Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2017-05-05
Page range: 1–165
Abstract views: 366
PDF downloaded: 1

The black flies (Diptera: Simuliidae) of Vietnam

Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur, 50603 Malaysia
Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur, 50603 Malaysia
Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur, 50603 Malaysia
Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur, 50603 Malaysia
Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur, 50603 Malaysia
Tropical Infectious Diseases Research and Education Centre, University of Malaya, Kuala Lumpur, 50603 Malaysia
National Institute of Food Control, Ministry of Health, Hanoi, Vietnam (presently, The national office in southern region, Ministry of Science and Technology, Government of Vietnam, No 31, Han Thuyen street, District 1, Ho Chi Minh city, Vietnam)
Entomology Program, Clemson University, Clemson, South Carolina, 29634-0310, USA
black fly Simulium new species fauna Vietnam Diptera

Abstract

The biodiversity of black flies (Diptera: Simuliidae), which are biting insects of medical and veterinary importance, is strikingly high in Southeast Asian countries, such as Indonesia, Malaysia, Philippines and Thailand. In 2013, we began to explore the fauna of black flies in Vietnam, which has so far been poorly studied. In this monograph, the wealth of the biodiversity of black flies in Vietnam is also confirmed on the basis of the results of our recent investigations, though limited to five provinces in the country.

      Morphotaxonomic studies of black flies obtained from Sapa, Lao Cai Province, northern Vietnam, in 2014 and Nghe An Province, northern Vietnam, in 2015, and reexaminations of black flies collected from Tam Dao, Vinh Phuc Province, northern Vietnam, in 2013, Thua Thien Hue Province, central Vietnam, in 2014, and Lam Dong Province, southern Vietnam, in 2014, were conducted. A total of 22 species are described as new, including one in the newly recorded subgenus Montisimulium Rubtsov, and three species are recognized as new records from Vietnam. This investigation brings the number of species of black flies known in Vietnam to 70, all of which are assigned to the genus Simulium Latreille, and are placed in four subgenera (25 in Gomphostilbia Enderlein, one in Montisimulium, seven in Nevermannia Enderlein, and 37 in Simulium Latreille s. str.). The numbers of species-groups recognized include seven in Gomphostilbia, three in Nevermannia and nine in Simulium, indicating a high diversity of putative phylogenetic lineages. New species include S. (G.) sanchayense sp. nov. (= the species formerly regarded as S. (G.) brinchangense Takaoka, Sofian-Azirun & Hashim), S. (S.) lowi sp. nov. (= the species formerly regarded as S. (S.) brevipar Takaoka & Davies), S. (S.) fuscicoxae sp. nov. [= the species formerly regarded as S. (S.) rufibasis Brunetti (in part)], S. (S.) suoivangense sp. nov. [= morphoform ‘b’ of the S. (S.) tani Takaoka & Davies (complex)]. Newly recorded species are S. (G.) parahiyangum Takaoka & Sigit, S. (N.) maeaiense Takaoka & Srisuka, and S. (S.) doipuiense Takaoka & Choochote (complex) [= the species formerly regarded as S. (S.) rufibasis Brunetti (in part)]. The substitute name, S. (S.) huense, is given for the species that was described under the name of S. (S.) cavum from southern Vietnam. A redescription of the female, male, pupa and larva of S. (G.) asakoae Takaoka & Davies is presented, and the female and larva of S. (G.) hongthaii Takaoka, Sofian-Azirun & Ya’cob are described for the first time. Keys to 10 subgenera in the Oriental Region and all 70 species recorded from Vietnam are provided for females, males, pupae and mature larvae.

      As investigations extend nationwide in all the provinces in Vietnam, more new species and records are expected to be discovered. It is hoped that this monograph will be useful as a baseline taxonomic reference for future studies of black flies in Vietnam and neighbouring countries.

