Hostname: page-component-586b7cd67f-dsjbd Total loading time: 0 Render date: 2024-11-20T07:26:08.844Z Has data issue: false hasContentIssue false

Biting midges (Diptera: Ceratopogonidae) as indicators of biostratigraphy, ecological reconstructions and identification of amber deposits

Published online by Cambridge University Press:  15 January 2018

Ryszard Szadziewski*
Affiliation:
Department of Invertebrate Zoology and Parasitology, University of Gdańsk, Wita Stwosza 59, 80-308 Gdańsk, Poland. Email: [email protected]

Abstract

Biting midges (Diptera: Ceratopogonidae) are a large family of flies that commononly appear in Lower Cretaceous to Miocene strata, with over 280 fossil species (4.3 % of the family), belonging to 49 genera (26 extant; 23 extinct). Morphological characters used in the identification of fossil genera and species are identical to those used in studies of extant Ceratopogonidae and, as a result, their potential indicative value is reliable. Two relictual extant genera, Leptoconops and Austroconops, reported from Lower Cretaceous Lebanese amber, are at least 125 million years old. Certain ceratopogonid genera are indicative for the Lower Cretaceous, Upper Cretaceous, Eocene or Miocene. A morphological character indicative for the Upper Cretaceous and Cenozoic is macrotrichia on the wing membrane of adults. Indicator species and genera are reviewed for all amber deposits. Eocene Baltic amber contains the best known fauna of biting midges, with 109 named species. Some genera are indicative of aquatic and semiaquatic habitats (predaceous genera, subfamily Ceratopogoninae), forests with rotting trees (Forcipomyia), sandy sea shore habitats (Leptoconops), a cold boreal climate (Ceratopogon) or warm climates (Nannohelea, Austroconops, Leptoconops, Meunierohelea, Metahelea). Females require a protein-rich meal and are well known for feeding on the blood of vertebrates, but many feed on other things, so this information can help with the interpretation of palaeoenvironments. Washingtonhelea taimyrica Szadziewski, 1996, described from Siberian amber, is transferred to the fossil genus Palaeobrachypogon: P.taimyricus (Szadziewski, 1996), comb. nov. For Serromyia alphea, mistakenly redescribed and illustrated from Eocene Bitterfeld amber (= Baltic amber) by Szadziewski (1993), a new name – Serromyiaerrata Szadziewski, nom. nov. – is proposed.

Type
Articles
Copyright
Copyright © The Royal Society of Edinburgh 2018 

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

5. References

Azar, D., Granier, B. & Maksoud, S. 2015. Latest news on the age of the fossiliferous Early Cretaceous amber of Lebanon. World Amber Council 2015, 6568. Gdańsk.Google Scholar
Boesel, M. W. 1937. Order Diptera, Family Chironomidae, In Carpenter, F. M., Folsom, J. W., Essig, E. O., Kinsey, A. C., Brues, C. T., Boesel, M. W. & Ewing, H. E. Insects and arachnids from Canadian amber. University of Toronto Studies, Geological Series 40, 4455. Toronto: The University Library; Published by the Librarian. 62 pp.Google Scholar
Borkent, A. 1995. Biting midges in the Cretaceous amber of North America (Diptera: Ceratopogonidae). Leiden: Ba ckhuys Publishers. 237 pp.Google Scholar
Borkent, A. 1996. Biting midges from Upper Cretaceous New Jersey amber (Ceratopogonidae: Diptera). American Museum Novitates 3159. 29 pp.Google Scholar
Borkent, A. 1997. Upper and Lower Cretaceous biting midges (Ceratopogonidae: Diptera) from Hungarian and Austrian amber and the Koonwarra fossil bed of Australia. Stuttgarter Beiträge zur Naturkunde, Series B 249. 10 pp.Google Scholar
