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Chironomidae

The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many species superficially resemble mosquitoes, but they lack the wing scales and elongated mouthparts of the Culicidae.

Chironomidae
Temporal range: Triassic–Present
Male Chironomus plumosus
Scientific classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Diptera
Suborder: Nematocera
Infraorder: Culicomorpha
Superfamily: Chironomoidea
Family: Chironomidae
Newman, 1834[1]
Subfamilies

See text

Two lake flies observed in Neenah, Wisconsin, after the yearly hatch in Lake Winnebago

The name Chironomidae stems from the Ancient Greek word kheironómos, "a pantomimist".

Common names and biodiversity edit

This is a large taxon of insects. Some estimates of the species numbers suggest well over 10,000 world-wide.[2] Males are easily recognized by their plumose antennae. Adults are known by a variety of vague and inconsistent common names, largely by confusion with other insects. For example, chironomids are known as "lake flies" in parts of Canada and Lake Winnebago, Wisconsin, but "bay flies" in the areas near the bay of Green Bay, Wisconsin. They are called "sand flies," "muckleheads,"[3] "muffleheads,"[4] "Canadian soldiers,"[5] or "American soldiers"[6] in various regions of the Great Lakes area. They have been called "blind mosquitoes" or "chizzywinks" in Florida.[7] In Kansas, they are known as "midges."[8] However, they are not mosquitoes of any sort, and the term "sandflies" generally refers to various species of biting flies unrelated to the Chironomidae.

The group includes the wingless Belgica antarctica, the largest terrestrial animal of Antarctica.[9][10]

Their larvae produce silk, and Chironomus has been studied as an alternative source of silk other than the silk moth, as it is possible to extract it without killing the animal (Ahimsa silk).[11]

The biodiversity of the Chironomidae often goes unnoticed because they are notoriously difficult to identify and ecologists usually record them by species groups. Each morphologically distinct group comprises a number of morphologically identical (sibling) species that can only be identified by rearing adult males or by cytogenetic analysis of the polytene chromosomes. Polytene chromosomes were originally observed in the larval salivary glands of Chironomus midges by Balbiani in 1881. They form through repeated rounds of DNA replication without cell division, resulting in characteristic light and dark banding patterns which can be used to identify inversions and deletions which allow species identification.

Behavior and description edit

Chironomids as a group are very diverse in their life histories, and exhibit a variety of behaviors during their development. Due to their species ambiguity and this diversity of behaviors, there is much controversy in research on their feeding habits, particularly as adults.

Many reference sources in the past century or so have repeated the assertion that the chironomidae do not feed as adults, but an increasing body of evidence contradicts this view. Adults of many species do, in fact, feed. The natural foods reported include fresh fly droppings, nectar, pollen, honeydew, and various sugar-rich materials.

The question whether feeding is of practical importance has by now been clearly settled for some Chironomus species, at least; specimens that had fed on sucrose flew far longer than starved specimens, and starved females longer than starved males, which suggested they had eclosed with larger reserves of energy than the males. Some authors suggest the females and males of certain species apply the resources obtained in feeding differently. Generally, Males expend the extra energy on flight, while females use their food resources to achieve longer lifespans. The respective strategies should be compatible with maximal probability of successful mating and reproduction in those species that do not mate immediately after eclosion, particularly in species that have more than one egg mass maturing, the less developed masses being oviposited after a delay. Such variables also would be relevant to species that exploit wind for dispersal, laying eggs at intervals. Chironomids that feed on nectar or pollen may well be of importance as pollinators, but current evidence on such points is largely anecdotal. However, the content of protein and other nutrients in pollen, in comparison to nectar, might well contribute to the females' reproductive capacities.[2]

Larvae of some species are bright red in color due to a hemoglobin analog; these are often known as "bloodworms".[12] Their ability to capture oxygen is further increased by making undulating movements.[13]

Adults can be pests when they emerge in large numbers. They may cause difficulty during driving if they collide with the windshield, creating an opaque coating which obscures the driver's vision.[14] They can damage paint, brick, and other surfaces with their droppings. When large numbers of adults die, they can build up into malodorous piles. They can provoke allergic reactions in sensitive individuals.[15] These allergic reactions have been shown to be caused by the haemolglobins that are primarily found in Chironomidae larval stages.[2]

Ecology and distribution edit

Chironomids are highly versatile species that can tolerate a very wide range of environmental stresses. They are found in high abundances in many of the habitats they live in, and represent important food sources for a variety of organisms.[2] In addition, they are found representing a significant number of symbiotic relationships with other aquatic insects, terrestrial insects, and some plants.[16]

They are distributed, and in most cases found in high abundance globally. They are found in wide variety of habitats, from the glaciated areas of the tallest mountains, to the deepest bodies of freshwater.[2]

Habitats edit

Aquatic habitats edit

Larval stages of the Chironomidae can be found in almost any aquatic or semiaquatic habitat. In fact, in many freshwater aquatic habitats, especially polluted ones, chironomids are often one of the most abundant insects found. Aquatic habitats can be marine or freshwater, the latter including treeholes, bromeliads, interstitial and benthic zones, as well as man-made sewage and artificial containers.[17] They can also be found inhabiting plant held waters, intertidal zones, interstitial zones. Many species of chironomids are found residing in sediments or benthic levels of water bodies, where Dissolved oxygen is very low.[2]

A number of chironomid species inhabit marine habitats. Midges of the genus Clunio are found in the intertidal zone, where they have adjusted their entire life cycle to the rhythm of the tides. This made the species Clunio marinus an important model species for research in the field of chronobiology.[18]

Terrestrial habitats edit

Chironomid larvae can also be found in some terrestrial habitats. Many species that are terrestrial are found living in soil as a dominant part of soil fauna community, particularly in wet soil habitats but also in agricultural land and in early stages of succession. Some species will use humic soils to develop as larvae on land; usually using decaying vegetation or sometimes living vegetation to survive. One genus of chironomids, Camptocladius, are known to develop as larvae in cow dung. There are also some that are known to burrow deep into soil in cases of temporary aquatic habitats or droughts.[17]

Chironomid adults are predominately found in terrestrial habitats.[2]

Roles in the ecosystem edit

Chironomidae have variable feeding ecology: most species feed on algae and other small soil organisms they can filtrate.[19][20] Some commensal species feed off of algae on their hosts, which also provides the benefit of protection and additional mobility; particularly when their chosen host site is another predatory larval species. Some species are parasitic, and pierce the integument of its host in order to feed off of its hemolymph.[21]

Larvae and pupae are important food items for fish, such as trout, banded killifish, and sticklebacks, and for many other aquatic organisms as well such as newts. Many aquatic insects, such as various predatory hemipterans in the families Nepidae, Notonectidae, and Corixidae eat Chironomidae in their aquatic phases. Additionally, predatory water beetles in families such as the Dytiscidae and Hydrophilidae have been found to feed on larval chironomids. Flying midges in their adult stage are eaten by fish and insectivorous birds, such as swallows and martins. They are also thought to be an especially important food source for tufted duck chicks during their first few days of life. They also are preyed on by bats and flying predatory insects, such as Odonata and dance flies.