References

  1. Adler, P.H. & Crosskey, R.W. (2016) World Blackflies (Diptera: Simuliidae): A Comprehensive Revision of the Taxonomic and Geographical Inventory [2016]. 126 pp. Available from: https://www.clemson.edu/cafls/biomia/pdfs/blackflyinventory.pdf (accessed 10 May 2016)

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

    Adler, P.H., Takaoka, H., Sofian-Azirun, M., Low, V.L., Ya’cob, Z., Chen, C.D., Lau, K.W. & Pham, X.D. (2016) Vietnam, a hotspot for chromosomal diversity and cryptic species in black flies (Diptera: Simuliidae). PLoS ONE, 11 (10), e0163881.
    https://doi.org/10.1371/journal.pone.0163881

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

    Brunetti, E. (1911) New Oriental Nematocera. Record of Indian Museum, 4, 282–288.
    https://doi.org/10.5962/bhl.part.10539

    Cai, R., An, J.Y. & Li, C.P. (2008) A new species of the subgenus Simulium from Sichuan, China (Diptera: Simuliidae). Acta Parasitologica Medica and Entomologica Sinica, 15 (2), 100–102.

    Chen, H.B. (2003) Notes on Chinese members of the Simulium striatum-group, with descriptions of three new sepceis (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 28 (2), 336–342.

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

    Chen, H.B., Luo, H.B. & Yang, M. (2006) First list of blackflies with two new species from Shennongjia, Hubei Province, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 31 (4), 874–879.

    Chen, H.B., Xiu, J.F. & Zhang, C.L. (2012) A survey of blackflies with three new species from Kuankuoshui, Guizhou, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 37 (2), 382–388.

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

    Chen, H.B., Zhang, C.L. & Bi, G.H. (2004) Descriptions of three new species of Simulium (subg. Nevermannia) from Hunan Province, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 29 (2), 365–371.

    Chen, H.B., Zhang, C.L. & Huang, L. (2005) A new species of Simulium (Nevermannia) from Sichuan Province, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 30, 625–627.

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

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

    Chen, H.B., Zhang, C.L. & Yang, M. (2003b) Descriptions of two new species of the subgenus Nevermannia Enderlein and subgenus Simulium Latreille from Hainan Island, Chaina (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 28, 745–750.

    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.
    https://doi.org/10.7601/mez.56.335

    Crosskey, R.W. (1969) A Re-classification of the Simuliidae (Diptera) in Africa and Its Islands. In: Bulletin of the British Museum Natural History. Entomology. Supplement 14. United States Department of Agriculture, London, pp. 1–194 pp.

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

    Crosskey, R.W. & Howard, T.H. (1997) A New Taxonomic and Geographical Inventory of World Black Flies (Diptera: Simuliidae). Natural History Museum, London, 144 pp.

    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.
    https://doi.org/10.1080/00305316.1973.10434099

    Datta, M. (1974) Some black flies (Diptera: Simuliidae) of the subgenus Simulium Latreille (s. str.) from the Darjeeling area (India). Oriental Insects, 8 (1),15–27.
    https://doi.org/10.1080/00305316.1974.10434435

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

    Edwards, F.W. (1933) Diptera Nematocera from Mount Kinabalu. Journal of Federated Malay State Museum, 17, 223–296.

    Fukuda, M., Choochote, W., Bain, O., Aoki, C. & Takaoka, H. (2003) Natural infections with filarial larvae in two species of black flies (Diptera: Simuliidae) in northern Thailand. Japanese Journal of Tropical Medicine and Hygiene, 31 (2), 99–102.
    https://doi.org/10.2149/tmh1973.31.99

    Huang, L., Zhang, C.L. & Chen, H.B. (2013) A new species of the genus Simulium from Sichuan, China (Diptera: Simuliidae). Acta Zootaxonomica Sinica, 38 (2), 368–371.

    Huang, Y.T. & Takaoka, H. (2008) Simulium (Simulium) pingtungense, a new black fly (Diptera: Simuliidae) from Taiwan. Medical Entomology and Zoology, 59 (4), 309–317.
    https://doi.org/10.7601/mez.59.309

    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.
    https://doi.org/10.1051/parasite/2008152121

    Lewis, D.J. (1974) Man-biting Simuliidae (Diptera) of Northern India. Israel Journal of Entomology, 9, 23–53.

