name="Kathirithamby">{{cite journal | last1 = Kathirithamby | first1 = Jeyaraney | last2 = Hughes | first2 = David | year = 2006 | title = Description and biological notes of the first species of Xenos (Strepsiptera: Stylopidae) parasitic in Polistes carnifex F. (Hymenoptera: Vespidae) in Mexico | url = http://ento.psu.edu/publications/DH17 | journal = Zootaxa | volume = 1104 | pages = 35–45 | doi = 10.11646/zootaxa.1104.1.3 }}</ref> These [[obligate parasite]]s infect the developing wasp larvae in the nest and are present within the abdomens of female wasps when they hatch out. Here they remain until they thrust through the cuticle and pupate (males) or release infective first-[[instar]] larvae onto flowers (females). These larvae are transported back to their nests by foraging wasps.<ref name="Hughes other">{{cite journal |author1=Hughes, D. P. |author2=Beani, L. |author3=Turillazzi, S. |author4=Kathirithamby, J. |year=2003 |title=Prevalence of the parasite Strepsiptera in ''Polistes'' as detected by dissection of immatures |journal=Insectes Sociaux |volume=50 |issue=1 |pages=62–68 |doi=10.1007/s000400300010 |s2cid=9691419 }}</ref>

=== Cladogram ===
After:<ref name=":0" />
{{clade|{{clade
   |1=†[[Protoxenidae]]
   |2={{clade
      |1=†[[Cretostylopidae]]
      |2=†[[Phthanoxenidae]]
      |3={{clade   
          |1=†[[Mengeidae]]
          |2=[[Bahiaxenidae]]
          |3={{clade 
             |1=[[Mengenillidae]]
             |label2=[[Stylopidia]] 
             |2={{clade 
                |1=[[Corioxenidae]]
                |label2=[[Stylopiformia]] 
                |2={{clade 
                   |1=[[Bohartillidae]]
                   |2={{clade 
                      |1=[[Halictophagidae]]
                      |2={{clade 
                         |1=[[Elenchidae]]
                         |2={{clade 
                            |1=†[[Protelencholacidae]]
                            |2={{clade 
                               |1=[[Myrmecolacidae]]
                               |2={{clade 
                                  |1=[[Callipharixenidae]]
                                  |2={{clade 
                                     |1=[[Xenidae]]
                                     |2=[[Stylopidae]]
                                     }}
                                  }}
                               }}
                            }}
                         }}
                      }}
                  }}
               }}
            }}
         }}
      }}
   }}|style=font-size:100%;line-height:80%|label1=Strepsiptera}}