Ceratopogonidae

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Antliophora
Ordo: Diptera
Subordo: Nematocera
Infraordo: Culicomorpha
Superfamilia: Chironomoidea

Familia: Ceratopogonidae
Subfamiliae (5 + 1†): Austroconopinae - Ceratopogoninae - Dasyheleinae - Forcipomyiinae - Leptoconopinae - †Lebanoculicoidinae
Overview of genera (107 + 6†)

• Afrohelea – Afrostilobezzia – Agilihelea – Allohelea – Alluaudomyia – Amerohelea – Ankylohelea – Archiaustroconops – Atrichopogon – Atyphohelea – Austroconops – Austrohelea – Austrosphaeromias – Baeodasymyia – Baeohelea – Bahiahelea – Bezzia – Boreohelea – Borkenthelea – Bothahelea – Bothamia – Brachypogon – Cacaohelea – Calyptopogon – Camptopterohelea – Capehelea – Cecidomyia – Ceratoculicoides – Ceratopogon – Chelohelea – Clastrieromyia – Clinohelea – Crispomyia – Culicoides – Dasyhelea – Diaphanobezzia – Dibezzia – Downeshelea – Echinohelea – Fanthamia – Fittkauhelea – Forcipomyia – Groganhelea – Hebetula – Heteroceratopogon – Heterohelea – Heteromyia – Homohelea – Indobezzia – Isthmohelea – Jenkinshelea – Johannsenomyia – Kolenohelea – Lanatomyia – Lanehelea – Lasiohelea – Leehelea – Leptoconops – Leptohelea – Luciamyia – Mackerrasomyia – Macropeza – Macrurohelea – Mallochohelea – Metacanthohelea – Metahelea – Meunierohelea – Minyohelea – Monohelea – Nannohelea – Nemoromyia – Neobezzia – Neoculicoides – Neohelea – Neosphaeromias – Neurobezzia – Neurohelea – Nilobezzia – Niphanohelea – Notiohelea – Notoceratopogon – Pachyhelea – Palaeobrachypogon – Palpomyia – Parabezzia – Paradasyhelea – Parastilobezzia – Paryphoconus – Pellucidomyia – Phaenobezzia – Probezzia – Protoculicoides – Pseudostilobezzia – Rhynchohelea – Schizohelea – Schizonyxhelea – Serromyia – Sinhalohelea – Sphaerohelea – Sphaeromias – Spinellihelea – Stenoxenus – Stilobezzia – Stiloculicoides – Tetrabezzia – Washingtonhelea – Xenohelea – †Archiaustroconops – †Fossileptoconops – †Jordanconops – †Lebanoculicoides – †Mantohelea – †Minyohelea

[source: Catalogue of Life: 2013 Annual Checklist, except fossils, minus (1) Tipula
]

Check: †Archiculicoides
Name

Ceratopogonidae Newman, 1834: 388
References

Newman, E. 1834. Attempted division of British insects into natural orders. The Entomological Magazine 2: 379–431. BHL.
Ayala, M.M., Spinelli, G.R., Funes, A. & Ronderos, M.M. 2015: Additional notes on biting midges from the subtropical forest of northeastern Argentina (Diptera: Ceratopogonidae). Zootaxa 3941(1): 79–90. DOI: 10.11646/zootaxa.3941.1.3. Reference page.
Borkent, A. 2011: World species of biting midges (Diptera: Ceratopogonidae). [electronic publication, last updated: February 10, 2011] Download PDF
Borkent, A. 2014. The pupae of the biting midges of the World (Diptera: Ceratopogonidae), with a generic key and analysis of the phylogenetic relationships between genera. Zootaxa 3879(1): 1–327. DOI: 10.11646/zootaxa.3879.1.1 Reference page.
Borkent, A. 2015: New combinations and changes in the classification of Ceratopogonidae (Diptera, biting midges). Zootaxa 3972(4): 599–600. DOI: 10.11646/zootaxa.3972.4.12. Preview (PDF) Reference page.
Borkent, A.; Grogan, W.L., jr. 2009: Catalog of the New World biting midges north of Mexico (Diptera: Ceratopogonidae). Zootaxa, 2273: 1–48. Abstract & excerpt
Dominiak, P. & Alwin, A. & Giłka, W. 2014. New records of predaceous midges from the Middle East, with the description of two new species (Diptera: Ceratopogonidae). Zootaxa 3753(2): 133–145. DOI: 10.11646/zootaxa.3753.2.3 Reference page.
Dominiak, P. & Mathieu, B. 2015. Serromyia diabolica, a new biting midge species from Lebanon (Diptera: Ceratopogonidae). Zootaxa 3946(3): 436–444. DOI: 10.11646/zootaxa.3946.3.10. Preview (PDF). Reference page.
Li, C-Q., Bellis, G., Wu, X-X. & Li, J-H. 2019. A new species of Nilobezzia Kieffer (Diptera, Ceratopogonidae) from the mangrove forest of Hainan Island, China. ZooKeys, 893: 135–141.DOI: 10.3897/zookeys.893.39032 Open access Reference page.
Szadziewski, R. & Dominiak, P. 2015. Afrostilobezzia, a new genus of predatory biting midges from the Afrotropical Region (Diptera: Ceratopogonidae). Zootaxa 3941(3): 445–450. DOI: 10.11646/zootaxa.3941.3.11 Preview (PDF). Reference page.
Szadziewski, R., Dominak, P. & Filatov, S. 2015. Two new species of predatory biting midges of the genus Alluaudomyia from Europe and the Canary Islands (Diptera: Ceratopogonidae). Zootaxa 4039(2): 345–358. DOI: 10.11646/zootaxa.4039.2.8 Preview (PDF). Reference page.
Torreias, S.R.S., Ferreira-Keppler, R.L. & Ronderos, M.M. 2013 (Online) 2014 (Print). Biting midges (Ceratopogonidae: Diptera) present in aquatic macrophytes from wetlands of Marchantaria Island, Iranduba, Central Amazonia, Brazil. Journal of Natural History 48(1–2): 109–122. DOI: 10.1080/00222933.2013.791934 Reference page.
Yu, Y.-X., Yan, G., Liu, G.-P. & Liu, Z.-J. 2013. Three new record species and a new species of biting midge from China (Diptera, Ceratopogonidae). Acta zootaxonomica Sinica 38(2): 372–376. Abstract. Reference page.

