Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2012-12-10
Page range: 1–42
Abstract views: 246
PDF downloaded: 1

Morphotaxonomic revision of Simulium (Gomphostilbia) (Diptera: Simuliidae) in the Oriental Region

Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur 50603, Malaysia
Diptera black fly Simuliidae Gomphostilbia taxonomy classification Oriental Region

Abstract

Simulium (Gomphostilbia) Enderlein, the third largest (206 named species included) in the genus Simulium Latreille s. l., is one of the two most abundant and diverse subgenera in the Oriental Region. To provide a classification scheme to facilitate morphological identification of the species within this subgenus, its diagnostic characters are redefined, and nine known species-groups within it are reviewed. Based on putative lineages explored by using certain adult morphological characters, seven more species-groups are proposed: asakoae, darjeelingense, epistum, gombakense, heldsbachense, hemicyclium and palauense species-groups, while the trirugosum species-group is merged in the varicorne species-group. Subgroups are also introduced to represent apparently different lineages within certain species-groups based on certain pupal morphological characters: two in the banauense species-group, seven in the batoense species-group redefined, four in the ceylonicum species-group redefined, four in the epistum species-group, two in the hemicyclium species-group, two in the sherwoodi species-group and four in the varicorne species-group redefined. A new checklist of species of the subgenus Gomphostilbia, and a key to all 15 species-groups within it are provided. The eastward expansion of the geographical distribution of the subgenus Gomphostilbia is inferred on the basis of the more frequent occurrence of apomorphic characters of certain adult and pupal morphological features in insular species-groups than in continental species-groups. A preliminary attempt using a cladistic analysis of morphological characters shows that among 10 subgenera examined, Gomphostilbia has a sister-taxon relationship with the Australasian subgenus Morops Enderlein, and this clade, together with the Central-Western Pacific subgenus Inseliellum Rubtsov, is positioned closest to the most derived clade formed by Daviesellum Takaoka & Adler and Simulium Latreille s. str.

References

  1. Adler, P.H. & Crosskey, R.W. (2012) World Blackflies (Diptera: Simuliidae): A Comprehensive Revision of the Taxonomic and Geographical Inventory [2012]. 119 pp., http://entweb.clemson.edu/biomia/pdfs/blackflyinventory.pdf [accessed on April 10, 2012]

    Adler, P.H., Currie, D.C. & Wood, D.M. (2004) The Black Flies (Simuliidae) of North America. xv+941 pp., Cornell University Press, Ithaca, New York, USA.

    An, J.Y. & Chen, H.B. (2007) Studies of subgenus Gomphostilbia on divide into groups (Diptera: Simuliidae). Acta Parasitologia and Medical Entomologia Sinica, 11, 110–113. (Text in Chinese with an English abstract).

    An, J.Y., Hao, B.S. & Mai, Z.Q. (1990a) Description of Simulium (Gomphostilbia) pingxiangensis sp. nov. from Guangxi Zhuangzu Aut. Reg. China. Control of blood-sucking dipteran insects, 2, 100–102. (Text in Chinese with an English abstract).

    An, J.Y., Zhang, Y.Z. & Deng, C.Y. (1990b) Survey of Simuliidae (Diptera) from Xizang, China. Control of blood-sucking dipteran insects, 2, 103–109. (Text in Chinese with an English abstract).

    Audley-Charles, M.G. (1981) Geological history of the region of Wallace’s line. p. 24–35. In: Whitmore, T.C. ed. Wallace’s line and plate tectonics. 91 pp., Clarendon Press, Oxford, UK.

    Bentinck, W. (1955) The black flies of Japan and Korea (Diptera: Simuliidae). 23 pp., Contributions of the 406th Medical General Laboratory, US Department of the Army, Tokyo, Japan.

    Borah, S., Rahman, I., Goswami, S., Manab Deka, M. & Takaoka, H. (2012) Notes on black flies (Diptera: Simuliidae) from North-East India: New records of five species from Arunachal Pradesh and taxonomic reviews of two species from Assam. Tropical Biomedicine, 29, 92–97.

    Brunetti, E. (1911) New Oriental Nematocera. Record of Indian Museum, 4, 259–316.

