Revision of the New World Ceratoculicoides Wirth & Ratanaworabhan (Diptera, Ceratopogonidae, Ceratopogonini)

  • Andrew Fasbender Rhithron Associates Inc., 33 Fort Missoula Road, Missoula, MT 59847, USA
Keywords: Biting midges, predaceous midges, phylogeny, Nearctic, Neotropical


The New World species of the genus Ceratoculicoides Wirth & Ratanaworabhan are described, illustrated and keyed in both sexes. Ceratoculicoides borkenti sp. nov., C. confusus sp. nov., C. grogani sp. nov., C. pacificus sp. nov. and C. propinquus sp. nov. are described, with C. confusus being the first record of the genus from South America (Colombia). Ceratoculicoides blantoni is a junior synonym of C. virginianus and the identity of C. longipennis remains unclear as the male morphospecies associated in taxonomic literature with the female type series is not conspecific. A morphological phylogeny of all extant species in the genus is presented and characters discussed. The moravicus species group is recognized for a clade of species with the lateral margins of the aedeagus straight or concave.


Borkent A. 1992. A new key to some genera of Ceratopogonini in the Holarctic (Diptera: Cerato-pogonidae). Entomologica Scandinavica 22: 433–436.

Borkent A. 1995. Biting Midges in the Cretaceous Amber of North America (Diptera: Ceratopogonidae). Backhuys Publishers, Leiden.

Borkent A. 2014. The pupae of the biting midges of the world (Diptera: Ceratopogonidae), with a generic key and analysis of the phylogenetic relationships between genera. Zootaxa 3879 (1): 1–327.

Borkent A. 2017. Ceratopogonidae. In: Kirk-Spriggs A.H. & Sinclair B.J. (eds) Manual of Afrotropical Diptera. Volume 2. Nematocerous Diptera and lower Brachycera. Suricata 5: 733–814.

Borkent A. & Dominiak P. 2020. Catalog of the biting midges of the world (Diptera: Ceratopogonidae). Zootaxa 4787 (1): 1–377.

Borkent A. & Grogan W.L. 2009. Catalog of the New World biting midges north of Mexico (Diptera: Ceratopogonidae). Zootaxa 2273 (1): 1–48.

Borkent A. & Wirth W.W. 1997. World species of biting midges (Diptera: Ceratopogonidae). Bulletin of the American Museum of Natural History 233: 1–257.

Borkent A., Spinelli G.R. & Grogan W.L. 2009. Ceratopogonidae (biting midges, purrujas). In: Brown B.V., Borkent A., Cumming J.M., Wood D.M., Woodley N.E. & Zumbado M.A. (eds) Manual of Central American Diptera. Volume 1: 407–435. NRC Research Press, Ottawa.

Borkent A., Brown B.V., Adler P.H., Amorim D.D., Barber K., Bickel D., Boucher S., Brooks S.E., Burger J., Burington Z.L., Capellari R.S., Costa D.N.R., Cumming J.M., Curler G., Dick C.W., Epler J.H., Fisher E., Gaimari S.D., Gelhaus J., Grimaldi D.A., Hash J., Hauser M., Hippa H., Ibáñez-Bernal S., Jaschhof M., Kameneva E.P., Kerr P.H., Korneyev V., Korytkowski C.A., Kung G., Kvifte G.M., Lonsdale O., Marshall S.A., Mathis W.N., Michelsen V., Naglis S., Norrbom A.L., Paiero S., Pape T., Pereira-Colavite A., Pollet M., Rochefort S., Rung A., Runyon J.B., Savage J., Silva V.C., Sinclair B.J., Skevington J.H., Stireman J.O., Swann J., Vilkamaa P., Wheeler T., Whitworth T., Wong M., Wood D.M., Woodley N., Yau T., Zavortink T.J. & Zumbado M.A. 2018. Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest: Why inventory is a vital science. Zootaxa 4402 (1): 53–90.

Cumming J.M. & Wood D.M. 2017. Adult morphology and terminology. In: Kirk-Spriggs A.H. & Sinclair B.J. (eds) Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4: 89–133.

Delécolle J. & Schiegg K. 1998. Contribution à l'étude des cératopogonidés de Suisse. Révision et redescription des espèces paléarctiques du genre Ceratoculicoides Wirth & Ratanaworabhan, 1971 (Diptera, Nematocera). Bulletin de la Société entomologique de France 103: 273–286.

Downes J.A. & Wirth W.W. 1981. Ceratopogonidae. In: McAlpine J.F., Peterson B.V., Shewell G.E., Teskey H.J., Vockeroth J.R. & Wood D.M. (eds) Manual of Nearctic Diptera. Vol. 1: 393–421. Research Branch Agriculture Canada, Ottawa.

