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Type: Monograph
Published: 2008-07-30
Page range: 1–100
Abstract views: 44
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Review of the Neotropical blackfly subgenus Chirostilbia Enderlein (Diptera: Simuliidae) based on adults and pupal morphology

Diptera Division, Simuliidae and Onchocerciasis Research Programme, Department of Entomology, The Natural History Museum, Cromwell Road, SW7 5BD, London, UK
Diptera Division, Simuliidae and Onchocerciasis Research Programme, Department of Entomology, The Natural History Museum, Cromwell Road, SW7 5BD, London, UK
Laboratório de Simulídeos e Oncocercose, Instituto Oswaldo Cruz, Rio de Janeiro, RJ, Brasil
Laboratório de Simulídeos e Oncocercose, Instituto Oswaldo Cruz, Rio de Janeiro, RJ, Brasil
Diptera Simuliidae black fly Neotropical region taxonomy Brazil genus Simulium subgenus Chirostilbia

Abstract

The species of the subgenus Chirostilbia are reviewed based on the adult and pupal morphology. All main taxonomic characters are fully illustrated with colour digital images and keys for species identification are given. Taxonomic discussions for each species and summaries of their distribution and biology are also provided. Thirteen species are now considered to be valid in the subgenus Chirostilbia. Since they are similar morphologically no separation into species groupshas been attempted. Lectotypes are designated for the following species: S. distinctum Lutz and Trichodagmia lutziana Enderlein [= S. pertinax Kollar]. Simulium nilesi Rambajan is transferred from its synonymy with S. perflavum Roubaud in the subgenus Psilopelmia Enderlein to Chirostilbia as a junior synonym of S. subpallidum Lutz. The thoracic pattern of the male of S. empascae is described and illustrated for the first time.

References

  1. Andrade, C. F. S. & Branco, A. C. Jr. (1991) Susceptibilidade de populaces de Simulium (Chirostilbia) pertinax Kollar, 1832 (Culicomorpha, Simuliidae) ao temephos e a um formulado a base de Bacillus thuringiensis var. israelensis Revista de Saúde Pública, 25, 367–370.

    Adler, P.H. & Crosskey, R.W. (2008) World blackflies (Diptera: Simuliidae): a fully revised edition of the taxonomic and geographical inventory. http://entweb.clemson.edu/biomia/pdfs/blackflyinventory.pdf/[ last accessed 7.June.2008]

    Araújo-Coutinho, C.J.P.C. & Lacey, L.A. (1990) Field evaluation of Bacillus thuringiensis var. israelensis for control of black flies in the north littoral zone of Brazil’s São Paulo State. Bulletin of PAHO, 24, 307–313.

    Araújo-Coutinho, C.J.P.C., Cunha, A.B.P.V., Serra-Freire, N.M. & Pinto de Mello, R. (2003) Evaluation of the impact Bacillus thuringiensis serovar israelensis and temephos, used for the control of Simulium (Chirostilbia) pertinax Kollar, 1832 (Diptera, Simuliidae) on the associated entomofauna, Paraty, State of Rio de Janeiro, Brazil. Memórias do Instituto Oswaldo Cruz, 98, 697–702.

    Branco, A. C.Jr. (1999) Effects of Polydispyrenia simulii (Microspora; Duboscqiidae) on development of the gonads of Simulium pertinax (Diptera; Simuliidae). Memórias do Instituto Oswaldo Cruz, 94, 421–424.

    Campos, J. & Andrade, C. F. S. (2002) Resisténcia a inseticidas em populaces de Simulium (Diptera, Simuliidae). Cadernos de Saúde Pública, 18, 661–671.

    Campos, J., Andrade, C.F.S. & Recco-Pimentel, S.M. (2001) Chromosomal comparisons among and within populations of Simulium (Chirostilbia) pertinax (Diptera, Simuliidae). Memórias do Instituto Oswaldo Cruz, 96, 365–369.

    Campos, J., Recco-Pimentel, S. M., Andrade, C. F. S. (1996) Polytene chromosome analysis of a population of Simulium pertinax (Diptera: Simuliidae). Brazilian Journal of Genetics, 19, 47–52.