    Cai, R., An, J.Y. & Li, C.P. (2005) A new species of Simulium (Gomphostilbia) from Xinjiang, China (Diptera: Simuliidae). Acta Parasitologia and Medical Entomologia Sinica, 12, 177–179.

    Chen, H.B. (2001) Studies on black flies of Hainan Island II. Records of subgenus Morops Enderlein in China with description of a new species (Diptera: Simuliidae). Acta Zoologica Sinica, 26, 565–568.

    Chen, H.B. & An, J.Y. (2003) The blackflies of China (Diptera: Simuliidae). 448 pp., Science Press, Beijing, China. (Text in Chinese).

    Chen, J.Y. & Cao, Y.C. (1983) New species and new records of Eusimulium from China (Diptera: Simuliidae). Acta Entomologica Sinica, 26, 229–232. (Text in Chinese with an English abstract).

    Chen, H.B. & Chen, H. (2000) A new species of Simulium (Gomphostilbia) from Hainan Island, China. Special issue of Chinese Society of Entomology for 21st century, 129–133.

    Chen, H.B. & Zhang, C.L. (2001) Studies on blackflies of Hainan Island, China. I. Description of three new species of Simulium (Gomphostilbia) (Diptera: Simuliidae). Acta Zootaxonomica Sinica, 26, 361–368.

    Chen, H.B. & Zhang, C.L. (2003) Review of taxonomy of blackflies from Xishuangbanna, China, with a description of a new species (Diptera: Simuliidae). Acta Zootaxonomica Sinica, 28, 542–545.

    Chen, H.B. & Zhang, C.L. (2004) A new species of Simulium (Gomphostilbia) from Yunnan Province, China (Diptera: Simuliidae). Acta Parasitologia and Medical Entomologia Sinica, 11, 87–90.

    Chen, H.B., Zhang, C.L. & Yang, M. (2003) Checklist of Guizhou blackflies with descriptions of a new species (Diptera: Simuliidae). Guizhou Science, 21, 46–50.

    Chen, H.B, Zhang, J.Q. & Wen, X.J. (2000) Two new species of Simulium (Gomphostilbia) from Fanjing Mountain, China (Diptera, Simuliidae). Entomologia Sinica, 7, 21–28.

    Chen, H.B, Zhang, J.Q. & Zhang, C.L. (2007) Two new species of the genus Simulium (Gomphostilbia) from Guangxi, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 32, 782–786.

    Choochote, W., Takaoka, H., Fukuda, M., Otsuka, Y., Aoki, C. & Eshima, N. (2005) Seasonal abundance and daily flying activity of black flies (Diptera: Simuliidae) attracted to human baits in Doi Inthanon National Park, northern Thailand. Medical Entomology and Zoology, 56, 335–348.

    Chubareva, L.A. (1996) Morphological and karyotypical characters of two blackfly species of the genus Gomphostilbia Enderlein (Diptera, Simuliidae) from Tadzhikistan. Entomological Review, 76, 89–93.

    Craig, D.A. (1997) A taxonomic revision of the Pacific black fly subgenus Inseliellum (Diptera: Simuliidae). Canadian Journal of Zoology, 75, 855–904.

    Craig, D.A., Englund, R.A. & Takaoka, H. (2006) Simuliidae (Diptera) of the Solomon Islands: new records and species, ecology, and biogeography. Zootaxa, 1328, 1–26.

    Crosskey, R.W. (1967) The classification of Simulium Latreille (Diptera: Simuliidae) from Australia, New Guinea and the Western Pacific. Journal of Natural History, 1, 23–51.

    Crosskey, R.W. (1990) The Natural History of Blackflies. ix+711 pp., John Wiley & Sons Inc., Chichester, England.

    Datta, M. (1973) New species of black flies (Diptera: Simuliidae) of the subgenera Eusimulium Roubaud and Gomphostilbia Enderlein from the Darjeeling area, India. Oriental Insects, 7, 363–402.

    Datta, M. (1975) Simuliidae (Diptera) from Assam foot-hills, India. Japanese Journal of Sanitary Zoology, 26, 31–40.

    Datta, M. (1992) An overview of the Simuliidae (Diptera) of West Bengal, India. Journal of Bengal Natural History Society, New Series, 11, 41–62.

    Davies, D.M. & Győrkős, H. (1987) The Simuliidae (Diptera) of Sri Lanka. Descriptions of species in the subgenera Eusimulium and Gomphostilbia of the genus Simulium. Canadian Journal of Zoology, 65, 1483–1502.