Borkent, A. 2000a. Biting midges (Diptera: Ceratopogonidae) from Lower Cretaceous Lebanese amber with a discussion of the diversity and patterns found in other ambers. In Grimaldi, D. (ed.) Studies on fossils in amber, with particular reference to the Cretaceous of New Jersey, 355451. Leiden: Backhuys Publishers. 498 pp.Google Scholar
Borkent, A. 2000b. Further biting midges (Diptera: Ceratopogonidae) from Upper Cretaceous New Jersey amber. In Grimaldi, D. (ed.) Studies on fossils in amber, with particular reference to the Cretaceous of New Jersey, 453–72. Leiden: Backhuys Publishers. 498 pp.Google Scholar
Borkent, A. 2001. Leptoconops (Diptera: Ceratopogonidae), the earliest extant lineage of biting midge, discovered in 120–122 milion-year-old Lebanese amber. American Museum Novitates 3328. 11 pp.Google Scholar
Borkent, A. 2012. Further biting midges (Diptera: Ceratopogonidae) in Canadian Cretaceous amber. Canadian Entomologist 144, 758–66.Google Scholar
Borkent, A. 2016. World species of biting midges (Diptera: Ceratopogonidae). Available from: http://www.inhs.uiuc.edu/cee/FLYTREE/CeratopogonidaeCatalog.pdf (accessed July 2016).Google Scholar
Borkent, A., Coram, R. A. & Jarzembowski, E. A. 2013. The oldest fossil biting midge (Diptera: Ceratopogonidae) from the Purbeck Limestone Group (Lower Cretaceous) of southern Great Britain. Polish Journal of Entomology 82, 273–79.Google Scholar
Borkent, A. & Bissett, B. 1990. A revision of the Holarctic species of Serromyia Meigen (Diptera: Ceratopogonidae). Systematic Entomology 15, 153217.Google Scholar
Borkent, A. & Grogan, W. L. 1995. A revision of the genus Ceratopogon Meigen with a discussion of phylogenetic relationships, zoogeography, and bionomic divergence (Diptera: Ceratopogonidae). Memoirs of the Entomological Society of Washington 15, 1198.Google Scholar
Choufani, J., Azar, D., Perrichot, V., Soriano, C., Tafforeau, P. & Nel, A. 2011. The genus Leptoconops Skuse (Diptera: Ceratopogonidae) in early Cretaceous Charentese amber. Palaeobiodiversity and Palaeoenvironments 91, 285–91.Google Scholar
Choufani, J., Perrichot, V., Girard, V., Garrouste, R., Azar, D. & Nel, A. 2013. Two new biting midges of the modern type from Santonian amber of France (Diptera: Ceratopogonidae). In Azar, D., Engel, M., Jarzembowski, E., Krogmann, L., Nel, A. & Santiago-Blay, J. (eds) Insect Evolution in an Amberiferous and Stone Alphabet. Proceedings of the 6th International Congress on Fossil Insects, Arthropods and Amber, 7395. Leiden: Brill. 210 pp.Google Scholar
Choufani, J., Azar, D. & Nel, A. 2014a. New biting midges from the Cretaceous amber of Lebanon (Diptera: Ceratopogonidae). Annales de la Société Entomologique de France (N.S.) 50, 272–85.Google Scholar
Choufani, J., Perrichot, V., Azar, D. & Nel, A. 2014b. New biting midges (Diptera: Ceratopogonidae) in late Cretaceous Vendean amber. Paleontological Contributions 10H, 3440.Google Scholar
Choufani, J., El-Halabi, W., Azar, D. & Nel, A. 2015. First fossil insect from Lower Cretaceous Lebanese amber in Syria (Diptera: Ceratopogonidae). Cretaceous Research 54, 106–16.Google Scholar
Clastrier, J. & Raccurt, C. 1979. Baeodasymyia modesta n. g., n. sp., de la République d'Haiti (Diptera, Ceratopogonidae). Annales de Parasitologie 54, 99104.Google Scholar
Cockerell, T. D. A. 1919. Insects in Burmese amber. The Entomologists 52, 241–43.Google Scholar
Edwards, F. W. 1929. Philippine nematocerous Diptera II. Notulae Entomologicae 9, 114.Google Scholar
Granier, B., Toland, C., Géze, R., Azar, D. & Maksoud, S. 2016. Some steps toward a new story for the Jurassic–Cretaceous transition in Mount Lebanon. Carnets de Géologie 16, 247–69.Google Scholar
