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Type: Monograph
Published: 2011-04-29
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A revision of the genera Pseudosmittia Edwards, 1932, Allocladius Kieffer, 1913, and Hydrosmittia gen. n. (Diptera: Chironomidae, Orthocladiinae)

Department of Entomology, University of Minnesota, 219 Hodson Hall, 1980 Folwell Avenue, Saint Paul, Minnesota USA 55108
Department of Natural History, Bergen Museum, University of Bergen, Pb 7800, N-5020 Bergen, Norway
Diptera Chironomidae Orthocladiinae Allocladius Hydrosmittia Pseudosmittia new genus new species new combinations phylogeny keys

Abstract

The genera Pseudosmittia Edwards, 1932, and Allocladius Kieffer, 1913, supported by parsimony analysis, can be divided into three groups, Pseudosmittia, Allocladius and Hydrosmittia gen. n. Generic diagnoses to all stages and both sexes of all three genera are given. The mostly Neotropical and Afrotropical Allocladius together with the mostly Holarctic and Afrotropical Hydrosmittia form the sister group of the more widespread Pseudosmittia. Allocladius in addition to some basal species is divided into a paraphyletic nearly Cosmopolitan fortispinatus group, a monotypic Afrotropical soemmei group, an Afrotropical and Palaearctic niger group, a Holarctic nanseni group, and a Neotropical, Afrotropical and Holarctic longicrus group; Hydrosmittia into an Afrotropical and Palaearctic brevicornis group and a mostly Holarctic ruttneri group. Pseudosmittia is divided into 13 species groups in phyletic sequence: gracilis [simplex + rotunda (topei (digitata (xanthostola + brachydicrana + insulsa)], [conjuncta (tokaraneoa + albipennis (brevifurcata (angusta group, divided into angusta, trilobata, lamasi, danconai, uncata and forcipata subgroups)]. The xanthostola and brachydicrana groups are with few exceptions known only from South Asia and the Indo-Pacific region, while most other groups are present in most regions.

        Keys to male and female imagines, pupae and larvae of all three genera are given. Type material of 180 species assigned to Pseudosmittia, Camptocladius v. d. Wulp, Spaniotoma Philippi, Smittia Holmgren, Mesosmittia Brundin, Orthosmittia Goetghebuer, Ancylocladius Sublette et Wirth, Allocladius Kieffer, and Lindebergia Tuiskunen, belonging to 15 museums have been re-examined, lectotypes designated, and new combinations and synonyms given. An additional 21 species are lost or could not be located, of these 5 are declared nomina dubia. Another 4 species were not available for study or not examined. Most of these could be placed in other genera or as synonyms of species in Pseudosmittia. One hundred and thirty species are treated, 37 of which are new, 93 species are completely redescribed in all available stages. The following new species are described: Allocladius bubatus, A. caspersi, A. deborae, A. hirticaudatus, A. luciniolus, A. soemmei, A. wangorum, Hydrosmittia aagaardi, H. annulata, H. falsicostata, H. soelii, H. tenuistylata, Pseudosmittia aculeathrix, P. acutilobata, P. carita, P. christmasensis, P. cristagata, P. cunealata, P. dolabrata, P. digitrienta, P. fusata, P. laticauda, P. legonensis, P. licina, P. longicornia, P. malickyi, P. navama, P. parifusata, P. parinavama, P. pedata, P. propetropis, P. pugnata, P. siamensis, P. spinispinata, P. tericristata, P. tokunagai, and P. unniae. Ten species of Allocladius (including 2 parthenogenetic), 4 species of Hydrosmittia (including 2 parthenogenetic), and 31 species of Pseudosmittia (including 3 parthenogenetic and 3 species known only from females) are known as female imagines; 5 species of Allocladius, 6 species of Hydrosmittia and 12 species of Pseudosmittia are known as pupae; and 4 species of Allocladius, 5 species of Hydrosmittia and 10 species of Pseudosmittia are known as larvae.

        Errors in previous publications are corrected and remarks on variation given. The genera treated contain at least 7 apparently obligate parthenogenetic species, while facultative parthenogenesis is found in at least one species and may be widespread.

        The species of Hydrosmittia probably all are truly aquatic, while Allocladius has species found in both fully aquatic environments and in moist earth. In Pseudosmittia there are no certain fully aquatic species. Most species appear to be semiaquatic to semiterrestrial or live in the marine intertidal zone. The insulsa, brachydicrana, and xanthostola groups contain mostly marine seashore species, and several groups have species, which larvae live both in semiaquatic freshwater localities and on the seashore. In the angusta group, however, only P. bifurcata appears to be intertidal.

