Culicomorpha

The Culicomorpha are an infraorder of Nematocera, including mosquitoes, black flies, and several extant and extinct families of insects.^[1] They originated 176 million years ago, in the Triassic period. There are phylogenetic patterns that are used to interpret bionomic features such as differences in the nature of blood-feeding by adult females, daytime or nighttime feeding by adult females, and occurrence of immature stages in aquatic habitats.^[1]

Culicomorpha
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Diptera
Suborder:	Nematocera
Infraorder:	Culicomorpha Hennig, 1948
Superfamilies
Culicoidea Chironomoidea See text for families.

Most adult, females lay their eggs on bodies of water. Some are restricted to very clean waters, but others can tolerate highly polluted environments.^[2]

Many adults transmit parasites or diseases that can be debilitating or fatal to humans, such as malaria and West Nile virus. Biting midges can transmit an extensive range of pathogens of veterinary importance, including Akabane virus, bovine ephemeral fever virus, Schmallenberg virus, African horse sickness virus, epizootic haemorrhagic disease virus, and bluetongue virus.^[3][4]

Classification

The Culicomorpha comprises eight extant families divided into two superfamilies. It was described based on phylogenetic analyses of morphological features.^[5][6][7]

• Superfamily Culicoidea
  • Dixidae - meniscus midges
  • Corethrellidae - frog-biting midges
  • Chaoboridae – phantom midges
  • Culicidae – mosquitoes
• Superfamily Chironomoidea
  • Thaumaleidae – solitary midges
  • Simuliidae – black flies and buffalo gnats
  • Ceratopogonidae – biting midges
  • Chironomidae – nonbiting midges

The monophyly of Culicomorpha and Culicoidea have been confirmed in subsequent morphological and molecular studies, but several studies have found Chironomoidea to be paraphyletic.^[6][7][8] A morphological study in 2012, using characters from all life stages (egg, larva, pupae and adult), found that Chironomidae branched first and a sister relationship between Culicoidea and a clade of Simuliidae, Ceratopogonidae and Thamaleidae. A new superfamily, Simulioidea, was proposed for the latter clade and restricted Chironomoidea to Chironomidae.^[1] A phylogenomic analysis in 2018 also found a paraphyletic Chironomoidea and a close relationship between Simuliidae and Thamaleidae, but in this study Ceratopoginidae grouped with Chironomidae.^[8] The phylogenetic trees below illustrate the different arrangements.

Traditional morphological^[5]

Culicomorpha Chironomoidea Thaumaleidae Simuliidae Ceratopogonidae Chironomidae Culicoidea Dixidae Corethrellidae Chaoboridae Culicidae

Revised morphological^[1]

Culicomorpha Chironomoidea Chironomidae Simulioidea Ceratopogonidae Thaumaleidae Simuliidae Culicoidea Dixidae Corethrellidae Chaoboridae Culicidae

Phylogenomic^[8]

Culicomorpha Chironomoidea Chironomidae Ceratopogonidae Simulioidea Thaumaleidae Simuliidae Culicoidea Dixidae Corethrellidae Chaoboridae Culicidae

Extinct families

• Asiochaoboridae (Upper Jurassic)
• Architendipedidae (Upper Triassic)
• Protendipedidae (Middle Jurassic)
• Mesophantasmatidae (Middle Jurassic)