Tabanidae

Splayed deer fly Chrysops caecutiens, copyright Hectonichus.

Belongs within: Tabanomorpha.
Contains: Diachlorini, Tabanini, Pangoniini, Scionini, Bouvieromyiini.

The Tabanidae, horse flies, are a group of medium-sized to large flies, females of which commonly feed on vertebrate blood. Males and females of this family are readily distinguished by the eyes being holoptic in males but distinctly separated in females.

Characters (from Pechuman & Teskey 1981): Moderate to large flies, 6-30 mm long, without bristles. Head large; eyes large, often brightly patterned, holoptic in male, separated by frons in female. Antenna porrect and composed of scape, pedicel, and flagellum; flagellum usually consisting of a larger basal portion and terminal annulations usually numbering four to eight but sometimes as few as two. Palpus composed of two segments; basal palpal segment usually rather small; other palpal segment usually rather long and curved downward in female, often inflated near base. Proboscis stout and rigid; mandible and maxilla of female often styliform and adapted for piercing, sometimes absent in species that are not bloodsuckers. Thorax large; notopleural lobe prominent; scutellum without spines or macrotrichia. Legs rather stout; apical spurs present on mid tibia, absent on fore tibia, present or absent on hind tibia; tarsi each with three pads formed by pulvilliform empodium located between two pulvilli. Wing venation with as many as 11 veins reaching margin (Sc, four branches of R, three branches of M, CuA₁, CuA₂, A₁), C extending around wing, cells br, bm, and d conspicuous, and cell cup usually closed near wing margin; calypter large. Abdomen broad, with seven segments usually visible; terminalia of both sexes usually inconspicuous; male with gonocoxites fused with sternite 9, with gonostylus single or partially divided, aedeagus and associated pair of slender filamentous recurved aedeagal tines enclosed in a sheath apparently derived from parameres, tergite 9 entire or divided, tergite l0 absent, cercus flattened, round to oval or somewhat pointed; female usually with tergite l0 divided, with sternite 8 shield-shaped, and with cercus one-segmented. Larva fusiform or more or less expanded posteriorly or anteriorly, comprising 11 body segments and completely retractile head capsule; head capsule elongate, subcylindrical, tapering anteriorly, features including semitubular cranium, paired tentorial arms attached anteriorly to inner dorsal surfaces of cranium and appearing to be free posteriorly, three-segmented antennae, median down-curved labrum, mandibular-maxillary sclerites including maxillary palpi, and labium closing head capsule ventrally; mandibles each with central canal for excretion of paralytic enzyme, linked with subdorsal brushes of spines that are erected to anchor head within host when mandibles strike downward; body without projecting appendages on thoracic segments, with annular rings of small tubercles and crenulate frills or with prolegs on abdominal segments; respiratory system functionally metapneustic.

<==Tabanidae B09
    |  i. s.: Japenoides BLM02
    |         Mesopangonius RL09
    |         Pangonia A71
    |           |--P. lasiophtalma Boisduval 1835 B35
    |           `--P. longirostris A71
    |--Scepsidinae [Scepsidini] M54
    |    |--Adersia oestroides M54
    |    |--Scepsis nivalis M54
    |    |--Braunsiomyia M54
    |    `--Lesneus M54
    |--Tabaninae PT81
    |    |--Diachlorini PT81
    |    |--Tabanini PT81
    |    `--Haematopotini PT81
    |         |--Heptatoma M54
    |         |--Hippocentrum M54
    |         `--Haematopota PT81
    |              |--H. americana PT81
    |              `--H. pluvialis WT11
    |--Pangoniinae PT81
    |    |  i. s.: Palaepangonius GE05
    |    |         Eopangonius GE05
    |    |         Scaptiella aperta M54
    |    |--Pangoniini PT81
    |    |--Scionini PT81
    |    `--Philolichini M54
    |         |--Philoliche M54
    |         |--Ommatiosteres M54
    |         |--Metaphara M54
    |         |--Nuceria M54
    |         |--Stenophara M54
    |         |--Dorcaloemus compactus M54
    |         |--Phara M54
    |         |--Subpangonia M54
    |         `--Buplex M54
    |              |--B. albifacies M54
    |              `--B. suavus M54
    `--Chrysopsinae PT81
         |--Bouvieromyiini PT81
         |--Rhinomyzini M54
         |    |--Tabanocella denticornis M54
         |    |--Orgizomyia M54
         |    |--Thriambeutes M54
         |    |--Sphecodemyia M54
         |    |--Gastroxides ater M54
         |    |--Rhinomyza M54
         |    |--Thaumastocera M54
         |    `--Guyona M54
         `--Chrysopsini PT81
              |--Nemorius M54
              |--Melissomorpha indiana M54
              |--Neochrysops globosus PT81
              |--Silvius PT81
              |    |  i. s.: S. vituli M54
              |    |--S. (Assipala) ceras PT81
              |    |--S. (Griseosilvius) B09
              |    |    |--S. (G.) atitlanensis B09
              |    |    `--S. (G.) quadrivittatus PT81
              |    `--S. (Zeuximyia) philipi PT81
              `--Chrysops B09
                   |--C. caecutiens M54
                   |--C. cincticornis PT81
                   |--C. dimidiata A71
                   |--C. discalis A71
                   |--C. excitans PT81
                   |--C. furcatus PT81
                   |--C. (Liochrysops) hyalinus PT81
                   |--C. marmoratus K01
                   |--C. pikei PT81
                   |--C. scalaratus B09
                   `--C. silacea A71

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[BLM02] Blagoderov, V. A., E. D. Lukashevich & M. B. Mostovski. 2002. Order Diptera Linné, 1758. The true flies (=Muscida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 227–240. Kluwer Academic Publishers: Dordrecht.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[B09] Burger, J. F. 2009. Tabanidae (horse flies, deer flies, tabanos). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 495–507. NRC Research Press: Ottawa.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[M54] Mackerras, I. M. 1954. The classification and distribution of Tabanidae (Diptera). I. General review. Australian Journal of Zoology 2 (3): 431–454.

