================== Tipulidae | | =============== Blephariceromorpha | | | |============== Axymyiidae | | | | ====== Culicimorpha <<===| | | ===| ===| === Tanyderidae |=====| ===| | | === Ptychopteridae | | | ========= Psychodomorpha | =============== Bibionomorpha ?=== BrachyceraTree from Wood and Borkent (1989)
Containing clade: Endopterygota
The earliest fossil flies are known from the Upper Triassic of the Mesozoic geological period, some 225 million years ago (Evenhuis, 1995). Since that time they have diversified to become one of the largest groups of organisms. There have been about 120,000 species of flies formally described by scientists; thus about 1 in every 10 animals described is a fly. An equal number of species may await description and most of these will be found in environments that remain to be studied intensively, such as tropical forests.
Flies are holometabolous insects, that is their life cycle involes a major change in form from a soft-bodied, wingless larval stage to a hardened, winged adult.
Larval flies have a variety of common names, such as wriggler and maggot. Fly larvae have an enormous variety of feeding habits, and individual species often have very precise requirements. Many consume decaying organic matter, or are predacious, and a large proportion are parasitic on other insects and other organisms. Adult flies are almost always free-living and fly during the day. They typically consume liquid food such as nectar and other plant exudates, or often decomposing organic matter.
Figure 1. Life Stages of the house fly, Musca domestica
Because of the reliance on the forewings for flight, the mesothorax has become enlarged to contain the enormous flight muscles, and the pro- and metathorax are correspondingly reduced.
The mouthparts of flies are also characteristically suctorial and many have large fleshy pads with drainage canals termed pseudotracheae for efficient liquid uptake. Some flies have mouthparts modified for stabbing and piercing other insects, such as the predatory robber-flies (Asilidae) and dance flies (Empididae). Mosquitoes and some other ectoparasitic groups have mouthparts modified for piercing the skin of a vertebrate host and removing blood and other fluids.
Larval Diptera are typically small, pale and soft-bodied. They lack true legs and move by peristaltic waves of muscular contraction through the body. The larvae of most species of flies have a reduced head capsule and all that remains are the mandibles and some associated sclerites which are collectively called the cephalopharyngeal skeleton.
Dipteran pupae have non-functional mandibles (adecticous), and may have the appendages free from the body (exarate), or glued to the body (obtect). If exarate, the pupa is concealed inside the hardened skin (puparium) of the last larval instar.
Figure 3. Olbiogaster sackeni, family Anisopodidae, Nematocera.© 1996 C. R. Nelson
Figure 4. Milesia scutellata, a flower fly, family Syrphidae, Aschiza, Cyclorrhapha.© 1996 C. R. Nelson
Figure 5. Paracantha sp., a fruit fly, family Tephritidae, Schizophora, Cyclorrhapha.© 1996 C. R. Nelson
The addition of data from broad-based comparative morphological studies of both adult and immature stages (for example, Courtney 1991; Sinclair 1992; Ovchinnikova 1989; Oosterbroek and Courtney 1995) and also from DNA sequences will be critical in the reformulation of dipteran classification. The pages at this web site will document the areas of agreement, outstanding difficulties, and research being conducted to derive a new classification. These are exciting times for students of dipteran classification.
Bickel, D. J. 1982. Diptera. In: S. P. Parker (ed.). Synopsis and Classification of Living Organisms, Vol. 2. McGraw-Hill, New York, pp. 563-599. Courtney, G. W. 1991. Phylogenetic analysis of the Blephariceromorpha, with special reference to mountain midges (Diptera: Deuterophlebiidae). Systematic Entomology 16(2): 137-172. Cumming, J. M., B. J. Sinclair, and D.M Wood. 1995. Homology and phylogenetic implications of male genitalia in Diptera-Eremoneura. Entomologica Scandinavica 26: 120-151. Ennos, A. R. 1989. Comparative functional morphology of the wings of Diptera. Zoological Journal Of The Linnean Society 96(1): 27-48. Ferrar, P. 1987. A guide to the breeding habits and immature stages of Diptera Cyclorrhapha. Entomonograph 8 (1-2), 907 pp. Leiden. Griffiths, G.C.D. 1972. The phylogenetic classification of Diptera Cyclorrhapha, with special reference to the male postabdomen. Series entomologica 8, 340pp. The Hague. Griffiths, G.C.D. 1994. Relationships among the major subgroups of Brachycera (Diptera): A critical review. The Canadian Entomologist, 126:861-880. Griffiths, G.C.D. 1996. Review of papers on the male genitalia of Dipteraby D.M. Wood and associates. Studia Dipterologica 3: 107-123. Grimaldi, D and J. Cumming. 