Only in Allocladius a cool southern transantarctic Gondwanian vicariance may have taken place, but combined with dispersal to West Asia, further to the Euro-Mediterranean and East Asian Regions, and across a Beringian Region to North America. A more northern warm Gondwanian connection between West Africa and East South America could be present but not common in Pseudosmittia. The dominating distribution pattern in Pseudosmittia appear to be caused by direct dispersal, in America across the Caribbean and the Central American lowland, in the Indian Ocean and the Pacific across oceans by floating debris and vegetation.

References

  1. Adam, J.I. & Sæther, O.A. (1999) Revision of the genus Nilothauma Kieffer, 1921 (Diptera: Chironomidae). Entomologica scandinavica, Supplement, 56, 1–107.

    Adams, C.G., Gentry, A.W. & Whybrow, P.J. (1983) Dating the terminal Tethyan Event (pp.237–298). In Meulenkamp, J. E. (Ed). Reconstruction of marine palaeontological environments. Utrecht Micropalaeontological Bulletin, 30, 1–298.

    Albu, P. (1968) Pseudosmittia mathildae sp. n. (Diptera, Chironomidae. Annales Zoologici Fennici, 5, 4–5.

    Amorim, D.S., Santos, C.M.D. & de Oliveira, S. (2009) Allochronic taxa as an alternative model to explain circumantarctic disjunctions. Systematic Entomology, 34, 2–9.

    Andersen, F. Søgaard (1937) Über die Metamorphose der Ceratopogoniden und Chironomiden Nordost-Grönlands. Meddelelser om Grønland, 116, 1–95.

    Andersen, T. & Mendes, H.F. (2007) Five enigmatic new orthoclad genera from Brazil (Diptera: Chironomidae, Orthocladiinae). In: Andersen, T. (Ed.) Contributions to the systematic and ecology of aquatic Diptera. A tribute to Ole A. Sæther. The Caddis Press, Columbus, Ohio, pp. 17–52.

    Andersen, T. & Mendes, H.F. (2010) Order Diptera family Chironomidae (with the exception of the tribe Tanytarsini). In: van Harten, A. (Ed.) Arthropod fauna of the UAE Vol.3, pp. 564–598.

    Andersen, T. & Sæther, O.A. (1993) Lerheimia, a new genus of Orthocladiinae from Africa (Diptera: Chironomidae). Spixiana, 16, 105–112.

    Andersen, T. & Sæther, O.A. (1994) Colosmittia clavata gen. n., sp. n., a new orthoclad from the Usambara Mts, Tanzania (Diptera: Chironomidae). Journal of the Kansas Entomological Society, 66, 439–443.

    Andersen, T., Sæther, O.A. & Mendes, H.F. (2010) Neotropical Allocladius Kieffer, 1913, and Pseudosmittia Edwards, 1932 (Diptera: Chironomidae). Zootaxa, 2472, 1–77.

    Ashe, P. (1983) A catalogue of chironomid genera and subgenera of the world including synonyms (Diptera: Chironomidae). Entomologica scandinavica, Supplement, 17, 1–68.

    Ashe, P. & Cranston, P.S. (1990) Family Chironomidae. In: Sóos, Á. & Papp, L. (Eds.) Catalogue of Palaearctic Diptera. Vol. 2 Psychodidae - Chironomidae. Akadémia Kiadó, Budapest, pp. 113–355.

    Banarescu, P. (1991) Zoogeography of fresh waters. General distribution and dispersal of freshwater animals. Volume 1, AULA-Verlag GmbH, Wiesbaden, pp.1–158.

    Banarescu, P. (1992) Zoogeography of fresh waters. Distribution and dispersal of freshwater animals in North America and Eurasia. Volume 2, AULA-Verlag GmbH, Wiesbaden, pp. 159–1091.

    Banarescu, P. (1995) Zoogeography of fresh waters. Distribution and dispersal of freshwater animals in Africa, Pacific areas and South America. Volume 3, AULA-Verlag GmbH, Wiesbaden, pp. 1093–1617.

    Bremer, K. (1992) Ancestral areas: a cladistic reinterpretation of the center of origin concept. Systematic Biology, 41, 436–445.

    Brooks, D.R. (1990) Parsimony analysis in historical biogeography and coevolution: methodological and theoretical update. Systematic Zoology, 39, 14–30.

    Brooks, D., Dowling, A.P.G., van Veller, M.G.P. & Hoberg, E.P. (2004) Ending a decade of deception: a valiant failure, a not-so-valiant failure, and a success story. Cladistics, 20, 32–46.