[PT81] Pechuman, L. L., & H. J. Teskey. 1981. Tabanidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 463–478. Research Branch, Agriculture Canada.

[RL09] Ren, D., C. C. Labandeira, J. A. Santiago-Blay, A. Rasnitsyn, C. Shih, A. Bashkuev, M. A. V. Logan, C. L. Hotton & D. Dilcher. 2009. A probable pollination mode before angiosperms: Eurasian, long-proboscid scorpionflies. Science 326: 840–847.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated: 13 May 2018.

Stratiomyini

Odontomyia sp., copyright Muhammad Mahdi Karim.

Belongs within: Stratiomyidae.

The Stratiomyini are a group of soldier flies with the antenna lacking an apical stylus, and vein A₁ in the wing straight. They may be divided between tribes Stratiomyini sensu stricto and Odontomyiini based on whether the antenna has five (Stratiomyini) or six (Odontomyiini) flagellomeres (James 1981) but this division is not recognised below, following Woodley (2009). Members of the Stratiomyini often resemble bees or wasps and may frequent flowers as adults (Woodley 2009).

Characters (from James 1981, as Stratiomyini + Odontomyiini): Antennal flagellum with five or six flagellomeres, not aristate. Scutellum with spines (sometimes minute). Wing with crossvein m-cu present; cell d present; A₁ not at all sinuous. Abdomen at least three-quarters as wide as long, usually patterned.

<==Stratiomyini [Odontomyiini] W09
    |--Psellidotus johnsoni [=Labostigmina johnsoni] W09
    |--Panamamyia silbergliedi W09
    |--Stratiomyella nana W09
    |--Chloromelas W09
    |--Zuerchermyia [=Zuercheria (preoc.)] W09
    |    `--Z. bequaerti W09
    |--Anoplodonta W09
    |    |--A. fratella W09
    |    `--A. nigrirostris J81
    |--Promeranisa W09
    |    |--P. nasuta W09
    |    `--P. varipes W09
    |--Hoplitimyia W09
    |    |--H. constans W09
    |    |--H. inbioensis W09
    |    `--H. subalba W09
    |--Hedriodiscus W09
    |    |--H. binotatus W09
    |    |--H. dorsalis W09
    |    |--H. trivittatus W09
    |    `--H. truquii W09
    |--Stratiomys W09
    |    |--S. badia M90
    |    |--S. barbata W09
    |    |--S. chamaeleon L02
    |    |--S. constricta W09
    |    |--S. convexa W09
    |    |--S. longicornis [=Hirtea longicornis] L02
    |    |--S. norma W09
    |    |--S. potamida I92
    |    `--S. strigata L02
    `--Odontomyia W09
         |--O. cincta W09
         |--O. discolorata W09
         |--O. (Odontomyia) furcata [=Stratiomys (O.) furcata] G20
         |--O. hydroleon K01
         |--O. (Catatasina) interrupta J81
         |--O. limbata Macquart 1847 E12
         |--O. ornata K01
         |--O. pilimana W09
         |--O. scutellata CM91
         |--O. (Odontomyiina) virgo J81
         `--O. viridula K01

*Type species of generic name indicated

REFERENCES

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[I92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[J81] James, M. T. 1981. Stratiomyidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 497–511. Research Branch, Agriculture Canada.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[M90] McAlpine, J. F. 1990. Insecta: Diptera adults. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1211–1252. John Wiley & Sones: New York.

[W09] Woodley, N. E. 2009. Stratiomyidae (soldier flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 521–549. NRC Research Press: Ottawa.

Last updated: 1 January 2021.

Ceratopogonidae

Female Leptoconops, copyright Alice Abela.

Belongs within: Culicomorpha.
Contains: Forcipomyia, Culicoidini, Ceratopogonini, Sphaeromiini.

The Ceratopogonidae, biting midges, are a cosmopolitan group of small flies, a number of species of which have females that feed on vertebrate blood. Four subfamilies are recognised, the most divergent of which includes the genus Leptoconops which has eyes widely separated on top of the head (vs closely approximated in other subfamilies) and lacks the crossvein r-m in the wing (Downes & Wirth 1981).

Characters (from Downes & Wirth 1981): Small flies, 1–6 mm long, slender to moderately robust. Antenna with 11–13 flagellomeres, showing marked sexual dimorphism; female with five distal flagellomeres elongated and differing from proximal flagellomeres in many details; male with pedicel greatly enlarged and containing a specialised Johnston's organ, flagellomeres 1–8 usually bearing whorls of long setae giving characteristic plumose appearance. Proboscis about as long as head; palpus slightly longer. Female with biting and sucking mouthparts, consisting of a rigid labrum, blade-like mandibles and laciniae, and a hypopharynx, each typically with apical armature; females in non-biting species and all males with proboscis weaker and armature reduced. Palpus five-segmented; segments 1 and 2 sometimes poorly developed; segment 3 with sensory organ enclosed in a pit or on surface; segments 4 and 5 occasionally fused. Wings held horizontally at rest one above the other, narrower in male than in female; apex evenly rounded. Venation characterized by compact radial system lying closely behind costal margin and meeting margin before apex of wing, often near or even before the midpoint. R₁ and Rs typically short, with Rs branching into R₂₊₃, and R₄₊₅; R₂₊₃ running forward and rejoining R₁ to form cell r₁ (usually termed first radial cell); R₄₊₅ meeting C farther out to form cell r₂₊₃; one or both of these cells sometimes lost either by closure (approximation of upper and lower veins) or by becoming confluent through loss of R₂₊₃. Strong and characteristic crossvein r-m present. Posterior veins relatively weak; M₂ sometimes incomplete basally or absent; crossvein m-cu absent; cubital fork well-developed; CuP and A weak, not reaching wing margin. Legs moderately long; claws usually similar, average in size; male with small claws that are usually bifid at tips. Abdomen l0-segmented; spiracles present on segments 2–7; segment 1 somewhat reduced; tergites 1 and 2 sometimes fused. Sternite 8 in female with slightly bilobate hind margin extending below opening of spermathecal duct; segment 9 with small tergite fused laterally to narrow band-like sternite, sternite usually dividing in midline around spermathecal opening; segment l0 small; cercus rounded, on small dorsolateral basal plate. Segment 8 in male with normal tergite and sternite; tergite 9 usually enlarged and prolonged backward to cover other elements of terminalia; gonopod two-segmented, conspicuous; aedeagus extending back from sternite 9 and articulated laterally with base of gonocoxite, often triangular with sclerotized lateral arms or ventral plate; paired parameres usually present above aedeagus. Early stages usually found in moist or aquatic habitats; larva apneustic, often elongate in form, and a strong swimmer; head usually sclerotized; mandible strong, toothed, not apposed; pharyngeal apparatus conspicuous internally, with two strongly diverging arms and series of combs apparently serving to reduce and sort food; short collar between head and thorax; three thoracic and nine abdominal segments well-defined; no functional spiracles.

