Editing Mecoptera (section)

==Evolution and phylogeny==

===Taxonomic history===
The European scorpionfly was named ''[[Panorpa communis]]'' by [[Linnaeus]] in [[10th edition of Systema Naturae|1758]].<ref>[[Linnæus]], Carolus.  1758. ''[[Systema naturae]] per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis''. Laurentii Salvii, Holmiae [= Stockholm].  Vol. Tomus I,  Editio decima, reformata: i–ii, 1–824.</ref>
The Mecoptera were named by [[Alpheus Hyatt]] and [[Jennie Maria Arms Sheldon|Jennie Maria Arms]] in 1891.<ref name="HyattArms1891">{{cite journal |last1=Hyatt |first1=Alpheus |last2=Arms |first2=Jennie Maria |title=A novel diagrammatic representation of the orders of insects |journal=Psyche: A Journal of Entomology |date=1891 |volume=6 |issue=177 |pages=11–13 |doi=10.1155/1891/39454|doi-access=free }}</ref> The name is from the [[Greek language|Greek]], ''mecos'' meaning long, and ''ptera'' meaning wings.<ref>{{Cite book|title=A Dictionary of Entomology|journal=Nature|volume=91|issue=2267|url=https://books.google.com/books?id=9IcmCeAjp6cC&pg=PA858|year=2011|isbn=978-1-84593-542-9|page=858|bibcode=1913Natur..91S.134.|doi=10.1038/091134c0|s2cid=3947767}}</ref>
The families of Mecoptera are well accepted by taxonomists but their relationships have been debated. In 1987, R. Willman treated the Mecoptera as a [[clade]], containing the Boreidae as sister to the Meropeidae,<ref>{{cite journal |author=Willman, R. |date=1987 |title=The phylogenetic system of the Mecoptera |journal=Systematic Entomology |volume=12 |issue=4 |pages=519–524 |doi=10.1111/j.1365-3113.1987.tb00222.x|bibcode=1987SysEn..12..519W |s2cid=86349146 }}</ref> but in 2002 [[Michael F. Whiting]] declared the Mecoptera so-defined as paraphyletic, with the Boreidae as sister to another order, the [[Siphonaptera]] (fleas).<ref name=Whiting2002/>

===Fossil history===

Among the earliest members of the Mecoptera are the [[Nannochoristidae]] of [[Upper Permian]] age. Fossil Mecoptera become abundant and diverse during the [[Cretaceous]], for example in China,<ref name="MartillBechly2007">{{cite book |last1=Martill|first1=David M. |last2=Bechly |first2=Günter |last3=Loveridge |first3=Robert F. |title=The Crato Fossil Beds of Brazil: Window into an Ancient World |url=https://books.google.com/books?id=5ZBh_-QvX2MC&pg=PA367 |year=2007 |publisher=Cambridge University Press |isbn=978-1-139-46776-6 |pages=367–369}}</ref> where panorpids such as ''Jurassipanorpa'', hangingflies (Bittacidae and Cimbrophlebiidae), Orthophlebiidae, and Cimbrophlebiidae have been found.<ref>{{cite journal|last1=Wang |first1=Chen |last2=Shih |first2=Chungkun|last3=Ren|first3=Dong |title=A New Fossil Hangingfly (Mecoptera: Cimbrophlebiidae) from the Early Cretaceous of China |journal=Acta Geologica Sinica - English Edition |date=2014 |volume=88 |issue=1 |pages=29–34 |doi=10.1111/1755-6724.12180|bibcode=2014AcGlS..88...29W |s2cid=129796533 }}</ref><ref>{{cite journal |last1=Ding |first1=He|last2=Shih |first2=Chungkun |last3=Bashkuev |first3=Alexei |last4=Zhao |first4=Yunyun |last5=Ren |first5=Dong |title=The earliest fossil record of Panorpidae (Mecoptera) from the Middle Jurassic of China |journal=ZooKeys |date=2014 |issue=431|pages=79–92 |doi=10.3897/zookeys.431.7561|pmid=25152669|pmc=4141175|doi-access=free|bibcode=2014ZooK..431...79D }}</ref>