1999. Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History. 239:1-124. Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschaftsbeziehungen. Beitrage zur Entomologie 8: 505-688. Hennig, W. 1973. Diptera. In: W. Kukenthal (ed.) Handbuch der Zoologie, IV: Arthropoda. de Gruyter, New York, pp. 1-337. King, D. G. 1991. The origin of an organ: Phylogenetic analysis of evolutionary innovation in the digestive tract of flies (Insecta: Diptera). Evolution 45(3): 568-588. Krivosheina, N. P. 1991. Phylogeny of lower Brachycera (Diptera): A new view. Acta Entomologica Bohemoslovaca 88(2): 81-92. Krzeminski, W. 1992. Triassic and Lower Jurassic stage of Diptera evolution. Mitteilungen Der Schweizerischen Entomologischen Gesellschaft 65(1-2): 39-59. Lawrence, D. 1992. The Making of a Fly, Blackwell Scientific, Inc., Oxford. McAlpine, J.F. 1989. Phylogeny and classification of the Muscomorpha. In: McAlpine J.F., Wood D.M. (eds.)Manual of Nearctic Diptera 3. Research Branch, Agriculture Canada, Monograph 32:1397-1518. McAlpine, J.F., B.V. Peterson, G.E. Shewell, H.J. Teskey, J.R. Vockeroth, and D.M. Wood (eds.). 1981, 1987. Manual of Nearctic Diptera, Vol. 1 & 2. Research Branch, Agriculture Canada, Monographs 27 & 28. McAlpine, J.F and D.M. Wood (eds.). Manual of Nearctic Diptera, Vol. 3. Research Branch, Agriculture Canada, Monograph 32. Michelsen, V. 1996. Neodiptera: New insights into the adult morphology and higher level phylogeny of Diptera (Insecta). Zoological Journal of the Linnean Society 117: 71-102. Nagatomi, A. 1991. History of some families of Diptera, chiefly those of the lower Brachycera (Insecta: Diptera). Bulletin Of The Biogeographical Society Of Japan 46(1-22): 21-38. Nagatomi, A. 1992. Notes on the phylogeny of various taxa of the orthorrhaphous Brachycera (Insecta: Diptera). Zoological Science 9(4): 843-857. Oosterbroek, P. and G. Courtney. 1995. Phylogeny of the nematocerous families of Diptera (Insecta). Zoological Journal of the Linnean Society 115:267-311. Oosterbroek, P. and B. Theowald. 1991. Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera): With an index to the literature except Tipulidae. Tijdschrift Voor Entomologie 134(2): 211-267. Ovchinnikova, O.G. 1989. Musculature of the male genitalia of Brachycera-Orthorrhapha (Diptera). Trudy Zoologicheskogo Instituta Akademiya Nauk SSSR 190: 1-166. Pape, T. 1992. Phylogeny of the Tachinidae family-group (Diptera: Calyptratae). Tijdschrift Voor Entomologie 135(1): 43-86. Sinclair, B. J. 1992. A phylogenetic interpretation of the Brachycera (Diptera) based on the larval mandible and associated mouthpart structures. Systematic Entomology 17(3): 233-252. Sinclair, B.J., Cumming, J.M. and D.M. Wood. 1994. Homology and phylogenetic implications of the male genitalia in Diptera-Lower Brachycera. Entomologica Scandinavica 24: 407-432. Wada, S. 1991. Morphological evidence for the direct sister group relationship between the Schizophora and the Syrphoidea (Aschiza) in the phylogenetic systematics of the Cyclorrhapha (Diptera: Brachycera). Journal Of Natural History 25(6): 1531-1570. Wiegmann, B.M., C. Mitter, and F.C. Thompson. 1993. Evolutionary origin of the Cyclorrhapha (Diptera): tests of alternative morphological hypotheses. Cladistics, 9:41-81. Wood, D. M. and A. Borkent 1989. Phylogeny and classification of the Nematocera. In: McAlpine J.F., Wood, D.M. (eds.) Manual of nearctic Diptera 3. Research Branch, Agriculture Canada, Monograph 32: 1333-1370. Woodley, N.E. 1989. Phylogeny and classification of the 'Orthorrhaphous' Brachycera. In: McAlpine J.F., Wood D.M. (eds.)Manual of Nearctic Diptera 3. Research Branch, Agriculture Canada, Monograph 32:1371-1395. Yeates, D.K. 1994. The cladistics and classification of the Bombyliidae (Diptera: Asiloidea). Bulletin of the American Museum of Natural History 219: 1-191. Yeates, D. K. and B. M. Wiegmann. 1999. Congruence and controversy: Toward a higher-level phylogeny of Diptera. Annual Review of Entomology 44: 397-428. Zatwarnicki, T. 1996. A new reconstruction of the origin of the eremoneuran hypopygium and its implications for classification (Insecta: Diptera). Genus 7:103-175.
Brian M. Wiegmann
Department of Entomology, Box 7613, North Carolina State University, Raleigh, NC 27695
David K. Yeates
Department of Entomology, University of Queensland, Brisbane, Qld 4072, Australia
Correspondence regarding this page should be directed to Brian M. Wiegmann, at firstname.lastname@example.org.
Page copyright © 1996 Brian M. Wiegmann and David K. Yeates
Last saved 16 January 1997
Title Figure 1. Tipula (Lunatipula) sp., a crane fly, family Tipulidae.
Title Figure 2. Condylostylussp., a long-legged fly, family Dolichopodidae.
Title Figure 3. Calodexia sp., a parasitoid of army ants from Costa Rica, family Tachinidae.