Belongs within: Culicomorpha.
Contains: Forcipomyia, Culicoidini, Ceratopogonini, Sphaeromiini, Stilobezziini, Palpomyiini.
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). Forcipomyiinae have well-developed empodia, at least in females, and strongly curved claws (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 [Ceratopogonina, Heleidae, Leptoconopidae]
|--Dasyhelea [Dasyheleinae]BSG09
| |--D. antiquaS02
| |--D. cinctaBSG09
| `--D. pseudoincisurataBSG09
|--ForcipomyiinaeBSG09
| |--ForcipomyiaBSG09
| `--AtrichopogonBSG09
| |--A. (Atrichopogon) levisBSG09
| `--A. polydactylusDW81
|--Leptoconops Skuse 1889S89 [LeptoconopinaeBSG09]
| | i. s.: L. bequaertiBSG09
| | L. succineus Szadzieweski 1988H02
| |--L. (Leptoconops)DW81
| | |--*L. (L.) stygius Skuse 1889S89
| | `--L. (L.) torrensDW81
| |--L. (Brachyconops) californiensisDW81
| `--L. (Megaconops) floridensisDW81
`--CeratopogoninaeBSG09
|--CulicoidiniBSG09
|--CeratopogoniniBSG09
|--SphaeromiiniBSG09
|--StilobezziiniDW81
|--PalpomyiiniBSG09
|--StenoxeniniBSG09
| |--StenoxenusBSG09
| | |--S. coomaniBSG09
| | `--S. johnsoniDW81
| `--ParyphoconusDW81
| |--P. angustipennisBSG09
| `--P. sonorensisDW81
`--HeteromyiiniBSG09
|--Heteromyia Say 1825S89
| `--H. fasciataBSG09
|--Neurohelea nigraDW81
|--Neurobezzia Wirth & Ratanaworabhan 1972WR72
| `--*N. granulosa (Wirth 1952) [=Bezzia granulosa]WR72
|--Clinohelea bimaculataBSG09
`--PellucidomyiaBSG09
|--P. ugandaeBSG09
`--P. wirthiDW81
Ceratopogonidae incertae sedis:
AustroconopsGE05
|--A. fossilisGE05
`--A. macmillaniGE05
Fittkauhelea amazonicaBSG09
Cacaohelea youngiBSG09
Baeohelea nanaBSG09
Leptohelea micronyxBSG09
Nannohelea bourioniBSG09
Schizonyxhelea forattiniiBSG09
AlloheleaBSG09
|--A. johannseniBSG09
`--A. neotropicaBSG09
Parastilobezzia leeiBSG09
Downeshelea stoneiBSG09
Baeodasymyia dominicanaBSG09
SpinelliheleaBSG09
MacruroheleaCM70
AcanthoheleaCM70
XenoheleaCM70
DibezziaCM970
Archiaustroconops alavensis Szadziewski & Arillo 1998PR13
StyloconopsCM70
Eopalpomyitis Hong 2002H02
`--*E. unca (Hong 1981) [=Palpomyia unca]H02
Sinopogonites Hong 2002H02
`--*S. eocenius Hong 2002H02
HartomyiaH02
PseudohezziaH02
AtriculicoidesH02
ProtoculicoidesH02
CeratopalpomyiaP92
EoheleaP92
|--E. petrunkevitchi Szadziewski 1988P92
`--E. sinuosaP92
FossiheleaP92
GedanoheleaP92
MantoheleaP92
MeunieroheleaP92
PhysoheleaP92
WirthoheleaP92
Oecacta Poey 1853S89
Pachyleptus Walker 1856S89
Psychophaena Philippi 1865S89
Didymophleps Weyenberg 1883S89
Psamathiomya Deby 1889S89
*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.
[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.
[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.
[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.
[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.
[S89] Skuse, F. A. A. 1889. Diptera of Australia. Part VI.—The Chironomidae. Proceedings of the Linnean Society of New South Wales, series 2, 4 (2): 215–311, pls 11–14.
[WR72] Wirth, W. W., & N. C. Ratanaworabhan. 1972. A new genus of biting midge from California related to Neurohelea Kieffer (Diptera, Ceratopogonidae). Pan-Pacific Entomologist 48 (4): 244–245.