Editing Fly (section)

=== Relationships between subgroups and families ===

[[File:Fossil insect Diptera, Brachycera in Baltic amber. Age 50 Mill. years (the Lower Eocene).jpg|thumb|left|upright=0.7 |Fossil [[brachycera]]n in [[Baltic amber]]. [[Lower Eocene]], c. 50 million years ago]]

The first true dipterans known are from the [[Middle Triassic]] (around 240 million years ago), and they became widespread during the Middle and [[Late Triassic]].<ref>{{cite book |last1=Blagoderov |first1=V. A. |last2=Lukashevich |first2=E. D. |last3=Mostovski |first3=M. B. |editor1-last=Rasnitsyn |editor1-first=A. P. |editor2-last=Quicke |editor2-first=D. L. J. |year=2002 |title=History of Insects |chapter=Order Diptera Linné, 1758. The true flies |publisher=[[Kluwer Academic Publishers]] |isbn=978-1-4020-0026-3 |chapter-url=http://palaeoentomolog.ru/New/diptera.html}}</ref> [[Flowering plant|Modern flowering plants]] did not appear until the [[Cretaceous]] (around 140 million years ago), so the original dipterans must have had a different source of nutrition other than [[nectar]]. Based on the attraction of many modern fly groups to shiny droplets, it has been suggested that they may have fed on [[Honeydew (secretion)|honeydew]] produced by [[Homoptera|sap-sucking bugs]] which were abundant at the time, and dipteran mouthparts are well-adapted to softening and lapping up the crusted residues.<ref>{{cite journal |last1=Downes |first1=William L. Jr. |author2=Dahlem, Gregory A. |year=1987 |title= Keys to the Evolution of Diptera: Role of Homoptera |journal=[[Environmental Entomology]] |volume=16 |issue=4 |pages=847–854 |doi=10.1093/ee/16.4.847 }}</ref> The [[basal clade]]s in the Diptera include the [[Deuterophlebiidae]] and the enigmatic [[Nymphomyiidae]].<ref name="Wiegmann2011">{{cite journal |last1=Wiegmann |first1=B. M. |year=2011 |title=Episodic radiations in the fly tree of life |journal=[[PNAS]] |volume=108 |issue=14 |pages=5690–5695 |doi=10.1073/pnas.1012675108 |first2=M. D. |last3=Winkler |first3=I. S. |last4=Barr |first4=N. B. |last5=Kim |first5=J.-W. |last6=Lambkin |first6=C. |last7=Bertone |first7=M. A. |last8=Cassel |first8=B. K. |last9=Bayless |first9=K. M. |last10=Heimberg |first10=A. M. |last11=Wheeler |first11=B. M. |last12=Peterson |first12=K. J. |last13=Pape |first13=T. |last14=Sinclair |first14=B. J. |last15=Skevington |first15=J. H. |last16=Blagoderov |first16=V. |last17=Caravas |first17=J. |last18=Kutty |first18=S. N. |last19=Schmidt-Ott |first19=U. |last20=Kampmeier |first20=G. E. |last21=Thompson |first21=F. C. |last22=Grimaldi |first22=D. A. |last23=Beckenbach |first23=A. T. |last24=Courtney |first24=G. W. |last25=Friedrich |first25=M. |last26=Meier |first26=R. |last27=Yeates |first27=D. K. |pmid=21402926 |pmc=3078341 |display-authors=6 |last2=Trautwein |bibcode=2011PNAS..108.5690W |doi-access=free }}</ref> Three episodes of [[evolutionary radiation]] are thought to have occurred based on the fossil record. Many new species of lower Diptera developed in the [[Triassic]], about 220 million years ago. Many lower Brachycera appeared in the [[Jurassic]], some 180 million years ago. A third radiation took place among the [[Schizophora]] at the start of the [[Paleogene]], 66 million years ago.<ref name="Wiegmann2011" />

The phylogenetic position of Diptera has been controversial. The monophyly of [[holometabolous]] insects has long been accepted, with the main orders being established as Lepidoptera, Coleoptera, Hymenoptera and Diptera, and it is the relationships between these groups which has caused difficulties. Diptera is widely thought to be a member of [[Mecopterida]], along with Lepidoptera (butterflies and moths), Trichoptera (caddisflies), Siphonaptera (fleas), Mecoptera (scorpionflies) and possibly [[Strepsiptera]] (twisted-wing flies). Diptera has been grouped with Siphonaptera and Mecoptera in the Antliophora, but this has not been confirmed by molecular studies.<ref>{{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 |year=2012 |publisher=[[Columbia University Press]] |isbn=978-0-231-50170-5 |pages=4–6}}</ref>

[[File:Lutzomyia adiketis.jpg|thumb|right|upright=0.7 |Fossil [[nematocera]]n in Dominican amber. Sandfly, ''[[Lutzomyia adiketis]]'' ([[Psychodidae]]), [[Early Miocene]], c. 20 million years ago]]