    Davies, D.M. & Győrkős, H. (1988) The Simuliidae (Diptera) of Sri Lanka. Description of a new species of Simulium (Morops). Canadian Journal of Zoology, 66, 605–610.

    Dickerson, R.E. (1928) Distribution of life in the Philippines. 322 pp., Philippine Bureau of Sciences Monograph 21, Manila, Philippines.

    Edwards, F.W. (1925) Diptera Nematocera from Dutch East Indies. Treubia, 6, 154–157.

    Edwards, F.W. (1928) Diptera Nematocera from the Federated Malay States Museums. Journal of Federated Malay State Museums, 14, 1–139.

    Edwards, F.W. (1934) The Simuliidae (Diptera) of Java and Sumatra. Deutsche Limnologische Sunda-Expedition. 91, Archives of Hydrobiology, 13, 92–138.

    Hadi, U.K., Takaoka, H. & Aoki, C. (1995) Larval salivary gland chromosomes of the blackfly, Simulium (Gomphostilbia) yaeyamaense (Diptera: Simuliidae) from Ryukyu Islands, Japan. Japanese Journal of Sanitary Zoology, 46, 235–239.

    Hadi, U.K., Takaoka, H., Kondo, K. & Hirai, H. (1996) Larval salivary gland chromosomes of the blackfly, Simulium (Gomphostilbia) sundaicum (Diptera: Simuliidae) from Java, Indonesia. Medical Entomology and Zoology, 47, 55–61.

    Huang, Y.T. & Takaoka, H. (2008) A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Taiwan. Medical Entomology and Zoology, 59, 171–179.

    Huang, Y.T., Takaoka, H., Otsuka, Y. & Aoki, C. (2011) Simulium (Gomphostilbia) taitungense, a new species of black fly (Diptera: Simuliidae) from Taiwan, with description of the male of S. (G.) tuenense Takaoka. Tropical Biomedicine, 28, 577–588.

    Ishii, Y., Choochote, W., Bain, O., Fukuda, M., Otsuka, Y. & Takaoka, H. (2008) Seasonal and diurnal biting activities and zoonotic filarial infections of two Simulium species (Diptera: Simuliidae) in northern Thailand. Parasite, 15, 121–129.

    Jitklang, S. & Kuvangkadilok, C. (2007) A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from sourthern Thailand, with description of its polytene chromosomes. Studia dipterologica, 14, 1–7.

    Jitklang, S., Kuvangkadilok, C., Baimai, V., Takaoka, H., & Adler, P.H. (2008) Cytogenetics and morphotaxonomy of the Simulium (Gomphostilbia) ceylonicum species group (Diptera: Simuliidae) in Thailand. Zootaxa, 1917, 1–28.

    Kang, Z., Zhang, C.L. & Chen, H.B. (2007) A new species of Simulium (Gomphostilbia) from Jiangxi Province, China (Diptera: Simuliidae). Acta Parasitologia and Medical Entomologia Sinica, 14, 185–187.

    Kuvangkadilok, C., Lualon, U. & Baimai, V. (2008) Cytotaxonomy of Simulium siamense Takaoka and Suzuki (Diptera: Simuliidae) in Thailand. Genome, 51, 972–987.

    Kuvangkadilok, C. & Takaoka, H. (2000) Taxonomic notes on Simuliidae (Diptera) from Thailand: Description of a new species and new distributional records of nine known species. Japanese Journal of Tropical Medicine and Hygiene, 28, 167–175.

    Lewis, D.J. (1964) Diptera from Nepal. Simuliidae. Bulletin of the British Museum (Natural History), 15, 291–294.

    Long, Z.M, An, J.Y. , Hao, B.S., Yan, G. & Yuan, W.H. (1994) Two new blackflies of Simulium (Gomphostilbia) from Hainan Island, China (Diptera, Simuliidae). Chinese Journal of Vector Biology and Control, 5, 405–409. (Text in Chinese with an English abstract).

    Mackerras, M.J. & Mackerras, I.M. (1950) Notes on Australasian Simuliidae (Diptera). II. Proceedings of Linnean Society of New South Wales, 125, 167–187.