Goloboff P.A. & Catalano S.A. 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32: 221–238.

Goloboff P.A., Farris J.S. & Nixon K.C. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24: 774–786.

Gosseries J. 1989. Replacement of some junior primary homonyms in the Diptera. Insect Nomenclature 1: 1–4.

Grogan W.L. & Borkent A. 1992. Sinhalohelea, a new genus of predaceous midge from Sri Lanka (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 94: 314–319.

Hadley A. 2008. CombineZP. Available at [archived]

Havelka P. 1976. Limnologische und systematische Studien an Ceratopogoniden (Diptera: Nematocera). Beiträge zur Entomologie 26: 211–305.

Havelka P. 1980. Zwei neue Gnitzen (Diptera: Ceratopogonidae) aus dem Annaberger Back bei Bonn. Decheniana 133: 86–92.

Hogue C.L. & Bedoya Ortiz E. 1989. The net-winged midge fauna (Diptera: Blephariceridae) of Antioquia Department, Colombia. Contributions in Science, Natural History Museum of Los Angeles County 413: 1–57.

Huerta H. & Borkent A. 2005. A new species and first record of Ceratoculicoides Wirth & Ratana-worabhan from the Neotropical region and new species and records of Brachypogon Kieffer from Mexico (Diptera: Ceratopogonidae). Folia Entomológica Mexicana 44: 111–119.

ICZN. 1999. International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London.

Knoz J. 1987. Description of a new species of Ceratoculicoides (Diptera, Ceratopogonidae) from Czechoslovakia and a review of the genus. Acta Entomologica Bohemoslovaca 84: 388–392.

Letunic I. & Bork P. 2019. Interactive Tree Of Life (iTOL) v4: recent updates and new developments. Nucleic Acids Research 47 (W1): gkz239.

Maddison W.P. & Maddison D.R. 2019. Mesquite: a modular system for evolutionary analysis. Ver. 3.61. Available from [accessed 17 May 2023]

McKie B.G. & Cranston P.S. 2005. Size matters: systematic and ecological implications of allometry in the responses of chironomid midge morphological ratios to experimental temperature variations. Canadian Journal of Zoology 83: 553–568.

Remm H. 1967. On the fauna of Ceratopogonidae (Diptera) of the Caucasus [in Russian]. Tartu Riikliku Ulikooli Toimetised 194: 3–37.

Stur E. & Ekrem T. 2011. Exploring unknown life stages of Arctic Tanytarsini (Diptera: Chironomidae) with DNA barcoding. Zootaxa 2743 (1): 27–39.

Szadziewski R. 1988. Biting midges (Diptera, Ceratopogonidae) from Baltic amber. Polskie Pismo Entomologiczne 57: 3–283.

Szadziewski R. & Knoz J. 2002. New synonyms of European biting midges (Diptera: Ceratopogonidae). Annales Zoologici 52: 249–251.

Szadziewski R., Krzywiński J. & Giłka W. 1997. Diptera Ceratopogonidae, biting midges. In: Nilsson A.N. (ed.) Aquatic Insects of North Europe–A Taxonomic Handbook: 244–263. Apollo Books, Stenstrup, Denmark.

Wilkening A.J., Kline D.L. & Wirth W.W. 1985. An annotated checklist of the Ceratopogonidae (Diptera) of Florida with a new synonymy. The Florida Entomologist 68: 511–537.

Wirth W.W. 1951. New species and records of Virginia Heleidae. Proceedings of the Entomological Society of Washington 53: 313–326.

Wirth W.W. 1952. The Heleidae of California. University of California Publications in Entomology 9: 95–266.

Wirth W.W. 1965. Family Ceratopogonidae. In: Stone A., Sabrosky C.W., Wirth W.W., Foote R.H. & Coulson J.R. (eds) A Catalog of the Diptera of America North of Mexico: 121–142. U.S. Government Printing Office, Washington D.C.

Wirth W.W. & Grogan W.L. 1988. The predaceous midges of the world (Diptera: Ceratopogonidae; tribe Ceratopogonini). Flora and Fauna Handbook 4: 1–160.

Wirth W.W. & Ratanaworabhan N.C. 1971. Ceratoculicoides, a new genus related to Ceratopogon Meigen (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 73: 170–177.

How to Cite
Fasbender, A. (2023). Revision of the New World Ceratoculicoides Wirth & Ratanaworabhan (Diptera, Ceratopogonidae, Ceratopogonini). European Journal of Taxonomy, 875, 159-202.
Research article