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.

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 [Dasyheleinae] DW81
    |    |--D. antiqua S02
    |    |--D. cincta BSG09
    |    `--D. pseudoincisurata DW81
    |--Leptoconops [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 DW81
         |    |    |--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. Ottawa, NRC Research Press.

[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.

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