<==Ceratopogonidae [Heleidae]
    |--Dasyhelea BSG09 [Dasyheleinae DW81]
    |    |--D. antiqua S02
    |    |--D. cincta BSG09
    |    `--D. pseudoincisurata DW81
    |--Leptoconops BSG09 [Leptoconopinae DW81]
    |    |  i. s.: L. bequaerti BSG09
    |    |--L. (Leptoconops) torrens DW81
    |    |--L. (Brachyconops) californiensis DW81
    |    `--L. (Megaconops) floridensis DW81
    |--Forcipomyiinae DW81
    |    |--Forcipomyia DW81
    |    `--Atrichopogon DW81
    |         |--A. (Atrichopogon) levis DW81
    |         `--A. polydactylus DW81
    `--Ceratopogoninae DW81
         |--Culicoidini DW81
         |--Ceratopogonini DW81
         |--Sphaeromiini DW81
         |--Stenoxenini DW81
         |    |--Stenoxenus DW81
         |    |    |--S. coomani DW81
         |    |    `--S. johnsoni DW81
         |    `--Paryphoconus DW81
         |         |--P. angustipennis DW81
         |         `--P. sonorensis DW81
         |--Heteromyiini DW81
         |    |--Heteromyia fasciata DW81
         |    |--Neurohelea nigra DW81
         |    |--Neurobezzia granulosa DW81
         |    |--Clinohelea bimaculata DW81
         |    `--Pellucidomyia DW81
         |         |--P. ugandae BSG09
         |         `--P. wirthi DW81
         |--Palpomyiini DW81
         |    |--Pachyhelea pachymera DW81
         |    |--Bezzia DW81
         |    |    |--B. nobilis BSG09
         |    |    `--B. setulosa DW81
         |    |--Palpomyia BSG09
         |    |    |--P. lacustris BSG09
         |    |    |--P. lineata DW81
         |    |    `--P. plebeja BSG09
         |    `--Phaenobezzia DW81
         |         |--P. maya BSG09
         |         |--P. opaca DW81
         |         `--P. pistiae DW81
         `--Stilobezziini DW81
              |--Echinohelea lanei DW81
              |--Serromyia femorata DW81
              |--Parabezzia DW81
              |    |--P. bystraki BSG09
              |    `--P. petiolata DW81
              |--Monohelea DW81
              |    |--M. hieroglyphica DW81
              |    `--M. (Schizohelea) leucopeza DW81
              `--Stilobezzia DW81
                   |--S. antennalis DW81
                   |--S. (Eukraiohelea) elegantula DW81
                   `--S. (Neostilobezzia) lutea DW81

Ceratopogonidae incertae sedis:
  Austroconops GE05
    |--A. fossilis GE05
    `--A. macmillani GE05
  Fittkauhelea amazonica BSG09
  Cacaohelea youngi BSG09
  Baeohelea nana BSG09
  Leptohelea micronyx BSG09
  Nannohelea bourioni BSG09
  Schizonyxhelea forattinii BSG09
  Allohelea BSG09
    |--A. johannseni BSG09
    `--A. neotropica BSG09
  Parastilobezzia leei BSG09
  Downeshelea stonei BSG09
  Baeodasymyia dominicana BSG09
  Amerohelea BSG09
    |--A. frontispina BSG09
    `--A. galindoi BSG09
  Clastrieromyia dycei BSG09
  Spinellihelea BSG09
  Macrurohelea CM91
  Acanthohelea CM91
  Xenohelea CM91
  Dibezzia CM91
  Archiaustroconops alavensis Szadziewski & Arillo 1998 PR13
  Styloconops CM70

*Type species of generic name indicated

REFERENCES

[BSG09] Borkent, A., G. R. Spinelli & W. L. Grogan Jr. 2009. Ceratopogonidae (biting midges, purrujas). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 407–435. NRC Research Press: Ottawa.

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[DW81] Downes, J. A., & W. W. Wirth. 1981. Ceratopogonidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 393–421. Research Branch, Agriculture Canada.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[PR13] Peris, D., & E. Ruzzier. 2013. A new tribe, new genus, and new species of Mordellidae (Coleoptera: Tenebrionoidea) from the Early Cretaceous amber of Spain. Cretaceous Research 45: 1–6.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

Last updated: 7 May 2018.

Tabanini

Band-eyed brown horsefly Tabanus bromius, copyright Entomart.

Belongs within: Tabanidae.

The Tabanini is a cosmopolitan group of horse flies, most diverse in the Northern Hemisphere. Some species of Tabanus grow quite large, up to three centimetres in length.

Characters (from Pechuman & Teskey 1981): Scape barely longer than wide. Basicosta with macrotrichia similar in appearance and density to those on C. Hind tibia without apical spurs. Male gonostylus truncate. Ends of spermathecal ducts in female with mushroom-like expansions.

