Abstract
Six new species are described as males: P. (Pe.) botiense, P. (Pe.) botosaneanui, P. (Pe.) camposense, P. (Pe.) intuber, P. (Pe.) reei, and P. (Pe.) tissamaharense. One additional species is described as female imago. P. (Pe.) pseudotritum and P. (Pe.) utonaiprimum are synonyms of P. (Pe.) tigrinum, and P. (Pe.) takorageheum and P. (Pe.) famibeceum synonyms of P. (Pe.) furusecundum. Five species misidentified as Pentapedilum and one as Polypedilum s. str. have been transferred, four to Polypedilum s. str. [P. (P.) inaijeum Sasa, Kitami et Suzuki, P. (P.) iriodeeum Sasa et Suzuki, P. (P.) jokasecundum Sasa et Ogata, and P. (P.) parasordens Ree], one to Ainuyusurika [A. yakuabeum (Sasa et Suzuki)] and one to P. (Uresi- pedilum). P. (Pe.) pseudotritum and P. (Pe.) utonaiprimum are synonymized with P. (Pe.) tigrinum, and P. (Pe.) tokarageheum and P. (Pe.) famibeceum synonymized with P. (Pe.) furudosecundum. Subgeneric diagnoses are given for all stages and sexes of the subgenus Pentapedilum Kieffer. Keys are provided for all stages and sexes of all available material from all zoogeographical regions. The phylogenetic analysis, which included some species of Polypedilum s. str. and all available Pentapedilum indicate that Pentapedilum is likely monophyletic, while Polypedilum s. str. is polyphyletic. The subgenus Pentapedilum can be divided into three relatively distinct apical groups and the remaining species in a basal Hennigian comb when a parsimony analysis is done. The convexum group consists of one Afrotropical species, and five High, East and South Asian species with one of them also occurring in the Indo-Pacific. The sordens group consists of 13 species from Asia, the Indo-Pacific and Africa. The apparently paraphyletic uncinatum group consists of 9 species from the Holarctic and Oriental regions. The zoogeographical distribution does not support any vicariance patterns, but indicates that the present distribution is caused by dispersal. Diagnostic characters and keys to the males of Ainuyusurika Sasa et Shirasaki is given and Pentapedilum yakuabeum Sasa et Suzuki and Polypedilum (P.) yakucedeum Sasa et Suzuki assigned to the genus. The genus appears to be close to Phaenopsectra Kieffer and is known from Japan only.References
Ashe, P. (1981) A taxonomic note of the genus Polypedilum Kieffer (Diptera: Chironomidae) with an earlier date for the genus and designation as a type-species. Aquatic Insects, 3, 57–59.
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. (1990) Zoogeography of Fresh Waters. General Distribution and Dispersal of Freshwater Animals. AULA, Wiesbaden, pp. 1–512.
Banarescu, P. (1991) Zoogeography of Fresh Waters. Distribution and Dispersal of Freshwater Animals in North America and Eurasia. AULA, Wiesbaden, pp. 513–1092.
Banarescu, P. (1995) Zoogeography of Fresh Waters. Distribution and Dispersal of Freshwater Animals in Africa, Pacific Areas and South America. AULA, Wiesbaden, pp. 1093–1617.
Bolton, M. (1991) The identity of Chironomini Genus C (Diptera: Chironomidae) in Pinder and Reiss (1986). Entomological News, 102, 125–126.
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.R., van Veller, M.G.P. & McLennan, D.A. (2001) How to do BPA, really. Journal of Biogeography, 28, 345–358.
Brooks, D.R., 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.
Chernovskii, A.A. (1949) Opredelitel’ lichinok komarov semeistva Tendipedidae. Opredeliteli po faune SSSR, 31, 1–186.
Cranston, P.S. (1996) Identification guide to the Chironomidae of south Wales. Australian Water Technologies Pty Ltd, West Ryde, NSW. AWT Identification Guide, 1, 1–376.
Cranston, P.S., Dillon, M.E., Pinder, L.C.V. & Reiss, F. (1989) The adult males of Chironominae (Diptera, Chironomidae) of the Holarctic region - keys and diagnoses. Entomologica scandinavica, Supplement, 34, 353–502.
Edwards, F.W. (1929) British non-biting midges (Diptera: Chironomidae). Transactions of the Royal Entomological Society of London, 77, 279–439.
