Forms, Functions and Names

  • Graham E. Rotheray
Part of the Zoological Monographs book series (ZM, volume 4)


Cyclorrhaphan larval potential to provide character data is poorly realised and the main aim of this chapter is to substantiate this claim. It does this by introducing some major sources of variation, such as the larval head and the shape and ornamentation of the body. The cyclorrhaphan larval head is one of the most problematic and controversial of structures within larval Diptera, but by including lower cyclorrhaphan taxa and combining morphological with movement analyses the head is shown to be resolvable and not as enigmatic as it seems. In fact the head is a major source of diversity. Underpinning the features of the cyclorrhaphan head are developments in the head pump which is a component of the Diptera groundplan, but highly developed in cyclorrhaphans, and for cyclorrhaphans, pumping is the chief means of transferring food into the alimentary tract. The whole body is involved in feeding and locomotion with modularised sections performing specific functions. These functions are supported by diversified body shapes and ornamentation which combine to form an astonishing array of specialised feeding and locomotor mechanisms. The distinguishing features of cyclorrhaphan larvae are also evaluated and synapomorphies assessed.


  1. Andries M (1912) Zur systematik, biologie und entwicklung von Microdon Meigen. Z Wiss Zool 103:300–361Google Scholar
  2. Anthon H (1943) Der Kopfhau der Larven einiger nematoceren dipterenfamilien: rhyphidae, trichoceridae, psychodidae und ptychopteridae. Spolia Zool Mus Hauniensis 3:1–61Google Scholar
  3. Berrigan D, Pepin DJ (1995) How maggots move: allometry and kinematics of crawling in larval Diptera. J Insect Physiol 41:329–337CrossRefGoogle Scholar
  4. Brauns A (1954) Terricole Dipterenlarven. Musterschmidt Wissenschaftlicher Verlag, BerlinGoogle Scholar
  5. Brown BV, Borkent A, Cumming JM, Wood DM, Woodley NE, Zumbado MA (2009/2010) Manual central American Diptera, vols 1–2. National Research Council of Canada, Research Press, Ottawa, ONGoogle Scholar
  6. Campos-Ortega JA, Hartenstein V (1997) The embryonic development of Drosophila melanogaster. Springer, BerlinCrossRefGoogle Scholar
  7. Chandler PJ (2001) The flat-footed flies (Diptera: Opetiidae and Platypezidae) of Europe. Fauna Ent Scand 36:1–276Google Scholar
  8. Collins KP, Wiegmann BM (2002) Phylogenetic relationships of the lower Cyclorrhapha (Diptera: Brachycera) based on 28S rDNA sequences. Insect Syst Evol 33:445–456CrossRefGoogle Scholar
  9. Cook EF (1949) The evolution of the head in the larvae of the Diptera. Microent 14:1–57Google Scholar
  10. Courtney GW, Sinclair BJ, Meier R (2000) Morphology and terminology of Diptera larvae. In: Papp L, Darvas B (eds) Contributions manual Palaearctic Diptera, vol 1. Science Herald, Budapest, pp 85–161Google Scholar
  11. de Meijere JCH (1900) Ueber die Larva von Lonchoptera. Ein Beitrag zur Kenntnis der cyclorraphen Dipterenlarven. Zool Jahrb (Ab Syst) 14:87–132Google Scholar
  12. Disney H (1994) Scuttle flies: the Phoridae. SpringerGoogle Scholar
  13. Dowding VM (1967) The function and ecological significance of the pharyngeal ridges occurring in the larvae of some cyclorrhaphous Diptera. Parasitol (Cam) 57:371–388CrossRefGoogle Scholar
  14. Ferrar P (1987) A guide to the breeding habits and immature stages of Diptera Cyclorrhapha. Entomonograph 8:1–907Google Scholar
  15. Green CH, Burnet B, Connolly KJ (1983) Organisation and patterns of inter- and intraspecific variation in the behaviour of Drosophila larvae. Anim Behav 31:282–291CrossRefGoogle Scholar
  16. Günther MN, Nettesheim G, Shubeita GT (2016) Quantifying and predicting Drosophila larvae crawling phenotypes. Sci Rep 10:1–10. CrossRefGoogle Scholar