They form an important fraction of the macro zoobenthos of most freshwater ecosystems. They are highly tolerant to low dissolved oxygen levels and changing salinity levels, both of which are often resultant from human pollution.[17] Thus, The Chironomidae are important as indicator organisms, i.e., the presence, absence, or quantities of various species in a body of water can indicate whether pollutants are present.[17]

Also, their fossils are widely used by palaeolimnologists and paleoentomologists as potential indicators of past environmental changes, including past climatic variability.[22] Although, the results of these tests are often controversial, and there are disagreements as to how beneficial they can be in the paleoentomology world.[17]

Symbiotic relationships edit

A significant portion of the Chironomidae larvae have been reported in commensal relationships with other organisms in their ecosystems. Although little is known about host choice and preference, it's been reported widely that Ephemeroptera are typically their most commonly reported host sites.[23]

Larval Chironomids often partake in commensal behaviors as a result of limited mobility and defensive morphology.[2] Although they do not have appendages designed for swimming, and most free-living movement is done through undulations, Chironomids do possess strong appendages that help them grasp onto a host.[24]

Commensalism in chironomids edit

Larval Commensal chironomids have been known to compete for space on the host. This competition happens on an interspecies level, but also with other commensal and or phoretic organisms. Chironomids have been reported to compete with ciliated protozoan Ephemera danica, although this competition has resulted in niche-partitioning on the hosts body. These species do not compete for food, but rather for ideal spaces on the host. In these competitive relationships, Chironomids generally choose larger host body sizes, while the protozoans might prefer smaller hosts. In host sites that contained both species, potential micro niches were observed, as these organisms were observed to deliberately space themselves out. Generally however, it seemed that chironomids beat out the protozoan in abundance on a host.[24]

A number of biotic and abiotic factors may affect host choice in chironomids. There is evidence that this choice is pressured by hydrological factors. A higher number of chironomids associating phoretically with Ephemeroptera during a rainy season. In the dry season, host choices seemed to diversify a great deal, with many hosts only being found in a dry season. It has been suggested that this may be due to an influx of vegetative debris and detritus that characterizes a rainy season in aquatic environments, which would cause an increase in associated taxa, such as Ephemeroptera.[23]

There have been reports of phoretic associates with predatory aquatic insects as well in chironomids. These relationships, although uncommon, represent more steady hosts for chironomids if they are accessible. Phoresy on a predatory host has been suggested to mean a more mobile, and protective host, resulting in an easier development stage for the commensal organism.[23]

Stress tolerance edit

Heat tolerance edit

Some chironomids can withstand high temperatures without the need to desiccate and pause development. A Japanese species of chironomid, Tokunagayusurika akasumi, have adapted to aestivation in sediments below their habitats in the event of high temperatures.[16]

Cold tolerance edit

Chironomids as a whole are a relatively cold-tolerant group. Overwintering of species is found in almost all subfamilies of chironomidae. Most of them cocoon during their pupation to overwinter, but this is not found in all individuals within a species. It is hypothesized that this overwintering difference in behavior occurs when temperature are at their lowest, and perhaps has to do with the presence of ice cover in the habitat.[16]

Pollution tolerance edit

The Chironomidae are important as indicator organisms, i.e., the presence, absence, or quantities of various species in a body of water can indicate whether pollutants are present. Also, their fossils are widely used by palaeolimnologists as indicators of past environmental changes, including past climatic variability.[25] Contemporary specimens are used by forensic entomologists as medico-legal markers for the postmortem interval assessment.[26]

Anhydrobiosis and other stresses edit

Anhydrobiosis is the ability of an organism to survive in the dry state. Anhydrobiotic larvae of the African chironomid Polypedilum vanderplanki can withstand prolonged complete desiccation (reviewed by Cornette and Kikawada[27]). These larvae can also withstand other external stresses including ionizing radiation.[28] The effects of anhydrobiosis, gamma ray and heavy-ion irradiation on the nuclear DNA and gene expression of these larvae were studied by Gusev et al.[28] They found that larval DNA becomes severely fragmented both upon anhydrobiosis and irradiation, and that these breaks are later repaired during rehydration or upon recovery from irradiation. An analysis of gene expression and antioxidant activity suggested the importance of removal of reactive oxygen species as well as the removal of DNA damages by repair enzymes. Expression of genes encoding DNA repair enzymes increased upon entering anhydrobiosis or upon exposure to radiation, and these increases indicated that when DNA damages occurred, they were subsequently repaired. In particular, expression of the Rad51 gene was substantially up-regulated following irradiation and during rehydration.[28] The Rad51 protein plays a key role in homologous recombination, a process required for the accurate repair of DNA double-strand breaks.

Many species of chironomids have adapted high tolerances to desiccation, particularly those inhabiting temporary aquatic habitats or places where droughts frequently occur. Anhydrobiosis, which is the resultant of complete desiccation, is the loss of most or all water in the body. Although its more common in terrestrial species, tolerance for drought within the Chironomidae is similar in both terrestrial and aquatic species.[16] A species from Africa was found with extreme tolerance to desiccation. This species was able to completely desiccate, and within 48 hours movement was found after rehydration. This same species is able to withstand this pause in development for an extended period of time, with records of it surviving up to 17 years completely dehydrated.[16]

Chironomids and humans edit

Chironomids can be both beneficial and nuisance to humans. In many areas of the world, anglers and aquarium hobbyists use them as bait or a food source, particularly bloodworms. They are also historically important food sources for some communities in Africa.[17]

Some Chironomid larvae thrive in very heavily polluted habitats. These habitats are often man-made, and result in little to no biodiversity, which implies no natural predators present to keep midge populations at natural levels. These conditions often worsen emergence events, which also take place near human dominated habitats.[2]

Subfamilies and genera edit

The family is divided into 11 subfamilies: Aphroteniinae, Buchonomyiinae, Chilenomyinae, Chironominae, Diamesinae, Orthocladiinae, Podonominae, Prodiamesinae, Tanypodinae, Telmatogetoninae, and Usambaromyiinae.[29][30] Most species belong to Chironominae, Orthocladiinae, and Tanypodinae. Diamesinae, Podonominae, Prodiamesinae, and Telmatogetoninae are medium-sized subfamilies with tens to hundreds of species. The remaining four subfamilies have fewer than five species each.

