NRE516 - Aquatic Entomology
General Lab Notes on the Aquatic Diptera

Notes on Morphology/Taxonomy
Synonomies
List of Taxa Presented
Useful References

Notes on Morphology/Taxonomy

Nearctic Diptera includes 108 families with an estimated 25,000 to 30,000 species. Only 29 families include aquatic members. A complete description of all Nearctic Diptera is found in McAlpine et al. (1981, 1987) from which the following discussion is derived. Two suborders are presently recognized, Nematocera and Brachycera, which differ among the larvae primarily in terms of reduction of the head. Most Nematocera have a well developed, fully exposed head capsule with mandibles usually bearing teeth and operating in a horizontal or oblique plane (i.e. eucephalic head). An exception to this is the Tipulidae (see below) which has a head capsule reduced posteriorly and partially retracted into the thorax. The tipulid mandibles vary from stout and toothed, adapted for chewing, to slender and sickle-shaped, for piercing. But they are almost always opposed to each other and move in a horizontal plane. Most Brachycera (Orthorrhapha), on the other hand, have a head capsule, more or less, reduced posteriorly and partially retracted within the thorax, with sickle-shaped mandibles operating in a vertical plane (i.e. a hemicephalic head). Some of the Brachycera (Cyclorrhapha or infraorder Muscamorpha) have further reduction and retraction of the head capsule into the thorax combined with the development of an internal pharyngeal skeleton (an acephalic head).

The number of recognizable body segments in the larvae is most commonly 12 in Nematocera and 11 in the Brachycera. Of these three, three are always thoracic and the remainder abdominal. Any variance from these numbers is to decrease the number of abdominal segments in the Nematocera and to increase it in the Brachycera. Critical to the identification of the families of Diptera larvae is: 1) the extent of development of the head and whether head is exposed or retracted into the thorax, 2) body shape, 3) the location and number of prolegs, 4) the presence the location of spiracles, and 5) the presence of terminal appendages which sometimes serve a respiratory function. Generic identification frequently requires removal of the head (esp. eucephalic dipterans) which are then cleared and pressed on a slide to expose the mouth structure and musculature. In some cases larval keys do not exist due to the lack of consistent diagnostic features and limited description of various genera.

Synonomies

Many current names for Diptera trace their origins back to Meigen's 1803 work. However, Hendel discovered in 1908 that Meigen published in 1800 an obscure paper for many of the same taxa, but had used different names. Many American and Russian authors followed Hendel, and based on the rule of priority, used Meigen's 1800 names. However, most other dipterists had been using, and continued to use, the Meigen names of 1803. Despite some bitter infighting among researchers over the issue (particularly that of Henry Townes), the issue was not resolved until the 1960's when the governing body for animal taxonomy, the ICZN, ruled to suppress the Meigen 1800 names. They based this ruling principally because of common usage, but also because Meigen had not described and designated type species in association with any of the 1800 names, whereas in 1803 there were species descriptions, and further Meigen himself did not subsequently use the 1800 names.

Ceratopogonidae = Helidae
Athericidae = Rhagionidae
Chaoboridae = Culicidae (in part)
Culicidae
    Culiseta = Theobaldia
    Toxorhynchites = Megarhinus
Tipulidae
    Leptotarsus = Longurio
    Erioptera includes Trimicra
    Hexatoma includes Eriocera
    Limonia includes Geranomyia
Chironomidae = Tendipedidae
    Tanypus = Pelopia
    Thalassosmittia = Sauderia
    Chironomus = Tendipes
    Dicrotendipes = Limnochironomus
    Lauterborniella = Zavreliella
    Paralauterborniella = Apedilum
    Phaenopsectra = Segentia, Lenzia
    Tanytarsus = Calpsectra
Ptychopteridae = Liriopeidae
Stratiomyiidae
    Odontomyia = Eulalia
    Oxycera = Hermione
    Stratiomys = Stratiomyia
Ephydridae
    Hydropyrus = Ephydra
Sciomyzidae = Tetanoceridae = Tetanoceratidae
Scathophagidae = Scatophagidae = Scopeumatidae = Cordiluridae = Anthomyiidae (in part)
Muscidae = Anthomyiidae (in part)
Sarcophagidae
    Fletcherimyia = Blaesoxipha

List of Taxa Presented

List of Chironomidae Subfamilies and Tribes

Useful References

General Diptera

Friedrich, M. and D. Tautz. 1997. Evolution and phylogeny of the Diptera: a molecular phylogenetic analysis using 28s rDNA sequences. Systematic Biology 46:674-698

McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood (Coords). 1981. Manual of Nearctic Diptera, Vol. 1. Res. Branch. Agric. Can. Monogr. 27. 674 pp.

McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth, and D. M. Wood (Coords). 1987. Manual of Nearctic Diptera, Vol. 2. Res. Branch. Agric. Can. Monogr. 28.

Michelsen, V. 1996. Neodiptera: new insights into the adult morphology and higher level phylogeny of Diptera (Insecta). Zoological Journal of the Linnean Society 117:71-102.

Miller, B.R., M.B. Crabtree and H.M. Savage. 1997. Phylogenetic relationships of the Culicomorpha inferred from 18s and 5.8s ribosomal DNA sequences (Diptera, Nematocera). Insect Molecular Biology 6(2):105-114.

Oosterbroek, P., and G. Courtney. 1995. Phylogeny of the nematocerous families of Diptera (Insecta). Zoological Journal of the Linnean Society 115:267-311.

Pawlowski, J., R. Szadziewski, D. Kmieciak, J. Fahrni, G. Bittar. 1996. Phylogeny of the infraorder Culicomorpha (Diptera: Nematocera) based on 28S RNA gene sequences. Systematic Entomology 21(2): 167-178.

Papp, L. and B. Darvas (Eds). 1997. Contributions to a manual of Palaearctic Diptera. Volume 2. Science Herald, Budapest.

Skarlato, O. A., ed. 1985. Systematics of Diptera (Insecta). Ecological and morphological principles. Oxonian Press, New Delhi, India.

Stark, J., J. Bonacum, J. Remsen, and R. DeSalle. 1999. The evolution and development of dipteran wing veins: a systematic approach. Annual Review of Entomology 44.

Yeates, D.K. and B.M. Wiegmann. 1999. Congruence and controversy: toward a higher-level phylogeny of Diptera. Annual Review of Entomology 44:397-428.

Nematocera
Blephariceridae and Deuterophlebiidae

Courtney, G. W. 1991. Phylogenetic analysis of the Blephariceromorpha, with special reference to mountain midges (Diptera: Deuterophlebiidae). Systematic Entomology 16:137-172.

Courtney, G.W. 2000. A.1. Family Blephariceridae. In L. Papp & B. Darvas (editors). Contributions to a Manual of Palaearctic Diptera. Appendix. Science Herald, Budapest. [in press]

Hogue, C. L. 1987. Blephariceridae. Series editor: G. C. D. Griffiths, Flies of the Nearctic Region, Volume II (Archaeodiptera and Oligoneura), Part 4. E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), Stuttgart.

Ceratopogonidae

Borkent, A. and W.W. Wirth. 1997. World Species of Biting Midges (Diptera: Ceratopogonidae). Bulletin of the American Museum of Natural History 233, 257 pp.

Wilkening, A. J., D. L. Kline, and W. W. Wirth. 1985. An annotated checklist of the Ceratopogonidae (Diptera) of Florida with a new synonymy. Fl. Entomol. 68:511-537.

Wirth, W. W., A. L. Dyce, and B. V. Peterson. 1985. An atlas of wing photographs, with a summary of the numerical characters of the Nearctic species of Culicoides Diptera: Ceratopogonidae). Contributions to the American Entomological Institute 22:1-46.

Chironomidae (those otherwise not cited in Hilsenhoff 1995)

Boesel, M. W., and R. W. Winner. 1980. Corynoneurinae of northeastern United States, with a key to adults and observations on their occurrence in Ohio (Diptera: Chironomidae). Ohio Journal of Science 85:245-262.

Brundin, L. 1956. Zur Systematik der Orthocladiinae (Diptera: Chironomidae). Report of the Institute of Freshwater Research, Drottningholm 37:5-185.

Chernovskii, A. A. 1961. Identification of larvae of the midge family Tendipedidae. Translation E. Lees, ed. K. E. Marshall. National Lending Library for Science and Technology: Boston Spa, Yorkshire, UK.

Coffmam, W. P. 1979. Neglected characters in pupal morphology as tools in taxonomy and phylogeny. Entomogica Scandinavia Supplement 10:37-46.