  17. Hanslik U, Scoofs A, Niederegger S, Heinzel H-G, Spief R (2010) The thoracic muscular system and Its innervation in third instar Calliphora vicina larvae. 1. Muscles of the pro- and mesothorax and the pharyngeal complex. J Morphol 271:960–968PubMedGoogle Scholar
  18. Harrison DA, Cooper RL (2003) Characterization of development, behavior and neuromuscular physiology in the phorid fly, Megaselia scalaris. Comp Biochem Physiol A Mol Integr Physiol 136:427–439CrossRefGoogle Scholar
  19. Hartley JC (1963a) The cephalopharyngeal apparatus of syrphid larvae and its relationship to other Diptera. Proc Zool Soc Lond 141:261–280CrossRefGoogle Scholar
  20. Hartley JC (1963b) A taxonomic account of the larvae of some British Syrphidae. Proc Zool Soc Lond 136:505–573CrossRefGoogle Scholar
  21. Headrick DH, Goeden RD (1996) The biology of nonfrugivorous tephritid fruit flies. Annu Rev Entomol 43:217–241CrossRefGoogle Scholar
  22. Henneguy LF (1904) Les Insectes. Masson et cie, ParisGoogle Scholar
  23. Hennig W (1952) Die Larvenformen der Dipteren, vol 3. Akademie-Verlag, BerlinGoogle Scholar
  24. Hennig W (1973) Diptera (Zweiflügler). In: Kükenthal W (ed) Handbuch der Zoologie, eine Naturgeschichte der Stämme des Tierreiches IV:1–337Google Scholar
  25. Hennig W (1976) Das hypopygium von Lonchoptera lutea Panzer und die phylogenetischen verwandtschaftsbeziehungen der Cyclorrhapha (Diptera). Stuttgart: Beit Naturk 283:1–63Google Scholar
  26. Holloway BA (1976) A new bat-fly family from New Zealand (Diptera: Mystacinobiidae). NZ J Zool 3:279–301CrossRefGoogle Scholar
  27. Jürgens G, Lehmann R, Schardin M, Nüsslein-Volhard C (1986) Segmental organisation of the head in the embryo of Drosophila melanogaster. Roux’s Arch Dev Biol 195:359–377CrossRefGoogle Scholar
  28. Keilin D (1915) Recherches sur les larves de Dipteres Cyclorrhaphes. Bull Sci Fr Bel 49:15–198Google Scholar
  29. Kohsaka H, Okusawa S, Itakura Y, Fushiki A, Nose A (2012) Development of larval motor circuits in Drosophila. Devel Growth Differ 54:408–419CrossRefGoogle Scholar
  30. Lambkin C, Sinclair BJ, Pape T, Courtney GW, Skevington JH, Meier R, Yeates DK, Blagoderov V, Wiegmann BM (2013) The phylogenetic relationships among infraorders and superfamilies of Diptera based on morphological evidence. Syst Entomol 38:164–179CrossRefGoogle Scholar
  31. Ludwig CE (1949) Embryology and morphology of the larval head of Calliphora erythrocephala Meigen. Microentomology 14:75–111Google Scholar
  32. Mahmoud M, Bahgat M, Zalat S, Dewedar A (1999) Eristalinus larvae and the role of bacteria in their feeding. J Union Arab Biologists Cairo Zool 11A:417–433Google Scholar
  33. Matsuda R (1965) Morphology and evolution of the insect head. Mem Am Ent Soc 4:1–334Google Scholar
  34. McAlpine JF (1989) Phylogeny and classification of the Muscomorpha. In: McAlpine JF, Wood DM (eds) Manual Nearctic Diptera. Research Branch Agriculture Canada, Ottawa; Monograph No. 32, vol 3, pp 1397–1502Google Scholar
  35. Menees JH (1962) The skeletal elements of the gnathocephalon and its appendages in the larvae of higher Diptera. Ann Entomol Soc Am 55:607–616CrossRefGoogle Scholar
  36. Miller D (1933) The buccopharyngeal mechanism of a blow fly larva (Calliphora quadrimaculata Swed.). Parasitol (Cam) 24:591–597Google Scholar
  37. Morge G (1967) Die Lonchaeidae und Pallopteridae Österreichs und der angrenzenden Gebiete 2 Teil Pallopteridae. Naturkund Jahr Stadt Linz 13:141–212Google Scholar
  38. Neugart C, Schneeberg K, Beutel RG (2009) The morphology of the larval head of Tipulidae (Diptera, Insecta) – the dipteran groundplan and evolutionary trends. Zool Anz 248:213–235CrossRefGoogle Scholar
  39. Nye IWB (1958) The external morphology of Dipterous larvae occurring in the Gramineae of Britain. Trans R Ent Soc Lond 110:411–487CrossRefGoogle Scholar