 
Chironomidae sp. female on flower of Euryops sp. Damage caused by beetles in family Meloidae.
 
Chironomidae larva, about 1 cm long, the head is right: The magnified tail details are from other images of the same animal.
 
Chironomid "bloodworm" larva showing the characteristic red color, about 40× magnification: The head is towards the upper left, just out of view.

References edit

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  3. ^ "Muckleheads[permanent dead link]" from Andre's Weather World (Andre Bernier, staff at WJW-TV), June 2, 2007.
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  25. ^ Walker, I. R. 2001. Midges: Chironomidae and related Diptera. pp. 43-66, In: J. P. Smol, H. J. B. Birks, and W. M. Last (eds). Tracking Environmental Change Using Lake Sediments. Volume 4. Zoological Indicators. Kluwer Academic Publishers, Dordrecht.
  26. ^ González Medina A, Soriano Hernando Ó, Jiménez Ríos G (2015). "The Use of the Developmental Rate of the Aquatic Midge Chironomus riparius (Diptera, Chironomidae) in the Assessment of the Postsubmersion Interval". J. Forensic Sci. 60 (3): 822–826. doi:10.1111/1556-4029.12707. hdl:10261/123473. PMID 25613586. S2CID 7167656.
  27. ^ Cornette R, Kikawada T (June 2011). "The induction of anhydrobiosis in the sleeping chironomid: current status of our knowledge". IUBMB Life. 63 (6): 419–29. doi:10.1002/iub.463. PMID 21547992.
  28. ^ a b c Gusev O, Nakahara Y, Vanyagina V, Malutina L, Cornette R, Sakashita T, Hamada N, Kikawada T, Kobayashi Y, Okuda T (2010). "Anhydrobiosis-associated nuclear DNA damage and repair in the sleeping chironomid: linkage with radioresistance". PLOS ONE. 5 (11): e14008. Bibcode:2010PLoSO...514008G. doi:10.1371/journal.pone.0014008. PMC 2982815. PMID 21103355.
  29. ^ J.H. Epler. 2001. Identification manual for the larval Chironomidae (Diptera) of North and South Carolina 2005-12-14 at the Wayback Machine. North Carolina Department of Environment and Natural Resources.
  30. ^ Armitage, P., Cranston, P.S., and Pinder, L.C.V. (eds.) (1994) The Chironomidae: Biology and Ecology of Non-biting Midges. Chapman and Hall, London, 572 pp.
  31. ^ Ekrem, Torbjørn. . Archived from the original on 2009-03-18. Retrieved 2009-04-30.
  32. ^ Makarchenko, Eugenyi A. (2005). (PDF). Zootaxa. 1084: 59–64. doi:10.11646/zootaxa.1084.1.5. Archived from the original (PDF) on 2016-03-04. Retrieved 2009-04-03.
  33. ^ Caldwell, Broughton A.; Soponis, Annelle R. (1982). "Hudsonimyia Parrishi, a New Species of Tanypodinae (Diptera: Chironomidae) from Georgia" (PDF). The Florida Entomologist. 65 (4): 506–513. doi:10.2307/3494686. ISSN 0015-4040. JSTOR 3494686. Retrieved 2009-04-20.
  34. ^ Zorina, Oksana V. (2007). "Olecryptotendipes, a new genus in the Harnischia complex (Diptera: Chironomidae) from the Russian Far East" (PDF). In Andersen, T. (ed.). Contributions to the Systematics and Ecology of Aquatic Diptera—A Tribute to Ole A. Sæther. The Caddis Press. pp. 347–351.[permanent dead link]
  35. ^ Halvorsen, Godtfred A. (1982). "Saetheriella amplicristata gen. n., sp. n., a new Orthocladiinae (Diptera: Chironomidae) from Tennessee". Aquatic Insects. 4 (3): 131–136. Bibcode:1982AqIns...4..131H. doi:10.1080/01650428209361098. ISSN 1744-4152.
  36. ^ Andersen, Trond; Sæther, Ole A. (January 1994). "Usambaromyia nigrala gen. n., sp. n., and Usambaromyiinae, a new subfamily among the Chironomidae (Diptera)". Aquatic Insects. 16 (1): 21–29. Bibcode:1994AqIns..16...21A. doi:10.1080/01650429409361531. ISSN 1744-4152.