<==Tabanini PT81
    |--Styporhamphis M54
    |--Atylotus incisuralis PT81
    |--Abatylotus M54
    |--Phyrta M54
    |--Dasyrhamphis M54
    |--Efflatounanus M54
    |--Agkistrocerus megerlei PT81
    |--Hamatabanus carolinensis PT81
    |--Whitneyomyia beatifica PT81
    |--Lophotabanus M54
    |--Taeniotabanus M54
    |--Euancala M54
    |--Ancala M54
    |--Poeciloderas quadripunctatus B09
    |--Therioplectes M54
    |    |--T. tarandinus K01
    |    `--T. tropicus K01
    |--Hybomitra PT81
    |    |--H. epistates PT81
    |    |--H. lasiophthalma M90
    |    `--H. rhombica M54
    `--Tabanus Linnaeus 1758 L58
         |--T. albipes L02
         |--T. americanus M90
         |--T. antarcticus Linnaeus 1758 L58
         |--T. atratus WT11
         |--T. bovinus Linnaeus 1758 L58
         |--T. bromius Linnaeus 1758 L58
         |--T. caecutiens Linnaeus 1758 L58
         |--T. calens Linnaeus 1758 L58
         |--T. ceylonicus M70b
         |--T. exaestuans Linnaeus 1758 L58
         |--T. fervens Linnaeus 1758 L58
         |--T. (Glaucops) fratellus PT81
         |--T. fuscicostatus A71
         |--T. lineola PT81
         |--T. marginalis T90
         |--T. mexicanus Linnaeus 1758 L58
         |--T. micans A71
         |--T. occidentalis Linnaeus 1758 L58
         |--T. particaecus M70a
         |--T. pluvialis Linnaeus 1758 L58
         |--T. reinwardtii T90
         |--T. similis PT81
         |--T. striatus A71
         |--T. subsimilis PT81
         |--T. sudeticus K01
         |--T. sulcifrons M90
         |--T. tarandinus Linnaeus 1758 L58
         |--T. texanus BM76
         `--T. tropicus Linnaeus 1758 L58

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[B09] Burger, J. F. 2009. Tabanidae (horse flies, deer flies, tabanos). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 495–507. NRC Research Press: Ottawa.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M54] Mackerras, I. M. 1954. The classification and distribution of Tabanidae (Diptera). I. General review. Australian Journal of Zoology 2 (3): 431–454.

[M70a] Mackerras, I. M. 1970a. Skeletal anatomy. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 3–28. Melbourne University Press.

[M70b] Mackerras, I. M. 1970b. Composition and distribution of the fauna. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 187–203. Melbourne University Press.

[M90] McAlpine, J. F. 1990. Insecta: Diptera adults. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1211–1252. John Wiley & Sones: New York.

[PT81] Pechuman, L. L., & H. J. Teskey. 1981. Tabanidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 463–478. Research Branch, Agriculture Canada.

[T90] Teskey, H. J. 1990. Insecta: Diptera larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1253–1276. John Wiley & Sones: New York.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated: 13 May 2018.

Tipulinae

Tipula oleracea, copyright David Nicholls.

Belongs within: Tipulidae.

The Tipulinae are a group of mostly large crane flies, with wing lengths commonly over 10 mm. The legs are particularly long and slender in members of the genera Dolichopeza, Megistocera and Brachypremna; other genera have stouter albeit still long legs (Alexander & Byers 1981). The cosmopolitan genus Tipula includes about 2500 species, often with an opaque, pruinose coloration, characterised by a relatively long subcostal vein meeting R₁ beyond the origin of Rs (Gelhaus 2009).

Characters (from Alexander & Byers 1981): Size large; body length commonly over 8–10 mm; wing commonly over 10 mm, usually much larger. Terminal segment of palpus elongate; nasus usually distinct. Flagellum commonly with eleven segments. Wing with Sc₁ usually atrophied or incomplete; CuA slightly constricted at branching of CuA₁ and CuA₂. Larva with spiracular disc bordered by six (rarely eight) usually subconical lobes usually arranged with two dorsally, two dorsolaterally, and two below spiracles; these lobes sometimes short and blunt or sclerotised and hook-like.