    Liu, Z.J. & An, J.Y. (2009) A new species of Simulium (Diptera: Simuliidae) from Shaanxi, China. Entomotaxonomica, 31 (2), 143–146.

    Low, V.L., Takaoka, H., Pramual, P., Adler, P.H., Ya’cob, Z., Huang, Y.T., Pham, X.D., Ramli, R., Chen, C.D., Wannaket, A. & Sofian-Azirun, M. (2016) Delineating taxonomic boundaries in the largest species complex of black flies (Simuliidae) in the Oriental Region. Scientific Report, 6, 20346.
    https://doi.org/10.1038/srep20346

    Luo, H.B., Yang, M. & Chen, H.B. (2010) A new species of genus Simulium from Xingdou mountain, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 35 (3), 472–474.

    Ogata, K. (1956) Notes on Simuliidae of the Ryukyu Islands (Diptera). Japanese Journal of Medical Science and Biology, 9, 59–69.
    https://doi.org/10.7883/yoken1952.9.59

    Ogata, K. (1966) Additional notes on Simuliidae of the Ryukyu Islands (Diptera). Kontyu, 34, 123–130.

    Otsuka, Y., Takaoka, H., Aoki, C. & Choochote, W. (2003) Phylogenetic analysis of the subgenus Himalayum within the genus Simulium s.l. (Diptera: Simuliidae) using mitochondrial 16S rRNA gene sequences. Medical Entomology and Zoology, 54 (1), 113–120.
    https://doi.org/10.7601/mez.54.113

    Pham, X.D. (1998) New records of six black fly species (Diptera: Simuliidae) from Vietnam. Medical Entomology and Zoology, 49 (2), 121–123.
    https://doi.org/10.7601/mez.49.121

    Pham, X.D. (1999) Additional records of three black fly species (Diptera: Simuliidae) from Vietnam. Medical Entomology and Zoology, 50 (4), 335–336.
    https://doi.org/10.7601/mez.50.335

    Puri, I.M. (1932a) Studies on Indian Simuliidae. Part II. Descriptions of males, females, and pupae of Simulium rufibasis Brunetti, its variety fasciatum nov. var. and of three new species from the Himalayas. Indian Journal of Medical Research, 19, 899–913.

    Puri, I.M. (1932b) Studies on Indian Simuliidae. Part III. Descriptions of males, females, and pupae of Simulium griseifrons Brunetti (1911), and of four new species with striped thorax. Indian Journal of Medical Research, 19, 1125–1143.

    Puri, I.M. (1932c) Studies on Indian Simuliidae. Part IV. Descriptions of two new species from North-East India Simulium howletti sp. n. and Simulium hirtipannus sp. n., with a note on S. ornatum Meigen. Indian Journal of Medical Research, 20, 505–514.

    Puri, I.M. (1932d) Studies on Indian Simuliidae. Part V. Species and varieties of the striatum series. Indian Journal of Medical Research, 20, 515–532.

    Puri, I.M. (1933a) Studies on Indian Simuliidae. Part VI. Descriptions of males, females, and pupae of two new species from Palni Hills and of male and pupa of S. tenuitarsus sp. n. from Bengal Terai. Indian Journal of Medical Research, 20, 803–812.

    Puri, I.M. (1933b) Studies on Indian Simuliidae. Part VII. Descriptions of larva, pupa and female of Simulium nodosum sp. nov. with an appendix dealing with S. novolineatum nov. nom. (= S. lineatum Puri). Indian Journal of Medical Research, 20, 813–817.

    Puri, I.M. (1933c) Studies on Indian Simuliidae. Part VIII. Descriptions of larvae, pupae, males and females of S. aureohirtum Brunetti and S. aureum Fries. Indian Journal of Medical Research, 21 (1), 1–9.

    Rubtsov, I.A. (1956) Blackflies (Simuliidae). Fauna of the USSR. 2nd Edition. No. 61, Diptera 6 (6). Acad. Nauk SSSR, Moscow and Leningrad, 859 + [1] pp. [translated from Russian by Amrind Publ. Co. Pot. Ltd., Delhi, India, 1989]

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

    Shiraki, T. (1935) Simuliidae of the Japanese Empire. Memory of Faculty of Science and Agriculture Taihoku Imperial University, 16, 1–90.