Additional references

Spinelli, G. & Wolff, M.I.. 2016. FAMILY CERATOPOGONIDAE. In Wolff, M.I., Nihei, S.S. & Carvalho, C.J.B. de (eds.), Catalogue of Diptera of Colombia. Zootaxa 4122(1): 98–141. DOI: 10.11646/zootaxa.4122.1.15. Reference page.

Vernacular names
suomi: Polttiaiset
日本語: ヌカカ科
한국어: 등애모깃과, 좀모깃과

Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally 1–3 millimetres (5⁄128–15⁄128 in) in length. The family includes more than 5,000 species,[1] distributed worldwide, apart from the Antarctic and the Arctic.

Ceratopogonidae are holometabolous, meaning their development includes four life stages: egg, larva, pupa, and imago or adult.[2] Most common species in warmer climates will take about two to six weeks to complete a life cycle. Both adult males and females feed on nectar. Most females also feed on the blood of vertebrates, including humans, to get protein for egg-laying. Their bites are painful, and can cause intensely itchy lesions.[3] Their mouthparts are well-developed for cutting the skin of their hosts. Some species prey on other insects.

Larvae need moisture to develop, but also air and food. They are not strictly aquatic or terrestrial.[3]

Some species within the biting midges are thought to be predatory on other small insects. Particularly mosquito larvae have been investigated as common prey for biting midges in the genus Bezzia. For example, experiments have been conducted on the species Bezzia nobilis that suggest their reliance on mosquito larvae as one source of prey.[4][5] They can also be hematophagous parasites of invertebrates, depending on whether the bloodsucking attack is fatal.[6]

Like other blood sucking flies, Culicoides species can be vectors of disease-causing pathogens. Among diseases transmitted are the parasitic nematodes Mansonella, bluetongue disease, African horse sickness, epizootic hemorrhagic disease, arboviruses,[7] and nonviral animal pathogens.[8]

Historically, numbers were managed with the insecticide DDT[3] as with Leptoconops torrens populations in California. They can be trapped by luring them with carbon dioxide. Most midges are small enough to pass through ordinary insect window screening. They can be repelled with DEET,[3] oil of Eucalyptus, or Icaridin. Their larvae have also been shown to be susceptible to treatment with commercially available preparations of Bacillus thuringiensis israelensis.[9]

Subfamilies

The Leptoconopinae is a subfamily of biting midges.[10] The larvae are recognized by their unique sclerites of the head, and by their mouthparts.

The Forcipomyiinae are a subfamily of biting midges. In this subfamily, both anterior and posterior prolegs are present on the larvae. Larvae are both terrestrial and aquatic, and feed primarily on algae and fungi. Some species are important pollinators of tropical crops such as the cocoa bean.

Larvae of species in the Dasyheleinae subfamily are characterized by an anal segment with retractile posterior prolegs. Larvae are aquatic and adults do not feed on vertebrate blood, nor do they prey on other insects. They take nectar only, an unusual feeding behavior within the Ceratopogonidae.