<==Tipulinae [Tipulini] G09
    |--Megistocera longipennis G09
    |--Brachypremna dispellens G09
    |--Clytocosmus helmsi CM91
    |--Ptilogyna ramicornis CM91
    |--Stygeropsis A16
    |--Plusiomyia CM70
    |--Macromastix CM70
    |--Pselliophora G09
    |    |--P. margarita G09
    |    `--P. xanthopimplina Enderlein 1921 [incl. P. quadrivittata Edwards 1921] A40
    |--Holorusia AB81
    |    |--H. hespera G09
    |    `--H. rubiginosa AB81
    |--Dolichopeza AB81
    |    |--D. americana WT11
    |    `--D. subalbipes TW05
    |--Prionocera AB81
    |    |--P. dimidiata AB81
    |    |--P. gracilistyla HP68
    |    `--P. parrii G09
    |--Ctenacroscelis A40
    |    |--C. clavipes Edwards 1921 A40
    |    |--C. calliergon Alexander 1940 A40
    |    `--C. serratus A40
    |--Dictenidia A40
    |    |--D. luteicostalis A40
    |    |    |--D. l. luteicostalis A40
    |    |    `--D. l. longisector Alexander 1940 A40
    |    `--D. stalactitica Alexander 1940 A40
    |--Ischnotoma G09
    |    |  i. s.: I. eburnea G09
    |    |         I. rubriventris CM91
    |    |--I. (Ischnotoma) fagetorum G09
    |    |--I. (Icriomastix) jujuyensis G09
    |    `--I. (Neotipula) maya G09
    |--Ozodicera G09
    |    |--O. (Ozodicera) G09
    |    |    |--O. (O.) cinereipennis G09
    |    |    `--O. (O.) eurystyla G09
    |    `--O. (Dihexaclonus) G09
    |         |--O. (D.) macracantha G09
    |         `--O. (D.) panamensis G09
    |--Zelandotipula G09
    |    |--Z. associans G09
    |    |--Z. furcifera G09
    |    |--Z. gracilipes G09
    |    |--Z. laevis G09
    |    |--Z. novarae G09
    |    `--Z. sinuosa G09
    |--Leptotarsus G09 [incl. Longurio AB81]
    |    |  i. s.: L. imperatorius CM91
    |    |         L. testaceus G09
    |    |--‘Longurio’ (Longurio) fulvus Edwards 1916 A40
    |    |--L. (Pehlkea) G09
    |    |    |--L. (P.) colombianus G09
    |    |    `--L. (P.) regina G09
    |    `--L. (Tanypremna) G09
    |         |--L. (T.) incompletus G09
    |         `--L. (T.) opilio G09
    |--Ctenophora AB81
    |    |  i. s.: C. apicata G09
    |    |         C. perjocosa Alexander 1940 A40
    |    |         C. pselliophoroides A40
    |    |--C. (Ctenophora) bimaculata [=Tipula (C.) bimaculata] G20
    |    |--C. (Tanyptera) AB81
    |    |    |--‘Tanyptera’ antica Alexander 1938 A40
    |    |    `--C. (T.) dorsalis AB81
    |    `--C. (Phoroctenia) vittata AB81
    |         |--C. v. vittata AB81
    |         `--C. v. angustipennis AB81
    |--Nephrotoma Meigen 1803 (see below for synonymy) A16
    |    |--*N. dorsalis A16 [=Tipula (Nephrotoma) dorsalis G20]
    |    |--N. australasiae ZS10
    |    |--‘Pachyrrhina’ crocata K01
    |    |--N. eucera [=Pachyrrhina eucera] A16
    |    |--N. ferruginea G09
    |    |--N. flavescens V09
    |    |--‘Pachyrrhina’ histrio K01
    |    |--N. incurva A16
    |    |--N. lugens A16
    |    |--N. macrocera A16
    |    |--N. pedunculata A16
    |    |--N. penumbra Alexander 1916 A16
    |    |--‘Pachyrrhina’ scurra K01
    |    `--N. vittula A16
    `--Tipula Linnaeus 1758 A16
         |  i. s.: T. alta Doane 1912 A16
         |         T. angulata Loew 1864 [incl. T. decora Doane 1901] A16
         |         T. angustipennis Loew 1863 A16
         |         T. annulata Linnaeus 1758 L58
         |         T. apicalis Loew 1863 A16
         |         T. appendiculata Loew 1863 A16
         |         T. atrata Linnaeus 1758 L58
         |         T. australis Doane 1901 A16
         |         T. balioptera Loew 1863 A16
         |         T. bella Loew 1863 A16
         |         T. besselsi Osten Sacken 1876 A16
         |         T. bicornis Forbes 1891 A16
         |         T. brevicornis Linnaeus 1758 L58
         |         T. californica [=Pachyrrhina californica] A16
         |         T. ‘californica’ Doane 1912 non Pachyrrhina californica A16
         |         T. canadensis Loew 1864 A16
         |         T. carinifrons HP68
         |         T. cayuga Alexander 1916 A16
         |         T. centralis Loew 1864 A16
         |         T. cincta Loew 1863 A16
         |         T. cincticornis A16
         |         T. collaris [=Pachyrrhina collaris] A16
         |         T. contaminata Linnaeus 1758 L58
         |         T. cornicina Linnaeus 1758 L58
         |         T. crocata Linnaeus 1758 L58
         |         T. culiciformis L02
         |         T. cunctans Say 1823 [incl. T. casta Loew 1863, T. infuscata Loew 1863] A16
         |         T. dietziana Alexander 1916 A16
         |         T. discolor Loew 1863 A16
         |         T. fallax Loew 1863 A16
         |         T. febrilis Linnaeus 1758 L58
         |         T. fenestrarum L02
         |         T. flavescens Linnaeus 1758 L58
         |         T. florilega Linnaeus 1758 L58
         |         T. fragilis Loew 1863 [incl. T. suspecta Loew 1863] A16
         |         T. fraterna Loew 1864 A16
         |         T. fuliginosa (Say 1823) [=Ctenophora fuliginosa; incl. T. speciosa Loew 1863] A16
         |         T. fungorum L02
         |         T. fuscipennis Meigen 1818 A16
         |         T. grata Loew 1863 A16
         |         T. hebes Loew 1863 A16
         |         T. hermannia Alexander 1916 [=T. fasciata Loew 1863 non Linnaeus 1767] A16
         |         T. hiemalis L02
         |         T. hortorum Linnaeus 1758 L58
         |         T. ignobilis Loew 1863 A16
         |         T. imperfecta Alexander 1916 A16
         |         T. irrorata F92
         |         T. jejuna A16
         |         T. johnsoniana Alexander 1916 A16
         |         T. juniperina Linnaeus 1758 L58
         |         T. kennicotti Alexander 1916 A16
         |         T. latipennis Loew 1864 A16
         |         T. littoralis Linnaeus 1758 L58
         |         T. loewiana Alexander 1916 A16
         |         T. longicornis Linnaeus 1758 L58