    Mahe, M., Ma, D. & An, J.Y. (2003) Two new species of Simulium (Gomphostilbia) from China (Diptera: Simuliidae). Acta Parasitologia and Medical Entomologia Sinica, 10, 113–117. (Text in Chinese with an English abstract).

    Ogata, K. (1956) Notes on Simuliidae of the Ryukyu Islands (Diptera). Japanese Journal of Medical Science and Biology, 9, 59–69.

    Ogata, K. & Sasa, M. (1954) Taxonomic notes on Simuliidae or black flies of Japan, with special references on the subgenera Eusimulium Roubaud and Nevermannia Enderlein (Diptera). Japanese Journal of Experimental Medicine, 24, 325–333.

    Ogata, K. & Sasa, M. (1955) Keys to the adult females and pupae of Japanese Simuliidae and notes on the control of black fly larvae by insecticide (Studies on black fly 6). Japanese Journal of Sanitary Zoology, 6, 10–18.

    Otsuka, Y., Aoki, C., Saito, K., Hadi, U.K., Suzuki, H. & Takaoka, H. (2001) Phylogenetic analyses of a blackfly subgenus Simulium (Nevermannia) based on mitochondrial 16S ribosomal RNA gene sequences. Japanese Journal of Tropical Medicine and Hygiene, 29, 261–266.

    Otsuka, Y., Aoki, C., Choochote, W., De Las Llagas, L. & Takaoka, H. (2007) Phylogenetic analysis of three subgenera: Asiosimulium, Daviesellum and Wallacellum, of the genus Simulium s. l. endemic in the Oriental Region. Medical Entomology and Zoology, 58, 329–333.

    Phasuk, J., Chanpaisaeng, J., Adler, P.H. & Courtney, G.W. (2005) Chromosomal and morphological taxonomy of larvae of Simulium (Gomphostilbia) (Diptera: Simuliidae) in Thailand. Zootaxa, 1052, 49–60.

    Phayuhasena, S., Colgan, D.J., Kuvangkadilok, C., Pramual, P. & Baimai, V. (2010) Phylogenetic relationships among the black fly species (Diptera: Simuliidae) of Thailand based on multiple gene sequences. Genetica, 138, 633–648.

    Pramual, P. & Kuvangkadilok, C. (2012) Integrated cytogenetic, ecological, and DNA barcode study reveals cryptic diversity in Simulium (Gomphostilbia) angulistylum (Diptera: Simuliidae). Genome, 55, 1–12.

    Pramual, P. & Tangkawanit, U. (2008) A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Northeastern Thailand. Medical Entomology and Zoology, 59, 297–303.

    Pramual, P. & Wongpakam, K. (2011) Cytogenetics of Simulium siamense Takaoka and Suzuki, 1984 (Diptera: Simuliidae) in northern Thailand. Aquatic Insects, 33, 171–184.

    Pramual, P., Wongpakam, K. & Adler, P.H. (2011) Cryptic biodiversity and phylogenetic relationships revealed by DNA barcoding of Oriental black flies in the subgenus Gomphostilbia (Diptera: Simuliidae). Genome, 54, 1–9.

    Procunier, W.S., Shelley, A.J. & Arzube, M. (1987) Cytological identification of Simulium oyapockense manabi form (Diptera: Simuliidae): a potential vector of onchocerciasis in Ecuador. Tropical Medicine and Parasitology, 38, 71.

    Rothfels, K. (1989) Speciation in black flies. Genome, 32, 500–509.

    Rubtsov, I.A. (1959–1964) Simuliidae (Melusinidae). In: Lindner, Die Fliegen der Palaearktischen Region, 14, 1–689.

    Shrestha, S. & Takaoka, H. (2009) New records of ten species of black flies (Diptera: Simuliidae) from Nepal. Medical Entomology and Zoology, 60, 253–258.

    Smart, J. & Clifford, E.A. (1965) Simuliidae (Diptera) of the territory of Papua and New Guinea. Pacific Insects, 7, 505–619.

    Smart, J. & Clifford, E.A. (1969) Simuliidae (Diptera) of Sabah (British North Borneo). Zoological Journal of Linnean Society, 48, 9–47.

    Swofford, D.L. (1998) PAUP*, Phylogenetic Analysis Using Parsimony (*and other methods) version 4. Sinauer Associates.