Extinct Mecoptera species may have been important pollinators of early [[gymnosperm]] seed plants during the late [[Middle Jurassic]] to mid–[[Early Cretaceous]] periods before other pollinating groups such as the bees evolved. These were mainly wind-pollinated plants, but fossil mecopterans had siphon-feeding apparatus that could have fertilized these early gymnosperms by feeding on their [[nectar]] and [[pollen]]. The lack of iron enrichment in their fossilized [[probosces]] rules out their use for drinking blood. Eleven species have been identified from three families, [[Mesopsychidae]], [[Aneuretopsychidae]], and [[Pseudopolycentropodidae]] within the clade [[Aneuretopsychina]]. Their lengths range from {{convert|3|mm|in|abbr=on}} in ''Parapolycentropus burmiticus'' to {{convert|28|mm|in|abbr=on}} in ''Lichnomesopsyche gloriae''. The proboscis could be as long as {{convert|10|mm|in|abbr=on}}. It has been suggested that these mecopterans transferred pollen on their mouthparts and head surfaces, as do [[bee flies]] and [[hoverflies]] today, but no such associated pollen has been found, even when the insects were finely preserved in [[Eocene]] [[Baltic amber]]. They likely pollinated plants such as [[Caytoniaceae]], [[Cheirolepidiaceae]], and [[Gnetales]], which have ovulate organs that are either poorly suited for wind pollination or have structures that could support long-proboscid fluid feeding.<ref name="Ollerton">{{cite journal | last1 = Ollerton | first1 = J. | last2 = Coulthard | first2 = E. | year = 2009 | title = Evolution of Animal Pollination | journal = Science | volume = 326 | issue = 5954| pages = 808–809 | doi = 10.1126/science.1181154 | pmid = 19892970 | bibcode = 2009Sci...326..808O | s2cid = 856038 }}</ref><ref name="Ren">{{cite journal | last1 = Ren | first1 = D. | last2 = Labandeira | first2 = C.C. | last3 = Santiago-Blay | first3 = J.A. | last4 = Rasnitsyn | first4 = A. | last5 = Shih | first5 = C.K. | last6 = Bashkuev | first6 = A. | last7 = Logan | first7 = M.A. | last8 = Hotton | first8 = C.L. | last9 = Dilcher | first9 = D. | year = 2009 | title = Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies | journal = Science | volume = 326 | issue = 5954| pages = 840–847 | doi = 10.1126/science.1178338 | pmid = 19892981 | pmc = 2944650 | bibcode = 2009Sci...326..840R }}</ref>  The Aneuretopsychina were the most diverse group of mecopterans in the Latest Permian, taking the place of the [[Permochoristidae]], to the [[Middle Triassic]].  During the [[Late Triassic]] through the [[Middle Jurassic]], Aneuretopsychina species were gradually replaced by species from the [[Parachoristidae]] and Orthophlebiidae.  Modern mecopteran families are derived from the Orthophlebiidae.<ref name="Bashkuev2011b">{{Cite book |last1=Bashkuev |first1=A. S. |year=2011 |chapter=The earliest Mesopsychidae and revision of the family Mesopanorpodidae (Mecoptera)|title=Advances in the Systematics of Fossil and Modern Insects: Honouring Alexandr Rasnitsyn |series=ZooKeys 130 |pages=263–279|isbn=978-954-642-609-3 |doi=10.3897/zookeys.130.1611|pmid=22259282 |pmc=3260765 |doi-access=free|publisher=PenSoft|editor-last1=Shcherbakov|editor-first1=D.|editor-last2=Engel|editor-first2=M.|editor-last3=Sharkey|editor-first3=M.}}</ref>