    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.
    https://doi.org/10.7601/mez.27.385

    Takaoka, H. (1977) Studies on blackflies of the Nansei Islands, Japan (Simuliidae; Diptera). III. On six species of the subgenus Simulium Latreille. Japanese Journal of Sanitary Zoology, 28 (2), 193–217.
    https://doi.org/10.7601/mez.28.193

    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. Japan Society For The Promotion of Science, Tokyo, xi + 199 pp.

    Takaoka, H. (1989) Further observations on the autogeny of Simulium aureohirtum Brunetti (Diptera: Simuliidae) in the Ryukyu Islands. Japanese Journal of Sanitary Zoology, 40 (1), 21–28.
    https://doi.org/10.7601/mez.40.21

    Takaoka, H. (1996) Description of a new species of Simulium (Simulium) from Sabah, Malaysia (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 24 (3), 157–161.
    https://doi.org/10.2149/tmh1973.24.157

    Takaoka, H. (2001a) Simulium (Simulium) weji sp. nov. (Diptera: Simuliidae) from Thailand. Japanese Journal of Tropical Medicine and Hygiene, 29 (4), 349–354.
    https://doi.org/10.2149/tmh1973.29.349

    Takaoka, H. (2001b) Two new and three newly recorded species of black flies (Diptera: Simuliidae) from Sabah, Malaysia. Japanese Journal of Tropical Medicine and Hygiene, 29 (2), 221–230.

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

    Takaoka, H. (2006) A new species of Simulium (Simulium) (Diptera: Simuliidae) from Luzon Island, Philippines. Medical Entomology and Zoology, 57 (2), 99–103.
    https://doi.org/10.7601/mez.57.99

    Takaoka, H. (2012) Morphotaxonomic revision of Simulium (Gomphostilbia) (Diptera: Simuliidae) in the Oriental Region. Zootaxa, 3577, 1–42.

    Takaoka, H. & Adler, P.H. (1997) A new subgenus, Simulium (Daviesellum), and a new species, S. (D.) courtneyi, (Diptera: Simuliidae) from Thailand and Peninsular Malaysia. Japanese Journal of Tropical Medicine and Hygiene, 25 (1), 17–27.
    https://doi.org/10.2149/tmh1973.25.17

    Takaoka, H. & Choochote, W. (2002) Taxonomic notes on the griseifrons species-group in Simulium (Simulium) (Diptera: Simuliidae) from Thailand: Descriptions of two new species and description of the male, pupa and larva of S. (S.) digrammicum Edwards. Japanese Journal of Tropical Medicine and Hygiene, 30 (2), 115–132.
    https://doi.org/10.2149/tmh1973.30.115

    Takaoka, H. & Choochote, W. (2004a) Two new species of Simulium (Simulium) (Diptera: Simuliidae) from Thailand. Tropical Medicine and Health, 32 (1), 31–36.
    https://doi.org/10.2149/tmh.32.31

    Takaoka, H. & Choochote, W. (2004b) A list of and keys to black flies (Diptera: Simuliidae) in Thailand. Tropical Medicine and Health, 32 (2), 189–197.
    https://doi.org/10.2149/tmh.32.189

    Takaoka, H. & Choochote, W. (2004c) Taxonomic notes on the griseifrons species-group of Simulium (Simulium) (Diptera: Simuliidae) in Northern Thailand. Tropical Medicine and Health, 32 (4), 311–327.
    https://doi.org/10.2149/tmh.32.311

    Takaoka, H. & Choochote, W. (2005a) Discovery of two more new species of Simulium (Montisimulium) (Diptera: Simuliidae) in Doi Inthanon National Park, Chiang Mai, Thailand. Tropical Medicine and Health, 33 (4), 209–215.
    https://doi.org/10.2149/tmh.33.209