         |         T. longiventris Loew 1863 A16
         |         T. lunata Linnaeus 1758 L58
         |         T. macrocephala Linnaeus 1758 L58
         |         T. macrolabis Loew 1864 A16
         |         T. macrosterna [=Pachyrrhina macrosterna] A16
         |         T. mainensis Alexander 1916 [incl. T. laevigata ms] A16
         |         T. mandan Alexander 1916 A16
         |         T. marci Linnaeus 1758 L58
         |         T. maxima W01
         |         T. megaura Doane 1901 A16
         |         T. mingwe Alexander 1916 A16
         |         T. monilis Linnaeus 1758 L58
         |         T. morrisoni Alexander 1916 A16
         |         T. motatrix Linnaeus 1758 L58
         |         T. nigra Linnaeus 1758 L58
         |         T. nobilis [=Pachyrrhina nobilis] A16
         |         T. notata Linnaeus 1758 L58
         |         T. pachyrhinoides Alexander 1916 A16
         |         T. pallida Loew 1863 A16
         |         T. palustris Linnaeus 1758 L58
         |         T. parshleyi Alexander 1916 A16
         |         T. pectinicornis Linnaeus 1758 L58
         |         T. penicillata Alexander 1916 A16
         |         T. penobscot Alexander 1916 A16
         |         T. perlongipes Johnson 1909 A16
         |         T. phalaenoides Linnaeus 1758 L58
         |         T. piliceps Alexander 1916 A16
         |         T. pinnicornis Linnaeus 1758 L58
         |         T. plumosa Linnaeus 1758 L58
         |         T. polymera [=Pachyrrhina polymera] A16
         |         T. praecisa Loew 1872 A16
         |         T. pratensis Linnaeus 1758 L58
         |         T. pubera Loew 1864 A16
         |         T. putris Linnaeus 1758 L58
         |         T. quaylii AB81
         |         T. rangiferina Alexander 1916 A16
         |         T. regelationis Linnaeus 1758 L58
         |         T. replicata Linnaeus 1758 L58
         |         T. rostrata L02
         |         T. seminole Alexander 1916 A16
         |         T. septentrionalis Loew 1863 A16
         |         T. serrulata Loew 1864 A16
         |         T. serta Loew 1863 A16
         |         T. subfasciata Loew 1863 A16
         |         T. submaculata Loew 1863 [incl. T. bidens ms] A16
         |         T. sulphurea A16
         |         T. taughannock Alexander 1916 A16
         |         T. tephrocephala Loew 1864 A16
         |         T. ternaria Loew 1864 A16
         |         T. terrestris Linnaeus 1758 L58
         |         T. tesselata Loew 1863 A16
         |         T. tremula Linnaeus 1758 L58
         |         T. trinidadensis [=Pachyrrhina trinidadensis] A16
         |         T. triton Alexander 1916 A16
         |         T. trivittata G09
         |         T. tuscarora Alexander 1916 [incl. T. hamata ms] A16
         |         T. umbrosa Loew 1863 A16
         |         T. unimaculata [=Pachyrrhina unimaculata] A16
         |         T. valida Loew 1863 A16
         |         T. variegata Linnaeus 1758 L58
         |         T. varipennis I92
         |         T. vernalis K01
         |         T. versicolor Loew 1863 A16
         |         T. vibratoria Linnaeus 1758 L58
         |--T. (Tipula) G09
         |    |--T. (T.) oleracea Linnaeus 1758 G09, L58
         |    `--T. (T.) paludosa AB81
         |--T. (Acutipula) A40
         |    |--T. (A.) graphiptera A40
         |    |--T. (A.) megaleuca A40
         |    |--T. (A.) phaeoleuca Alexander 1940 A40
         |    `--T. (A.) stenacantha Alexander 1937 A40
         |--T. (Angarotipula) illustris G09
         |--T. (Bellardina) cydippe G09
         |--T. (Cinctotipula Alexander 1916) A16
         |    `--T. (*C.) algonquin Alexander 1916 A16
         |--T. (Eumicrotipula) G09
         |    |--T. (E.) armillatus G09
         |    |--T. (E.) balloui G09
         |    |--T. (E.) exilis G09
         |    |--T. (E.) ligulipenicillata G09
         |    |--T. (E.) michoacana G09
         |    |--T. (E.) monilifera G09
         |    `--T. (E.) virgulata G09
         |--T. (Formotipula) A40
         |    |--T. (*F.) holoserica A40
         |    |--T. (F.) spoliatrix A40
         |    `--T. (F.) vindex Alexander 1940 A40
         |--T. (Lunatipula) G09
         |    |--T. (L.) dorsimacula G09
         |    |--T. (L.) monticola Alexander 1916 G09, A16
         |    `--T. (L.) triplex G09
         |--T. (Microtipula) [incl. Nephrotomodes] G09
         |    |--T. (M.) amazonica G09
         |    |--T. (M.) armatipennis G09
         |    |--T. (M.) aymara G09
         |    |--T. (M.) contemplata G09
         |    |--T. (M.) smilodon G09
         |    `--T. (M.) smithi G09
         |--T. (Nesotipula) pribilovia AB81
         |--T. (Nippotipula) AB81
         |    |--T. (N.) abdominalis AB81
         |    `--T. (N.) sinica Alexander 1935 A40
         |--T. (Oreomyza) A40
         |    |--T. (O.) famula Alexander 1935 A40
         |    `--T. (O.) tetramelania Alexander 1935 A40
         |--T. (Rhiphidia) maculata Linnaeus 1758 G20, L58
         |--T. (Schummelia) A40
         |    |--T. (S.) angustiligula A40
         |    |    |--T. a. angustiligula A40
         |    |    `--T. a. mokanensis Alexander 1940 A40
         |    |--T. (S.) spectata Alexander 1940 A40
         |    `--T. (S.) variicornis A40
         |--T. (Tipulodinodes) lacteipes AB81
         |--T. (Trichotipula Alexander 1916) A16
         |    `--T. (*T.) oropezoides A16
         |--T. (Triplicitipula) flavoumbrosa S96
         |--T. (Vestiplex) bifida Alexander 1921 A40
         `--T. (Yamatotipula) G09
              |--T. (Y.) caloptera Loew 1863 AB81, A16
              |--T. (Y.) eluta Loew 1863 G09, A16
              |--T. (Y.) nova Walker 1848 A40
              |--T. (Y.) noveboracensis G09
              |--T. (Y.) poliocephala Alexander 1921 A40
              |--T. (Y.) strepens Loew 1863 G09, A16
              `--T. (Y.) tricolor G09

