Ceratopogoninae
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Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed worldwide, apart from the Antarctic and the
Arctic The Arctic ( or ) is a polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenland), Finland, Iceland, N ...
. Ceratopogonidae are
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
, meaning their development includes four life stages:
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
,
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
,
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...
, and
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the imaginal stage, the stage in which the insect attains maturity. It follows the f ...
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
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
s, including humans, to get protein for egg-laying. Their bites are painful, and can cause intensely itchy
lesions A lesion is any damage or abnormal change in the tissue of an organism, usually caused by disease or trauma. ''Lesion'' is derived from the Latin "injury". Lesions may occur in plants as well as animals. Types There is no designated classifi ...
. 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. Particularly
mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e 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 Hematophagy (sometimes spelled haematophagy or hematophagia) is the practice by certain animals of feeding on blood (from the Greek words αἷμα ' "blood" and φαγεῖν ' "to eat"). Since blood is a fluid tissue rich in nutritious p ...
parasites of invertebrates, depending on whether the bloodsucking attack is fatal. Like other blood sucking flies, ''
Culicoides ''Culicoides'' is a genus of biting midges in the family Ceratopogonidae. There are over 1000 species in the genus,Connelly, C. RBiting midges: ''Culicoides'' spp.Featured Creatures, Entomology and Nematology Department, University of Florida IF ...
'' species can be vectors of disease-causing
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
s. Among diseases transmitted are the parasitic nematodes ''
Mansonella ''Mansonella'' is a genus of parasitic nematodes. It includes three species that are responsible for the disease mansonelliasis: ''Mansonella ozzardi'', ''M. perstans'', and ''M. streptocerca''. Species There are nine species: * ''Mansonella ba ...
'', bluetongue disease,
African horse sickness African horse sickness (AHS) is a highly infectious and deadly disease caused by ''African horse sickness virus''. It commonly affects horses, mules, and donkeys. It is caused by a virus of the genus ''Orbivirus'' belonging to the family ''Reov ...
,
epizootic hemorrhagic disease Epizootic hemorrhagic disease (EHD) is a hemorrhagic disease of white-tailed deer (''Odocoileus virginianus'') caused by an infection of a virus from the genus ''Orbivirus'' subsequently called ''Epizootic hemorrhagic disease virus'' (EHDV). It is ...
,
arbovirus Arbovirus is an informal name for any virus that is transmitted by arthropod vectors. The term ''arbovirus'' is a portmanteau word (''ar''thropod-''bo''rne ''virus''). ''Tibovirus'' (''ti''ck-''bo''rne ''virus'') is sometimes used to more sp ...
es, and nonviral animal pathogens. Historically, numbers were managed with the insecticide
DDT Dichlorodiphenyltrichloroethane, commonly known as DDT, is a colorless, tasteless, and almost odorless crystalline chemical compound, an organochloride. Originally developed as an insecticide, it became infamous for its environmental impacts. ...
as with '' Leptoconops torrens'' populations in California. They can be trapped by luring them with
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
. Most midges are small enough to pass through ordinary insect window screening. They can be repelled with DEET, oil of ''
Eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as e ...
'', or
Icaridin Icaridin, also known as picaridin, is an insect repellent which can be used directly on skin or clothing. It has broad efficacy against various arthropods such as mosquitos, ticks, gnats, flies and fleas, and is almost colorless and odorless. A s ...
. 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
sclerite A sclerite (Greek , ', meaning " hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly ...
s 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 A pollinator is an animal that moves pollen from the male anther of a flower to the female stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are the maj ...
of tropical crops such as the
cocoa bean The cocoa bean (technically cocoa seed) or simply cocoa (), also called the cacao bean (technically cacao seed) or cacao (), is the dried and fully fermented seed of ''Theobroma cacao'', from which cocoa solids (a mixture of nonfat substance ...
. 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 The Purbeck Group is an Upper Jurassic to Lower Cretaceous lithostratigraphic group (a sequence of rock strata) in south-east England. The name is derived from the district known as the Isle of Purbeck in Dorset where the strata are exposed in ...
of
Dorset Dorset ( ; archaically: Dorsetshire , ) is a county in South West England on the English Channel coast. The ceremonial county comprises the unitary authority areas of Bournemouth, Christchurch and Poole and Dorset. Covering an area of , ...
, England dating to the
Berriasian In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago ...
, around 142 million years ago.


Systematics

Basal lineages * †''Lebanoculicoides'' Szadziewski 1996
Lebanese amber Lebanese amber is fossilized resin found in Lebanon and southwest Syria. It dates back approximately 130-125 million years to the Barremian of the Early Cretaceous. It formed on what was then the northern coast of Gondwana, believed to be a tropi ...
,
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is precede ...
, Spanish amber,
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
* Subfamily Leptoconopinae Noe 1907 ** †''Archiaustroconops'' Szadziewski 1996 Durlston Formation, United Kingdom, Berriasian, Lebanese amber, Barremian, Jordanian amber, Spanish amber, Albian,
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
, Myanmar, Cenomanian ** '' Austroconops'' Wirth and Lee 1958 Barremian-Present ** †''Fossileptoconops'' Szadziewski 1996 Lebanese amber, Barremian ** †''Jordanoconops'' Szadziewski 2000 Jordanian amber, Albian ** ''
Leptoconops ''Leptoconops'' (black gnat) is a midge genus in the family Ceratopogonidae. It has a mostly tropical or subtropical distribution worldwide, but some species occur as far north as Moscow region in Russia and the Yukon Territory in Canada. This ...
'' Skuse 1889 Barremian-Present ** †''Minyohelea'' Borkent 1995 Austrian amber,
Hauterivian The Hauterivian is, in the geologic timescale, an age in the Early Cretaceous Epoch or a stage in the Lower Cretaceous Series. It spans the time between 132.9 ± 2 Ma and 129.4 ± 1.5 Ma (million years ago). The Hauterivian is preceded by the ...
Lebanese amber, Barremian, Canadian amber,
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
* †''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 The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. ...
*''
Dasyhelea ''Dasyhelea'' is a genus of biting midges and the only genus of its subfamily, the Dasyheleinae. Larvae of species in this genus are characterized by an anal segment with retractile posterior prolegs. Larvae are aquatic animal, aquatic and adults ...
'' Kieffer 1913 *Subfamily Forcipomyiinae **'' Atrichopogon'' Kieffer 1906 **''
Forcipomyia ''Forcipomyia'' is a genus of biting midges in the subfamily Forcipomyiinae Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed w ...
'' Meigen, 1818 *†''Adelohelea'' Borkent 1995 Hungarian amber, Santonian, Canadian amber, Campanian *†''Heleageron'' Borkent 1995
New Jersey amber New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of t ...
,
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...
Canadian amber, Campanian *†''Alautunmyia'' Borkent 1996
New Jersey amber New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of t ...
,
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...


Gallery

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 ''Forcipomyia'' is a genus of biting midges in the subfamily Forcipomyiinae Ceratopogonidae is a family of flies commonly known as no-see-ums, or biting midges, generally in length. The family includes more than 5,000 species, distributed w ...
'' sp. sucks
hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...
from '' Nemophora metallica'' File:Palpomyiini penstemon.jpg, Palpomyiini caught by sticky hairs of penstemon


References

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