Cranston, P. S., and D. D. Judd. 1987. Metriocnemus (Diptera: Chironomidae) – an ecological survey and description of a new species. Journal of the New York Entomological Society 95(4):534-546.

___________., and D. R. Oliver. 1988. Aquatic xylophagous Orthocladiinae – systematics and ecology. Spixiana Supplement 14:143-154.

Curry, L. L. 1958. Larvae and pupae of the species of Crytpochironomus (Diptera) in Michigan. Limnology and Oceanography 3:427-442.

Doughman, J. S. 1983. A guide to the larvae of the Nearctic Diamesinae (Diptera: Chironomidae). The genera Boreoheptagyia, Protanypus, Diamesa and Pseudokiefferiella. U.S. Geological Survey, Water-Resources Investigations Report 83-4006:1-58.

___________. 1985. Annotated key to the genera of the tribe Diamesini (Diptera: Chironomidae). Description of the female and immature of Potthastia iberica Tosio, and keys to the known species of Potthastia. University of Alaska IWR (Institute of Water Resources) Series IWR-107.

Epler, J. H.  1995. Identification manual for the larval Chironomidae (Diptera) of Florida, revised edition. Department of Environmental Protection, Florida. Comment: This is available as a PDF document free-of-charge.

Ferrington, L. C. Jr. 1983. Kansas Chironomidae. 3. The Harnischia complex. Technical Publication of the State Biological Survey of Kansas 13:48-62.

Fittkau, E. J., and D. A. Murray. 1988. Bethbilbeckia floridensis: a new genus and species of Macropelopiini from the South Eastern Nearctic. Spixiana Supplement 14:253-259.

Gouin, F. J. 1959. Morpholoyg of the larval head of some Chironomidae (Diptera, nematocera). Smithsonian Miscellaneous Collections 137:175-201.

Jacobsen, R. E. 1992. Descriptions of the larvae of four Nearctic species of Epoicocladius (Diptera: Chironomidae) with a redescription of Epoicocladius ephemerae (Kieffer). Netherlands Journal of Aquatic Ecology 26:145-155.

Kowalyk, H. E. 1985. The larval cephalic setae in the Tanypodinae (Diptera: Chironomidae) and their importance in generic determinations. Canadian Entomologist 117(1):67-106.

Lindeberg, B., and T. Wiederholm. 1979. Notes on the taxonomy of European species of Chironomus (Diptera: Chironomidae). Entomologica Scandinavia Supplement 10:99-116.

Maschwitz, D. E., and E. F. Cook. 2000. Revision of the Nearctic species of the genus Polypedilum Kieffer (Diptera: Chironomidae) in the subgenera P. (Polypedilum) Kieffer and P. (Uresipedilum) Oyewo and Sæther. Bulletin of the Ohio Biological Survey, New Series 12(3).

Mozley, S. C. 1970. Neglected charactersin larval morphology as tools in taxonomy and phylogeny of Chironomidae (Diptera). Entomologica Scandinavia Supplement 10:27-36.

Mozley, S. C. 1971. Maxillary and premental patterns in Chironominae and Orthocladiinae (Diptera, Chironomidae). Canadian Entomologist 103:298-305.

Oliver, D. R. 1959. Some Diamesini (Chironomidae) from the Nearctic and palaearctic. Entomologie Tidskrift 80:48-64.

___________. 1977. Bicintus-group of the genus Cricotopus van der Wulp (Diptera: Chironomidae) in the Nearctic with a description of a new species. Journal of the Fisheries Research Board of Canada 34:98-104.

___________. 1985. Review of Xylotopus Oliver and description of Irisobrilla n. gen. (Diptera: Chironomidae). The Canadian Entomologist 117:1093-1110.

___________., and M. E. Roussel. 1982. The larvae of Pagastia Oliver (Diptera: Chironomidae) with descriptions of three Nearctic species. The Canadian Entomologist 114:849-854.

___________., and M. E. Roussel. 1983. Redescription of Brillia Kieffer (Diptera: Chironomidae) with descriptions of Nearctic species. The Canadian Entomologist 115:257-279.

Pinder, L. C. V. 1986. Biology of freshwater Chironomidae. Annual Review of Entomology 31:1-24.

Roback, S. S. 1971. The adults of the subfamily Tanypodinae (=Pelopiinae) in North America (Diptera: Chironomidae). Monograph of the Academy of Natural Science Philadelphia 17:1-410.