Nephrotoma Meigen 1803 [incl. Pachyrrhina Macquart 1834, Pales Meigen 1800 (n. n.) non Robineau-Desvoidy 1830] A16

*Type species of generic name indicated

REFERENCES

[A16] Alexander, C. P. 1916. New or little-known crane-flies from the United States and Canada: Tipulidae, Diptera, Part 2. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (3): 458–514, pls 16–21.

[A40] Alexander, C. P. 1940. New or little-known Tipulidae from eastern China. Notes d'Entomologie Chinoise 8 (1): 1–28.

[AB81] Alexander, C. P., & G. W. Byers. 1981. Tipulidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 153–190. Research Branch, Agriculture Canada.

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[F92] Fan Z. 1992. Key to the Common Flies of China 2^nd ed. Science Press: Beijing.

[G09] Gelhaus, J. K. 2009. Tipulidae (crane flies, tipúlidos). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 193–236. NRC Research Press: Ottawa.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[HP68] Holmes, R. T., & F. A. Pitelka. 1968. Food overlap among coexisting sandpipers on northern Alaskan tundra. Systematic Zoology 17 (3): 305–318.

[I92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[S96] Smith, D. R. 1996. Trigonalyidae (Hymenoptera) in the eastern United States: seasonal flight activity, distribution, hosts. Proceedings of the Entomological Society of Washington 98 (1): 109–118.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240–257.

[V09] Verdcourt, B. (ed.) 2009. Additions to the Wild Fauna and Flora of the Royal Botanic Gardens, Kew. XXVI. Miscellaneous records. Kew Bulletin 64 (1): 183–194.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

[W01] Wohltmann, A. 2001. Closely related species of Parasitengonae (Acari: Prostigmata) inhabiting the same areas: features facilitating coexistence. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 121–135. CSIRO Publishing: Melbourne.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3^rd ed. Reed New Holland: Sydney.

Last updated: 27 December 2020.

Tipulidae

Orimarga mirabilis, copyright James V. Remsen.

Belongs within: Tipulomorpha.
Contains: Dicranota, Limnophilini, Cylindrotominae, Tipulinae, Limonia, Dicranomyia.

The Tipulidae, crane flies, are a cosmopolitan group of mostly slender, long-legged flies that differ from all other flies in the presence of a V-shaped suture dividing the mesoscutum, and usually two complete anal veins in the wings (if present). Some classifications divide the Tipulidae between three families Tipulidae, Cylindrotomidae and Limoniidae; not only is the broader sense of Tipulidae used here but it seems possible that the subfamily 'Limoniinae' is paraphyletic to the Cylindrotominae and Tipulinae (e.g. Wiegmann et al. 2011). The Pediciini, historically included in the Limoniinae, are characterised by hairy eyes with short hairs between the ommatidia (Alexander & Byers 1981).

Characters (from Alexander & Byers 1981): Typically slender-bodied. Ocelli absent. Mesonotum with transverse V-shaped suture. Wing elongate, rather narrow. Legs very long and slender, breaking readily at suture between trochanter and femur.

<==Tipulidae [Architipulidae, Eoasilidae, Limnobiinae, Limoniidae, Tipuloidea] G09
    |--Pediciini [Pediciidae] AB81
    |    |--Ula elegans WT11
    |    |--Ornithodes AB81
    |    |--Nasiternella hyperborea AB81
    |    |--Nipponomyia O98
    |    |--Dicranota G09
    |    `--Pedicia AB81
    |         |  i. s.: P. albovitta A16
    |         |         P. hannai HP68
    |         |--P. (Pedicia) bellamyana AB81
    |         `--P. (Tricyphona) AB81
    |              |--‘Tricyphona’ calcar A16
    |              |--‘Tricyphona’ inconstans A16
    |              `--P. (T.) protea AB81
    `--+--Limnophilini WT11
       `--+--Cylindrotominae WT11
          `--+--Tipulinae WT11
             `--Limoniinae [Lechriinae] O98
                  |--Lechria [Lechriini] CM91
                  `--Limoniini AB81
                       |--Limonia G09
                       |--Elliptera tennessa AB81
                       |--Thaumastoptera hynesi AB81
                       |--Atypophthalmus (Atypophthalmus) umbratus G09
                       |--Dicranomyia G09
                       |--Tonnoiromyia CM91
                       |--Dicranoptycha AB81
                       |    |--D. germana AB81
                       |    `--D. phallosomica Alexander 1937 A40
                       |--Antocha WT11
                       |    |--A. obtusa WT11
                       |    `--A. saxicola AB81
                       |--Helius (Helius) AB81
                       |    |--H. (H.) albitarsis G09
                       |    |--H. (H.) darlingtonae G09
                       |    `--H. (H.) flavipes G09
                       |--Geranomyia CM91
                       |    |--G. canadensis G09
                       |    |--G. recondita G09
                       |    `--G. subvirescens G09
                       |--Rhipidia (Rhipidia) G09
                       |    |--R. (R.) lecontei G09 [=Limonia (Rhipidia) lecontei AB81]
                       |    |--R. (R.) multiguttata G09
                       |    |--R. (R.) subcostalis G09
                       |    `--R. (R.) willistoniana G09
                       `--Orimarga AB81
                            |--O. (Orimarga) G09
                            |    |--O. (O.) attenuata G09
                            |    |--O. (O.) nigroapicalis G09
                            |    `--O. (O.) niveitarsis G09
                            `--O. (Diotrepha) G09
                                 |--O. (D.) flavescens G09
                                 `--O. (D.) mirabilis G09

Tipulidae incertae sedis:
  Aphrophila TE03
  Architipula WT11
  Styringomyia G09
    |--S. maya G09
    `--S. paulista G09
  Amphineurus M81
  Paralimnophila setulicornis CM91
  Limnobia A16
    |--L. cinctipes A16
    |--L. indigena A16
    |--L. rivosa (Linnaeus 1758) G20, L58 [=Tipula rivosa G20]
    `--L. solitaria A16
  Rhypholophus A16
    |--R. meigeni A16
    `--R. rubellus A16
  Adelphomyia minuta A16
  Oropeza venosa A16
  Xiphura frontalis A16
  Semnotes imperatoria CM70
  Dapanoptera O98
  Libnotes O98

*Type species of generic name indicated

REFERENCES

[A16] Alexander, C. P. 1916. New or little-known crane-flies from the United States and Canada: Tipulidae, Diptera, Part 2. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (3): 458–514, pls 16–21.