    Brooks, D., van Veller, M.G.P. & McLennan, D.A. (2001) How to do BPA, really. Journal of Biogeography, 28, 345–358.Brundin, L. (1947) Zur Kenntnis der schwedischen Chironomiden. Arkiv för Zoologi, 39A, 1–95.

    Brundin, L. (1956) Zur Systematik der Orthocladiinae (Dipt., Chironomidae). Institute of Freshwater Research, Drottningholm, Report, 37, 5–185.

    Caspers, N. (1990) Pseudosmittia duplicata sp. n. und Pseudosmittia forcipata (Goetghebuer, 1921) aus China (Diptera, Chironomidae). Entomofauna, 11(12), 217–225.

    Caspers, N. & Reiss, F. (1989) Die Chironomidae der Türkei. Teil I: Podonominae, Diamesinae, Prodiamesinae, Orthocladiinae (Diptera, Nematocera, Chironomidae). Entomofauna, 10, 105–160.

    Cranston, P.S. (1996) Flying the southern hemisphere (abstract). Proceedings of the XX international congress of entomology, 1996, 78.

    Cranston, P.S. & Martin, J. (1989) 26. Family Chironomidae. In Evenhuis N. L. (Ed.), Catalog of the Diptera of the Australasian and Oceanic Region. Bishop Museum Special Publications, 86, 252–274.

    Cranston, P.S. & Oliver, D.R. (1988) Additions and corrections to the Nearctic Orthocladiinae (Diptera: Chironomidae). Canadian Entomologist, 120, 425–462.

    Edwards, F.W. (1922) Results of the Oxford University Expedition to Spitsbergen, 1921. Annals and Magazine of Natural History, (9) 10, 193–215.

    Edwards, F.W. (1926) On marine Chironomidae (Diptera) with descriptions of a new genus and four new species from Samoa. Proceedings of the Zoological Society of London, 51, 779–806.

    Edwards, F.W. (1927) Diptera Nematocera from the South Pacific collected by the “St. George” Expedition, 1925. Annals and Magazine of Natural History, (9), 20, 236–244.

    Edwards, F.W. (1928) Nematocera. Insects of Samoa, 6, 23–102.

    Edwards, F.W. (1929) British non-biting midges (Diptera, Chironomidae). Transactions of the Entomological Society of London, 77, 279–430.

    Edwards, F.W. (1931) Chironomidae. In: Diptera of Patagonia and South Chile. Part II. Fascicle 5. The British Museum (Natural History), London, pp. 233–331.

    Edwards, F.W. (1932) Recent literature. Faune de France: 23. Diptères: Chironomidae, IV. Par M. Goetghebuer. Paris (Lechevalier), 1932. The Entomologist, 65, 140–141.

    Edwards, F.W. (1933) Mycetophilidae, Culicidae and Chironomidae and additional records of Simuliidae, from the Marqesas Islands. Bernice P. Bishop Museum Bulletin, 114: 85–92.

    Edwards, F.W. (1937) Chironomidae (Diptera) collected by Prof. Aug. Thienemann in Swedish Lapland. Annals and Magazine of Natural History, (10) 20, 140–148.

    Ferrington, L.C.jr., Kavanaugh, R.G., Schmidt, F. J. & Kavanaugh, J.L. (1995) Habitat separation among Chironomidae (Diptera) in Big Springs. Journal of the Kansas Entomological Society, 68(2) suppl., 1995, 152–165.

    Ferrington, L.C.jr. & Sæther, O.A. (1995a) Physoneura, a new genus of Orthocladiinae from Patagonia and South Chile (Diptera: Chironomidae). Aquatic Insects 17, 57–63.

    Ferrington, L.C.jr. & Sæther, O.A. (1995b) Afrotropical species of Parakiefferiella Thienemann, with a review of chironomids with palpal projections (Diptera: Chironomidae). Pp. 369–377 in: P.S. Cranston (ed.); Midges: from genes to ecotypes. CSIRO, Canberra, 482 pp.

    Ferrington, L.C.jr. & Sæther, O.A. (2006) Rhagosmittia and Trondia, two new genera of Orthocladiinae from Oceania and Australia (Diptera: Chironomidae). Aquatic Insects, 28: 243–250.

    Freeman, P. (1953) Chironomidae from western Cape Province.- I. Proceedings of the royal entomological Society in London, 22, 201–213.

    Freeman, P. (1954) Chironomidae from western Cape Province.- IV. Proceedings of the royal entomological Society in London, 22, 201–213.

    Freeman, P. (1955) Diptera (Nematocera): Chironomidae. South African animal Life, 2, 361–381.

    Freeman, P. (1956) A study of the Chironomidae (Diptera) of Africa south of the Sahara. Part II. Bulletin of British Museum (Natural History), Entomology, 4, 287–368.

