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Reproduction

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Adult non-biting midges often form mating swarms, either in the air near oviposition sites, or "skating" on the surface of water. These swarms are composed mostly of males, and may serve to attract females.

In most species, eggs are laid in gelatinous masses on the water surface or on emergent vegetation. In some species, females lay their eggs in or under the water. Adult chironomids usually only live for a few days or weeks, and so reproduction is a single concerted effort. Most species breed seasonally. A very few species are reported to be parthenogenic, most have male and female adults

Key Reproductive Features: semelparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); parthenogenic ; sexual ; fertilization (Internal ); oviparous

No male investment. Female investment is in provisioning eggs and producing a protective gel mass for them.

Parental Investment: pre-fertilization (Provisioning)

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Life Cycle

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Like all flies, the Chironomidae are holometabolous, and undergo metamorphosis in their life cycle. Adult females lay eggs in aquatic habitats. The larvae that hatch from these are often planktonic in their first instar, floating in the water column and feeding on microscopic particles in the water. After their first molt, larvae of most species descend to the bottom and remain benthic through the rest of the larval stage (usually four instars). The larvae transforms into a pupa, which often stays within a shelter or cocoon while it transforms into an adult. When it's time to emerge, the pupa swims to the surface, and the adult pulls itself out of its old skin.

Development - Life Cycle: metamorphosis

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Comprehensive Description

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The Chironomidae is a large and diverse family of flies. They are commonly known as "non-biting midges." There are over 20,000 species known world-wide, including 2,000 in the Nearctic. Adult midges are relatively small (1-20 mm long), with narrow bodies and long legs. They are often confused with mosquitos, but no members of this family are blood-feeders (hence the "non-biting" part of the common name). Adults, if they feed at all, feed on nectar or similar substances. Midge larvae are nearly all aquatic or sub-aquatic, and are a very important part of many freshwater ecosystems. Both in numbers and in diversity, they are often the largest group of primary consumers in these systems. Species of Chironomidae can be found in an enormous variety of aquatic habitats, from brackish estuaries to pools in tree-holes, and from low-oxygen lake sediments to fast-flowing mountain streams.

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Associations

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Chironomids are the most diverse and abundant macroinvertebrates in most of the aquatic ecosystems they inhabit (and they inhabit most aquatic ecosystems). Most natural ponds, lakes and streams are home to 50-100 different species of non-biting midges. Collectively, they play a vital role in freshwater ecosystems as primary consumers. They harvest an enormous amount of energy from detritus and are one of the major supports for animal communities in these systems.

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Trophic Strategy

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The many thousands of chironomid species have many different feeding habits. Most species feed on small particles of organic debris, but the size of particles varies, some shred bits of dead wood and leaves, some gather smaller particles, some even filter tiny particles suspended in the water. Some of these detritivores also collect algae cells, and some species are herbivores, specialize in feeding on algae. Other herbivores are "miners" tunneling in larger vascular plants. There are some fungivore chironomids as well, eating spores and grazing on hyphae. A few species are simple predators, often attacking other chironomid species.

Primary Diet: detritivore

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Distribution

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Species of chironomid midges are found in moist or wet habitats in all major landmasses of the world, including Antarctica, and most islands.

Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native ); antarctica (Native ); oceanic islands (Native )

Other Geographic Terms: cosmopolitan

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Habitat

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Midge larvae occur in all kinds of benthic freshwater habitats, including the bottoms of streams, rivers, lakes, ponds, and temporary pools, also wetlands such as marshes and swamps. Some breed in isolated damp habitats such as tree-holes, pitcher plants, patches of moist soil, even dung pats. The "blood midges" or "bloodworms" are species of midges with hemoglobin in their hemolymph, which allows them to survive in low-oxygen (and often heavily-polluted) habitats. Adults rarely disperse far from the larval habitat.

Habitat Regions: temperate ; tropical ; polar ; terrestrial ; freshwater

Terrestrial Biomes: forest ; rainforest

Aquatic Biomes: benthic ; lakes and ponds; rivers and streams; temporary pools; brackish water

Wetlands: marsh ; swamp ; bog

Other Habitat Features: riparian ; estuarine ; intertidal or littoral

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Life Expectancy

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Lifespan varies greatly between and within species in the Chironomidae. Individual growth and development rates are strongly influenced by temperature and other environmental factors. Many temperate species live for a year, surviving the winter as larvae. Some species are known to complete entire life-cycles in a few weeks, if temperatures are warm and food is abundant.

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Morphology

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Adults are small (1-20 mm long, most less than 10 mm), slim, long-legged flies. They resemble, and are often confused with, mosquitoes (Culicidae), but unlike mosquitoes, they do not bite, and have no scales on their wings. Many species rest on their hind two pairs of legs, and hold their forelegs out in front of them. In most species, adult males have plumose antennae that are much larger than the females (these are probably used to locate females). Most species are dark-colored, usually brown or black.

Larvae are elongate and cylindrical, with distinct segmentation and a hard sclerotized head capsule that cannot be retracted into the body. They have no true legs, but do have a pair of unjointed "prolegs" on the first segment of the thorax. The presences of this pair of prolegs, the absence of true legs, and the structure of the head are good distinguishing marks for identifying larvae in the Chironomidae. Color varies widely among larvae, most are tan or brown, but some are whitish, some are green. Larvae of a number of species in the subfamily Chironominae have the hemoglobin in their circulatory fluid, which helps them survive in low-oxygen habitats. These larvae are pinkish or red when alive, and are often called "blood midges."

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: sexes shaped differently

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Associations

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Non-biting midges are so abundant in so many freshwater habitats that practically every kind of predator in these habitats feeds on them at some stage of their life cycle. Midges try to avoid predation by limiting their activity during daylight, and larvae and pupae take refuge in tunnels that they build in sediment. Many species are cryptically colored.