___________. 1986. The immature chironomids of the eastern United States VIII. Pentaneurini – Genus Nilotanypus, with the description of a new species from Kansas. Proceedings of the Academy of Natural Science Philadelphia 138(2):159-209.

___________. 1986. The immature chironomids of the eastern United States VII. Pentaneurini – Genus Monopelopia, with redescription of the male adults and description of some Neotropical material. Proceedings of the Academy of Natural Science Philadelphia 138(2):350-365.

___________. 1987. The immature chironomids of the eastern United States. IX. Pentaneurini – Genus Labrundinia with the description of some Neotropical material. Proceedings of the Academy of Natural Science Philadelphia 139:159-209.

___________., and K. J. Tennessen. 1978. The immature stages of Djalmabastista pulcher [=Procladius (Calotanypus) pulcher (Joh.). Proceedings of the Academy of Natural Science Philadelphia 130:11-20.

Rossaro, B. 1984. Stilocladius Rossaro, 1979 reconsidered, with descriptions of the female and larva of S. montanus Rossaro (Diptera: Chironomidae, Orthocladiinae). Entomologica scandinavica 15:185-191.  

Sæther, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chironomidae). Bulletin of the Fisheries Research Board of Canada 170:1-154.

___________. 1971. Notes on general morphology and terminology of the Chironomidae (Diptera). Canadian Entomologist 103:1237-1260.

___________. 1973. Taxonomy and ecology of three new species of Monodiamesa Kieffer, with keys to Nearctic and Palaearctic species of the genus (Diptera: Chironomidae). Journal of the Fisheries Research Board of Canada  30:665-679.

___________. 1975. Nearctic and Palaearctic Heterotrissocladius (Diptera: Chironomidae).  Bulletin of the Fisheries Research Board of Canada 193.

___________. 1976. Revision of Hydrobaenus, Trissocladius, Zalutschia, Paratrissocladius, and some related genera (Diptera: Chironomidae).  Bulletin of the Fisheries Research Board of Canada 195.

___________. 1977. Taxonomic studies of hicornomidae: nanocladius, Pseudochironomus, and teh Harnischia complex. Bulletin of the Fisheries Research Board of Canada 196. 

___________. 1985. Review of the genus Rheocricotopus Thienemann & Harnisch, 1932, with the description of three new species (Diptera, Chironomidae). Spixiana Supplement 11:59-108.

___________. 1988. Metriocnemus van der Wulp: a new species and a revision of species described by Meigen, Zetterstedt, Stæger, Holmgren, Lundström and Strenzke (Diptera: Chironomidae). Entomologica scandinavica 19:393-430.

___________. 1990. A review of the genus Limnophyes Eaton from the Holarctic and Afrotropical regions (Diptera: Chironomidae, Orthocladiinae). Entomologica scandinavica Supplement 35:1-139.

___________. 2000. Phylogeny of the subfamilies of Chironomidae (Diptera). Systematic Entomology 25:393-403.

___________., and J. E. Sublette. 1983. A review of the genera Doithrix n. gen., Georthocladius Strenzke, Parachaetocladius Wükler, and Pseudorthocladius Goetghebuer (Diptera: Chironomidae, Orthocladiinae). Entomologica scandinavica Supplement 20:1-100.

___________., Ashe, P. & Murray, D. E. 2000. Family Chironomidae. Pp. 113-334 in: Papp, L. and Darvas, B. (eds): Contributions to a Manual of Palaearctic Diptera (with special reference to the flies of economic importance). Vol. 4.  A.6- Science Herald, Budapest.

Soponis, A. R. 1977. A revision of the Nearctic species of Orthocladius (Orthocladius) Van der Wulp (Diptera: Chironomidae). Memoirs of the Entomological Society of Canada, No. 102.

___________. 1990. A revision of the Holarctic species of Orthocladius (Euorthocladius) (Diptera: Chironomidae). Spixiana Supplement 13.

Webb, D. W. 1982. Smittia lasiops (Malloch): a redescription of the adults with a description of the immature stages (Diptera: Chironomidae). Proceedings of the Entomological Society of Washington 84:468-474.

___________. and A. School. 1985. Identification of larvae of European species of Chironomus (Diptera: Chironomidae) by morphological characters. Systematic Entomology 10(3):353-372.