    Takaoka, H. (1973) Descriptions of 2 new species of blackflies, Simulium (Gomphostilbia) tokarense and S. (Eusimulium) morisonoi (Diptera: Simuliidae), from the Tokara Islands, Japan. Japanese Journal of Sanitary Zoology, 23, 201–207.

    Takaoka, H. (1976) Studies on blackflies of the Nansei Islands, Japan (Simuliidae; Diptera). II. On six species of the subgenera, Gomphostilbia Enderlein, Morops Enderlein, Odagmia Enderlein and Gnus Rubzov, with the description of Simulium (Gomphostilbia) okinawense sp. nov. Japanese Journal of Sanitary Zoology, 27, 385–398.

    Takaoka, H. (1979) The black flies of Taiwan (Diptera: Simuliidae). Pacific Insects, 20, 365–403.

    Takaoka, H. (1983) The Blackflies (Diptera: Simuliidae) of the Philippines. xi + 119 pp., Japan Society For The Promotion of Science, Tokyo, Japan.

    Takaoka, H. (1989) Notes on blackflies (Diptera: Simuliidae) from Myanmar (formerly Burma). Japanese Journal of Tropical Medicine and Hygiene, 17, 243–257.

    Takaoka, H. (1991) A revisionary note on the blackfly, hitherto called "Simulium (Gomphostilbia) batoense Edwards, 1934" from the Ryukyu Islands, Japan (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 19, 265–273.

    Takaoka, H. (1994) A new blackfly species of Simulium (Gomphostilbia) from Solomon Islands, South Pacific (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 22, 103–108.

    Takaoka, H. (1995) The Simuliidae (Diptera) from Bougainville Island, Papua New Guinea. Japanese Journal of Tropical Medicine and Hygiene, 23, 239–252.

    Takaoka, H. (1996) The geographical distribution of the genus Simulium Latreille in the Oriental and Australasian Regions. Japanese Journal of Tropical Medicine and Hygiene, 24, 113–124.

    Takaoka, H. (2000a) Taxonomic notes on Simulium gombakense (Diptera: Simuliidae) from Peninsular Malaysia: descriptions of male and pupa, and subgeneric transfer from Morops to Gomphostilbia. Japanese Journal of Tropical Medicine and Hygiene, 28, 111–114.

    Takaoka, H. (2000b) A new black-fly species of Simulium (Morops) from Luzon Island, Philippines (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 28, 361–364.

    Takaoka, H. (2001a) Two new and three newly recorded species of black flies (Diptera: Simuliidae) from Sabah, Malaysia. Japanese Journal of Tropical Medicine and Hygiene, 29, 221–230.

    Takaoka, H. (2001b) Description of two new species of black flies (Diptera: Simuliidae) from Sarawak, Malaysia. Japanese Journal of Tropical Medicine and Hygiene, 29, 243–252.

    Takaoka, H. (2003) The Black Flies (Diptera: Simuliidae) of Sulawesi, Maluku and Irian Jaya. xxii + 581 pp., Kyushu University Press, Fukuoka, Japan.

    Takaoka, H. (2005) A new species of Simulium (Gomphostilbia) from Luzon Island, Philippines (Diptera: Simuliidae). Medical Entomology and Zoology, 56, 211–218.

    Takaoka, H. (2008a) Simulium (Gomphostilbia) hoiseni sp. nov. (Diptera: Simuliidae): a new species from Peninsular Malaysia. Medical Entomology and Zoology, 59, 9–14.

    Takaoka, H. (2008b) Four new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Sarawak, Malaysia. Medical Entomology and Zoology, 59, 181–211.

    Takaoka, H. (2009a) Three new species of the Simulium (Gomphostilbia) banauense species-group (Diptera: Simuliidae) in the Philippines. Medical Entomology and Zoology, 60: 65–79.

    Takaoka, H. (2009b) Three new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Sabah and Sarawak, Malaysia. Medical Entomology and Zoology, 60, 97–112.

    Takaoka, H. & Choochote, W. (2004) A list of and keys to black flies (Diptera: Simuliidae) in Thailand. Tropical Medicine and Health, 32, 189–197.

    Takaoka, H. & Choochote, W. (2005a) A new subgenus and a new species of Simulium s. l. (Diptera: Simuliidae) from Thailand. Medical Entomology and Zoology, 56, 33–41.