    Cavados, C.F.G., Fonseca, R.N., Chaves, J.Q., Araújo-Coutinho, C.J.P.C. & Rabinovitch, L. (2005) A new black fly isolate of Bacillus thuringiensis autoagglutinating strain highly toxic to Simulium pertinax (Kollar) (Diptera, Simuliidae) larvae. Memórias do Instituto Oswaldo Cruz, 100, 795–797.

    Cerqueira, N. & Barbosa de Almeida, F. (l970) Simuliidae de Amazônia. V. Descrição de Simulium septentrionale sp. n. (Diptera, Nematocera). Boletim do INPA (Manaus), Patologia Tropical, 2, 1–11.

    Coscarón, S. (1981) Notas sobre simulidos Neotropicales XI. Sobre el subgénero Simulium (Chirostilbia) Enderlein, con la descripción de dos species nuevas del S.E. del Brasil (Diptera, Insecta). Revista de la Sociedad Entomológica Argentina, 40, 157–164.

    Coscarón, S. (1982) Notas sobre simulidos Neotropicales X. Sobre un grupo peculiar de especies del genero Simulium del S.E. de Brasil, Paraguay y N.O. de Argentina (Diptera-Insecta). Revista de la Sociedad Entomológica Argentina, 41, 65–76.

    Coscarón, S. (1987) El género Simulium Latreille en la región neotropical: análisis de los grupos supraespecíficos, especies que los integran y distribución geográfica (Simuliidae, Diptera). 112 pp. Museu Paraense Emílio Goeldi, Belém, Brazil.

    Coscarón, S. (1991) Simuliidae. Fauna de Agua Dulce de la República Argentina 38 (Insecta Diptera) (2). FECIC, Buenos Aires, Argentina, 304 pp.

    Coscarón, S. & Coscarón-Arias, C.L. (1997) Cladistic analysis of the subgenera Inaequalium, Psaroniocompsa, and Chirostilbia of the genus Simulium, with comments on their distribution. Revista de la Sociedad Entomológica Argentina, 56, 109–121.

    Coscarón & Coscarón-Arias (2007) Neotropical Simuliidae (Diptera: Insecta). In Adis, J., Arias, J.R., Rueda-Delgado, G. & K.M.Wantzen, eds., Aquatic Biodiversity in Latin America (ABLA). Vol. 3. Pensoft, Sofia-Moscow, 685 pp.

    Coscarón, S. & Wygodzinsky, P. (1972) Notas sobre simulidos neotropicales. Sobre tres especies de jejenes con hembras de color claro del S. E. del Paraguay y del N. E. de Argentina (Simuliidae-Diptera). Revista del Museo de La Plata (N.S.) Sección Zoología, 11, 203–231.

    Crosskey, R.W. (1988) An annotated checklist of the world blackflies (Diptera: Simuliidae). Pp. 425–520. In Kim, K.C. & Merritt, R.W., eds., Blackflies: Ecology, population management, and an annotated world list. The Pennsylvania State University, University Park.

    Crosskey, R. W. & Howard, T.M. (1997): A new taxonomic and geographical inventory of world blackflies (Diptera: Simuliidae). The Natural History Museum, London, 144 pp.

    Crosskey, R.W. & Howard, T.M. (2004) A revised taxonomic and geographical inventory of world blackflies (Diptera: Simuliidae). The Natural History Museum, London, 82 pp. [web version: http://www.nhm.ac.uk/entomology/projects/blackflies/Inventory.pdf] (last accessed 19 September 2007).

    D'Andretta Jr, C. (1954) Sobre Simulium distinctum Lutz, 1910 (Diptera: Simuliidae). Anais da Facultade é Odontología da Universidade de São Paulo, 12, 63–66.

    D’Andretta, C, Jr. & D’Andretta, M.A.V. (1950) Especies Neotropicais da família Simuliidae Schiner (Diptera Nematocera) VI- Redescrição de Simulium pertinax Kollar, 1832. Papéis Avulsos, Departamento do Zoologia, São Paulo, 9, 193–213.

    Eaton, D.P., Diaz, L.A., Hans-Filho, G., Santos, V. D., Aoki, V., Friedman, H., Rivitti, E.A., Sampaio, S.A.P., Gottlieb, M.S., Giudice, G. J., Lopez, A. & Cupp, E. W. (1998) Comparison of black fly species (Diptera: Simuliidae) on an Amerindian reservation with a high prevalence of fogo selvagem to neighbouring disease-free sites in the state of Mato Grosso do Sul, Brazil. Journal of Medical Entomology, 35, 120–131.

