Culicomorpha

{{Automatic taxobox

| image = AnophelesGambiaemosquito.jpg

| taxon = Culicomorpha

| authority = Hennig, 1948

| subdivision_ranks = Superfamilies

| subdivision =

Culicoidea

Chironomoidea

See text for families.

}}

The Culicomorpha are an infraorder of Nematocera, including mosquitoes, black flies, and several extant and extinct families of insects.{{cite journal |last=Borkent |first=Art |title=The pupae of Culicomorpha : morphology and a new phylogenetic tree |journal=Zootaxa |volume=3396 |issue=1 |year=2012 |url=https://mosquito-taxonomic-inventory.myspecies.info/sites/mosquito-taxonomic-inventory.info/files/Borkent%202012.pdf |doi=10.11646/zootaxa.3396.1 |isbn=978-1-86977-957-3 |oclc=807999306}} 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.

Most adult, females lay their eggs on bodies of water. Some are restricted to very clean waters, but others can tolerate highly polluted environments.{{Cite book |url=https://www.worldcat.org/oclc/899273856 |title=Ecology and General Biology |date=2015 |publisher=Academic Press |others=James H. Thorp, D. Christopher Rogers |isbn=978-0-12-385027-0 |edition=4th |location=London |oclc=899273856}}

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.{{Cite journal |last1=Carpenter |first1=Simon |last2=Groschup |first2=Martin H. |last3=Garros |first3=Claire |last4=Felippe-Bauer |first4=Maria Luiza |last5=Purse |first5=Bethan V. |date=October 2013 |title=Culicoides biting midges, arboviruses and public health in Europe |journal=Antiviral Research |volume=100 |issue=1 |pages=102–113 |doi=10.1016/j.antiviral.2013.07.020 |issn=1872-9096 |pmid=23933421|doi-access=free }}{{Cite journal |last1=Carpenter |first1=S. |last2=Veronesi |first2=E. |last3=Mullens |first3=B. |last4=Venter |first4=G. |date=April 2015 |title=Vector competence of Culicoides for arboviruses: three major periods of research, their influence on current studies and future directions |url=https://pubmed.ncbi.nlm.nih.gov/26470451 |journal=Revue Scientifique et Technique (International Office of Epizootics) |volume=34 |issue=1 |pages=97–112 |doi=10.20506/rst.34.1.2347 |issn=0253-1933 |pmid=26470451|doi-access=free }}

Classification

The Culicomorpha comprises eight extant families divided into two superfamilies. It was described based on phylogenetic analyses of morphological features.{{cite book |first1=D. M. |last1=Wood |first2=A. |last2=Borkent |chapter=Phylogeny and Classilication of the Nematocera |editor-first1=J.F. |editor-last1=McAlpine |editor-first2=D.M. |editor-last2=Woods |year=1989 |title=Manual of Nearctic Diptera, vol. 3. |publisher=Ottawa: Agriculture Canada Research Branch |url=https://publications.gc.ca/collections/collection_2016/aac-aafc/A54-3-32-eng.pdf}}{{cite journal |last1=Yeates |first1= David K. |last2=Weigmann |first2=Brian M |last3=Courtney |first3=Greg W. |last4=Meier |first4=Rudolf |last5=Lambkins |first5=Christine |last6=Pape |first6=Thomas |title=Phylogeny and systematics of Diptera: Two decades of progress and prospects |year=2007 |journal=Zootaxa |volume=1668 |pages=565–590 |doi=10.11646/zootaxa.1668.1.27 }}{{cite journal |last1=Bertone |first1=Matthew A. |last2=Courtney |first2=Gregory W. |last3=Wiegmann |first3=Brian M. |title=Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes |year=2008 |journal=Systematic Entomology |volume=33 |issue=4 |pages=668–687 |doi=10.1111/j.1365-3113.2008.00437.x |s2cid=54781592 |url=http://online-keys.net/sciaroidea/add01/Bertone_et_al_Nematocera_2008.pdf }}

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.{{cite journal |last1=Narayanan Kutty |first1=Sujatha |last2=Wong |first2=Wing Hing |last3=Meusemann |first3=Karen |last4=Meier |first4=Rudolf |last5=Cranston |first5=Peter S. |title=A phylogenomic analysis of Culicomorpha (Diptera) resolves the relationships among the eight constituent families |year=2018 |journal=Systematic Entomology |volume=43 |issue=3 |pages=434–446 |doi=10.1111/syen.12285 |s2cid=90255287 |doi-access=free }} 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. 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. The phylogenetic trees below illustrate the different arrangements.

{{clade gallery

|caption1=Traditional morphological

|caption2=Revised morphological

|caption3=Phylogenomic

|cladogram1=

{{clade

|label1=Culicomorpha

|1={{clade

|label1=Chironomoidea

|1={{clade

|1=Thaumaleidae

|2={{clade