Epler, J.H. (2001) Identification Manual for the Larval Chironomidae (Diptera) of North and South Carolina. A guide to the taxonomy of the midges of the southeastern United States, including Florida. North Carolina Department of Environment and Natural Resources, Raleigh, NC, and St. Johns River Water Management District, Palatka, FL., Special Publication SJ2001-SP13, 526 pp.
Freeman, P. (1954) Chironomidae (Diptera) from Western Cape Province. III. Proceedings of the Royal Entomological Society of London, Series B, 23, 17–25.
Freeman, P. (1955a) Chironomidae (Diptera, Nematocera). Exploration du Parc National Albert, Mission G. F. De Witte, 83, 3–41.
Freeman, P. (1955b) Diptera (Nematocera): Chironomidae. South African animal Life, 2, 361–381.
Freeman, P. (1958) A study of the Chironomidae (Diptera) of Africa South of the Sahara. Part IV. Bulletin of the British Museum of Natural History, Entomology, 6, 263–363.
Freeman, P. (1961) The Chironomidae (Diptera) of Australia. British Museum (Natural History), London, 731 pp.
Freeman, P. & Cranston, P.S. (1980) 11. Family Chironomidae. In: Crosskey, R.W. (Ed.), Catalogue of the Diptera of the Afrotropical Region. Trustees of The British Museum (Natural History), London, pp. 175–202.
Ghonaim, M.F., Ali, A. & Osheibah, A. (2005) Polypedilum (Diptera: Chironomidae) from Egypt with description of a new species. Aquatic Insects, 27, 221–230.
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. (1942) Ceratopogonidae et Chironomidae nouveaux ou peu connus d’Europe. Bulletin du Musée Royal d’Histoire Naturelle de Belgique, 18(46), 1–16.
Guha, D.K. & Chaudhuri, P.K. (1981) Record of three genera of Chironominae (Chironomidae: Diptera) from India. Bulletin of the Zoological Survey of India, 3, 159–165.
Guha, D.K. & Chaudhuri, P.K. (1985) Pentapedilum robusticeps - a new species from India. Bulletin of the Zoological Survey of India, 3, 195–197.
Harrison, A.D. (2000) Four new genera and species of Chironomidae (Diptera) from southern Africa. Aquatic Insects, 22, 219–236.
Harrison, A.D. (2001) Chironomidae (Diptera) in the Albany Museum, Part 1. Annals of the Eastern Cape Museums, 2, 9–18.
Hashimoto, H. (1983) Pentapedilum (Diptera, Chironomidae) from Japan, with description of a new species. Kontyu, Tokyo, 51, 17–24.
Johannsen, O.A. (1905) Aquatic nematocerous Diptera II. Chironomidae. In: Needham, J.G., Morton, K.J. & Johannsen, O.A. (Eds), May flies and midges of New York. New York State Museum Bulletins, 86, 76–327, pl. 16–37.
Johannsen, O.A. (1932) Chironominae of the Malayan subregion of the Dutch East Indies. Archiv für Hydrobiologie, Supplement, 11, 503–552.
Johannsen, O.A. (1934) New species of North American Ceratopogonidæ and Chironomidæ. New York Entomological Society Journal, 42, 343–352.
Kawai, K., Ohsugi, T. & Imabayashi, H. (2006) Morphological and genetical relationships among the Japanese chironomid species of the genus Polypedilum. Abstracts Volume, Sixth International Congress of Dipterology, Fukuoka, Japan, p. 131.
Kieffer, J.J. (1909) Diagnoses de nouveaux Chironomides d’Allemagne. Bulletin de la Société d'Histoire naturelle de Metz, 26, 37–56.
Kieffer, J.J. (1911) Nouvelle descriptions des Chironomides obtenues d’éclosion. Bulletin de la Société d'Histoire naturelle de Metz, 27, 1–60.
Kieffer, J.J. (1912) H. Sauter’s Formosa-Ausbeute. Tendipedidae (Chironomidae) (Dipt.). Supplementa entomologica, 1, 27–43.
Kieffer, J.J. (1913) Nouveaux Chironomides (Tendipédides) d'Allemagne. Bulletin de la Société d'Histoire naturelle de Metz, 28, 7–35.
Kieffer, J.J. (1921a) Chironomides nouveaux ou peu connus de la région paléarctique. Bulletin de la Société d’Histoire Naturelle de Metz, 29, 51–109.