  40. Papp L, Darvas B (eds) (1998, 2000) Contributions to a manual of Palaearctic Diptera. Vols 1–3. Budapest, Science HeraldGoogle Scholar
  41. Pratt HS (1901) The embryonic history of the imaginal discs in Melophagus ovinus L., together with an account of the earlier stages in the development of the insect. Proc Boston Soc Nat Hist 29:241–272Google Scholar
  42. Roberts MJ (1969) Structure of the mouthparts of the larvae of the flies Rhagio and Sargus in relation to feeding habits. J Zool (Lond) 159:381–398CrossRefGoogle Scholar
  43. Roberts MJ (1970) The structure of the mouthparts of syrphid larvae (Diptera) in relation to feeding habits. Acta Zool Stockh 51:43–65CrossRefGoogle Scholar
  44. Roberts MJ (1971a) The structure of the mouthparts of some calypterate dipteran larvae in relation to their feeding habits. Acta Zool 52:171–188CrossRefGoogle Scholar
  45. Roberts MJ (1971b) On the locomotion of cyclorrhaphan maggots (Diptera). J Nat Hist 5:583–590CrossRefGoogle Scholar
  46. Rotheray EL (2013) Differences in ecomorphology and microhabitat use of four saproxylic larvae (Diptera, Syrphidae) in Scots pine stump rot holes. Ecol Entomol 38:219–229CrossRefGoogle Scholar
  47. Rotheray GE (2016) Improving knowledge of the cyclorrhaphan larva (Diptera). J Nat Hist 50:2169–2198CrossRefGoogle Scholar
  48. Rotheray GE, Gilbert F (1999) Phylogeny of Palaearctic Syrphidae (Diptera): evidence from larval stages. Zool J Linnean Soc 127:1–112CrossRefGoogle Scholar
  49. Rotheray GE, Gilbert F (2008) Phylogenetic relationships and the larval head of the lower Cyclorrhapha (Diptera). Zool J Linnean Soc 153:287–323CrossRefGoogle Scholar
  50. Rotheray GE, Gilbert F (2011) The natural history of hoverflies. Forrest Text, CardiganGoogle Scholar
  51. Rotheray GE, Lyszkowski R (2015) Diverse mechanisms of feeding and movement in Cyclorrhaphan larvae (Diptera). J Nat Hist 49:2139–2211CrossRefGoogle Scholar
  52. Rotheray GE, Wilkinson G (2015) Trophic structure and function in the larva of predatory muscid flies (Diptera, Muscidae). Zoomorphology 134:553–563CrossRefGoogle Scholar
  53. Rotheray GE, Chandler PJ, Gilbert F (2004) Final stage larvae and puparia of Platypezidae (Diptera). Insect Syst Evol 35:79–105CrossRefGoogle Scholar
  54. Rotheray GE, Hancock EG, Marcos-Garcia M (2007) Neotropical Copestylum (Diptera, Syrphidae) breeding in bromeliads (Bromeliaceae) including 22 new species. Zool J Linnean Soc 150:267–317CrossRefGoogle Scholar
  55. Schneeberg K, Beutel RG (2014) The evolution of head structures in lower Diptera. Sci Open Res.
  56. Schoeller J (1964) Recherches descriptives er experimentales sur la cephalogenese de Calliphora etythrocephala (Meigen), au cours des developpements embryonnaire et postembryonnaire. Archs Zool Expo Gen 103:1–216Google Scholar
  57. Schremmer F (1951) Die Mundteile der Brachycerenlarven and der Kopfbau der Larve von Stratiomys chamaeleon. L Osterr Zool Z 3:326–397Google Scholar
  58. Schremmer F (1956) Funktionsmophologische Srudien an Diprerenlarven. Verh. Dt. Zool. Ges 1956:301–305Google Scholar
  59. Sinclair BJ (1992) A phylogeneric interpretation of the Brachycera (Diptera) based on the larval mandible and associated mouthpart structures. Syst Entomol 17:233–252CrossRefGoogle Scholar
  60. Sinclair BJ, Cumming JM (2006) The morphology, higher-level phylogeny and classification of the Empidoidea (Diptera). Zootaxa 1180:1–172CrossRefGoogle Scholar
  61. Skidmore P (1985) The biology of the Muscidae of the world. Junk, DordrechtGoogle Scholar
  62. Smith KGV (1966) The larva of Thecophora occidensis, with comments upon the biology of Conopidae (Diptera). J Zool (Lond) 149:263–276CrossRefGoogle Scholar
  63. Smith KGV (1989) An introduction to the immature stages of British flies. Handbks Ident Br Insects 10:1–280Google Scholar