Anti-predator Adaptations: cryptic

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Chironomidae

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

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. An example of mosquito-resembling species is Tokunagayusurika akamusi.

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

Common names and biodiversity

This is a large taxon of insects; some estimates of the species numbers suggest well over 10 000 world-wide.[1] 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",[2] "muffleheads",[3] "Canadian soldiers",[4] or "American soldiers"[5] in various regions of the Great Lakes area. They have been called "blind mosquitoes" or "chizzywinks" in Florida.[6] 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.[7][8]

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

Larval stages of the Chironomidae can be found in almost any aquatic or semiaquatic habitat, including treeholes, bromeliads, rotting vegetation, soil, and in sewage and artificial containers. They form an important fraction of the macro zoobenthos of most freshwater ecosystems. They are often associated with degraded or low-biodiversity ecosystems because some species have adapted to virtually anoxic conditions and are dominant in polluted waters. Larvae of some species are bright red in color due to a hemoglobin analog; these are often known as "bloodworms".[9] Their ability to capture oxygen is further increased by their making undulating movements.[10]

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.[1]

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 apply the resources obtained in feeding differently. 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, and in particular 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.[1]

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.[11] 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.[12]

Ecology

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. So do predatory water beetles in families such as the Dytiscidae and Hydrophilidae. The flying midges 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.

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.[13] Contemporary specimens are used by forensic entomologists as medico-legal markers for the postmortem interval assessment.[14]

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.[15]

Anhydrobiosis and stress resistance

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[16]). These larvae can also withstand other external stresses including ionizing radiation.[17] 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.[17] 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.[17] The Rad51 protein plays a key role in homologous recombination, a process required for the accurate repair of DNA double-strand breaks.

Subfamilies and genera

The family is divided into 11 subfamilies: Aphroteniinae, Buchonomyiinae, Chilenomyinae, Chironominae, Diamesinae, Orthocladiinae, Podonominae, Prodiamesinae, Tanypodinae, Telmatogetoninae, and Usambaromyiinae.[18][19] 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.

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

C. Annularius

Chironomus annularius (commonly known as bayflies or muffleheads) is a species of non-biting midge in the family Chironomidae. It is usually found in regions with bodies of fresh water but can be found in almost every environment. It tends to form "hotspots" around specific areas. The species is distinguished by the size of its chromosomes and the lack of a proboscis.

References

  1. ^ a b c Armitage, P. D.; Cranston, P. S.; Pinder, L. C. V. (1995). The Chironomidae: biology and ecology of non-biting midges. London: Chapman & Hall. ISBN 978-0-412-45260-4.
  2. ^ "Muckleheads" from Andre's Weather World (Andre Bernier, staff at WJW-TV), June 2, 2007.
  3. ^ "You don't love muffleheads, but Lake Erie does", Sandusky Register, May 24, 2010.
  4. ^ Galbincea, Barb, "Canadian Soldiers Invade Rocky River", The Plain Dealer, Cleveland.com, June 18, 2014, accessed June 3, 2015.
  5. ^ "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.
  6. ^ Chizzywinks are Blind Mosquitos by Dan Culbert of the University of Florida, August 17, 2005
  7. ^ 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.
  8. ^ Luke Sandro & Juanita Constible. "Antarctic Bestiary — Terrestrial Animals". Laboratory for Ecophysiological Cryobiology, Miami University. Archived from the original on 23 December 2008. Retrieved December 9, 2008.
  9. ^ W.P. Coffman and L.C. Ferrington Jr. 1996. Chironomidae. pp. 635-754. In: R.W. Merritt and K.W. Cummins, eds. An Introduction to the Aquatic Insects of North America. Kendall/Hunt Publishing Company.
  10. ^ Int Panis, L; Goddeeris, B.; Verheyen, R (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.
  11. ^ McConnaughey, Janet (June 19, 2019). "The Swarm: Billions of skeeter lookalikes plague New Orleans". Associated Press. Retrieved September 5, 2019.
  12. ^ A. Ali. 1991. Perspectives on management of pestiferous Chironomidae (Diptera), an emerging global problem. Journal of the American Mosquito Control Association 7: 260-281.
  13. ^ 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.
  14. ^ 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.
  15. ^ Kaiser, Tobias S.; Poehn, Birgit; Szkiba, David; Preussner, Marco; Sedlazeck, Fritz J.; Zrim, Alexander; Neumann, Tobias; Nguyen, Lam-Tung; Betancourt, Andrea J. (2016). "The genomic basis of circadian and circalunar timing adaptations in a midge". Nature. 540 (7631): 69–73. doi:10.1038/nature20151. PMC 5133387. PMID 27871090.
  16. ^ 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.
  17. ^ 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. doi:10.1371/journal.pone.0014008. PMC 2982815. PMID 21103355.
  18. ^ 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.
  19. ^ 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.
  20. ^ Ekrem, Torbjørn. "Systematics and biogeography of Zavrelia, Afrozavrelia and Stempellinella (Diptera: Chironomidae)". Archived from the original on 2009-03-18. Retrieved 2009-04-30.
  21. ^ 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. Retrieved 2009-04-03.
  22. ^ 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.
  23. ^ 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.
  24. ^ Halvorsen, Godtfred A. (1982). "Saetheriella amplicristata gen. n., sp. n., a new Orthocladiinae (Diptera: Chironomidae) from Tennessee". Aquatic Insects. 4 (3): 131–136. doi:10.1080/01650428209361098. ISSN 1744-4152.
  25. ^ 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. doi:10.1080/01650429409361531. ISSN 1744-4152.

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Chironomidae: Brief Summary

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

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. An example of mosquito-resembling species is Tokunagayusurika akamusi.

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

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