    Freeman, P. (1961) The Chironomidae of Australia. Australian Journal of Zoology, 9, 611–737.

    Freeman, P. (1962) Chironomidae from the Batu Caves, Malaysia (Diptera: Nematocera). Pacific Insects, 4, 129–131.

    Freeman, P. & Cranston, P.S. (1980) 11. Family Chironomidae. Pp. 175–202 in Crosskey, R. W. (Ed.) Catalogue of the Diptera of the Afrotropical Region, Trustees of The British Museum (Natural History), London, 1437 pp.

    Goetghebuer, M. (1913) Description de Chironomides nouveaux récoltes en Belgique. Annales biologie lacustre, 6, 148–172.

    Goetghebuer, M. (1921) Chironomides de Belgique et spécialement de la zone des Flandres. Mémoires du Musée Royal d’Histoire Naturelle de Belgique, 8, 1–211.

    Goetghebuer, M. (1931) Ceratopogonidae et Chironomidae nouveaux d’Europe. Bulletin & Annales de la Société Royale d’Entomologique de Belgique, 71, 211–218.

    Goetghebuer, M. (1932) Diptères (Nématocères), Chironomidae IV. Orthocladiinae, Corynoneurinae, Clunioninae, Diamesinae. Faune de France, 23, 1–204.

    Goetghebuer, M. (1933) Chironomides du Groenland oriental, du Svalbard et de la terre de Francois Joseph. Skrifter om Svalbard og Ishavet, 53, 19–31.

    Goetghebuer, M. (1934) Note sur un nouveau Chironomide brachyptère. Bulletin & Annales de la Société Royale d’Entomologique de Belgique, 74, 388–390.

    Goetghebuer, M. (1940–1950) Subfamilie Orthocladiinae. In Lindner, E. (Ed.) Die Fliegen der Palaearktischen Region, 13(g), pp. 1–208, pl. i–xxiv.

    Goetghebuer, M. (1941) Ceratopogonidae et Chironomidae nouveaux d’Allemagne. Archiv für Hydrobiologie, 38, 288–290.

    Goetghebuer, M. (1950) Ceratopogonidae et Chironomidae nouveaux ou peu connus d’Europe (Treizième note). Bulletin de la Institute Royal des Sciences Naturelles de Belgique, 25(14), 1– 8.

    Hamilton, A.C. (1982) Environmental history of East Africa: a study of the Quaternary. Academic Press, London, xiv+1–328 pp.

    Hardy, D.E. (1960) Diptera: Nematocera-Brachycera (except Dolichopodidae). In: Zimmermann, E.C. (Ed.) Insects of Hawaii 10, Chironomidae 94–165.

    Holmgren, A.E. (1869) Bidrag til Kännedomen om Beeren Eilands och Spetzbergens Insekt-Fauna. Kungliga Svenska Vetenskapsakademiens Handlingar, 8, 1–55.

    Johannsen, O.A. (1946) Some new species of nematocerous Diptera from Guam. Bulletin of Bernice P. Bishop Museum, 189, 187–193.

    Kieffer J.J. (1911) The Percy Slade Trust expedition to the Indian Ocean in 1905, under the leadership of Mr J. Stanley Gardiner. Vol. 3. No. XV. - Diptera, Chironomidae der Seychellen-Inseln, aus der Sammlung von Mr H. Scott. Transactions of the Linnéan Society of London (2nd Ser. Zool.), 14, 331–366.

    Kieffer, J.J. (1913) Chironomidae et Cecidomyidae. Résult scientifique de Voyage de Ch, Allaud et R. Jeannel en Afrique Oriental (19111912). Insectes Diptères, 1, 1–43

    Kieffer, J.J. (1918) Chironomides d´Afrique et d´Asie conservé au Muséum National Hongrois de Budapest. Annales historico-naturales Musei nationalis Hungarici, 16, 31–136.

    Kieffer, J.J. (1921a) Chironomides de Courlande. Annales de la Société Scientifique de Bruxelles, 40, 275–298.

    Kieffer, J.J. (1921b) Chironomides nouveaux ou peu connus de la région paléarctique. Bulletin de la Société d’Histoire Naturelle de Metz, 29, 51–109.

    Kieffer, J.J. (1924) Quelques chironomides nouveaux et remarquables du Nord de l’Europe. Annales de la Société Scientifique de Bruxelles, 43, 390–397.

    Kieffer, J.J. (1926) Chironomiden der 2. Fram-Expedition (1898–1902). Norsk Entomologisk Tidsskrift, 2 [1925], 78–89.

    ICZN (1999) International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, xxix + 306 pp.