Webb, C. J., A. Scholl, H. M. Ryser. 1985. Comparative morphology of the larval ventromental plates of European species of Chironomus (Diptera: Chironomidae). Systematic Entomology 10(3):373-385.

Culicidae

Wilmot, T. R., D. S. Zeller, and R. W. Merritt. 1992. A key to container-breeding mosquitoes in Michigan (Diptera: Culicidae), with notes on their biology. The Great Lakes Entomologist 25(3):137-148.

Nymphomyiidae

Courtney, G. W.1994. Biosystematics of the Nymphomyiidae (Insecta: Diptera): life history, morphology, and phylogenetic relationships. Smithsonian Contributions to Zoology Number 550.

Simuliidae

Alder, P. H., K. C. Kim. 1986. The black flies of Pennsylvania (Simuliidae, Diptera): Bionomics, Taxonomy, and Distribution. Bull. Penn. State Univ. 856:88 pp.

Fredeen, F. J. H. 1985. The black flies (Diptera: Simuliidae) of Saskatchewan. Sask. Cult. Recr. Mus. Nat. Hist. Contrib. (Regina) 8:41 pp. + 31 plates.

Moulton, J. K. 1998. Reexamination of Simulium (Psilopelmia) Enderlein (Diptera: Simuliidae) of America North of Mexico. Proceedings of the Entomological Society of Washington 100(1):50-71.

____________., and P. H. Adler. 1995. Revision of the Simulium jenningsi species-group (Diptera: Simuliidae). Transactions of the American Entomological Society 121(1-2):1-57.

Peterson, B. V. 1993. The black flies of the genus Simulium, subgenus Psilopelmia (Diptera: Simuliidae), in the contiguous United States. Journal of the New York Entomological Society 101(3):301-390.

Tipulidae

Gelhaus, J. K. 1985. Larvae of the crane fly genus Tipula in North America (Diptera: Tipulidae). The University of Kansas Science Bulletin 53(3):121-182.

Brachycera
Empididae

MacDonald, J. F. 1989. Review of Nearctic Metachela Coquillett, with description of a new species (Diptera: Empididae; Hemerodromiinae). Proceedings of the Entomological Society of Washington 91(4):513-522.

____________. 1993. Review of the genus Neoplasta Coquillett of America north of Mexico (Diptera: Empididae; Hemerodromiinae). Proceedings of the Entomological Society of Washington 95(3):351-376.

____________. 1993. Review of the genus Chelipoda Coquillett of America north of Mexico (Diptera: Empididae; Hemerodromiinae). Proceedings of the Entomological Society of Washington 95(3):327-350.

____________. 1994. Review of the Nearctic species of the genus Chelifera Macquart (Diptera: Empididae; Hemerodromiinae). Proceedings of the Entomological Society of Washington 96(2):236-275.

Ephydridae

Rao, N., and W. N. Mathis. 1995. A revision of the shore-fly genus Rhysophora Cresson (Diptera: Ephydridae). Proceedings of the Entomological Society of Washington 97(3):666-688.

Sciomyzidae

Knutson, L., R. E. Orth, and R. Rozkošný. 1990. New North American Colobaea, with a preliminary analysis of related genera (Diptera: Sciomyzidae). Proceedings of the Entomological Society of Washington 92(3):483-492.

Syrphidae

Rotheray, G. and F. Gilbert. 1999. Phylogeny of Palaearctic Syrphidae (Diptera): evidence from larval stages. Zoological Journal of the Linnean Society 127:1-112.

Thompson, F.C. 1999. A key to the genera of the flower flies (Diptera: Syrphidae) of the Neotropical Region including descriptions of new genera and species and a glossary of taxonomic terms. Contributions on Entomology, International 3(3):321-378.

Tabanidae

Burger, John F. 1995. Catalog of Tabanidae (Diptera) of North America North of Mexico. Contrib. Entomol., International 1(1): 1-100.

Fairchild, Graham B. and John F. Burger. 1994. A Catalog of the Tabanidae (Diptera) of the Americas South of the United States. Memoirs of the American Entomological Institute 55. 249pp.

Teskey, H.J. 1990. The insects and arachnids of Canada. Part 16. The horse flies and deer flies of Canada and Alaska: Diptera: Tabanidae. Research Branch Agriculture Canada Publication 1838: 1-381.

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