    Takaoka, H. & Choochote, W. (2005b) Two new species of black flies (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 56, 319–334.

    Takaoka, H. & Choochote, W. (2006a) A new species of the subgenus Simulium (Asiosimulium) (Diptera: Simuliidae) from Thailand. Medical Entomology and Zoology, 57, 45–48.

    Takaoka, H. & Choochote, W. (2006b) A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 57, 229–233.

    Takaoka, H. & Craig, D.A. (1999) Taxonomic and ecological notes on Simulium (Gomphostilbia) palauense (Diptera: Simuliidae) from Palau, Micronesia, with redescriptions of adults and descriptions of the pupa and mature larva. Japanese Journal of Tropical Medicine and Hygiene, 27, 195–201.

    Takaoka, H. & Davies, D.M. (1995) The Black Flies (Diptera: Simuliidae) of West Malaysia. viii + 175 pp, Kyushu University Press, Fukuoka, Japan.

    Takaoka, H. & Davies, D.M. (1996) The Black Flies (Diptera: Simuliidae) of Java, Indonesia. viii + 81 pp., Bishop Museum Bulletin in Entomology 6, Bishop Museum Press, Honolulu, U.S.A.

    Takaoka, H., Davies, D.M. & Dudgeon, D. (1995) Black flies (Diptera: Simuliidae) from Hong Kong: taxonomic notes with descriptions of two new species. Japanese Journal of Tropical Medicine and Hygiene, 23, 189–196.

    Takaoka, H., Fukuda, M., Otsuka, Y., Aoki, C., Uni, S. & Bain, O. (2012a) Blackfly vectors of zoonotic onchocerciasis in Japan. Medical and Veterinary Entomology, (doi: 10.1111/j.1365-2915.2012.01023.x).

    Takaoka, H., Hadi, U.K. & Sigit, S.H. (2006) The black flies (Diptera: Simuliidae) of Flores and Timor, Indonesia. Medical Entomology and Zoology, 57, 1–26.

    Takaoka, H. & Okazawa, T. (1988) A revised list of Japanese blackflies (Simuliidae). Medical Entomology and Zoology, 39, 97–103. (Text in Japanese).

    Takaoka, H., Otsuka, Y., Choochote, W., Aoki, C., Hayakawa, H. & Thongsuhuan, S. (2010a) Descriptions of the male, pupa and larva of Simulium (Gomphostilbia) novemarticulatum (Diptera: Simuliidae) from Peninsular Malaysia and Thailand. Medical Entomology and Zoology, 61, 59–67.

    Takaoka, H., Otsuka, Y., Choochote, W. & Thongsuhuan, S. (2009a) Two new and one newly recorded species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from southern Thailand. Medical Entomology and Zoology, 60, 259–268.

    Takaoka, H. & Roberts, D.M. 1988: Notes on blackflies (Diptera: Simuliidae) from Sulawesi, Indonesia. Japanese Journal of Tropical Medicine and Hygiene, 16, 191–219.

    Takaoka, H. & Shrestha, S. (2010) New species of black flies (Diptera: Simuliidae) from Nepal. Zootaxa, 2731 (Monograph), 1–62.

    Takaoka, H. & Sigit, S.H. (1992) A new blackfly species of Simulium (Gomphostilbia) from Java, Indonesia (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 20, 135–142.

    Takaoka, H. & Sigit, S.H. (1997) Three new black fly species of Simulium (Diptera: Simuliidae) from Sumatra, Indonesia. Japanese Journal of Tropical Medicine and Hygiene, 25, 69–80.

    Takaoka, H., Sofian-Azirun, M. & Hashim, R. (2011a) Simulium (Gomphostilbia) sofiani, a new species of black fly (Diptera: Simuliidae) from Peninsular Malaysia. Tropical Biomedicine, 28, 389–399.

    Takaoka, H., Sofian-Azirun, M., Hashim, R. & Ya’cob, Z. (2011b) Two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Peninsular Malaysia, with keys to Peninsular Malaysian members of the Simulium ceylonicum species-group. ZooKeys, 118, 53–74.

    Takaoka, H., Sofian-Azirun, M., Hashim, R., Ya’cob, Z. & Chen, C.D. (2012b) Two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Peninsular Malaysia. Journal of Medical Entomology, 49, 803–812.