    Edwards, F.W. (1931) Simuliidae. Pp. 121–154 in Diptera of Patagonia and South Chile, Part II Nematocera (excluding crane-flies and Mycetophilidae). 331pp. British Museum (Natural History), London.

    Enderlein, G. (1921a) Das System der Kriebelmücken (Simuliidae). Deutsche Tierärztliche Wochenschrift, 29, 197–200. [Actual publication date was April 1921.]

    Enderlein, G. (1921b) Neue paläarktische Simuliiden. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin, 1921, 212–224. [Actual publication date was June 1921.]

    Enderlein, G. (1921c) Die systematische Gliederung der Simuliiden. Zoologischen Anzeiger, 3, 43–46.

    Enderlein, G. (1934) Weiterer Ausbau des Systems der Simuliiden. (Dip.). Deutsche Entomologische Zeitschrift, 1933, 273–292.

    Enderlein, G. (1935) Neue Simuliiden, besonders aus Afrika. Sitzungsberichte der Gesellschaft naturforschender Freunde 1935, 358–364.

    Gil-Azevedo, L.H. & Maia-Herzog, M. (2004) Registros novos de ocorrência de seis especies de Simuliidae (Diptera) para o estado do Rio de Janeiro. Biotrópica, 4, 1–3.

    Hamada, N. & Pepinelli, M. (2004) Simulium (Chirostilbia) bifenestratum (Diptera, Simuliidae), a new black-fly species from Atlantic Forest, State of São Paulo, Brazil. Memórias do Instituto Oswaldo Cruz, 99, 45–52.

    Hamada, N., Hernández, L.M., Luz, S.L.B. & Pepinelli, M. (2006) Simulium (Chirostilbia) jefersoni, a new species of black fly (Diptera: Simuliidae) from the state of Bahía, Brazil. Zootaxa, 1123, 21–37.

    Hernández, L.M. & Shelley, A.J. (2005) New specific synonymies and taxonomic notes on Neotropical blackflies (Diptera: Simuliidae). Zootaxa, 853, 1–46.

    Hernández, L.M. & Shelley, A.J., Luna Dias, A.P.A. & Maia-Herzog, M. (2005) Re-description of the female, male and pupa of Simulium itaunense D’Andretta & González B. (Diptera: Simuliidae). Zootaxa, 896, 1–14.

    Hernández, L.M., Shelley, A.J., Luna Dias, A.P.A. & Maia-Herzog, M. (2007) New specific synonymies and taxonomic notes in Neotropical Blackflies (Diptera: Simuliidae) belonging to the subgenera Chirostilbia Enderlein, Hemicnetha Enderlein, Inaequalium Coscarón & Wygodzinsky, Psaroniocompsa Enderlein and Psilopelmia Enderlein. Zootaxa, 1580, 1–80.

    International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature, 4thEdition, The International Trust for Zoological Nomenclature, xix + 273 pp.

    Kollar, V. (1832) Die vorzüglich lästigen Insecten Brasiliens. In: J.E.Pohl (ed.): Reise im Innern von Brasilien, 1, 101–119, Wien, 448pp.

    Knowlton, G.F. & Rowe, J.A. (1934) New blood flies from Utah (Simuliidae, Diptera). Annals Entomological Society of America, 27, 580–584.

    Lane, J. & Porto, C.E. (1939) Simulideos da região neotropical: o gênero Eusimulium. Boletim Biológico (n. s.) (São Paulo), 4, 168–176.

    Lane, J. & Porto, C.E. (1940) Simulideos da região neotrópical. III - Descrição de novas espécies dos generos Simulium e Eusimulium. Arquivos do Instituto Biologico, 11, 189–195.

    Lozovei, A.L., Cunha, M.C. I. & Oliveira, V.L.R. (1992) Controle fisico de simulideos (Diptera, Simuliidae) em vertedouros de açudes de piscicultura e no leito do rio Dom Rodrigo, Campo Largo, Paraná, Brasil. Arquivos de Biologia e Tecnologia, 35, 679–684.

    Lutz, A. (1910) Segunda contribuição para o conhecimento das especies brazileiras do genero "Simulium". Memórias do Instituto Oswaldo Cruz, 2, 213–267.

