Chironomidae

Chironomidae
Male Chironomus plumosus
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Diptera
Suborder:	Nematocera
Infraorder:	Culicomorpha
Superfamily:	Chironomoidea
Family:	Chironomidae
Genera
See text

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.

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 10000 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 Belgica antarctica, the largest terrestrial animal of Antarctica.

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 macrozoobenthos 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".^[7] Their ability to capture oxygen is further increased by their making undulating movements.^[8]

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 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.^[9]

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 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.^[10] Contemporary specimens are used by forensic entomologists as medico-legal markers for the postmortem interval assessment.^[11]

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.^[12]

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^[13]). These larvae can also withstand other external stresses including ionizing radiation.^[14] 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.^[14] 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.^[14] 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.^[15][16] Most species belong to Chironominae, Orthocladiinae, and Tanypodinae. Diamesinae, Podonominae, Prodiamesinae, and Telmatogetoninae are medium size 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.

Chironomidae larva showing the characteristic red color, about 40× magnification: The head is towards the upper left, just out of view

Aagaardia Sæther, 2000 Abiskomyia Edwards, 1937 Ablabesmyia Johannsen, 1905 Acalcarella Acamptocladius Brundin, 1956 Acricotopus Kieffer, 1921 Aedokritus Aenne Afrochlus Afrozavrelia Harrison, 2004[17] Allocladius Allometriocnemus Allotrissocladius Alotanypus Roback, 1971 Amblycladius Amnihayesomyia Amphismittia Anaphrotenia Anatopynia Johannsen, 1905 Ancylocladius Andamanus Antillocladius Sæther, 1981 Anuncotendipes Apedilum Townes, 1945 Aphrotenia Aphroteniella Apometriocnemus Sæther, 1984 Apsectrotanypus Fittkau, 1962 Archaeochlus Arctodiamesa Makarchenko, 1983[18] Arctopelopia Fittkau, 1962 Arctosmittia Asachironomus Asclerina Asheum Australopelopia Austrobrillia Austrochlus Austrocladius Axarus Roback 1980 Baeoctenus Baeotendipes Kieffer, 1913 Bavarismittia Beardius Beckidia Sæther 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 & Kirschbaum 1990 Cardiocladius Kieffer, 1912 Chaetocladius Kieffer, 1911 Chasmatonotus Chernovskiia Sæther 1977 Chilenomyia Chirocladius Chironomidae (genus) Chironominae Chironomini Chironomus Meigen, 1803 Chrysopelopia Cladopelma Kieffer, 1921 Cladotanytarsus Kieffer, 1921 Clinotanypus Kieffer, 1913 Clunio Haliday, 1855 Coelopynia Coelotanypus Coffmania Collartomyia Colosmittia Compteromesa Sæther 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 Sæther 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 & ZavÍel, 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, Sæther & Andrade-Morraye, 2005 Hahayusurika Halirytus Halocladius Hirvenoja, 1973 Hanochironomus Hanocladius Harnischia Kieffer, 1921

Harrisius Harrisonina Hayesomyia Murray & Fittkau, 1985 Heleniella Gouin, 1943 Helopelopia Roback, 1971 Henrardia Heptagyia Heterotanytarsus Spärck, 1923 Heterotrissocladius Spärck, 1923 Hevelius Himatendipes Hirosimayusurika Hudsonimyia Roback, 1979[19] 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 & Krüger, 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 & Bause, 1913 Lepidopelopia Lepidopodus Lerheimia Limaya Limnophyes Eaton, 1875 Lindebergia Linevitshia Lipiniella Shilova 1961 Lipurometriocnemus Lithotanytarsus Litocladius Andersen, Mendes & Sæther 2004 Ljungneria Lobodiamesa Lobomyia Lobosmittia Lopescladius Lunditendipes Lyrocladius Mendes & 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 Nandeca 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 Oleia Oliveridia Sæther, 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 & 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 & Bause, 1913 Paratendipes Kieffer, 1911 Paratrichocladius Thienemann, 1942 Paratrissocladius ZavÍel, 1937