Grimaldi, D. A. & Ross, A. J. 2017. Extraordinary Lagerstätten in Amber, with particular reference to the Cretaceous of Burma. In Fraser, N.C. & Sues, H.-D. (eds) Terrestrial Conservation Lagerstätten: Windows into the Evolution of Life on Land, 287342 Edinburgh: Dunedin Academic Press Ltd. 450 pp.Google Scholar
Grogan, W. L. & Szadziewski, R. 1988. A new biting midge from Upper Cretaceous (Cenomanian) amber of New Jersey (Diptera: Ceratopogonidae). Journal of Paleontology 62, 808–12.Google Scholar
Grogan, W. L. & Wirth, W. W 1979. A new neotropical genus of predaceous midges, with a key to the genera of Heteromyiini (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 81, 5159.Google Scholar
Grogan, W. L. & Wirth, W. W. 1980. Nannohelea, a new genus of biting midges of the tribe Ceratopogonini, related to Baeohelea Wirth and Blanton (Diptera: Ceratopogonidae). Journal of the Kansas Entomological Society 53, 373–85.Google Scholar
Haeselbarth, E. 1965. Phaenobezzia, a new genus of biting midges (Diptera: Ceratopogonidae), with a review of the African species. Zeitschrift für Angewandte Zoologie 52, 297324.Google Scholar
Hagan, D. V., Hassold, E., Kynde, B., Szadziewski, R., Thunes, K. H., Skartveit, J. & Grogan, W. L. 2000. Biting midges (Diptera: Ceratopogonidae) from forest habitats in Norway. Polish Journal of Entomology 69, 465–76.Google Scholar
Heyden, L. F. J. D. 1870. Fossile Dipteren aus der Braunkohle von Rott im Siebengebirge. Palaeontographica 17, 237–66.Google Scholar
Hong, Y.-C. 1981. Eocene fossil Diptera Insecta in amber of Fushun coalfield. Beijing: Geological Publishing House. 166 pp.Google Scholar
Hong, Y.-C., Guo, X.-R. & Ren, D. 2000. A new genus – Eopalpomyitis gen. nov. from Eocene Fushun amber and discussion of its taxonomic position. Acta Parasitologica et Medica Entomologica Sinica 7, 225–34.Google Scholar
Kalugina, N. S. 1991. [New Mesozoic Simuliidae and Leptoconopidae and blood-sucking origin in lower Dipterans.] Palaeontologicheskii Zhurnal 1991, 6980. [In Russian, with English summary.]Google Scholar
Kieffer, J. J. 1899. Description d'un nouveau genre et tableau des genres européens de la famille des Chironomides (Dipt.). Bulletin de la Société Entomologique de France 1899, 6670.Google Scholar
Kieffer, J. J. 1906. Diptera. Fam. Chironomidae. In Wytsman, P. (ed.) Genera Insectorum, Fasc. 42. Bruxelles: L. Desmet-Verteneuil. 78 pp.Google Scholar
Kieffer, J. J. 1911a. Description de nouveaux chironomides de l'Indian Museum de Calcutta. Records of the Indian Museum 6, 113–77.Google Scholar
Kieffer, J. J. 1911b. Nouvelles descriptions de chironomides obtenus d'éclosion. Bulletin de la Société d'Histoire Naturelle de Metz 27, 160.Google Scholar
Kieffer, J. J. 1913. Chironomidae et Cecidomyidae. In Alluaud, C. A. & Jeannel, R. (eds) Voyage de Ch. Alluaud et R. Jeannel en Afrique orientale (1911–1912). Resultats scientifiques. Diptera 5, 1351.Google Scholar
Kieffer, J. J. 1917. Chironomides d'Amérique conservés au Musée National Hongrois de Budapest. Annales Historico-Naturales Musei Nationalis Hungarici 15, 292364.Google Scholar
Kieffer, J. J. 1925. Diptères (Nématocères piqueurs): Chironomidae Ceratopogoninae. Faune de France 11, 1139.Google Scholar
Kodrul, T. M. 1999. Paleogene phytostratygraphy of the South Sakhalin. Trudy Geologitscheskogo Instituta, Russian Academy of Sciences 519, 1150.Google Scholar
Latreille, P. A. 1809. Genera crustaceorum et insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisque plurimis explicata. Vol. 4. Paris and Strasbourg. 399 pp.Google Scholar
Loew, H. 1850. Über den Bernstein und dies Bernsteinfauna. Program der Königlichen Realschule zu Meseritz, 44 pp.Google Scholar
Macfie, J. W. S. 1940. Ceratopogonidae (Diptera) from British Guiana and Trinidad. Part 2. Proceedings of the Royal Entomological Society of London (B) 9, 187–95.Google Scholar