Kieffer, J.J. (1921b) Chironomides de Courlande. Annales de la Société Scientifiques Bruxelles, 40, 275–298.
Kieffer, J.J. (1921c) Chironomides des Philippines et de Formose. Philippine Journal of Science, 18, 557–593.
Kieffer, J.J. (1922) Chironomides nouveaux ou peu connus de la région paléarctique. Annales de la Société Scientifiques Bruxelles, 42, 71–128.
Kieffer, J.J. (1923) Chironomides de l’Afrique équatoriale (3e partie). Annales de la Société entomologique de France, 92, 149–204.
Kieffer, J.J. (1924) Chironomides nouveaux ou rares de l’Europe centrale. Bulletin de la Société d’Histoire Naturelle de Metz, 30, 11–110.
Kieffer, J.J. (1926) Neue Chironomiden aus dem Deutschen Entomologischen Instituts zu Berlin-Dahlem. Entomologische Mitteilungen, 15, 103–106.
Kobayashi, T., Ohtaka, A. & Takahashi, T. (2003) The second record of ectoparasitic Chironomidae on Trichoptera from Japan, Polypedilum (Cerobregma) kamotertium Sasa, 1989. Spixiana, 26, 83–91.
Kulshresta, A.K. (1979) First record of the genus Pentapedilum Kieffer from India. Oriental Insects, 13(304), 303–306.
Langton, P.H. & Visser, H. (2003) Chironomidae exuviae - a key to pupal exuviae of the West Palaearctic Region. CD-ROM, Expert Center for Taxonomic Information, Amsterdam.
Lenz, F. (1937) Chironomariae aus Niederländisch Indien. Larven und Puppen. Archiv für Hydrobiologie, Supplement, 15, 1–29.
Maschwitz, D.E. & Cook, E.F. (2000) Revision of the Nearctic species of the genus Polypedilum Kieffer (Diptera: Chironomidae) in the subgenera P. (Polypedilum) and P. (Uresipedilum) Oyewo and Sæther. Bulletin of the Ohio Biological Survey, New Series, 12, 11–35.
Morrone, J.J. & Crisci, J.V. (1995) Historical biogeography: introduction to methods. Annual Review of Ecology and Systematics, 26, 373–401.
Moubayed, J. (1992) On the genus Polypedilum Kieffer, 1912 from Lebanon. Description of three new species. Spixiana, 15, 197–205.
Oliver, D.R., Dillon, M.E. & Cranston, P.S. (1990) A catalog of Nearctic Chironomidae. Research Branch Agriculture Canada, Publication 1857/B, 1–89.
Oyewo, E.A. &. Sæther, O.A. (1998) Revision of Afrotropical Polypedilum Kieffer subgenus Uresipedilum Sasa et Kikuchi, 1995 (Diptera: Chironomidae), with a review of the subgenus. Annales of Limnology, 34, 315–362.
Oyewo, E.A. & Jacobsen, R.E. (2007) Polypedilum (Pentapedilum) epleri, a new species from USA (Diptera: Chironomidae). In: Andersen. T. (Ed.), Contributions to the Systematics and Ecology of Aquatic Diptera - A Tribute to Ole A. Sæther. The Caddis Press, Columbus, Ohio, pp. 225–234.
Picado, C. (1913) Les Broméliacées épiphytes considérées comme milieu biologique. Bulletin Scientifique de la France et de la Belgique, 47, 280–295.
Pankratova, V.Ya. (1983) Lichinki i kukolki komarov podsemeistva Chironominae fauny SSSR (Diptera, Chironomidae = Tendipedidae). Opredeliteli po Faune SSSR, Isdavaemye Zoologicheskim Institutom Akademii Nauk SSSR, 134, 1–296.
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
Pinder, L.C.V. & Reiss, F. (1983) The larvae of Chironominae (Diptera, Chironomidae) of the Holarctic region - keys and diagnoses. Entomologica scandinavica, Supplement, 19, 293–435.
Pinder, L.C.V. & Reiss, F. (1986) The pupae of Chironominae (Diptera, Chironomidae) of the Holarctic region - keys and diagnoses. Entomologica scandinavica, Supplement, 28, 299–456.
Ree, H.I. (1992) Three new species of Chironomidae (Diptera) from Korea. Korean Journal of systematic Zoology, special Issue, 3, 129–138.