    Lutz, A. (1922) Zoologia Médica. Nematoceros hematofagos não pertecendo aos Culicideos. A Folha Médica, 12, 89–92.

    Maia-Herzog, M., Felippe-Bauer, M.L., Malaguti, R. & Carvalho Leite, T. C. (1988) A contribution to the study of Simulium and Culicoides of Rio de Janeiro: monthly incidence and biting activity. Memórias do Instituto Oswaldo Cruz, 83, 95–99.

    Magni, S.T. & Py-Daniel, V. (1989) Aegla platensis Schmitt, 1942 (Decapoda: Anomura) um predador de imaturos de Simuliidae (Diptera: Culicomorpha). Revista de Saúde Pública, 23, 258–259.

    Nunes de Mello, J.A. & Barbosa de Almeida, N. (1974) Simuliidae da Amazônia VII. Descrição de Simulium pseudoexiguum sp. n. (Diptera, Nematocera). Acta Amazônica, 4, 65–68.

    Pegoraro, R.A. (1987) Longevidade de Simulim (Chirostilbia) pertinax Kollar, 1832 (Diptera: Simuliidae) em ambiente controlado, com diferente dietas. Anais da Sociedade Entomológica do Brasil, 16, 315–324.

    Pegoraro, R.A. (1993) Ciclo biológico de Simulim (Chirostilbia) pertinax Kollar, 1832 (Diptera: Simuliidae). Anais da Sociedade Entomológica do Brasil, 22, 29–38.

    Pepinelli, M., Trivinho-Strixino, S. & Hamada, N. (2003a) Description of Simulium (Chirostilbia) friedlanderi Py-Daniel, 1987 (Insecta: Diptera: Simuliidae) female. Biota Neotrópica, 3, 1–6.

    Pepinelli, M., Trivinho-Strixino, S. & Hamada, N. (2003b) New records of Simuliidae (Diptera, Nematocera) in the state of São Paulo, Brazil. Revista Brasileira de Entomologia, 47, 653–655.

    Pepinelli, M., Hamada, N. & Trivinho-Strixino, S. (2006) Larval description of Simulium (Notolepria) cuasiexiguum and Simulium (Chirostilbia) obesum and new records of black fly species (Diptera: Simuliidae) in the States of São Paulo and Minas Gerais. Neotropical Entomology, 35, 698–704.

    Pinto, C. (1932) Simuliidae da America Central e do Sul. Reunión de la Sociedad Argentina de Patologia Regional del Norte [1931], 7, 661–763 + i [corrigenda]

    Pohl, J.E. (1832) Reise im Innern von Brasilien. Erster Theil [=Part I] xxx+448pp. Anton Strauss, Wien [=Vienna]. [pp.101–119 of this work form a section entitled “Die vorzűglich lästigen Insecten Brasiliens” in which the description of Simulium pertinax by Kollar is on p. 117. Kollar is not named as co-author of this section, but is named so on the version issued as a separate (see Pohl & Kollar, 1832). The legend to the plate occupies pp.118 and 119.]

    Pohl, I. [=J.] E. & Kollar, V. (1832) Brasiliens vorzűglich lästige Insecten. 20pp. Wien. [This work contains the description of Simulium pertinax by Kollar on p.19. It is extracted and re-paginated from pp. 101–119 in Pohl, 1832. It differs from the original in attributing co-authorship to Kollar on the title page and in the differently set legend (p.20) to the plate.]

    Py-Daniel, V. (1981) Algumas considerações sôbre Simuliidae (Diptera: Nematocera) II. Acta Amazônica 11, 171–181.

    Py-Daniel, V. (1987) Simuliidae (Diptera: Culicomorpha) no Brasil. V. Sobre o Simulium (Chirostilbia) friedlanderi sp. n. e revisão do Simulium (C.) laneportoi Vargas, 1941. Revista da Saúde Pública, 21, 331–341.

    Py-Daniel, V. (1989) Novas sinonímias e Correçeós em Simulium com a revalidação de S. pruinosum Lutz, 1904 [error for 1910] (Culicomorpha, Simuliidae). Revista de Saúde Pública, 23, 254–257.