Maksoud, S., Azar, D., Granier, B. & Géze, R. 2016. New data on the age of the Lower Cretaceous amber outcrops of Lebanon. Palaeoworld 26, 331–38.Google Scholar
Meigen, J. G. 1803. Versuch einer neuen Gattungseintheilung der europäischen zweiflügligen Insekten. Magazin für Insektenkunde 2, 259–81.Google Scholar
Meigen, J. G. 1818. Systematische Beschreibung der bekannten europäischen zweiflügeligen Insekten. Vol. 1. Aachen: Bei Friedrich Wilhelm Forstmann; Gedrukt bei Beaufort Sohn. xxxvi+333 pp.Google Scholar
Meunier, F. 1904. Monographie des Cecidomyidae, des Sciaridae, des Mycetophilidae et des Chironomidae de l'ambre de la Baltique. Annales de la Société Scientifique de Bruxelles, 28, 12264.Google Scholar
Penney, D. 2010. Dominican amber. In Penney, D. (ed.) Biodiversity of fossils in amber from the major world deposits, 2241. Manchester: Siri Scientific Press. 304 pp.Google Scholar
Pérez-de la Fuente, R., Delclòs, X., Peñalver, E. & Arillo, A. 2011. Biting midges (Diptera: Ceratopogonidae) from the Early Cretaceous El Solplao amber (N Spain). Cretaceous Research 32, 750–61.Google Scholar
Perrichot, V., Néraudeau, D., Nel, A. & Ploëg, G. 2007. A reassessment of the Cretaceous amber deposits from France and their palaeontological significance. African Invertebrates, 48, 213–27.Google Scholar
Petrunkevitch, A. 1957. Eohelea stridulans, n. gen., n. sp., a striking example of paramorphism in an amber biting-midge. Journal of Paleontology 31, 208–14.Google Scholar
Poinar, G. O. 2008. Leptoconops nosopheris sp. n. (Diptera: Ceratopogonidae) and Paleotrypanosoma burmanicus gen. n., sp. n. (Kinetoplastida: Trypanosomatidae), a biting midge – trypanosome vector association from the Early Cretaceous. Memorias do Instituto Oswaldo Cruz 103, 468–71.Google Scholar
Remm, H. 1976. Pozdnemelovye mokrecy (Diptera, Ceratopogonidae) iskopaemykh smol Khatangskoj vpadiny. Paleontologicheskiy Zhurnal 3, 107–16.Google Scholar
Ritzkowski, S. 1997. K-Ar-Altersbestimmungen der bernsteinführenden Sedimente des Samlandes (Paläogen, bezirk Kaliningrad). Metalla (Sonderheft), 66, 1923.Google Scholar
Say, T. 1825. American entomology, or descriptions of the insects of North America. Vol. 2. Philadelphia: Philadelphia Museum. 121 pp.Google Scholar
Shi, G., Grimaldi, D. A., Harlow, G. E., Wang, J., Wang, J., Yang, M., Lei, W., Li, Q. & Li, X. 2012. Age constraint on Burmese amber based on U–Pb dating of zircons. Cretaceous Research 37, 155–63.Google Scholar
Skuse, F. A. A. 1889. Diptera of Australia. Part VI. – The Chironomidae. Proceedings of the Linnean Society of New South Wales 4, 215311, pls. 11–14.Google Scholar
Solórzano-Kraemer, M. M. 2007. Systematic, palaeoecology and palaeobiogeography of the insect fauna from Mexican amber. Palaeontographica Abteilung A 282, 1133.Google Scholar
Solórzano-Kraemer, M. M. 2010. Mexican amber. In Penney, D. (ed.) Biodiversity of fossils in amber from the major world deposits, 4256. Manchester: Siri Scientific Press. 304 pp.Google Scholar
Sontag, E. & Szadziewski, R. 2011. Biting midges (Diptera: Ceratopogonidae) in Eocene Baltic amber from the Rovno region (Ukraine). Polish Journal of Entomology 80, 779800.Google Scholar
Standke, G. 2008. Bitterfelder Bernstein gleich Baltischer Bernstein? Eine geologische Raum-Zeit-Betrachtung und genetische Schlussfolgerungen. Exkursionsführer und Veröffentlichungen der Deutschen Gesellschaft für Geowissenschaften, 236, 1133.Google Scholar
Statz, G. 1944. Neue Dipteren (Nematocera) aus dem Oberoligocän von Rott. I. Teil VI. Familie: Tendipedidae (Zuck- oder Schwarmmücken). II. Teil. VII. Familie Heleidae (Gnitzen). III. Teil VIII. Familie Lycoriidae (Trauermücken). Palaeontographica (A) 95, 122–87.Google Scholar