    Kikuchi, M. & Sasa, M. (1990) Studies on the chironomid midges (Diptera, Chironomidae) of the Lake Toba area, Sumatra, Indonesia. Japanese Journal of Sanitary Zoology, 41(4), 291–329.

    Langton, P. (1991) A key to pupal exuviae of West Palaearctic Chironomidae. P.H. Langton, Huntingdon, UK, 1–386.

    Langton, P.H. & Pinder, L.C.V. (2007) Keys to the adult male Chironomidae of Britain and Ireland. Volume 1. Introductory text, keys, references, checklist and index; Volume 2. Illustrations of the hypopygia and a supplement identifying sixteen species recently recorded from Britain and Ireland. Freshwater Biological Association, Scientific Publication 64, 231 plus 168 pp.

    Lehmann. J. (1979) Chironomidae (Diptera) aus Fliessgewässern Zentralafrikas (Systematik, Ôkologie, Verbreitung und Produktionsbiologie). Teil I: Kivu-Gebiet, Ost D. R. Congo. Spixiana Supplement, 3, 144 pp.

    Lundbeck, W. (1898) Diptera groenlandica. Videnskabelige Meddelelser fra dansk naturhistorisk Forening, 5(10), 236–314.

    Maheshwari, G. & Maheshwari, G. (2002) Some Chironomidae from South and Middle Andaman Islands, India (Diptera: Chironomidae). Journal of the Bombay Natural History Society, 98(3), 406-421.

    Makarchenko, E.A. & Makarchenko, M.A. (2005) A new species of chironomids (Diptera, Chironomidae) of the Russian Far East (In Russian with English abstract). Vladimir Ya. Levanidov’s Biennial Memorial Meetings, 3, 374–382.

    Makarchenko, E.A. & Makarchenko, M.A. (2006) New or little-known chironomids of Orthocladiinae (Diptera, Chironomidae) from the Russian Far East. Russian entomological journal, 15 (1), 83–92.

    Makarchenko, E.A. & Makarchenko, M.A. (2007) New records of chironomids (Diptera, Chironomidae) in the Russian Far East. I. Subfamily Orthocladiinae. Eurasian entomological journal, 6 (3), 299–310.

    Makarchenko, E.A. & Makarchenko, M.A. (2008) Review of the genus Pseudosmittia Edwards (Diptera: Chironomidae: Orthocladiinae) from the Russian Far East. Russian Entomological Journal, 17(2), 215–226.

    Malloch, J.R. (1915) The Chironomidae, or midges, of Illinois, with particular reference to the species occurring in the Illinois River. Bulletin of the Illinois State Laboratory of Natural History, 10, 275–543.

    Marcuzzi, G. (1947) Descrizione di tre nuove specie di Smittia della Laguna di Venezia. Bolletino della Società Entomologica Italiana, 77, 9–13.

    Mendes, H. F. & Andersen, T. (2008) A review of Antillocladius Sæther and Litocladius Mendes, Andersen et Sæther, with the description of two new Neotropical genera (Diptera, Chironomidae, Orthocladiinae). Zootaxa, 1887, 1–75.

    Mendes, H.F. & Andersen, T. (2009a) Neotropical Nilothauma Kieffer, 1921, with the description of thirteen new species (Diptera: Chironomidae). Zootaxa, 2063, 1–45.

    Mendes, H.F. & Andersen, T. (2009b) First record of the genus Colosmittia Andersen & Sæther (Chironomidae: Orthocladiinae) from the Neotropical Region. - Neotropical Entomology, 38, 649–652.

    Messias, M.C. & Oliveira, S.J de (2000) On a new species of the genus Bryophaenocladius (Chironomidae: Orthocladiinae). In: Hoffrichter, O. (Ed) Late 20th century research on Chironomidae: an anthology from the 13th International Symposium on Chironomidae, Shaker, Aachen, pp. 189–191.

    Oliver, D.R. (1963) Entomological studies in the Lake Hazen area, Ellesmere Island, including lists of species of Arachnida, Collembola, and Insecta. Journal of the Arctic Institute of North America, 16, 175–180.

    Oliver, D.R. (1970) Designation and description of the lectotypes of the six Greenland Orthocladiinae (Dipt. Chironomidae) described by Lundbeck in 1898. Entomologica scandinavica, 1(2): 102–108.

    Oliver, D.R. & Dillon, M.E. (1994) Corrections and additions to "A catalog of Nearctic Chironomidae". Proceedings of the entomological Society of Washington, 96, 8–10.

    Paggi, A.C. (1993) Redescription of Pseudosmittia bilobulata (Edw.) n. comb. Spaniotoma (Smittia) bilobulata Edwards 1931] and description of P. neobilobulata sp. nov. (Diptera Chironomidae). Annales de Limnologie, 29 (2), 171–174.