Parochlus Enderlein, 1912 Parorthocladius Thienemann, 1935 Parvitergum Paucispinigera Pelomus' Pentaneura Pentaneurella Pentaneurini Pentapedilum Petalocladius Phaenopsectra Kieffer, 1921 Physoneura Pirara Platysmittia Sæther, 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 Sæther, 1969 Pterosis Qiniella Reissmesa Rheochlus Rheocricotopus Brundin, 1956 Rheomus Rheomyia Rheopelopia Fittkau, 1962 Rheosmittia Brundin, 1956 Rheotanytarsus Thienemann & Bause, 1913 Rhinocladius Riethia Robackia Sæther, 1977 Saetheria Jackson, 1977 Saetheriella Halvorsen, 1982[20] Saetherocladius Saetherocryptus Saetheromyia Saetherops Sasayusurika Schineriella Murray & Fittkau, 1988 Semiocladius Setukoyusurika Seppia Sergentia Kieffer, 1922 Shangomyia Shilovia Skusella Skutzia Smittia Holmgren, 1869 Stackelbergina Stelechomyia Stempellina Thienemann & 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 & Remmert, 1957 Thienemannia Kieffer, 1909 Thienemanniella Kieffer, 1911 Thienemannimyia Fittkau, 1957 Thienemanniola Tobachironomus Tokunagaia Sæther, 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 Sæther, 1982 Usambaromyia Andersen & Sæther, 1994[21] 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 Zuluchironomus

References

1. Armitage, P. D.; Cranston, P. S.; Pinder, L. C. V. (1995). The Chironomidae: biology and ecology of non-biting midges. London: Chapman & Hall. ISBN 0-412-45260-X.
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. 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.
8. 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: 61–67. doi:10.1007/BF00014132.
9. A. Ali. 1991. Perspectives on management of pestiferous Chironomidae (Diptera), an emerging global problem. Journal of the American Mosquito Control Association 7: 260-281.
10. 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.
11. 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.
12. Kaiser, Tobias S.; Poehn, Birgit; Szkiba, David; Preussner, Marco; Sedlazeck, Fritz J.; Zrim, Alexander; Neumann, Tobias; Nguyen, Lam-Tung; Betancourt, Andrea J. "The genomic basis of circadian and circalunar timing adaptations in a midge". Nature. doi:10.1038/nature20151.
13. 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.
14. 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.
15. J.H. Epler. 2001. Identification manual for the larval Chironomidae (Diptera) of North and South Carolina. North Carolina Department of Environment and Natural Resources.
16. 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.
17. Ekrem, Torbjørn. "Systematics and biogeography of Zavrelia, Afrozavrelia and Stempellinella (Diptera: Chironomidae)". Retrieved 2009-04-30.
18. 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. Retrieved 2009-04-03.
19. Caldwell, Broughton A.; Soponis, Annelle R. (1982). "Hudsonimyia Parrishi, a New Species of Tanypodinae (Diptera: Chironomidae) from Georgia" (PDF). The Florida Entomologist. Lutz, FL, USA: Florida Entomological Society. 65 (4): 506–513. doi:10.2307/3494686. ISSN 0015-4040. JSTOR 3494686. Retrieved 2009-04-20.
20. Halvorsen, Godtfred A. (1982). "Saetheriella amplicristata gen. n., sp. n., a new Orthocladiinae (Diptera: Chironomidae) from Tennessee". Aquatic Insects. Taylor & Francis. 4 (3): 131–136. doi:10.1080/01650428209361098. ISSN 1744-4152.
21. Andersen, Trond; Sæther, Ole A. (January 1994). "Usambaromyia nigrala gen. n., sp. n., and Usambaromyiinae, a new subfamily among the Chironomidae (Diptera)". Aquatic Insects. Taylor & Francis. 16 (1): 21–29. doi:10.1080/01650429409361531. ISSN 1744-4152.

External links

• Media related to Chironomidae at Wikimedia Commons
• 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
• Chironomidae at Australian Faunal Directory
• "Hydrilla tip mining midge". Featured Creatures. University of Florida Institute of Food and Agricultural Sciences.
• Diptera.info Images