Order Diptera (Classification and Phylogeny)

[Ken Kinman]

Dear All,
      The flies (Order Diptera) are a cladist's dream.  Certainly NOT because it is all that easy to study their phylogeny, but rather because it is more pectinate (asymmetric) in its evolution than most groups.  This makes the group one that is ideal for total cladification (which should make strict cladists very happy).  However EVEN in this case, whether it should be totally cladified depends on your philosophic leanings.
     Therefore, in this case I am presenting two identical phylogenies of Diptera, one with limited paraphyly and other with no apparent paraphyly at all (holophyly only). However, it is just my first attempt at classifying Order Diptera to family level, so the coding is obviously going to be particularly preliminary.  The placement of Tanyderidae and Ptychopteridae at the base of Diptera is rather unusual (rather than Tipulidae), but it seems a viable option to me at the present time, so I'm taking the opportunity to introduce this possibility (pushing the envelope or playing devil's advocate, however you wish to view it).  Just an educated hunch, but hopefully well worth considering and testing.
    The first classification is more traditional and eclectic.  It is only unusual for me in that I am using taxa intermediate between order and family.  It's a huge order and these paraphyletic divisions are still used by Christian Thompson (and the vast majority of dipterists), so it is my preference at the present time.  However, since this classification shows unusual levels of pectination (asymmetry), I am also following it with a totally cladistic classification of Order Diptera as well (although the TOTAL elimation of paraphyly seems to require putting Family Dolichopodidae into Empididae).  Both classifications show the same phylogeny, so feel free to use whichever one suits you.
     Either way one should note that the Schizophora (clade 3 within Cyclorrhapa) is still a broad polytomy (coded by repeated "A"s) that is unresolved in both classifications.  So much so that the radiation at the end of these classifications is presently only shown as superfamilial groups, but Schizophora are quite derived however you look at it (NOTE:  I would be very interested to know if anyone else might think, as I suspect, that Conopoidea might be sister group of all other Schizophora).  And some strict cladists may be horrified that I call Nematocera "primitive flies", but during the early evolution of the Diptera they were they ONLY flies in existence, so don't even bother protesting my use of the word "primitive", as the use of the term "primitive" is only a problem if it is equated with "inferior" (which I certainly do not).  But do feel free to challenge the details of my phylogeny (that's the kind of feedback I definitely will find useful).  :-)
      ----- Cheers, Ken Kinman

ORDER DIPTERA
  1  Nematocera% (primitive flies)
          1  Tanyderidae
          B  Ptychopteridae
          2  Thaumaleidae
          b  Simuliidae
          c  Ceratopogonidae
          d  Chironomidae
          B  Dixidae
          C  Corethrellidae
          D  Chaoboridae
          E  Culicidae
          3  Nymphomyiidae
          B  Deuterophlebiidae
          C  Blephariceridae
          4  Tipulidae (sensu lato)
          5  Psychodidae
          ?  Axymyiidae
          6  Perissommatidae
             Scatopsidae
             Canthyloscelidae
          7  Pachyneuridae
          B  Bibionidae
          C  Mycetophilidae
          D  Cecidomyiidae
          ?  Rangomaramidae
          E  Sciaridae
          8  Anisopodidae
          9  {{Brachycera}}
              (= Orthorrhapa + Cyclorrhapa)

 _1_ Orthorrhapa (primitive brachyceran flies)
          1  Pantophthalmidae
          B  Xylomyidae
          C  Stratiomyidae
          2  Vermileonidae
          3  Xylophagidae
          B  Ragionidae
          ?  Austroleptidae
          ?  Spaniidae
          C  Athericidae
          D  Tabanidae
          4  Acroceridae
          B  Nemestrinidae
          5  Bombyliidae
             Mythicomyiidae
             Asilidae
             Scenopinidae
             Therevidae
             Mydidae
             Apioceridae
             Apsilocephalidae
             Apystomyiidae
             Hilarimorphidae
             Ocoidae
          6  Empididae%
         _a_ Dolichopodidae
          7  {{Cyclorrhapa}}

 _1_ Cyclorrhapa (derived brachyceran flies)
          1  Platypezidae
          B  Opetiidae
          C  Lonchopteridae
          D  Ironomyiidae
          E  Phoridae
          2  Syrphidae
          B  Pipunculidae
          3  Conopoid families
             Tephritoid families
             Nerioid families
             Diopsoid famlies
          A  Lauxanioid families
             Sciomyzoid families
          A  Sphaeroceroid families
             Ephydroid families
             Opomyzoid families
             Carnoid families
          A  Hippoboscoid families
             Muscoid families
             Oestroid families

*************************************

  ORDER DIPTERA (paraphyly eliminated)
          1  Tanyderidae
          B  Ptychopteridae
          2  Thaumaleidae
          b  Simuliidae
          c  Ceratopogonidae
          d  Chironomidae
          B  Dixidae
          C  Corethrellidae
          D  Chaoboridae
          E  Culicidae
          3  Nymphomyiidae
          B  Deuterophlebiidae
          C  Blephariceridae
          4  Tipulidae (sensu lato)
          5  Psychodidae
          ?  Axymyiidae
          6  Perissommatidae
             Scatopsidae
             Canthyloscelidae
          7  Pachyneuridae
          B  Bibionidae
          C  Mycetophilidae (sensu lato)
          D  Cecidomyiidae
          ?  Rangomaramidae
          E  Sciaridae
          8  Anisopodidae
          9  Pantophthalmidae
          B  Xylomyidae
          C  Stratiomyidae
         10  Vermileonidae
         11  Xylophagidae
          B  Ragionidae
          ?  Austroleptidae
          ?  Spaniidae
          C  Athericidae
          D  Tabanidae
         12  Acroceridae
          B  Nemestrinidae
         13  Bombyliidae
             Mythicomyiidae
             Asilidae
             Scenopinidae
             Therevidae
             Mydidae
             Apioceridae
             Apsilocephalidae
             Apystomyiidae
             Hilarimorphidae
             Ocoidae
         14  Empididae (incl. Dolichopodids)
         15  Platypezidae
          B  Opetiidae
          C  Lonchopteridae
          D  Ironomyiidae
          E  Phoridae
         16  Syrphidae
          B  Pipunculidae
         17  Conopoid families
             Tephritoid families
             Nerioid families
             Diopsoid famlies
          A  Lauxanioid families
             Sciomyzoid families
          A  Sphaeroceroid families
             Ephydroid families
             Opomyzoid families
             Carnoid families
          A  Hippoboscoid families
             Muscoid families
             Oestroid families
**************************************
P.S.  Asiloidea (clade 5 in Orthorrhapa in the eclectic classification; and clade 13 in the totally cladified classification) is left uncoded.  But I hope to cladisticly code it when I have access to more literature.  I will also add fossil families when I have more information.  I've only been working on this phylogeny for one day, so more tinkering with the details are inevitable.