The Ceratopogoninae subfamily has elongated larvae without prolegs or hooks. Most larvae of this subfamily are predatory. Adults generally take vertebrate blood or attack other insects. Most females in the subfamily Ceratopogoninae feed on insects similar to them in size.[6]

The oldest known member of the family is Archiaustroconops besti from the Purbeck Group of Dorset, England dating to the Berriasian, around 142 million years ago.[11]
Systematics

Basal lineages[12]

†Lebanoculicoides Szadziewski 1996 Lebanese amber, Barremian, Spanish amber, Albian
Subfamily Leptoconopinae Noe 1907
†Archiaustroconops Szadziewski 1996 Durlston Formation, United Kingdom, Berriasian, Lebanese amber, Barremian, Jordanian amber, Spanish amber, Albian, Burmese amber, Myanmar, Cenomanian
Austroconops Wirth and Lee 1958 Barremian-Present
†Fossileptoconops Szadziewski 1996 Lebanese amber, Barremian
†Jordanoconops Szadziewski 2000 Jordanian amber, Albian
Leptoconops Skuse 1889 Barremian-Present
†Minyohelea Borkent 1995 Austrian amber, Hauterivian Lebanese amber, Barremian, Canadian amber, Campanian
†Archiculicoides Szadziewski 1996 Lebanese amber, Barremian
†Gerontodacus Borkent 2019 Lebanese amber, Barremian, Spanish amber, Albian
†Protoculicoides Boesel 1937 Burmese amber, Myanmar, Cenomanian, Canadian amber, Campanian
†Atriculicoides Remm 1976 Spanish amber, Albian Durtal amber, France, Cenomanian, Burmese amber, Myanmar, Cenomanian, Taimyr amber, Russia, Cenomanian, Santonian
Dasyhelea Kieffer 1913
Subfamily Forcipomyiinae
Atrichopogon Kieffer 1906
Forcipomyia Meigen, 1818
†Adelohelea Borkent 1995 Hungarian amber, Santonian, Canadian amber, Campanian
†Heleageron Borkent 1995 New Jersey amber, Turonian Canadian amber, Campanian
†Alautunmyia Borkent 1996 New Jersey amber, Turonian

References

Boorman, John (1993). "Biting midges (Ceratopogonidae)". Medical Insects and Arachnids. Springer. pp. 288–309. doi:10.1007/978-94-011-1554-4_7. ISBN 978-94-010-4679-4.
Beckenbach, Andrew T.; Borkent, Art (2003-04-01). "Molecular analysis of the biting midges (Diptera: Ceratopogonidae), based on mitochondrial cytochrome oxidase subunit 2". Molecular Phylogenetics and Evolution. 27 (1): 21–35. doi:10.1016/S1055-7903(02)00395-0. ISSN 1055-7903. PMID 12679068.
"common name: biting midges, no-see-ums, scientific name: Culicoides spp. (Insecta: Diptera: Ceratopogonidae)". Featured Creatures. University of Florida. Retrieved 20 September 2018.
Hribar, L. J. & G. R. Mullen.Predation by Bezzia larvae (Diptera: Ceratopogonidae) on mosquito larvae (Diptera: Culicidae). Entomol. News 102: 183-186.
Mogi, M. Insects and other invertebrate predators. Journal of the American Mosquito Control Association, 23(sp2):93-109 (2007). doi:10.2987/8756-971X(2007)23[93:IAOIP]2.0.CO;2.
Marshall, Stephen (2012). Flies: The Natural History and Diversity of Diptera. Richmond Hill, Ontario: Firefly Books Ltd. p. 85. ISBN 9781770851009.
Carpenter, Simon; Groschup, Martin H.; Garros, Claire; Felippe-Bauer, Maria Luiza; Purse, Bethan V. (2013). "Culicoides biting midges, arboviruses and public health in Europe". Antiviral Research. 100 (1): 102–113. doi:10.1016/j.antiviral.2013.07.020. ISSN 0166-3542. PMID 23933421.
Linley, J. R. (1985). "Biting midges (Diptera: Ceratopogonidae) as vectors of nonviral animal pathogens". Journal of Medical Entomology. 22 (6): 589–599. doi:10.1093/jmedent/22.6.589. ISSN 0022-2585. PMID 3908679.
"World Health Organization" (PDF). WHO.int. World Health Organization. Retrieved 1 April 2020.
Borkent, Art; Craig, Douglas A (23 August 2004). "Austroconops Wirth and Lee, a Lower Cretaceous genus of biting midges yet living in Western Australia: a new species, first description of the immatures and discussion of their biology and phylogeny (Diptera: Ceratopogonidae)". American Museum Novitates (3449): 1–2. doi:10.1206/0003-0082(2004)449<0001:AWALAL>2.0.CO;2. hdl:2246/2814.
Borkent, Art; Coram, Robert A.; Jarzembowski, Edmund A. (2013-12-01). "The oldest fossil biting midge (Diptera: Ceratopogonidae) from the Purbeck Limestone Group (Lower Cretaceous) of southern Great Britain". Polish Journal of Entomology / Polskie Pismo Entomologiczne. 82 (4): 273–279. doi:10.2478/v10200-012-0041-8. ISSN 0032-3780.
Borkent, Art (2019-02-25). "The Phylogenetic Relationships of Cretaceous Biting Midges, with a Key to All Known Genera (Diptera: Ceratopogonidae)". American Museum Novitates (3921): 1–48. doi:10.1206/3921.1. ISSN 0003-0082. S2CID 91737913.