<gallery mode="nolines">
File:Juracimbrophlebia ginkgofolia.jpg|''[[Juracimbrophlebia ginkgofolia]]'' ([[Cimbrophlebiidae]]) reconstruction, Jurassic; China
File:Jurassipanorpa sticta holotype.jpg|''[[Jurassipanorpa|Jurassipanorpa sticta]]'' ([[Panorpidae]]), Jurassic; China
File:Dinokanaga andersoni holotype SR 01-06-01.jpg|Forewing of ''[[Dinokanaga andersoni]]'' ([[Dinopanorpidae]]), [[Eocene]]; Washington state <!--File:Cimbrophlebia brooksi Holotype SR 06-20-05 B.jpg|''[[Cimbrophlebia|Cimbrophlebia brooksi]]'' ([[Cimbrophlebiidae]]), Eocene; Washington state--> <!--File:Eorpa elverumi SRUI 08-07-07 A Holotype.JPG|Forewing of ''[[Eorpa elverumi]]'' ([[Eorpidae]]), Eocene; Washington State--> 
</gallery>

===External relationships===
Mecoptera have special importance in the evolution of the insects. Two of the most important insect orders, [[Lepidoptera]] (butterflies and moths) and [[Diptera]] (true flies), along with [[Trichoptera]] (caddisflies), probably evolved from ancestors belonging to, or strictly related to, the Mecoptera. Evidence includes anatomical and biochemical similarities as well as [[transitional fossil]]s, such as ''[[Permotanyderus]]'' and ''[[Choristotanyderus]]'', which lie between the Mecoptera and Diptera. The group was once much more widespread and diverse than it is now, with four suborders during the [[Mesozoic]].<ref name=IIBD/>

It is unclear as of 2020 whether the Mecoptera form a single clade, or whether the [[Siphonaptera]] (fleas) are inside that clade, so that the traditional "Mecoptera" taxon is paraphyletic. However the earlier suggestion that the Siphonaptera are sister to the [[Boreidae]]<ref name=Whiting2008>{{cite journal |last1=Whiting |first1=Michael F. |author1-link=Michael F. Whiting|last2=Whiting |first2=Alison S. |last3=Hastriter |first3=Michael W. |last4=Dittmar |first4=Katharina |title=A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations |journal=Cladistics |date=2008 |volume=24 |issue=5 |pages=677–707 |doi=10.1111/j.1096-0031.2008.00211.x |citeseerx=10.1.1.731.5211  |s2cid=33808144 }}</ref><ref name=Whiting2002>{{cite journal |last1=Whiting |first1=Michael F. |author1-link=Michael F. Whiting|year=2002 |title=Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera |journal=Zoologica Scripta |volume=31 |issue=1 |pages=93–104 |url=http://www3.interscience.wiley.com/journal/118916123/abstract |archive-url=https://archive.today/20130105095659/http://www3.interscience.wiley.com/journal/118916123/abstract |url-status=dead |archive-date=2013-01-05 |doi=10.1046/j.0300-3256.2001.00095.x |s2cid=56100681 |url-access=subscription }}</ref><ref name=WiegmannYates2012>{{cite book |last1=Wiegmann |first1=Brian |last2=Yeates |first2=David K. |title=The Evolutionary Biology of Flies |url=https://books.google.com/books?id=rElP5sNn6IYC&pg=PA5 |year=2012 |publisher=Columbia University Press |isbn=978-0-231-50170-5 |page=5 |quote=Recently, a close affinity between Siphonaptera and Mecoptera has been convincingly demonstrated via morphology (Bilinski et al. 1998) and molecular data (Whiting 2002), rendering Mecoptera paraphyletic, but making the clade including Mecoptera and Siphonaptera monophyletic}}</ref> is not supported; instead, there is the possibility that they are sister to another Mecopteran family, the [[Nannochoristidae]]. The two possible trees are shown below:<ref name="Meusemann Trautwein 2020">{{cite bioRxiv |last1=Meusemann |first1=Karen |last2=Trautwein |first2=Michelle |last3=Friedrich |first3=Frank |last4=Beutel |first4=Rolf G. |last5=Wiegmann |first5=Brian M. |last6=Donath |first6=Alexander |last7=Podsiadlowski |first7=Lars |last8=Petersen |first8=Malte |last9=Niehuis |first9=Oliver |last10=Mayer |first10=Christoph |last11=Bayless |first11=Keith M. |last12=Shin |first12=Seunggwan |last13=Liu |first13=Shanlin |last14=Hlinka |first14=Ondrej |last15=Minh |first15=Bui Quang |last16=Kozlov |first16=Alexey |last17=Morel |first17=Benoit |last18=Peters |first18=Ralph S. |last19=Bartel |first19=Daniela |last20=Grove |first20=Simon |last21=Zhou |first21=Xin |last22=Misof |first22=Bernhard |last23=Yeates |first23=David K. |display-authors=5|title=Are Fleas Highly Modified Mecoptera? Phylogenomic Resolution of Antliophora (Insecta: Holometabola) |year=2020 |biorxiv=10.1101/2020.11.19.390666}}</ref>