    Takaoka, H. & Somboon, P. (2008) Eleven new species and one new record of black flies (Diptera: Simuliidae) from Bhutan. Medical Entomology and Zoology, 59, 213–262.

    Takaoka, H. & Srisuka, W. (2009) Simulium (Gomphostilbia) chiangdaoense sp. nov. (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 60, 269–276.

    Takaoka, H. & Srisuka, W. (2010a) Description of the female of Simulium (Gomphostilbia) kuvangkadilokae (Diptera: Simuliidae) from Thailand. Medical Entomology and Zoology, 61, 39–47.

    Takaoka, H. & Srisuka, W. (2010b) Simulium (Gomphostilbia) chayamaritae, a new species of black fly (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 61, 265–271.

    Takaoka, H. & Suzuki, H. (1984) The blackflies (Diptera: Simuliidae) from Thailand. Japanese Journal of Sanitary Zoology, 35, 7–45.

    Takaoka, H. & Suzuki, H. (1994) A new blackfly species of Simulium (Morops) from Solomon Islands, South Pacific (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 22, 207–210.

    Takaoka, H. & Suzuki, H. (1995) The Simuliidae (Diptera) from the Solomon Islands, South Pacific. Japanese Journal of Tropical Medicine and Hygiene, 23, 253–272.

    Takaoka, H. & Tenedero, V.F. (2007a) Two new and four newly recorded species of Simulium (Diptera: Simuliidae) from Mindoro Island, the Philippines. Medical Entomology and Zoology, 58, 29–43.

    Takaoka, H. & Tenedero, V.F. (2007b) Two new and one newly recorded species of Simulium (Diptera: Simuliidae) from Palawan Island, the Philippines. Medical Entomology and Zoology, 58, 207–228. .

    Takaoka, H. & Tenedero, V.F. (2008a) A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Palawan Island, the Philippines. Medical Entomology and Zoology, 59, 1–7.

    Takaoka, H. & Tenedero, V.F. (2008b) Simulium (Gomphostilbia) rarum sp. nov., a new species of black fly (Diptera: Simuliidae) from Palawan Island, the Philippines. Medical Entomology and Zoology, 59, 319–324.

    Takaoka, H. & Tenedero, V.F. (2008c) New species and record of black flies (Diptera: Simuliidae) from Samar, the Philippines. Medical Entomology and Zoology, 59, 325–336.

    Takaoka, H., Tenedero, V.F. & Bertuso, A.G. (2009b) Description of the male of Simulium (Gomphostilbia) binuanense (Diptera: Simuliidae) from Palawan Island, Philippines. Medical Entomology and Zoology, 60, 213–216.

    Takaoka, H., Thapa, S. & Henry, W. (2010b) Description of two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Darjeeling, India. Medical Entomology and Zoology, 61, 105–110.

    Thanwisai, A., Kuvangkadilok, C. & Baimai, V. (2006) Molecular phylogeny of black flies (Diptera: Simuliidae) from Thailand, using ITS2 rDNA. Genetica, 128, 177–204.

    Wen, X.J. & Chen, H.B. (2000) A new species of Simulium (Gomphostilbia) from China (Diptera: Simuliidae). Guizhou Science, 18, 112–115.

    Whitmore, T.C. (1981) Introduction. p. 1–2. In: Whitmore, T.C. ed. Wallace’s line and plate tectonics. 91 pp., Clarendon Press, Oxford, UK.

    Yankovsky, A.V. (2002) A key of the identification of blackflies (Diptera: Simuliidae) of Russia and adjacent countries (former USSR). 569 pp., Zoological Institute, Russian Academy of Sciences, Vol. 170. Sankt-Petersburg, Russia.

    Zhang, C.L. & Chen, H.B. (2004) Two new species of subg. Gomphostilbia of Simulium Latreille from Hunan Province, China (Diptera: Simuliidae). Acta Zootaxonomica Sinica, 29, 372–376.

    Zhang, J.Q., Gao, B., Cai, H.Z., Huang, E.J. & Chen, H.B. (2009) Notes on the blackflies of the Wui mountain, China with a new species description (Diptera: Simuliidae). Acta Parasitologia and Medical Entomologia Sinica, 16, 107–110.