    Paasivirta, L. (2007) Chironomid species in Finnish springs and their surroundings. In: Andersen, T. (Ed.) Contributions to the systematic and ecology of aquatic Diptera. A tribute to Ole A. Sæther. The Caddis Press, Columbus, Ohio, pp. 235–246.

    Pinder, L.C.V. (1978) A key to adult males of British Chironomidae. Part 1. The key; part 2. Illustrations of the hypopygia. Freshwater Biological Association, Scientific Publication 37, 169 pp., 189 figs.

    Pinho, L.C., Mendes, H.F. & Andersen, T. (2009) A review of Diplosmittia Sæther, with the description of four new Neotropical species (Diptera: Chironomidae). Studies on Neotropical Fauna and Environment, 2009, 1-20.

    Queiroz, A. de (2005) The resurrection of oceanic dispersal in historical biogeography. Trends in ecology and evolution, 20 (2), 68–73.

    Ronquist, F. (1994) Ancestral areas and parsimony. Systematic Biology, 43, 267–274.

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

    Sæther, O.A. (1977) Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys. Bulletin of the Fisheries Research Board of Canada,197, 211 pp.

    Sæther, O.A. (1980) Glossary of chironomid morphology terminology (Chironomidae: Diptera). Entomologica scandinavica Supplement, 14, pp. 1–51.

    Sæther, O.A. (1981) Orthocladiinae (Diptera: Chironomidae) from the British West Indies, with descriptions of Antillocladius n. gen., Lipurometriocnemus n. gen. Compterosmittia n. gen. and Diplosmittia n. gen. Entomologica scandinavica, Supplement 16, 1–46.

    Sæther, O.A. (1989) Phylogenetic trends and their evaluation in chironomids with special reference to orthoclads. Acta biologica debrecina, supplementum oecologica hungarica, 2, 53–75.

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

    Sæther, O.A. (2000) Zoogeographical patterns in the Chironomidae (Diptera).

    Verhandlungen der Internationalen Vereinigung für Theoretische und Angewandte Limnologie, 27, 290–302.

    Sæther, O.A. (2004) The Chironomidae (Diptera) of the Seychelles. Annales de Limnologie, 40, 285–308.

    Sæther, O.A. (2006) Japanese Pseudosmittia Edwards (Diptera: Chironomidae). Zootaxa, 1198: 21–51.

    Sæther, O.A. & Andersen, T. (1993) Lobosmittia, a new genus of orthoclads from Tanzania and Turkey (Diptera: Chironomidae). Tijdschrift voor entomologie, 136, 283–287.

    Sæther, O.A. & Andersen, T. (1995) Ionthosmittia caudiga n. gen. n. sp., a new orthoclad from the Usambara Mts, Tanzania (Diptera Chironomidae). Tropical Zoology, 8, 197–202.

    Sæther, O.A. & Andersen, T. (2011) Chironomidae from Gough, Nightingale and Tristan da Cunha islands, South Atlantic Ocean (Diptera). Zootaxa (in press)

    Sæther, O.A., Ashe, P. & Murray, D.E. (2000) Family Chironomidae. 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. Appendix A.6. – Science Herald, Budapest, pp. 113–334.

    Sæther, O.A. & Ekrem, T. (2003) Biogeography of Afrotropical Chironomidae. Cimbebasia, 19, 175–191.

    Sæther, O.A. & Ferrington, L.C.Jr. (1993) Redescription of Prosmittia jemtlandica (Brundin, 1947), with a review of the genus. Journal of Kansas Entomological Society, 66, 257–262.

    Sæther, O.A. & Ferrington, L.C.Jr (1997) A review of the genus Semiocladius Sublette & Wirth (Diptera: Chironomidae. Aquatic Insects, 19, 219–236.

    Sæther, O.A. & Ferrington, L.C.Jr (2003) Nomenclature notes on some orthoclads (Diptera: Chironomidae). Zootaxa, 322, 1–7.

    Sæther, O.A. & Kyerematen, R.A.K. (2001) Towards phylogeny and zoogeography of the genus Rheotanytarsus Thienemann et Bause, 1913 (Diptera: Chironomidae). Tijdschrift Voor Entomologie, 144, 73–117.

    Sæther, O.A. & Spies, M. (2004) Fauna Europaea: Chironomidae, Fauna Europaea version 1.1. htpp://www.faunaeur.org.

    Sæther, O.A., Sublette, J.E. & Willassen, E. (1984) Chironomidae (Diptera) from the 2nd Fram Expedition (1898–1902) to Arctic North America described by J. J. Kieffer. Entomologica scandinavica, 15, 249–275.