(a) Mecoptera (clades in boldface) is paraphyletic, containing Siphonaptera:<ref name="Meusemann Trautwein 2020"/><ref name="Tihelka Giacomelli 2020">{{cite journal |last1=Tihelka |first1=Erik |last2=Giacomelli |first2=Mattia |last3=Huang |first3=Di-Ying |last4=Pisani |first4=Davide |last5=Donoghue |first5=Philip C. J. |last6=Cai |first6=Chen-Yang |title=Fleas are parasitic scorpionflies |journal=Palaeoentomology |volume=3 |issue=6 |year=2020 |pages=641–653 |issn=2624-2834 |doi=10.11646/palaeoentomology.3.6.16|doi-access=free |hdl=1983/8d3c12c6-529c-4754-b59d-3abf88a32fc9 |hdl-access=free }}</ref>

{{clade
|label1=[[Antliophora]]
|1={{clade 
   |1=[[Diptera]] (true flies) [[File:Common house fly, Musca domestica.jpg|70px]] 
   |grouplabel2={{clade labels|label1=<span style="color:red;padding:1em;">Mecoptera</span> |top1=38%}}
   |2={{clade
       |1='''[[Pistillifera]]''' (scorpionflies, hangingflies, 400 [[Species|spp]].) [[File:Gunzesrieder Tal Insekt 3.jpg|90px]] |bar1=red
       |2={{clade
          |2={{clade
             |1='''[[Nannochoristidae]]''' (southern scorpionflies, 8 spp.)[[File:Nannochorista philpotti (cropped).jpg|85px]]&nbsp;  |bar1=red
             |2=[[Siphonaptera]] (fleas, 2500 spp.) [[File:British Entomologycutted Plate114.png|50px]]
             }}
          |1='''[[Boreidae]]''' (snow scorpionflies, 30 spp.) [[File:Boreus hiemalis2 detail.jpg|85px]]  |bar1=red
          }}
      }}
   }}
}}

(b) Mecoptera is monophyletic, sister to Siphonaptera:<ref name="Meusemann Trautwein 2020"/>

{{clade
|label1=[[Antliophora]]
|1={{clade 
   |1=[[Diptera]] (true flies) [[File:Common house fly, Musca domestica.jpg|70px]] 
   |2={{clade
       |label1='''Mecoptera'''
       |1={{clade
          |1={{clade
             |1=[[Pistillifera]] (scorpionflies, hangingflies, 400 [[Species|spp]].) [[File:Gunzesrieder Tal Insekt 3.jpg|90px]]
             |2=[[Boreidae]] (snow scorpionflies, 30 spp.) [[File:Boreus hiemalis2 detail.jpg|85px]]
             }}
          |2=[[Nannochoristidae]] (southern scorpionflies, 8 spp.)[[File:Nannochorista philpotti (cropped).jpg|85px]]
          }}
       |label2=[[Siphonaptera]] 
       |2= (fleas, 2500 spp.) [[File:Pulex irritans female ZSM.jpg|50px]]
      }}
   }}
}}