Stebner, F., Szadziewski, R., Rühr, P., Hammel, J. U., Kvifte, G. M., Singh, H. & Rust, J. 2016a. A fossil biting midge (Diptera: Ceratopogonidae) from early Eocene Indian amber with a complex pheromone evaporator. Scientific Reports 6, 34352.Google Scholar
Stebner, F., Szadziewski, R. & Wang, B. 2016b. Biting midges (Diptera: Ceratopogonidae) in Fushun amber reveal further biotic links between Eocene Asia and Europe. Palaeontologia Electronica 19.3.31A, 19.Google Scholar
Stebner, F., Szadziewski, R., Singh, H., Gunkel, S. & Rust, J. 2017. Ceratopogonidae from Cambay amber indicate that the Eocene biting midge fauna of the Indian Subcontinent was not isolated. PlosOne 12(3), e0173135.Google Scholar
Szadziewski, R. 1984. Niezwykłe narządy strydulacyjne u eoceńskich muchówek z rodziny Ceratopogonidae (Diptera). Wiadomości Entomologiczne 5, 3740.Google Scholar
Szadziewski, R. 1988. Biting midges (Diptera, Ceratopogonidae) from Baltic amber. Polish Journal of Entomology 58, 3283.Google Scholar
Szadziewski, R. 1990. Biting midges (Insecta: Diptera: Ceratopogonidae) from Sakhalin amber. Prace Muzeum Ziemi 41, 7781.Google Scholar
Szadziewski, R. 1993. Biting midges (Diptera, Ceratopogonidae) from Miocene Saxonian amber. Acta Zoologica Cracoviensia 35, 603–56.Google Scholar
Szadziewski, R. 1996. Biting midges from Lower Cretaceous amber of Lebanon and Upper Cretaceous Siberian amber of Taimyr (Diptera, Ceratopogonidae). Studia Dipterologica 3, 2386.Google Scholar
Szadziewski, R. 1998. A new species of the predaceous midge genus Metahelea from Baltic amber (Diptera: Ceratopogonidae). Polish Journal of Entomology 67, 245–53.Google Scholar
Szadziewski, R. 2000. Biting midges (Diptera: Ceratopogonidae) from the Lower Cretaceous amber of Jordan. Polish Journal of Entomology 69, 251–56.Google Scholar
Szadziewski, R. 2004. Biting midges (Diptera: Ceratopogonidae) from Burmese amber, Myanmar. Journal of Systematic Palaeontology 2, 115–21.Google Scholar
Szadziewski, R. 2005. The first fossil species in the predaceous tribe Sphaeromiini (Diptera: Ceratopogonidae). Polish Journal of Entomology 74, 363–68.Google Scholar
Szadziewski, R. 2008. Age and recent distribution of extant genera of Ceratopogonidae (Diptera) present in the fossil record. Alavesia 2, 87-99.Google Scholar
Szadziewski, R., Krzywiński, J. & Giłka, W. 1997. Diptera Ceratopogonidae, Biting Midges. In Nilsson, A. (ed.) Aquatic Insects of North Europe. A taxonomic handbook. Vol. 2. Odonata – Diptera, 243–63. Stenstrup, Denmark: Apollo Books. 440 pp.Google Scholar
Szadziewski, R., Giłka, W. & Sontag, E. 2007. First description of males from Eocene Baltic amber in the fossil genus Mantohelea (Diptera: Ceratopogonidae). Alavesia 1, 3740.Google Scholar
Szadziewski, R., Giłka, W. & Urbanek, A. 2015a. A blood sucking biting midge from Upper Cretaceous Burmese amber with a key to the determination of fossil species in the relictual genus Leptoconops Skuse (Diptera: Ceratopogonidae). Cretaceous Research 54, 255–59.Google Scholar
Szadziewski, R., Ross, A. & Giłka, W. 2015b. Further records of biting midges (Diptera: Ceratopogonidae) from Upper Cretaceous Burmese amber (Myanmar). Cretaceous Research 52, 556–61.Google Scholar
Szadziewski, R., Arillo, A., Urbanek, A. & Sontag, E. 2016. Biting midges of the extinct genus Protoculicoides Boesel from Lower Cretaceous amber of San Just, Spain and new synonymy in recently described fossil genera (Diptera: Ceratopogonidae). Cretaceous Research 58, 19.Google Scholar
Szadziewski, R. & Arillo, A. 1998. Biting midges (Diptera: Ceratopogonidae) from the Lower Cretaceous amber from Alava, Spain. Polish Journal of Entomology 67, 291-98.Google Scholar