    Py-Daniel, V. & Darwich, S.M. (1997) Simuliid “Borrachudo/Pium” control in Brazil. The British Simuliid Group Bulletin, 9, 7–10.

    Py-Daniel, V. & Moreira, G.R.P. (1988) Simuliidae (Diptera, Culicomorpha) no Brasil VII. Sobre o Simulium (Chirostilbia) empascae sp. n. IHERINGIA. Série Zoologia, 67, 77–86.

    Py-Daniel, V. & Moreira Sampaio, R.T. (1994a) Jalacingomyia gen.n. (Culicomorpha); a ressurreição de Gymnopaidinae; a eliminação do nivel tribal; apresentação de novos caracteres e redescrição dos estágios larval e pupal de Simulium columbaschense (Fabricius, 1787) (Diptera: Simuliidae). Memorias del CAICET, 4, 101–148.

    Py-Daniel, V. & Moreira Sampaio, R.T. (1994b) Actualização nomenclatural para Simuliidae Neotropical (Diptera, Culicomorpha, Simuliidae). Memórias del CAICET, 4, 149–156.

    Py-Daniel, V. & Moreira Sampaio, R.T. (1995) Gêneros e espécies de Simuliidae (Diptera: Culicomorpha) assinalados para o Brasil até 1995. Entomologia y Vectores, 5, 117–121.

    Py-Daniel, V. & Shelley, A.J. (1980) Revisão do Simulium spinibranchium Lutz, 1910 (Diptera: Simuliidae), com a primeira descrição dos adultos e larva, e redescrição da pupa. Acta Amazonica, 10, 213–223.

    Py-Daniel, V., Souza, M.A.T. & Caldas, E.P. (1988) Simuliidae (Diptera, Culicomorpha) no Brasil. III. Sobre o Simulium (Chirostilbia) riograndense sp. n. e revisão do Simulium (Chirostilbia) distinctum Lutz, 1910. INHERINGIA, Série Zoología, 67, 37–57.

    Rabinovitch, L., Cavados, C. F. G., Chaves, J. Q., Araujo-Coutinho, C. J. P. C. A., Zahner, V., Silva, K.R.A. & Seldin, L. (1999) A new strain of Bacillus thuringiensis serovar israelensis very active against blackfly larvae. Memórias do Instituto Oswaldo Cruz, 94, 683–685.

    Rambajan, I. (1979) A new species of (Simuliidae: Diptera) from the Lethem area, Rupununi District, Guyana, South America. Journal Guyana Museum and Zoo of the Royal Agricultural and Commercial Society , 45, 63–77.

    Ramírez Pérez, J. (1971) Distribución geográfica y revisión taxonómica de los simúlidos (Diptera: Nematocera) de Venezuela con descripción de diez especies nuevas. Acta Biológica Venezolana 7, 271–371.

    Ramírez Pérez, J. (1983) "Los jejenes de Venezuela". Simposio de Oncocercosis Americana. Caicet. Puerto Ayacucho, 15–17 Octubre, 193, iii + 156 pp.

    Ramírez Pérez, J., Yarzábal, L. & Peterson, B. (1982) La simuliofauna del Territorio Federal Amazonas (Venezuela). Publicación Científica, 1, 1–104. Proicet Amazonas.

    Santos Júnior, F.E.dos, Strieder, M.N., Fiorenting, G.L. & Neiss, U.G. (2007) Velocidade da água e a distribuição de larvas e pupas de Chirostilbia pertinax (Kollar). Revista Brasileira de Entomologia, 51, 62–66.

    Schrottky, C. (1909) Drei neue blutsaugende Dipteren aus Paraguay. Zeitschrift für wissenschaftliche Insektenbiologie, Berlin, 5, 61–63.

    Shelley, A.J. & Py-Daniel, V. (1981) Simuliidae of Goiás State and the Federal District of Brasília. 1. Simulium dekeyseri, new species. Memórias do Instituto Oswaldo Cruz, 76, 23–32.

    Shelley A.J., Hernández L.M., Davies, J.B. (2004) Blackflies (Diptera: Simuliidae) of southern Guyana with keys for the identification of adults and pupae- A Review. Memórias do Instituto Oswaldo Cruz, 99, 443–470.