Ree, H.I. & Kim, H.S. (1988) Studies on Korean Chironomidae (Diptera) III. Description of two unrecorded species from Korea and three new species. Korean Journal of systematic Zoology, special Issue, 2, 13–24.
Sæther, O.A. (1977) Female genitalia in Chironomidae and other Nematocera: morphology, phylogenies, keys. Bulletin of the Fisheries Research Board of Canada, 197, 1–209.
Sæther, O.A. (1980) Glossary of chironomid morphology terminology (Chironomidae: Diptera). Entomologica scandinavica, Supplement, 14, 1–51.
Sæther, O.A. (1990) A review of the genus Limnophyes Eaton from the Holarctic and Afrotropical regions. Entomologica scandinavica, Supplement, 35, 1–139.
Sæther, O.A. & Spies, M. (2004) Fauna Europaea: Chironomidae. Fauna Europaea version 1.1. Available from: htpp://www.faunaeur.org. (30 May 2008).
Sæther, O.A. & Sundal, A. (1999) Cerobregma, a new subgenus of Polypedilum Kieffer. With a tentative phylogeny of subgenera and species groups within Polypedilum (Diptera: Chironomidae). Journal of the Kansas Entomological Society, 71, 311–378.
Sasa, M. (1979) A morphological study of adults and immature stages of 20 Japanese species of the family Chironomidae (Diptera). Research Report from the National Institute for Environmental Studies, Japan, 7, 1–148.
Sasa, M. (1983) Studies on chironomid midges of the Tama River - Part 5. An observation on the distribution of Chironominae along the main stream in June, with description of 15 new species. Research Report from the National Institute for Environmental Studies, Japan, 43, 1–67.
Sasa, M. (1985) Studies on the chironomid collected from lakes in southern Kyushu (Diptera, Chironomidae). Research Report from the National Institute of Environmental Studies, Japan, 83, 25–99.
Sasa, M. (1988a) Studies on the chironomid midges collected from lakes and streams in the southern region of Hokkaido, Japan. Research Report from the National Institute of Environmental Studies, Japan, 121, 8–76.
Sasa, M. (1988b) Chironomid midges collected on the shores of lakes in the costal region of Abasiri, northern Hokkaido. Research Report from the National Institute of Environmental Studies, Japan, 121, 77–89.
Sasa, M. (1989) Chironomid midges of some rivers in western Japan. Toyama Prefectural Environmental Pollution Research Centre, 1989, 45–110.
Sasa, M. (1994) Additional information on the Chironomidae of Japan, 1994. Toyama Prefectural Environmental Pollution Research Centre, 1994, 28–67.
Sasa, M. (1996) 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), 16–112.
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. (1983) Chironomid midges of the tribe Chironomini recovered from sewage ditches, eutrophicated ponds and clear streams of the Ryukyu Islands, southern Japan. Japanese Journal of Sanitary Zoology, 34, 305–341.
Sasa, M. & Kikuchi, M. (1995) Chironomidae (Diptera) of Japan. University of Tokyo Press, Tokyo, 333 pp.
Sasa, M., Kitami, K. & Suzuki, H. (2001) Additional studies on the chironomid midges collected on the shore of Lake Inawashiro. Noguchi Hideyko Kinenkaii, (The Dr. Nogichi Memorial Hall), Inawashiro-machi, Fukushima-ken, Japan, pp. 1–38.
Sasa, M. & Ogata, T. (1999) Taxonomic studies on the chironomid midges collected from the Kurobe Municipal Sewage Treatment Plant. Medical Entomology and Zoology, 50, 85–104.
Sasa, M. & Okazawa, T. (1991) Studies on the chironomids of the Joganji River and other places. Part 1. Studies on the chironomids of the Joganji River, Toyama. Research Report from Toyama Prefectural Environmental Pollution Research Centre, 1991, 52–67.
Sasa, M. & Suzuki, H. (1995) The chironomid species collected on the Tokara Islands Kagoshima (Diptera). Japanese Journal of Sanitary Zoology, 46, 255–288.
Sasa, M. & Suzuki, H. (1999) Studies on the chironomid midges of Tsushima and Iki Islands, Western Japan. Part 1. Species of Chironominae collected on Tsushima. Tropical Medicine, 41, 1–53.
Sasa, M. & Suzuki, H. (2000a) Studies on the chironomid midges collected on Ishigaki and Iriomote Islands, southwestern Japan. Tropical Medicine, 42, 1–37.