External links edit

chironomidae, lake, redirects, here, species, genus, chaoborus, family, chaoboridae, chaoborus, informally, known, chironomids, nonbiting, midges, lake, flies, comprise, family, nematoceran, flies, with, global, distribution, they, closely, related, ceratopogo. Lake fly redirects here For species in the genus Chaoborus family Chaoboridae see Chaoborus The Chironomidae informally known as chironomids nonbiting midges or lake flies comprise a family of nematoceran flies with a global distribution They are closely related to the Ceratopogonidae Simuliidae and Thaumaleidae Many species superficially resemble mosquitoes but they lack the wing scales and elongated mouthparts of the Culicidae ChironomidaeTemporal range Triassic Present PreꞒ Ꞓ O S D C P T J K Pg NMale Chironomus plumosusScientific classificationDomain EukaryotaKingdom AnimaliaPhylum ArthropodaClass InsectaOrder DipteraSuborder NematoceraInfraorder CulicomorphaSuperfamily ChironomoideaFamily ChironomidaeNewman 1834 1 SubfamiliesSee textTwo lake flies observed in Neenah Wisconsin after the yearly hatch in Lake WinnebagoThe name Chironomidae stems from the Ancient Greek word kheironomos a pantomimist Contents 1 Common names and biodiversity 2 Behavior and description 3 Ecology and distribution 3 1 Habitats 3 1 1 Aquatic habitats 3 1 2 Terrestrial habitats 3 2 Roles in the ecosystem 3 2 1 Symbiotic relationships 3 2 1 1 Commensalism in chironomids 4 Stress tolerance 4 1 Heat tolerance 4 2 Cold tolerance 4 3 Pollution tolerance 4 4 Anhydrobiosis and other stresses 5 Chironomids and humans 6 Subfamilies and genera 7 References 8 External linksCommon names and biodiversity editThis is a large taxon of insects Some estimates of the species numbers suggest well over 10 000 world wide 2 Males are easily recognized by their plumose antennae Adults are known by a variety of vague and inconsistent common names largely by confusion with other insects For example chironomids are known as lake flies in parts of Canada and Lake Winnebago Wisconsin but bay flies in the areas near the bay of Green Bay Wisconsin They are called sand flies muckleheads 3 muffleheads 4 Canadian soldiers 5 or American soldiers 6 in various regions of the Great Lakes area They have been called blind mosquitoes or chizzywinks in Florida 7 In Kansas they are known as midges 8 However they are not mosquitoes of any sort and the term sandflies generally refers to various species of biting flies unrelated to the Chironomidae The group includes the wingless Belgica antarctica the largest terrestrial animal of Antarctica 9 10 Their larvae produce silk and Chironomus has been studied as an alternative source of silk other than the silk moth as it is possible to extract it without killing the animal Ahimsa silk 11 The biodiversity of the Chironomidae often goes unnoticed because they are notoriously difficult to identify and ecologists usually record them by species groups Each morphologically distinct group comprises a number of morphologically identical sibling species that can only be identified by rearing adult males or by cytogenetic analysis of the polytene chromosomes Polytene chromosomes were originally observed in the larval salivary glands of Chironomus midges by Balbiani in 1881 They form through repeated rounds of DNA replication without cell division resulting in characteristic light and dark banding patterns which can be used to identify inversions and deletions which allow species identification Behavior and description editChironomids as a group are very diverse in their life histories and exhibit a variety of behaviors during their development Due to their species ambiguity and this diversity of behaviors there is much controversy in research on their feeding habits particularly as adults Many reference sources in the past century or so have repeated the assertion that the chironomidae do not feed as adults but an increasing body of evidence contradicts this view Adults of many species do in fact feed The natural foods reported include fresh fly droppings nectar pollen honeydew and various sugar rich materials The question whether feeding is of practical importance has by now been clearly settled for some Chironomus species at least specimens that had fed on sucrose flew far longer than starved specimens and starved females longer than starved males which suggested they had eclosed with larger reserves of energy than the males Some authors suggest the females and males of certain species apply the resources obtained in feeding differently Generally Males expend the extra energy on flight while females use their food resources to achieve longer lifespans The respective strategies should be compatible with maximal probability of successful mating and reproduction in those species that do not mate immediately after eclosion particularly in species that have more than one egg mass maturing the less developed masses being oviposited after a delay Such variables also would be relevant to species that exploit wind for dispersal laying eggs at intervals Chironomids that feed on nectar or pollen may well be of importance as pollinators but current evidence on such points is largely anecdotal However the content of protein and other nutrients in pollen in comparison to nectar might well contribute to the females reproductive capacities 2 Larvae of some species are bright red in color due to a hemoglobin analog these are often known as bloodworms 12 Their ability to capture oxygen is further increased by making undulating movements 13 Adults can be pests when they emerge in large numbers They may cause difficulty during driving if they collide with the windshield creating an opaque coating which obscures the driver s vision 14 They can damage paint brick and other surfaces with their droppings When large numbers of adults die they can build up into malodorous piles They can provoke allergic reactions in sensitive individuals 15 These allergic reactions have been shown to be caused by the haemolglobins that are primarily found in Chironomidae larval stages 2 Ecology and distribution editChironomids are highly versatile species that can tolerate a very wide range of environmental stresses They are found in high abundances in many of the habitats they live in and represent important food sources for a variety of organisms 2 In addition they are found representing a significant number of symbiotic relationships with other aquatic insects terrestrial insects and some plants 16 They are distributed and in most cases found in high abundance globally They are found in wide variety of habitats from the glaciated areas of the tallest mountains to the deepest bodies of freshwater 2 Habitats edit Aquatic habitats edit Larval stages of the Chironomidae can be found in almost any aquatic or semiaquatic habitat In fact in many freshwater aquatic habitats especially polluted ones chironomids