===Internal relationships===
All the families were formerly treated as part of a single order, Mecoptera. The relationships between the families are, however, a matter of debate. The cladogram, from Cracraft and Donoghue 2004, places the Nannochoristidae as a separate order, with the Boreidae, as the sister group to the Siphonaptera, also as its own order. The [[Eomeropidae]] is suggested to be the sister group to the rest of the Mecoptera, with the position of the [[Bittacidae]] unclear. Of those other families, the [[Meropeidae]] is the most basal, and the relationships of the rest are not completely clear.<ref>{{cite book |last=Whiting |first=Michael F. |chapter=Phylogeny of the Holometabolous Insects |editor-last1=Cracraft|editor-first1=Joel|editor-last2=Donoghue|editor-first2=Michael J.|title=Assembling the Tree of Life|chapter-url=https://books.google.com/books?id=6lXTP0YU6_kC&pg=PA355|year=2004|publisher=Oxford University Press|isbn=978-0-19-972960-9|page=355}}</ref>
{{clade
|1={{clade
   |label1='''Nannomecoptera'''
   |1={{clade 
      |1=[[Nannochoristidae]]{{efn|name=Nannomecoptera|Some studies find Nannomecoptera as sister to the Boreidae+Siphonaptera clade.<ref name=Whiting2002/>}}
      }}
   |2={{clade
      |1={{clade
         |label1='''Neomecoptera'''
         |1={{clade
            |1=[[Boreidae]] (snow scorpionflies) [[File:Boreus hyemalis male.jpg|90px]]
            }}
         |2='''[[Siphonaptera]]''' (fleas) [[File:Pulex irritans female ZSM.jpg|65px]] 
         }}      
      |2={{clade
         |label1='''Mecoptera''' |sublabel1= ''sensu stricto''
         |1={{clade
            |1=[[Eomeropidae]] (mainly fossil ([[Triassic]] to present), 1 extant sp.) [[File:Notiothauma reedi.jpg|90px]]
            |2=(?) [[Bittacidae]]{{efn|name=Bittacidae}} (hangingflies) |state2=dotted
            |3={{clade
               |1=[[Meropeidae]] (earwigflies) [[File:Merope tuber dorsal view - ZooKeys-269-051-g007A.jpeg|90px]]  
               |2={{clade
                  |1=[[Choristidae]] (Australian scorpionflies) [[File:Chorista australis Gundaroo NSW Australia.jpg|90px]] 
                  |2=[[Apteropanorpidae]] (Tasmanian snow scorpionflies)
                  |3={{clade
                     |1={{clade
                        |1=(?)[[Bittacidae]]{{efn|name=Bittacidae}} (hangingflies) [[File:Hylobittacus apicalis Michigan.jpg|90px]] |state2=dotted|state1=dotted
                        |2=[[Panorpodidae]] (short-faced scorpionflies)
                        }}
                     |2=[[Panorpidae]] ([[Jurassic]] to present,  common scorpionflies) [[File:Panorpa communis V.jpg|70px]]
                     }}
                  }}
               }}
            }}
         }}
      }}  
   }}
}}
{{notelist|refs=
{{efn |name=Bittacidae|The position of the [[Bittacidae]] is unclear. Two alternative positions, either basal within Mecoptera ''sensu stricto'' or sister to Panorpodidae, are shown.}}
}}
[[File:Panorpa dubia (Mecoptera) anatomy.tiff|thumb|250px|Male ''[[Panorpa|Panorpa dubia]]''.<br/> A, Body in lateral view; B–D. male genital bulb and gonostyli. B, dorsal view; C, ventral view; D, lateral view. ep, epandrium; gcx, gonocoxite; gs, gonostylus; hv, hypovalva; hyp, hypandrium. Scale bars represent 3 mm in A, 1 mm in B–D]]