Sasa, M. & Suzuki, H. (2000b) Studies on the chironomid species collected on Yakushima Island, southwestern Japan. Tropical Medicine, 42, 53–134.
Sasa, M. & Suzuki, H. (2001) Studies on the chironomid species collected in Hokkaido in September, 2000. Tropical Medicine, 43, 1–38.
Sasa, M. & Suzuki, H. (2002) Studies on the species of family Chironomidae (Diptera) collected on Minamidaito Island, Okinawa, South Japan. Part 1. Tropical Medicine, 43(3/4), 61–92.
Skuse, F.A.A. (1889) Diptera of Australia. Part VI. The Chironomidae. Proceedings of the Linnean Society of New South Wales, 2, 215–311.
Sublette, J.E. & Sublette, M.S. (1983) Asheum, a replacement name in Chironomidae (Diptera). Entomological News, 94, 34.
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. (1916) Die Chironomidenfauna der Eifelmaare. Mit Beschreibung neuer Arten von. Prof. Dr. Kieffer. Verhandlungen des Naturhistorischen Vereines der Preussischen Rheinlands und Westfalens, 72, 1–58.
Tokunaga, M. (1940a) Ceratopogonidæ and Chironomidæ from the Micronesian Islands. With biological notes by Teiso Esaki. Philippine Journal of Science, 71, 205–229.
Tokunaga, M. (1940b) Chironomoidea from Japan (Diptera), XII. New or little-known Ceratopogonidæ and Chironomidæ. Philippine Journal of Science, 72, 255–311.
Tokunaga, M. (1964) Insects of Micronesia. Diptera: Chironomidae. Insects of Micronesia,12, 485–628.
Townes, H.K. (1937) Biology, misdetermination of fulvescens Johannsen. Annual Report, NY State Conservation Department, 27, supplementary, 171.
Townes, H.K. (1945) The Nearctic species of Tendipedini (Diptera: Tendipedidae (= Chironomidae)). American Midland Naturalist, 34, 1–206.
Vårdal, H., Bjørlo, A. & Sæther, O.A. (2002) Afrotropical Polypedilum subgenus Tripodura, with a review of the subgenus (Diptera: Chironomidae). Zoologica Scripta, 31, 331–402.
van Veller, M.G.P., Zandee, M. & Kornet, D.J. (1999) Two requirements for obtaining valid common patterns under different assumptions in vicariance biogeography. Cladistics, 15, 393–406.
van Veller, M.G.P., Kornet, D.J. & Zandee, M. (2000) Methods in vicariance biogeography: assessment of the implementations of assumptions 0, 1, and 2. Cladistics, 16, 319–345.
Walker, F. (1856) Insecta Britannica. Diptera. Vol. III. Lovell & Reeve, London, pp. i–xxiv + 1–352, pl. 21–30.
Wiley, E. (1987) Methods in vicariance biogeography. In: Hovenkamp, P., Gittenberger, E., Hennipman, E., Jong, R. de, Roos, M.C., Sluys, R. & Zandee, M. (Eds), Systematics and Evolution: a Matter of Diversity. Utrecht University, Utrecht, pp. 238–306.
Wiley, E. (1988a) Parsimony analysis and vicariance biogeography. Systematic Zoology, 37, 271–290.
Wiley, E. (1988b) Vicariance biogeography. Annual Review of Ecology and Systematics, 19, 513–542.
Wulp, F.M. van der (1874) Dipterologische aanteekeningen. Tijdschrift voor Entomologie, 17, 109–148.
Zhang, R. & Wang, X. (2004) Polypedilum (Uresipedilum) Oyewo and Sæther from China (Diptera: Chironomidae). Zootaxa, 565, 1–38.
Zhang, R. & Wang, X. (2005) Polypedilum (Cerobregma) Sæther & Sundal from China (Diptera: Chironomidae). Aquatic Insects 27, 47–55.
Zhang, R., Wang, X. & Sæther, O.A. (2006) Two unusual species of Polypedilum Kieffer (Diptera: Chironomidae) from Oriental China. Zootaxa, 1282, 39–48.
Zorina, O.V. & Makarchenko, E.A. (2001) Four new species of the genus Polypedilum Kieffer (Diptera: Chironomidae) from the Russian Far East. Studia Dipterologica, 7, 483–490.