    Sasa, M. (1984) Studies on chironomid midges in lakes of the Nikko National Park. Pt. II. Taxonomical and morphological studies on the chironomid species collected from lakes in the Nikko National Park. Research Report from the National Institute of Environmental Studies, Japan, 70, 16–215.

    Sasa, M. (1988) Studies on the chironomid midges of lakes in southern Hokkaido. – Studies on the chironomids collected from lakes and streams in the southern region of Hokkaido, Japan. Research Report from the National Institute of Environmental Studies, Japan, 121, 77–90.

    Sasa, M. (1990) Studies on the chironomid midges (Diptera, Chironomidae) of the Nansei Islands, Southern Japan. Japanese Journal of Experimental Medicine, 60, 111–165.

    Sasa, M. (1993) Studies on the chironomid midges (Yusurika) collected in Toyama and other areas of Japan. Part 5. The chironomids collected from lakes in the Aizu District (Fukushima). Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1993, 69–95.

    Sasa, M. (1994) Studies on the chironomid midges collected in Toyama and other areas of Japan. Part i. Additional information on the Chironomidae of Japan, 1994. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1994, 28–67.

    Sasa, M. (1996a) Seasonal distribution of the chironomids collected with light traps. – At the side of two lakes in the Toyama City Family Park. Report from Toyama Prefectural Environmental Pollution Research Centre, 1996 (March), 15–85.

    Sasa, M. (1996b) Studies on the Chironomidae of Japan, 1996. D. Studies on the chironomids additionally collected on the shore of Jinzu river, Toyama. Report from Toyama Prefectural Environmental Pollution Research Centre, 1996 (December), 61–84.

    Sasa, M. (1997) Studies on the Chironomidae of Japan. Part 1. Studies on the chironomids collected throughout the year in the Shofuku Garden, near the mouth of Kurobe River, Toyama. Bulletin from Toyama Prefectural Environmental Pollution Research Centre, 25, 14–69.

    Sasa, M. & Arakawa, R. (1994) Studies on the chironomid midges collected in Toyama and other areas of Japan, 1994. Part 3. Seasonal changes of chironomid species emerging from Lake Furudo. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1994, 88–109.

    Sasa, M. & Hasegawa, H. (1988) Additional records of the chironomid midges from the Ryukyu Islands, southern Japan (Diptera, Chironomidae). Japanese Journal of Sanitary Zoology, 39, 229–256.

    Sasa, M. & Kondo, S. (1993) Studies on the chironomid midges (Yusurika) collected in Toyama and other areas of Japan. Part 7. Additional chironomids recorded from the middle reaches of Kiso River. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1993, 98–106.

    Sasa, M. & Kawai, K. (1987) Studies on the chironomid midges of the stream Itachigawa, Toyama. Bulletin of Toyama Science Museum,10, 25–72.

    Sasa, M. & Okazawa, T. (1992 a) Studies on the chironomid midges (yusurika) of Kurobe River. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1992, 40–91.

    Sasa, M. & Okazawa, T. (1992b) Studies on the chironomid meidges (yusurika) of Toga-Mura, Toyama. Part 2. The subfamily Orthocladiinae. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1992, 92–204.

    Sasa, M. & Suzuki, H. (1993) Additional records of Chironomidae from Amami Island. Research Report from the Toyama Prefectural environmental Pollution Research Center, 1993, 110–124.

    Sasa, M. & Suzuki, H. (1995) The chironomid species collected on the Tokara Islands, Kagoshima (Diptera, Chironomidae). Japanese Journal of Sanitary Zoology,46, 255–288.

    Sasa, M. & Suzuki, H. (1997a) Studies on the Chironomidae collected from the Ogasawara Islands, southern Japan. Medical Entomology and Zoology, 48, 315–343.

    Sasa, M. & Suzuki, H. (1997b) Studies on the Chironomidae (Diptera, Insecta) collected in Mongolia. Japanese Journal of tropical medicine and hygiene, 25, 149–189.

    Sasa, M. & Suzuki, H. (1998) Studies on the chironomid midges collected in Hokkaido and northern Honshu. Tropical Medicine, 40, 9–43.

    Sasa, M. & Suzuki, H. (2000a) Studies on the chironomid species collected on Yakushima Island, southwestern Japan. Tropical Medicine, 42, 53–134.

    Sasa, M. & Suzuki, H. (2000b) Studies on the chironomid species collected at five localities in Hokkaido in September, 1988 (Diptera, Chironomidae). Tropical Medicine, 42, 175–199.