[A40] Alexander, C. P. 1940. New or little-known Tipulidae from eastern China. Notes d'Entomologie Chinoise 8 (1): 1–28.

[AB81] Alexander, C. P., & G. W. Byers. 1981. Tipulidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 153–190. Research Branch, Agriculture Canada.

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[G09] Gelhaus, J. K. 2009. Tipulidae (crane flies, tipúlidos). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 193–236. NRC Research Press: Ottawa.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[HP68] Holmes, R. T., & F. A. Pitelka. 1968. Food overlap among coexisting sandpipers on northern Alaskan tundra. Systematic Zoology 17 (3): 305–318.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M81] McAlpine, J. F. 1981. Morphology and terminology—adults. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 9–63. Research Branch, Agriculture Canada.

[O98] Oosterbroek, P. 1998. The Families of Diptera of the Malay Archipelago. Brill: Leiden.

[TE03] Taylor, R. C., & R. M. Ewers. 2003. The invertebrate fauna inhabiting tree holes in a red beech (Nothofagus fusca) tree. Weta 25: 24–28.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated; 27 December 2020.

Chironomus

Male buzzer midge Chironomus plumosus, copyright Didier Descouens.

Belongs within: Chironomidae.

Chironomus is a cosmopolitan genus of midges. It is distinguished from related genera by features of the male genitalia, with appendage 1 often curving or hooking or with a ventral apical knob and with a digitiform process arising distad from the microtrichiose base, and appendage 2 well developed with long setae distally. Abdominal tergites often bear dark spot and transverse bands, and the r-m area of the wing is often darkened. Larvae inhabit soft sediments and other underwater organic deposits. Depending on the species, they may require pristine to acidic or brackish water habitats (Spies et al. 2009).

Characters (from Oliver 1981): Male antenna with eleven flagellomeres. Antepronotal lobes not separated medially. Wing with crossvein r-m definitely oblique to R₄₊₅; crossvein m-cu absent. Apex of fore tibia with inner low rounded scale that projects slightly beyond similar outer scale; hind tibia with two combs composed of basally fused spines. Male terminalia with gonocoxite and gonostylus separate; appendage 1 with basal long setae, or rarely with long setae only on apical two-thirds, without enlarged setiferous base; appendage 2 subcylindrical, longer than appendage 1, with many apical setae.

<==Chironomus Meigen 1803 [incl. Tendipes Meigen 1800 (nom. rej.)] O81
    |--C. (Chironomus) SA09
    |    |--C. (C.) elegans G20
    |    `--C. (C.) staegeri SA09
    `--C. (Lobochironomus) montuosus SA09

Chironomus incertae sedis:
  C. acutiventris R86
    |--C. a. acutiventris R86
    `--C. a. bavaricus R86
  C. alternans CM91
  C. atroviridis O81
  C. australis CM91
  C. clavaticornis RD77
  C. frigidus T04
  C. intertinctus W70
  C. limnaei A71
  C. pedellus D56
  C. plumosus SA09 [=Tendipes plumosus CS77]
  C. prasinus BBM91
  ‘Tendipes’ reductus CS77
  C. riparius SA09
  C. silvestris [incl. C. silvestris var. ornatus] K01
  C. singulatus OV95
  C. stercorarius RD77
  C. tentans MG06
  C. tepperi WT11
  C. thummi B91
  C. venerabilis Etheridge & Olliff 1890 F71
  C. zealandicus B87

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[B91] Baker, R. A. 1991. Development and life-history strategies in mussel mites (Hydrachnellae: Unionicolidae). In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 65–73. Chapman & Hall: London.

[BBM91] Baker, R. A., S. Brett & N. Morley. 1991. Water mite larvae parasitic on insects—a study of larval structures and host relationships. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 493–496. SPB Academic Publishing: The Hague.

[B87] Boothroyd, I. K. G. 1987. Taxonomic composition and life cycles of Chironomidae (Diptera) in a northern New Zealand stream. Entomologica Scandinavica Supplement 29: 15–22.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[K01] Kertész, K. 1901. Legyek [Dipteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[MG06] Mallatt, J., & G. Giribet. 2006. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40: 772–794.

[O81] Oliver, D. R. 1981. Chironomidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 423–458. Research Branch, Agriculture Canada.

[OV95] Ovčarenko, N. A., & I. Vima. 1995. [Issia singulati sp. n. (Microsporidia, Telomyxidae): a new species of Microsporidia from Chironomus singulatus Meigen larvae from Kiev’s water reservoir]. Gidrobiologicheskii Zhurnal 31 (5): 78–83.

[R86] Reiss, F. 1986. Erster Nachtrag zur Chironomidenfauna Bayerns (Diptera, Chironomidae). Spixiana 9 (2): 175–178.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[SA09] Spies, M., T. Andersen, J. H. Epler & C. N. Watson Jr. 2009. Chironomidae (non-biting midges). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 1 pp. 437–480. NRC Research Press: Ottawa.

[T04] Trägårdh, I. 1904. Fauna Arctica: Eine Zusammenstellung der arktischen Tierformen, mit besonderer Berücksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre 1898 vol. 4 pt 1. Monographie der arktischen Acariden. Gustav Fischer: Jena.

[W70] White, M. J. D. 1970. Cytogenetics. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 72–82. Melbourne University Press.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated: 28 December 2021.