    Shelley, A.J., Hernández, L.M., Maia-Herzog, M., Luna Dias, A.P.A. & Luz, S.B. (2006) An interpretation of the morphological variation in the Simulium amazonicum species group (Diptera: Simuliidae) of Latin America. Zootaxa, 1274, 1–68.

    Shelley, A.J., Luna Dias, A.P.A. & Maia-Herzog, M. (1984) New specific synonymy in Neotropical Simulium s.l. (Diptera: Simuliidae). Memórias do Instituto Oswaldo Cruz, 79, 143–161.

    Shelley, A.J., Lowry, C.A., Maia-Herzog, M., Luna Dias, A.P.A. & Moraes, M.A.P. (1997) Biosystematic studies on the Simuliidae (Diptera) of the Amâzonia onchocerciasis focus of Brazil. Bulletin of the Natural History Museum (Entomology series), 66, 1–121.

    Shelley, A.J., Maia-Herzog, M., Lowry, C.A., Luna Dias, A.P.A., Garritano, P.R., Shelley, A., Camargo, M. & Carter, H.G. (2000) The Simuliidae (Diptera) of the secondary onchocerciasis focus at Minaçu in central Brazil. Bulletin of The Natural History Museum (Entomology series), 69, 171–221.

    Smart, J. (1945) The classification of the Simuliidae (Diptera). Transactions of the Royal Entomological Society (London), 95, 463–532.

    Stone, A. (1962) Notes on the types of some Simuliidae (Diptera) described by Enderlein. Annals of the Entomological Society of America, 55, 206–209.

    Stone, A. (1963) An annotated list of genus-group names in the family Simuliidae (Diptera). U.S. Department of Agriculture Technical Bulletin 1284, 1–28.

    Strieder, M.N., Corseuil, E. & Py-Daniel, V. (1992) Espécies do genero Simulium (Diptera, Simuliidae) ocorrentes no Rio Grande do Sul – Brasil, com chaves para sua identificação. Acta Biologica Leopoldensia, 14, 53–73.

    Vargas, L. (1941) Simulium lane-portoi n.n. (Simuliidae, Dipt.) y lista de simúlidos mexicanos. Revista del Instituto de Salubridad y Enfermedades Tropicales, 2, 115–122.

    Vargas, L. (1945) Simúlidos del Nuevo Mundo. — Monografía del Instituto de Salubridad y Enfermedades Tropicales, México, 1: vi+ 241 pp., 10 pls.

    Vargas, L. & Díaz Nájera, A. (1953) Nota sobre el examen de tipos de simulidos descritos por el Prof. G. Enderlein. Revista del Instituto de Salubridad y Enfermedades Tropicales, 13, 137–149+iii.

    Viviani, A.B.P & Araújo-Coutinho, C.J.P.C. (1999) Período de desenvolvimiento ovarianao de Simulium pertinax Kollar, 1832 (Diptera: Simuliidae) sob condições de laboratório. Entomología y Vectores, 6, 180–189.

    Vulcano, M.A. (1959) Descrição de Sumulium obesum, sp. n. (Diptera, Simuliidae). Papéis Avulsos do Departamento de Zoologia, São Paulo, 13, 351–360.

    Vulcano, M.A. (1967) A catalogue of the Diptera of the Americas south of the United States. 16. Family Simuliidae. Departamento de Zoologia, Secretaria da Agricultura, São Paulo, 16.1–16.44.

    Vulcano, M.A. (1981) Simuliidae [pp. 275–285]. In Aquatic Biota of Tropical South America. Part 1. Arthropoda. Ed. Hurlbert, S.H., Rodriguez, G. & Dias dos Santos, N. San Diego State University, California, 323pp.

    Werner, D. (1996a) Die Simuliiden-Typen der Dipteren-Sammlung des Zoologischen Museums Berlin (Diptera: Simuliidae). Mitteilengun aus dem Zoologischen Museum in Berlin, 72, 221–258.

    Werner, D. (1996b) Das Typenmaterial der Zweiflüger des Staatlichen Museums für Tierkunde Dresden (Insecta). Teil III: Diptera-Simuliidae. Staatliches Museum für Tierkunde Dresden, 57, 295–298.

    Wygodzinsky, P. (1953) Sobre algunos simúlidos Argentinos (Diptera). Anales del Instituto de Medicina Regional de la Universidad Nacional de Tucumán, 3, 293–320.