Szadziewski, R. & Arillo, A. 2003. The oldest fossil record of the extant subgenus Leptoconops (Leptoconops) (Diptera: Ceratopogonidae). Acta Zoologica Cracoviensia 46 (suppl.) Fossil Insects, 271–75.Google Scholar
Szadziewski, R. & Grogan, W. L. 1994. Biting midges from Dominican amber. I. A new fossil species of Baeodasymyia (Diptera: Ceratopogonidae). Proceedings of the Entomological Society of Washington 96, 219–29.Google Scholar
Szadziewski, R. & Grogan, W. L. 1996. Biting midges (Diptera: Ceratopogonidae) from Mexican amber. Polish Journal of Entomology 65, 291–95.Google Scholar
Szadziewski, R. & Grogan, W. L. 1997. Biting midges from Dominican amber. II. Species of the tribes Heteromyiini and Palpomyiini (Diptera: Ceratopogonidae). Memoirs of the Entomological Society of Washington 16, 254–60.Google Scholar
Szadziewski, R. & Grogan, W. L. 1998a. Biting midges from Dominican amber III. Species of the tribes Culicoidini and Ceratopogonini (Diptera: Ceratopogonidae). Insecta Mundi 12, 3946.Google Scholar
Szadziewski, R. & Grogan, W. L. 1998b. Biting midges from Dominican amber. IV. Species of the tribes Dasyheleini and Forcipomyiini (Diptera: Ceratopogonidae). Polish Journal of Entomology 67, 255–90.Google Scholar
Szadziewski, R. & Poinar, G.O. 2005. Additional biting midges (Diptera: Ceratopogonidae) from Burmese amber. Polish Journal of Entomology 74, 349–62.Google Scholar
Szadziewski, R. & Schlüter, T. 1992. Biting midges (Diptera: Ceratopogonidae) from Upper Cretaceous (Cenomanian) amber of France. Annales de la Société Entomologique de France (N.S.) 28, 7381.Google Scholar
Szadziewski, R. & Sontag, E. 2013. A new species of Forcipomyia from Paleocene Sakhalin amber (Diptera: Ceratopogonidae). Polish Journal of Entomology 82, 5962.Google Scholar
Szwedo, J. & Sontag, E. 2013. The flies (Diptera) say that amber from the Gulf of Gdańsk, Bitterfeld and Rovno is the same Baltic amber. Polish Journal of Entomology 82, 379–88.Google Scholar
Urbanek, A., Richert, M., Giłka, W. & Szadziewski, R. 2011. Morphology and histology of secretory setae in terrestrial larvae of biting midges of the genus Forcipomyia (Diptera: Ceratopogonidae). Arthropod Structure & Development 40, 485–94.Google Scholar
Urbanek, A., Piotrowicz, M., Szadziewski, R. & Giłka, W. 2014. Sensilla coeloconica ringed by microtrichia in host-seeking biting midges. Medical and Veterinary Entomology 28, 355–63.Google Scholar
Urbanek, A., Szadziewski, R. & Dominiak, P. 2015. Harmful touch: tarsal palisades as unique tools of predation in biting midges (Diptera: Ceratopogonidae). Zoomorphology 134, 457–68.Google Scholar
Weitschat, W. & Wichard, W. 2010. Baltic amber. In Penney, D. (ed.) Biodiversity of Fossils in Amber from the Major World Deposits, 80115. Manchester: Siri Scientific Press. 304 pp.Google Scholar
Wirth, W. W. 1962. A reclassification of the PalpomyiaBezzia–Macropeza groups, and a revision of the North American Sphaeromiini (Diptera, Ceratopogonidae). Annals of the Entomological Society of America 55, 272–87.Google Scholar
Wirth, W. W. & Grogan, W. L. 1988. The predaceous midges of the world (Diptera: Ceratopogonidae; tribe Ceratopogonini). Flora & Fauna Handbooks 4. Leiden: E. J. Brill. xv+160 pp.Google Scholar
Wirth, W. W. & Hubert, A. A. 1960. Camptopterohelea a new genus of Ceratopogonidae from the Philippines (Diptera). Fieldiana, Zoology 42, 8991.Google Scholar
Wirth, W. W. & Lee, D. J. 1958. Australasian Ceratopogonidae (Diptera, Nematocera). Part VIII: A new genus from Western Australia attacking man. Proceedings of the Linnean Society of New South Wales 83, 337–39.Google Scholar
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, 170–77.Google Scholar