    Sasa, M., Suzuki, H. & Sakai, T. (1998) Studies on the chironomid species collected on the shore of Shimanto River in April, 1998. Part 2. Description of additional species belonging to Orthocladiinae, Diamesinae and Tanypodinae. Tropical Medicine, 40, 99–147.

    Sasa, M., Watanabe, M. & Arakawa, R. (1992) Additional records of Chironomidae from Togamura, 1992. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1992, 231–246.

    Schmid, P.E. (1993) A key to the larval Chironomidae and their instars from Austrian Danube region streams and rivers. With particular reference to a numerical taxonomic approach. Part I. Diamesinae, Prodiamesinae and Orthocladiinae. Wasser und Abwasser Supplement, 3/93, 1–514.

    Skuse, F.A.A. (1889) Diptera of Australia. Part VI. The Chironomidae. Proceedings of the Linnean Society of New South Wales, 2, 215–311.

    Spies, M. & Reiss, F. (1996) Catalog and bibliography of Neotropical and Mexican Chironomidae (Insecta, Diptera). Spixiana Supplement, 22, 61–119.

    Spies, M. & Sæther, O.A. (2004) Notes and recommendations on taxonomy and nomenclature of Chironomidae (Diptera). Zootaxa, 752, 1–90.

    Strenzke, K. (1950) Systematik, Morphologie und Ökologie der terrestrischen Chironomiden. Archiv für Hydrobiologie, Supplement, 18, 207–414.

    Strenzke, K. (1960a) Terrestrische Chironomiden. XVIII. Pseudosmittia curticosta (Edw.) Abhandlungen der naturwissenschaftlichen Vereinigung von Bremen, 35 (3), 464–476.

    Strenzke, K. (1960b) Terrestrische Chironomiden XIX–XXIII (Diptera, Chironomidae. Deutsche Entomologische Zeitschrift, 7, 414–441.

    Strenzke, K. & Thienemann, A. (1942) Zwei neue Pseudosmittia-Arten (Diptera, Chironomidae) aus dem Gebiete der Lunzer Seen. Internationale Revue der Gesamten Hydrobiologie und Hydrographie, 42, 356–387.

    Sublette, J.E. & Wirth, W.W. (1972) New genera and species of West Indian Chironomidae (Diptera). The Florida Entomologist, 55 (1), 1–17.

    Swofford, D.L. (2002) PAUP. Phylogenetic Analyses using Parsimony (and Other Methods, 4.0 Beta. Computer program distributed by Sinauer Associates, Inc., Sunderland, MA.

    Thienemann, A. & Strenzke, K. (1940) Terrestrische Chironomiden VI. Pseudosmittia holsata, eine neue Art mit fakultativer Parthenogenese. Zoologischer Anzeiger, 132, 238–244.

    Thienemann, A. & Strenzke, K. (1941) Terrestrische Chironomiden IX. Euphaenocladius Th. Zoologischer Anzeiger, 133, 244–253.

    Tokunaga, M. (1936) Chironomidae from Japan (Diptera) VIII. Marine or seashore Spaniotoma, with description of the immature forms of Spaniotoma nemelione sp. nov. and Tanytarsus boodlææ Tokunaga. Phillipine Journal of Science, 60, 303–321.

    Tokunaga, M. (1939) Chironomidæ from Japan (Diptera), XI. New or little-known midges, with special reference to the metamorphosis of torrential species. Phillipine Journal of Science, 69, 297–346.

    Tokunaga, M. (1940) Chironomidæ from Japan (Diptera), XII. New or little-known Ceratopogonidae and Chironomidae. Phillipine Journal of Science, 72, 255–317.

    Tokunaga, M. (1964) Insects of Micronesia. Diptera: Chironomidae. Bernice P. Bishop Museum, 12, 485–628.

    Tuiskunen, J. (1984) Lindebergia bothnica, gen. n., sp. n. (Diptera, Chironomidae) from Finland. Annales Entomologici Fennicae, 50, 121–122.

    Wang, S. (1990) Pseudosmittia aizaiensis, a new species of Orthocladiinae (Diptera, Chironomidae) from Hunan Province of China. Chinese Journal of Oceanology and Limnology, 8, 273–279.

    Wang, X., Andersen, T. & Sæther O.A. (2006) Neotropical Bryophaenocladius Thienemann, 1934 (Diptera: Chironomidae).

    Willassen, E. & Cranston, P.S. (1986) Afrotropical montane midges (Diptera, Chironomidae, Diamesa). Zoological Journal of the Linnaean Society, 87, 91–123.

    Yamamoto, M. (2004) A catalog of Japanese Orthocladiinae (Diptera: Chironomidae). Makunagi, Acta Dipterologica, 21, 1–121.