Ceratopogonidae

Ceratopogonidae is a family of flies commonly known as no-see-ums, sand flies or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the Arctic. A 2025 study from Oxford University lists the subspecies Ceratopogonidae midges as "the most widely recognised and best-studied cocoa pollinators."

Ceratopogonidae are holometabolous, meaning their development includes four life stages: egg, larva, pupa, and imago or adult. 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 due to the body producing histamines against the proteins from the midges' saliva. 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.

Some species within the biting midges are thought to be predatory on other small insects. In particular, 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. They can also be hematophagous parasites of invertebrates, depending on whether the bloodsucking attack is fatal.

Like other bloodsucking 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, and nonviral animal pathogens.

Historically, numbers were managed with the insecticide DDT 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, oil of ''Eucalyptus'', or Icaridin. Their larvae have also been shown to be susceptible to treatment with commercially available preparations of '' Bacillus thuringiensis israelensis''.

Subfamilies

The Leptoconopinae is a subfamily of biting midges. 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.

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.

Systematics

Basal lineages

* †'' 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

Gallery

File:Ceratopogonidae.jpg, Larval stage of a ceratopogonid species
File:Atrichopogon - 2013-07-01.webm, ''Atrichopogon'' sp. on '' Oedemera virescens''
File:Ceratopogonidae.male.jpg, Ceratopogonid male
File:Leptoconops spp. from CSIRO.jpg, '' Leptoconops''
File:Ceratopogonidae midge sucking Sphodromantis blood IMG 3045ss.JPG, Ceratopogonid feeds on a mantis (The midge is on the front right femorotibial joint of the mantis, the mantis is eating a bee)
File:A-F-2012-07-22.ogv, A '' Forcipomyia'' sp. sucks hemolymph from '' Nemophora metallica''
File:Palpomyiini penstemon.jpg, Palpomyiini caught by sticky hairs of penstemon

References

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Nematocera families
Taxa named by Edward Newman
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