are often one of the most abundant insects found Aquatic habitats can be marine or freshwater the latter including treeholes bromeliads interstitial and benthic zones as well as man made sewage and artificial containers 17 They can also be found inhabiting plant held waters intertidal zones interstitial zones Many species of chironomids are found residing in sediments or benthic levels of water bodies where Dissolved oxygen is very low 2 A number of chironomid species inhabit marine habitats Midges of the genus Clunio are found in the intertidal zone where they have adjusted their entire life cycle to the rhythm of the tides This made the species Clunio marinus an important model species for research in the field of chronobiology 18 Terrestrial habitats edit Chironomid larvae can also be found in some terrestrial habitats Many species that are terrestrial are found living in soil as a dominant part of soil fauna community particularly in wet soil habitats but also in agricultural land and in early stages of succession Some species will use humic soils to develop as larvae on land usually using decaying vegetation or sometimes living vegetation to survive One genus of chironomids Camptocladius are known to develop as larvae in cow dung There are also some that are known to burrow deep into soil in cases of temporary aquatic habitats or droughts 17 Chironomid adults are predominately found in terrestrial habitats 2 Roles in the ecosystem edit Chironomidae have variable feeding ecology most species feed on algae and other small soil organisms they can filtrate 19 20 Some commensal species feed off of algae on their hosts which also provides the benefit of protection and additional mobility particularly when their chosen host site is another predatory larval species Some species are parasitic and pierce the integument of its host in order to feed off of its hemolymph 21 Larvae and pupae are important food items for fish such as trout banded killifish and sticklebacks and for many other aquatic organisms as well such as newts Many aquatic insects such as various predatory hemipterans in the families Nepidae Notonectidae and Corixidae eat Chironomidae in their aquatic phases Additionally predatory water beetles in families such as the Dytiscidae and Hydrophilidae have been found to feed on larval chironomids Flying midges in their adult stage are eaten by fish and insectivorous birds such as swallows and martins They are also thought to be an especially important food source for tufted duck chicks during their first few days of life They also are preyed on by bats and flying predatory insects such as Odonata and dance flies They form an important fraction of the macro zoobenthos of most freshwater ecosystems They are highly tolerant to low dissolved oxygen levels and changing salinity levels both of which are often resultant from human pollution 17 Thus The Chironomidae are important as indicator organisms i e the presence absence or quantities of various species in a body of water can indicate whether pollutants are present 17 Also their fossils are widely used by palaeolimnologists and paleoentomologists as potential indicators of past environmental changes including past climatic variability 22 Although the results of these tests are often controversial and there are disagreements as to how beneficial they can be in the paleoentomology world 17 Symbiotic relationships edit A significant portion of the Chironomidae larvae have been reported in commensal relationships with other organisms in their ecosystems Although little is known about host choice and preference it s been reported widely that Ephemeroptera are typically their most commonly reported host sites 23 Larval Chironomids often partake in commensal behaviors as a result of limited mobility and defensive morphology 2 Although they do not have appendages designed for swimming and most free living movement is done through undulations Chironomids do possess strong appendages that help them grasp onto a host 24 Commensalism in chironomids edit Larval Commensal chironomids have been known to compete for space on the host This competition happens on an interspecies level but also with other commensal and or phoretic organisms Chironomids have been reported to compete with ciliated protozoan Ephemera danica although this competition has resulted in niche partitioning on the hosts body These species do not compete for food but rather for ideal spaces on the host In these competitive relationships Chironomids generally choose larger host body sizes while the protozoans might prefer smaller hosts In host sites that contained both species potential micro niches were observed as these organisms were observed to deliberately space themselves out Generally however it seemed that chironomids beat out the protozoan in abundance on a host 24 A number of biotic and abiotic factors may affect host choice in chironomids There is evidence that this choice is pressured by hydrological factors A higher number of chironomids associating phoretically with Ephemeroptera during a rainy season In the dry season host choices seemed to diversify a great deal with many hosts only being found in a dry season It has been suggested that this may be due to an influx of vegetative debris and detritus that characterizes a rainy season in aquatic environments which would cause an increase in associated taxa such as Ephemeroptera 23 There have been reports of phoretic associates with predatory aquatic insects as well in chironomids These relationships although uncommon represent more steady hosts for chironomids if they are accessible Phoresy on a predatory host has been suggested to mean a more mobile and protective host resulting in an easier development stage for the commensal organism 23 Stress tolerance editHeat tolerance edit Some chironomids can withstand high temperatures without the need to desiccate and pause development A Japanese species of chironomid Tokunagayusurika akasumi have adapted to aestivation in sediments below their habitats in the event of high temperatures 16 Cold tolerance edit Chironomids as a whole are a relatively cold tolerant group Overwintering of species is found in almost all subfamilies of chironomidae Most of them cocoon during their pupation to overwinter but this is not found in all individuals within a species It is hypothesized that this overwintering difference in behavior occurs when temperature are at their lowest and perhaps has to do with the presence of ice cover in the habitat 16 Pollution tolerance edit The Chironomidae are important as indicator organisms i e the presence absence or quantities of various species in a body of water can indicate whether pollutants are present Also their fossils are widely used by palaeolimnologists as indicators of past environmental changes including past climatic variability 25 Contemporary specimens are used by