  64. Smith KGV, Van Someren C (1970) The identity of Physocephala bimarginipennis Karsh (Diptera, Conopidae) with nites on the immature stages and biology. J Nat Hist 4:439–446CrossRefGoogle Scholar
  65. Snodgrass RE (1935) Principles of insect morphology. McGraw Hill Book Co, New YorkGoogle Scholar
  66. Snodgrass RE (1947) The insect cranium and the ‘epicranial suture’. Smithsonian Misc Colls 107:1–56Google Scholar
  67. Snodgrass RE (1953) The metamorphosis of a fly’s head. Smithsonian Misc Colls 122:1–25Google Scholar
  68. Ståhls G, Hippa H, Rotheray G, Muona J, Gilbert F (2003) Phylogeny of Syrphidae (Diptera) inferred from combined analysis of molecular and morphological characters. Syst Entomol 28:433–450CrossRefGoogle Scholar
  69. Teskey HJ (1981) Morphology and terminology – larvae. In: McAlpine J, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR, Wood DM (eds) Manual Nearctic Diptera, vol 1. Research Branch Agriculture Canada, Hull, Quebec, pp 65–88Google Scholar
  70. Tkoč M, Tóthová A, Ståhls G, Chandler PJ, Vaňhara J (2016) Molecular phylogeny of flat-footed flies (Diptera: Platypezidae): main clades supported by new morphoogical evidence. Zool Scr 46:429–444CrossRefGoogle Scholar
  71. Trautwein MD, Wiegmann BM, Beutel R, Kjer KM, Yeates DK (2012) Advances in insect phylogeny at the dawn of the postgenomic era. Annu Rev Entomol 57:449–468CrossRefGoogle Scholar
  72. Turner FR, Mahowald AP (1979) Scanning electron microscopy of Drosophila melanogaster embryogenesis. III. Formation of the head and caudal segments. Develop Biol 68:96–109CrossRefGoogle Scholar
  73. Vaillant F (1948) les premiers stades de Liancalus virens Scop. (Dolichopodidae). Bull Soc Zol Fr 6:41–52Google Scholar
  74. Weise H (1938) Die Atmung der Larven und Puppen der Schwebfleigen aus der Verwandtshaft der Eristalinae unter Berücksichtigung ihrer Metamorphose. Z Wiss Zool 151:467–514Google Scholar
  75. Wiegmann BM, Trautwein MD, Winkler IS, Barra NB, Kima J-W, Lambkin C, Berton MA, Cassela BK, Bayless KM, Heimberg AM, Wheeler BM, Petersone KJ, Pape T, Sinclair BJ, Skevington JH, Blagoderov V, Caravask J, Narayanan Kutty SN, Schmidt-Ott U, Kampmeier GE, Thompson FC, Grimaldi DA, Beckenbach AT, Courtney GM, Friedrich M, Meier R, Yeates DK (2011) Episodic radiations in the fly tree of life. Proc Natl Acad Sci U S A 108:5690–5695CrossRefGoogle Scholar
  76. Wilkinson G, Rotheray GE (2017) Melanostoma scalare (Meigen) larvae (Diptera, Syrphidae) feed on Diptera larvae in leaf litter. Dipt Digest 24:53–60Google Scholar
  77. Wipfler B, Schneeberg K, Löffler A, Hünefeld F, Meier R, Beutel RG (2013) The skeletomuscular system of the larva of Drosophila melanogaster (Drosophilidae, Diptera) – a contribution to the morphology of a model organism. Arthop Struct Develop 42:47–68CrossRefGoogle Scholar
  78. Wood et al. (1981–1989) In: McAlpine JF, Peterson BV, Shewell GE, Teskey HJ, Vockeroth JR, Wood DM (eds) (1981, 1987, 1989) Manual Nearctic Diptera, vols 1–3. Research Branch, Agriculture Canada. MonographGoogle Scholar
  79. Yang AS (2001) Modularity, evolvability, and adaptive radiations: a comparison of the hemi- and holometabolous insects. Evol Dev 3:59–72CrossRefGoogle Scholar
  80. Yeates DK, Wiegmann BM (1999) Congruence and controversy: toward a higher-level classification of Diptera. Annu Rev Entomol 44:397–428CrossRefGoogle Scholar
  81. Ylioja T, Saranpää P, Roininen H, Rousi M (1998) Larval tunnels of Phytobia betulae (Diptera: Agromyzidae) in birch wood. J Econ Entomol 4:185–181Google Scholar

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© Springer Nature Switzerland AG 2019

Authors and Affiliations

  • Graham E. Rotheray
    • 1
  1. 1.National Museums of ScotlandEdinburghUK

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