Diptera were traditionally broken down into two suborders, [[Nematocera]] and [[Brachycera]], distinguished by the differences in antennae. The Nematocera are identified by their elongated bodies and many-segmented, often feathery antennae as represented by mosquitoes and crane flies. The Brachycera have rounder bodies and much shorter antennae.<ref>B.B. Rohdendorf. 1964. Trans. Inst. Paleont., Acad. Sci. USSR, Moscow, v. 100</ref><ref name=TOL>{{cite web |last1=Wiegmann |first1=Brian M. |last2=Yeates |first2=David K. |title=Diptera True Flies |url=http://tolweb.org/Diptera/8226 |website=Tree of Life |date=29 November 2007 |access-date=25 May 2016}}</ref> Subsequent studies have identified the Nematocera as being non-monophyletic with modern phylogenies placing the Brachycera within grades of groups formerly placed in the Nematocera. The construction of a phylogenetic tree has been the subject of ongoing research. The following cladogram is based on the FLYTREE project.<ref>{{cite web |last1=Yeates |first1=David K. |last2=Meier |first2=Rudolf |last3=Wiegmann |first3=Brian |title=Phylogeny of True Flies (Diptera): A 250 Million Year Old Success Story in Terrestrial Diversification |url=http://www.inhs.illinois.edu/research/flytree/flyphylogeny/ |website=Flytree |publisher=Illinois Natural History Survey |archive-url=https://web.archive.org/web/20151228210535/http://inhs.illinois.edu/research/flytree/flyphylogeny/ |url-status=dead |archive-date=28 December 2015 |access-date=24 May 2016}}</ref><ref>{{cite journal |last1=Yeates |first1= David K. |last2=Weigmann |first2=Brian M |last3=Courtney |first3=Greg W. |last4=Meier |first4=Rudolf |last5=Lambkins |first5=Christine |last6=Pape |first6=Thomas |title=Phylogeny and systematics of Diptera: Two decades of progress and prospects |year=2007 |journal=[[Zootaxa]] |volume=1668 |pages=565–590 |doi=10.11646/zootaxa.1668.1.27 }}</ref>

{{clade | style=font-size:100%;line-height:105%;
 |label1='''Diptera'''
 |grouplabel1={{clade labels|label1="[[Nematocera]]" |top1=16%}}
 |1={{clade |bar1=green
  |1={{clade 
    |1=[[Ptychopteromorpha]] (phantom and primitive crane-flies) [[File:Ptychoptera contaminata male Walker 1856 plate-XXVIII.png|60px]]
    |2=[[Culicomorpha]] (mosquitoes, blackflies and midges) [[File:Stegomyia fasciata.jpg|55px]]
   }}
  |2={{clade
    |1=[[Blephariceromorpha]] (net-winged midges, etc) [[File:Imago of Blepharicera fasciata as Asthenia fasciata in Westwood 1842, plate 94.png|60px]]|bar1=green
    |2={{clade
     |1=[[Bibionomorpha]] (gnats) [[File:Isoneuromyia annandalei.jpg|55px]]|bar1=green
     |2={{clade
       |1=[[Psychodomorpha]] (drain flies, sand flies, etc) [[File:Clogmia_clean.jpg|55px]]|bar1=green
       |2={{clade
        |1=[[Tipulomorpha]] (crane flies) [[File:Tipula oleracea icon.jpg|65px]]|bar1=green
        |label2='''[[Brachycera]]'''
        |2={{clade
          |label1=
          |1={{clade
           |1=[[Stratiomyomorpha]] (soldier flies, etc) [[File:Hermetia illucens f.jpg|66px]]
           |2={{clade
             |1=[[Xylophagomorpha]] (stink flies, etc) [[File:Coenomyia_ferruginea.png|55px]]
             |2=[[Tabanomorpha]] (horse flies, snipe flies, etc) [[File:Chrysops relicta f.jpg|66px]]
            }}
           }}
          |label2=[[Muscomorpha|Mus]]
          |2={{clade
           |1=[[Nemestrinoidea]] [[File:Acrocera globulus.png|66px]]
           |2={{clade
             |1=[[Asiloidea]] (robber flies, bee flies, etc) [[File:Asilidae icon.jpg|66px]]
             |label2=[[Eremoneura|Ere]]
             |2={{clade
              |1=[[Empidoidea]] (dance flies, etc) [[File:Drapetis_brevior_fbi.jpg|35px]]
              |label2=[[Cyclorrhapha|Cyc]]
              |2={{clade
                |1=[[Aschiza]] (in part)
                |2={{clade
                 |1=[[Phoroidea]] (flat-footed flies, etc) [[File:Platypeza_picta.png|55px]]
                 |2={{clade
                   |1=[[Syrphoidea]] (hoverflies) [[File:Syrphidae icon.jpg|55px]]
                   |label2=[[Schizophora|Sch]]
                   |2={{clade
                    |label1=[[Calyptratae|Cal]]
                    |1={{clade
                      |1=[[Hippoboscoidea]] (louse flies, etc) [[File:Nycteribiidae icon.jpg|55px]]
                      |2={{clade
                       |1=[[Muscoidea]] (house flies, dung flies, etc) [[File:Musca_domestica_female.jpg|55px]]
                       |2=[[Oestroidea]] (blow flies, flesh flies, etc) [[File:Sarcophaga_haemorrhoidalis_m.jpg|55px]]
                       }}
                     }}
                    |2=[[Acalyptratae]] (marsh flies, fruit flies, etc) [[File:Ceratitis capitata illustration.jpg|55px]]
                    }}
                  }}
                 }}
               }}
              }}
            }}
           }}
         }}
        }}
      }}
     }}
   }}
  }}
}}

{{smalldiv|1=
''Abbreviations used in the cladogram:''
* Cal=[[Calyptratae]]
* Cyc=[[Cyclorrhapha]] 
* Ere=[[Eremoneura]] 
* Mus=[[Muscomorpha]]
* Sch=[[Schizophora]] 
}}