forensic entomologists as medico legal markers for the postmortem interval assessment 26 Anhydrobiosis and other stresses edit Anhydrobiosis is the ability of an organism to survive in the dry state Anhydrobiotic larvae of the African chironomid Polypedilum vanderplanki can withstand prolonged complete desiccation reviewed by Cornette and Kikawada 27 These larvae can also withstand other external stresses including ionizing radiation 28 The effects of anhydrobiosis gamma ray and heavy ion irradiation on the nuclear DNA and gene expression of these larvae were studied by Gusev et al 28 They found that larval DNA becomes severely fragmented both upon anhydrobiosis and irradiation and that these breaks are later repaired during rehydration or upon recovery from irradiation An analysis of gene expression and antioxidant activity suggested the importance of removal of reactive oxygen species as well as the removal of DNA damages by repair enzymes Expression of genes encoding DNA repair enzymes increased upon entering anhydrobiosis or upon exposure to radiation and these increases indicated that when DNA damages occurred they were subsequently repaired In particular expression of the Rad51 gene was substantially up regulated following irradiation and during rehydration 28 The Rad51 protein plays a key role in homologous recombination a process required for the accurate repair of DNA double strand breaks Many species of chironomids have adapted high tolerances to desiccation particularly those inhabiting temporary aquatic habitats or places where droughts frequently occur Anhydrobiosis which is the resultant of complete desiccation is the loss of most or all water in the body Although its more common in terrestrial species tolerance for drought within the Chironomidae is similar in both terrestrial and aquatic species 16 A species from Africa was found with extreme tolerance to desiccation This species was able to completely desiccate and within 48 hours movement was found after rehydration This same species is able to withstand this pause in development for an extended period of time with records of it surviving up to 17 years completely dehydrated 16 Chironomids and humans editChironomids can be both beneficial and nuisance to humans In many areas of the world anglers and aquarium hobbyists use them as bait or a food source particularly bloodworms They are also historically important food sources for some communities in Africa 17 Some Chironomid larvae thrive in very heavily polluted habitats These habitats are often man made and result in little to no biodiversity which implies no natural predators present to keep midge populations at natural levels These conditions often worsen emergence events which also take place near human dominated habitats 2 Subfamilies and genera editThe family is divided into 11 subfamilies Aphroteniinae Buchonomyiinae Chilenomyinae Chironominae Diamesinae Orthocladiinae Podonominae Prodiamesinae Tanypodinae Telmatogetoninae and Usambaromyiinae 29 30 Most species belong to Chironominae Orthocladiinae and Tanypodinae Diamesinae Podonominae Prodiamesinae and Telmatogetoninae are medium sized subfamilies with tens to hundreds of species The remaining four subfamilies have fewer than five species each nbsp Chironomidae sp female on flower of Euryops sp Damage caused by beetles in family Meloidae nbsp Chironomidae larva about 1 cm long the head is right The magnified tail details are from other images of the same animal nbsp Chironomid bloodworm larva showing the characteristic red color about 40 magnification The head is towards the upper left just out of view Aagaardia Saether 2000 Abiskomyia Edwards 1937 Ablabesmyia Johannsen 1905 Acalcarella Acamptocladius Brundin 1956 Acricotopus Kieffer 1921 Aedokritus Aenne Afrochlus Afrozavrelia Harrison 2004 31 Allocladius Allometriocnemus Allotrissocladius Alotanypus Roback 1971 Amblycladius Amnihayesomyia Amphismittia Anaphrotenia Anatopynia Johannsen 1905 Ancylocladius Andamanus Antillocladius Saether 1981 Anuncotendipes Apedilum Townes 1945 Aphrotenia Aphroteniella Apometriocnemus Saether 1984 Apsectrotanypus Fittkau 1962 Archaeochlus Arctodiamesa Makarchenko 1983 32 Arctopelopia Fittkau 1962 Arctosmittia Asachironomus Asclerina Asheum Sublette amp Sublette 1983 Australopelopia Austrobrillia Austrochlus Austrocladius Axarus Roback 1980 Baeoctenus Baeotendipes Kieffer 1913 Bavarismittia Beardius Reiss amp Sublette 1985 Beckidia Saether 1979 Belgica Bernhardia Bethbilbeckia Biwatendipes Boreochlus Edwards 1938 Boreoheptagyia Brundin 1966 Boreosmittia Botryocladius Brillia Kieffer 1913 Brundiniella Brunieria Bryophaenocladius Thienemann 1934 Buchonomyia Fittkau 1955 Caladomyia Camposimyia Camptocladius van der Wulp 1874 Cantopelopia Carbochironomus Reiss amp Kirschbaum 1990 Cardiocladius Kieffer 1912 Chaetocladius Kieffer 1911 Chasmatonotus Chernovskiia Saether 1977 Chilenomyia Chirocladius Chironomidae Chironominae Chironomini Chironomus Meigen 1803 Chrysopelopia Cladopelma Kieffer 1921 Cladotanytarsus Kieffer 1921 Clinotanypus Kieffer 1913 Clunio Haliday 1855 Coelopynia Coelotanypus Coffmania Collartomyia Colosmittia Compteromesa Saether 1981 Compterosmittia Conchapelopia Fittkau 1957 Conochironomus Constempellina Brundin 1947 Corynocera Zetterstedt 1838 Corynoneura Winnertz 1846 Corynoneurella Brundin 1949 Corytibacladius Cricotopus van der Wulp 1874 Cryptochironomus Kieffer 1918 Cryptotendipes Lenz 1941 Cyphomella Saether 1977 Dactylocladius Daitoyusurika Demeijerea Kruseman 1933 Demicryptochironomus Lenz 1941 Denopelopia Derotanypus Diamesa Meigen in Gistl 1835 Diamesinae Dicrotendipes Kieffer 1913 Diplocladius Kieffer 1908 Diplosmittia Djalmabatista Fittkau 1968 Doithrix Doloplastus Doncricotopus Dratnalia Echinocladius Edwardsidia Einfeldia Kieffer 1924 Endochironomus Kieffer 1918 Endotribelos Epoicocladius Sulc amp ZavIel 1924 Eretmoptera Eukiefferiella Thienemann 1926 Eurycnemus van der Wulp 1874 Euryhapsis Oliver 1981 Eusmittia Fissimentum Fittkauimyia Fleuria Freemaniella Friederia Georthocladius Strenzke 1941 Gillotia Kieffer 1921 Glushkovella Glyptotendipes Kieffer 1913 Goeldichironomus Graceus Goetghebuer 1928 Gravatamberus Gressittius Guassutanypus Guttipelopia Fittkau 1962 Gymnometriocnemus Goetghebeur 1932 Gynnidocladius Gynocladius Mendes Saether amp Andrade Morraye 2005 Hahayusurika Halirytus Halocladius Hirvenoja 1973 Hanochironomus Hanocladius Harnischia Kieffer 1921 Harrisius Harrisonina Hayesomyia Murray amp Fittkau 1985 Heleniella Gouin 1943 HelopelopiaRoback 1971 Henrardia Heptagyia Heterotanytarsus Sparck 1923 Heterotrissocladius Sparck 1923 Hevelius Himatendipes Hirosimayusurika Hudsonimyia Roback 1979 33 Hydrobaenus Hydrosmittia Hyporhygma Ichthyocladius Fittkau 1974 Ikiprimus Ikisecundus Imparipecten Indoaxarus Indocladius Ionthosmittia Irisobrillia Kaluginia Kamelopelopia Kaniwhaniwhanus Kiefferophyes Kiefferulus Goetghebuer 1922 Knepperia Kloosia Kruseman 1933 Krenopelopia Fittkau 1962 Krenopsectra Krenosmittia Thienemann amp Kruger 1939 Kribiobius Kribiocosmus Kribiodosis Kribiopelma Kribiothauma Kribioxenus Kurobebrillia Kuschelius Labrundinia Fittkau 1962 Lappodiamesa Serra Tosio 1968 Lappokiefferiella Lapposmittia Larsia Fittkau 1962 Lasiodiamesa Kieffer 1924 Laurotanypus Lauterborniella Thienemann amp Bause 1913 Lepidopelopia Lepidopodus Lerheimia Limaya Limnophyes Eaton 1875 Lindebergia Linevitshia Lipiniella Shilova 1961 Lipurometriocnemus Lithotanytarsus Litocladius Andersen Mendes amp Saether 2004 Ljungneria Lobodiamesa Lobomyia Lobosmittia Lopescladius Lunditendipes Lyrocladius Mendes amp Andersen 2008 Macropelopia Thienemann 1916 Macropelopini Manoa Maoridiamesa Mapucheptagyia Maryella Mecaorus Megacentron Mesocricotopus Mesosmittia Brundin 1956 Metriocnemus van der Wulp 1874 Microchironomus Kieffer 1918 Micropsectra Kieffer 1909 Microtendipes Kieffer 1915 Microzetia Molleriella Mongolchironomus Mongolcladius Mongolyusurika Monodiamesa Kieffer 1922 Monopelopia Fittkau 1962 Murraycladius Nakataia Nandeva Nanocladius Kieffer 1913 Naonella Nasuticladius Natarsia Fittkau 1962 Neelamia Neobrillia Neopodonomus Neostempellina Neozavrelia Goetghebuer 1941 Nesiocladius Nilodorum Nilodosis Nilotanypus Kieffer 1923 Nilothauma Kieffer 1921 Nimbocera Notocladius Odontomesa Pagast 1947 Okayamayusurika Okinawayusurika Olecryptotendipes Zorina 2007 34 Oleia Oliveridia Saether 1980 Omisus Townes 1945 Onconeura Ophryophorus Oreadomyia Orthocladiinae Orthocladius van der Wulp 1874 Oryctochlus Oukuriella Pagastia Oliver 1959 Pagastiella Brundin 1949 Paraboreochlus Thienemann 1939 Parachaetocladius Parachironomus Lenz 1921 Paracladius Hirvenoja 1973 Paracladopelma Harnisch 1923 Paracricotopus Thienemann amp Harnisch 1932 Parakiefferiella Thienemann 1936 Paralauterborniella Lenz 1941 Paralimnophyes Brundin 1956 Paramerina Fittkau 1962 Parametriocnemus Goetghebuer 1932 Pamirocesa Paraborniella Parachironominae Paradoxocladius Paraheptagyia Paranilothauma Parapentaneura Paraphaenocladius Thienemann 1924 Paraphrotenia Parapsectra Reiss 1969 Parapsectrocladius Parasmittia Paratanytarsus Thienemann amp Bause 1913 Paratendipes Kieffer 1911 Paratrichocladius Thienemann 1942 Paratrissocladius ZavIel 1937 Parochlus Enderlein 1912 Parorthocladius Thienemann 1935 Parvitergum Paucispinigera Pelomus Pentaneura Pentaneurella Pentaneurini Pentapedilum Petalocladius Phaenopsectra Kieffer 1921 Physoneura Pirara Platysmittia Saether 1982 Plhudsonia Podochlus Podonomopsis Podonomus Polypedilum Kieffer 1912 Pontomyia Potthastia Kieffer 1922 Prochironomus Procladiini Procladius Skuse 1889 Prodiamesa Kieffer 1906 Propsilocerus Prosmittia Protanypus Kieffer 1906 Psectrocladius Kieffer 1906 Psectrotanypus Kieffer 1909 Pseudobrillia Pseudochironomus Malloch 1915 Pseudodiamesa Goetghebuer 1939 Pseudohydrobaenus Pseudokiefferiella Zavrel 1941 Pseudorthocladius Goetghebuer 1932 Pseudosmittia Goetghebuer 1932 Psilochironomus Psilometriocnemus Saether 1969 Pterosis Qiniella Reissmesa Rheochlus Rheocricotopus Brundin 1956 Rheomus Rheomyia Rheopelopia Fittkau 1962 Rheosmittia Brundin 1956 Rheotanytarsus Thienemann amp Bause 1913 Rhinocladius Riethia Robackia Saether 1977 Saetheria Jackson 1977 Saetheriella Halvorsen 1982 35 Saetherocladius Saetherocryptus Saetheromyia Saetherops Sasayusurika Schineriella Murray amp Fittkau 1988 Semiocladius Setukoyusurika Seppia Sergentia Kieffer 1922 Shangomyia Shilovia Skusella Skutzia Smittia Holmgren 1869 Stackelbergina Stelechomyia Stempellina Thienemann amp Bause 1913 Stempellinella Brundin 1947 Stenochironomus Kieffer 1919 Stictochironomus Kieffer 1919 Stictocladius Stictotendipes Stilocladius Rossaro 1979 Sublettea Sublettiella Sumatendipes Symbiocladius Kieffer 1925 Sympotthastia Pagast 1947 Syndiamesa Kieffer 1918 Synendotendipes Grodhaus 1987 Synorthocladius Thienemann 1935 Tanypodinae Tanypus Meigen 1803 Tanytarsini Tanytarsus van der Wulp 1874 Tavastia Telmatogeton Schiner 1866 Telmatopelopia Fittkau 1962 Telopelopia Tempisquitoneura Tethymyia Thalassomya Schiner 1856 Thalassosmittia Strenzke amp Remmert 1957 Thienemannia Kieffer 1909 Thienemanniella Kieffer 1911 Thienemannimyia Fittkau 1957 Thienemanniola Tobachironomus Tokunagaia Saether 1973 Tokunagayusurika Tokyobrillia Tosayusurika Townsia Toyamayusurika Tribelos Townes 1945 Trichochilus Trichosmittia Trichotanypus Kieffer 1906 Trissocladius Kieffer 1908 Trissopelopia Kieffer 1923 Trondia Tsudayusurika Tusimayusurika Tvetenia Kieffer 1922 Unniella Saether 1982 Usambaromyia Andersen amp Saether 1994 36 Virgatanytarsus Pinder 1982 Vivacricotopus Wirthiella Xenochironomus Kieffer 1921 Xenopelopia Fittkau 1962 Xestochironomus Xestotendipes Xiaomyia Xylotopus Yaeprimus Yaequartus Yaequintus Yaesecundus Yaetanytarsus Yaetertius Yama Zalutschia Lipina 1939 Zavrelia Kieffer 1913 Zavreliella Kieffer 1920 Zavrelimyia Fittkau 1962 Zelandochlus Zhouomyia ZuluchironomusReferences edit Sabrosky C W 1999 Family Group Names in Diptera amp Bibliograpgy PDF Myia 10 1 360 page 85 a b c d e f g h i Armitage P D Cranston P S Pinder L C V 1995 The Chironomidae biology and ecology of non biting midges London Chapman amp Hall ISBN 978 0 412 45260 4 Muckleheads permanent dead link from Andre s Weather World Andre Bernier staff at WJW TV June 2 2007 You don t love muffleheads but Lake Erie does Sandusky Register May 24 2010 Galbincea Barb Canadian Soldiers Invade Rocky River The Plain Dealer Cleveland com June 18 2014 accessed June 3 2015 Call Them Mayflies Not June Bugs Biologist Says University of Windsor Professor Dispels Mayfly Myths CBC News CBC ca May 29 2012 accessed June 3 2015 Chizzywinks are Blind Mosquitos by Dan Culbert of the University of Florida August 17 2005 Melgares Pat August 9 2022 Sorghum midge aphid found in Kansas High Plains Journal Retrieved April 7 2023 Usher Michael B Edwards Marion 1984 A dipteran from south of the Antarctic Circle Belgica antarctica Chironomidae with a description of its larva Biological Journal of the Linnean Society 23 1 19 31 doi 10 1111 j 1095 8312 1984 tb00803 x Luke Sandro amp Juanita Constible Antarctic Bestiary Terrestrial Animals Laboratory for Ecophysiological Cryobiology Miami University Archived from the original on 23 December 2008 Retrieved December 9 2008 Structural and physical analysis of underwater silk from housing nest composites of a tropical chironomid midge Contreras Ramos Atilano 2010 08 01 Richard W Merritt Kenneth W Cummins y Martin B Berg eds 2008 An introduction to the aquatic insects of North America 4a edicion Kendall Hunt Publishing Company Dubuque 1158 p CD con clave interactiva Revista Mexicana de Biodiversidad 81 2 doi 10 22201 ib 20078706e 2010 002 247 ISSN 2007 8706 Panis Luc Int Goddeeris Boudewijn Verheyen Rudolf January 1996 On the relationship between vertical microdistribution and adaptations to oxygen stress in littoral Chironomidae Diptera Hydrobiologia 318 1 3 61 67 doi 10 1007 BF00014132 ISSN 0018 8158 S2CID 27026595 Skeeter June 2020 06 19 Response to Reviewer 1 doi 10 5194 bg 2019 477 ac1 a href Template Cite web html title Template Cite web cite web a Missing or empty url help Ali Arshad Leckel Robert J Jahan Nusrad Al Shami Salman A Rawi Che Salmah MD March 2009 Laboratory and Field Investigations of Pestiferous Chironomidae Diptera in Some Man Made Wetlands in Central Florida USA Journal of the American Mosquito Control Association 25 1 94 99 doi 10 2987 08 5798 1 ISSN 8756 971X PMID 19432073 S2CID 33142033 a b c d e Tokeshi Mutsunori June 1993 On the evolution of commensalism in the Chironomidae Freshwater Biology 29 3 481 489 Bibcode 1993FrBio 29 481T doi 10 1111 j 1365 2427 1993 tb00782 x ISSN 0046 5070 a b c d e f McLachlan Athol Armitage P Cranston P S Pinder L C V September 1995 The Chironomidae The Biology and Ecology of Non biting Midges The Journal of Animal Ecology 64 5 667 Bibcode 1995JAnEc 64 667M doi 10 2307 5810 ISSN 0021 8790 JSTOR 5810 Kaiser Tobias S Poehn Birgit Szkiba David Preussner Marco Sedlazeck Fritz J Zrim Alexander Neumann Tobias Nguyen Lam Tung Betancourt Andrea J Hummel Thomas Vogel Heiko Dorner Silke Heyd Florian von Haeseler Arndt Tessmar Raible Kristin 2016 12 01 The genomic basis of circadian and circalunar timing adaptations in a midge Nature 540 7631 69 73 Bibcode 2016Natur 540 69K doi 10 1038 nature20151 ISSN 0028 0836 PMC 5133387 PMID 27871090 Lukesova Alena Frouz Jan 2007 Seckbach Joseph ed Soil and Freshwater Micro Algae as a Food Source for Invertebrates in Extreme Environments Algae and Cyanobacteria in Extreme Environments Cellular Origin Life in Extreme Habitats and Astrobiology vol 11 Dordrecht Springer Netherlands pp 265 284 doi 10 1007 978 1 4020 6112 7 14 ISBN 978 1 4020 6112 7 retrieved 2021 07 05 Frouz Jan Ali Arshad Lobinske Richard J 2004 Algal food selection and digestion by larvae of the pestiferous chironomid Chironomus Crassicaudatus under laboratory conditions Journal of the American Mosquito Control Association 20 4 458 461 ISSN 8756 971X PMID 15669393 Corkum Lynda D Ciborowski Jan J H eds 1995 Current directions in research on ephemeroptera Toronto Canadian Scholars Press ISBN 978 1 55130 075 7 Walker Ian R 2001 Midges Chironomidae and Related Diptera Tracking Environmental Change Using Lake Sediments Developments in Paleoenvironmental Research vol 4 Dordrecht Springer Netherlands pp 43 66 doi 10 1007 0 306 47671 1 3 ISBN 978 90 481 6034 1 retrieved 2023 10 27 a b c Rosa Beatriz F J Vescovi Martins Renato T Oliveira Vivian C de Alves Roberto da G December 2009 Phoretic association between larvae of Rheotanytarsus Diptera Chironomidae and genera of Odonata in a first order stream in an area of Atlantic Forest in southeastern Brazil Zoologia Curitiba 26 4 787 791 doi 10 1590 s1984 46702009000400025 ISSN 1984 4689 a b Grzybkowska Maria Szczerkowska Majchrzak Eliza Dukowska Malgorzata Leszczynska Joanna Przybylski Miroslaw 2016 01 01 Ephemera danica Ephemeroptera Ephemeridae As a Resource for Two Commensals Ciliated Protozoans Sessilida and Chironomids Diptera Journal of Insect Science 16 1 doi 10 1093 jisesa iew050 ISSN 1536 2442 PMC 7175965 PMID 28076285 Walker I R 2001 Midges Chironomidae and related Diptera pp 43 66 In J P Smol H J B Birks and W M Last eds Tracking Environmental Change Using Lake Sediments Volume 4 Zoological Indicators Kluwer Academic Publishers Dordrecht Gonzalez Medina A Soriano Hernando o Jimenez Rios G 2015 The Use of the Developmental Rate of the Aquatic Midge Chironomus riparius Diptera Chironomidae in the Assessment of the Postsubmersion Interval J Forensic Sci 60 3 822 826 doi 10 1111 1556 4029 12707 hdl 10261 123473 PMID 25613586 S2CID 7167656 Cornette R Kikawada T June 2011 The induction of anhydrobiosis in the sleeping chironomid current status of our knowledge IUBMB Life 63 6 419 29 doi 10 1002 iub 463 PMID 21547992 a b c Gusev O Nakahara Y Vanyagina V Malutina L Cornette R Sakashita T Hamada N Kikawada T Kobayashi Y Okuda T 2010 Anhydrobiosis associated nuclear DNA damage and repair in the sleeping chironomid linkage with radioresistance PLOS ONE 5 11 e14008 Bibcode 2010PLoSO 514008G doi 10 1371 journal pone 0014008 PMC 2982815 PMID 21103355 J H Epler 2001 Identification manual for the larval Chironomidae Diptera of North and South Carolina Archived 2005 12 14 at the Wayback Machine North Carolina Department of Environment and Natural Resources Armitage P Cranston P S and Pinder L C V eds 1994 The Chironomidae Biology and Ecology of Non biting Midges Chapman and Hall London 572 pp Ekrem Torbjorn Systematics and biogeography of Zavrelia Afrozavrelia and Stempellinella Diptera Chironomidae Archived from the original on 2009 03 18 Retrieved 2009 04 30 Makarchenko Eugenyi A 2005 A new species of Arctodiamesa Makarchenko Diptera Chironomidae Diamesinae from the Russian Far East with a key to known species of the genus PDF Zootaxa 1084 59 64 doi 10 11646 zootaxa 1084 1 5 Archived from the original PDF on 2016 03 04 Retrieved 2009 04 03 Caldwell Broughton A Soponis Annelle R 1982 Hudsonimyia Parrishi a New Species of Tanypodinae Diptera Chironomidae from Georgia PDF The Florida Entomologist 65 4 506 513 doi 10 2307 3494686 ISSN 0015 4040 JSTOR 3494686 Retrieved 2009 04 20 Zorina Oksana V 2007 Olecryptotendipes a new genus in the Harnischia complex Diptera Chironomidae from the Russian Far East PDF In Andersen T ed Contributions to the Systematics and Ecology of Aquatic Diptera A Tribute to Ole A Saether The Caddis Press pp 347 351 permanent dead link Halvorsen Godtfred A 1982 Saetheriella amplicristata gen n sp n a new Orthocladiinae Diptera Chironomidae from Tennessee Aquatic Insects 4 3 131 136 Bibcode 1982AqIns 4 131H doi 10 1080 01650428209361098 ISSN 1744 4152 Andersen Trond Saether Ole A January 1994 Usambaromyia nigrala gen n sp n and Usambaromyiinae a new subfamily among the Chironomidae Diptera Aquatic Insects 16 1 21 29 Bibcode 1994AqIns 16 21A doi 10 1080 01650429409361531 ISSN 1744 4152 External links edit nbsp Wikispecies has information related to Chironomidae nbsp Wikimedia Commons has media related to Chironomidae The Chironomid Home Page Chironomidae and Water Beetles of Florida Chironomidae Research Group University of Minnesota Family Chironomidae at Soil and Water Conservation Society of Metro Halifax Checklist of UK Recorded Chironomidae Chironomidae at Nomina Insecta Nearctica Chironomid Palaeoecology UBC Okanagan Archived 2022 05 18 at the Wayback Machine Chironomidae at Australian Faunal Directory Hydrilla tip mining midge Featured Creatures University of Florida Institute of Food and Agricultural Sciences Diptera info Images Retrieved from https en wikipedia org w index php title Chironomidae amp oldid 1213239469, wikipedia, wiki, book, books, library,

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