    Takaoka, H. & Choochote, W. (2005b) A new subgenus and a new species of Simulium s. l. (Diptera: Simuliidae) from Thailand. Medical Entomology and Zoology, 56, 33–41.
    https://doi.org/10.7601/mez.56.33_1

    Takaoka, H. & Choochote, W. (2005c) Two new species of Simulium (Simulium) (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 56 (2), 99–110.
    https://doi.org/10.7601/mez.56.99

    Takaoka, H. & Choochote, W. (2005d) Two new species of black flies (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 56 (4), 319–334.
    https://doi.org/10.7601/mez.56.319

    Takaoka, H. & Choochote, W. (2009) Description of the female and male of Simulium (Montisimulium) angkaense (Diptera: Simuliidae) from Thailand. Medical Entomology and Zoology, 60 (2), 113–118.
    https://doi.org/10.7601/mez.60.113

    Takaoka, H., Choochote, W., Aoki, C., Fukuda, M. & Bain, O. (2003) Black flies (Diptera: Simuliidae) attracted to humans and water buffalos and natural infections with filarial larvae, probably Onchocerca sp., in northern Thailand. Parasite, 10 (1), 3–8.
    https://doi.org/10.1051/parasite/2003101p3

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

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

    Takaoka, H., Fukuda, M., Otsuka, Y., Aoki, C., Uni, S. & Bain, O. (2012a) Blackfly vectors of zoonotic onchocerciasis in Japan. Medical and Veterinary Entomology, 26, 372–378.
    https://doi.org/10.1111/j.1365-2915.2012.01023.x

    Takaoka, H. & Hadi, U.K. (1991) Two new blackfly species of Simulium (Simulium) from Java, Indonesia (Diptera: Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 19 (4), 357–370.
    https://doi.org/10.2149/tmh1973.19.357

    Takaoka, H. & Huang, Y.T. (2006) A new species of Simulium (Simulium) (Diptera: Simuliidae) from Taiwan. Medical Entomology and Zoology, 57 (3), 219–227.
    https://doi.org/10.7601/mez.57.219

    Takaoka, H. & Kuvangkadilok, C. (1999) Four new species of black flies (Diptera: Simuliidae) from Thailand. Japanese Journal of Tropical Medicine and Hygien, 27 (4), 497–509.
    https://doi.org/10.2149/tmh1973.27.497

    Takaoka, H. & Mulla, M.S. (2000) Taxonomic notes on Simulium siripoomense from Thailand (Diptera: Simuliidae): descriptions of female and pupa, and change of its subgeneric status from Byssodon to Simulium s. str. Japanese Journal of Tropical Medicine and Hygiene, 28 (1), 25–28.
    https://doi.org/10.2149/tmh1973.28.25

    Takaoka, H. & Noda, S. (1979) Autogeny of the black fly Simulium (Eusimulium) aureohirtum (Diptera: Simuliidae). Journal of Medical Entomology, 15 (2), 183–184.
    https://doi.org/10.1093/jmedent/15.2.183

    Takaoka, H. & Roberts, D.M. (1988) Notes on blackflies (Diptera: Simuliidae) from Sulawesi, Indonesia. Japanese Journal of Tropical Medicine and Hygiene, 16, 191–219.
    https://doi.org/10.2149/tmh1973.16.191

    Takaoka, H. & Saito, K. (1996) A new species and new records of black flies (Diptera: Simuliidae) from Thailand. Japanese Journal of Tropical Medicine and Hygiene, 24 (3), 163–169.
    https://doi.org/10.2149/tmh1973.24.163

    Takaoka, H. & Saito, K. (2007) A species list of black flies (Diptera: Simuliidae) in Japan. Yugaiseibutsu, 4, 44–60. [in Japanese]

    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 (2), 135–142.
    https://doi.org/10.2149/tmh1973.24.163

    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 (2), 69–80.
    https://doi.org/10.2149/tmh1973.25.69

    Takaoka, H., Sofian-Azirun, M., Hashim, R. & Ya’cob, Z. (2011) 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.
    https://doi.org/10.3897/zookeys.118.1552

    Takaoka, H., Sofian-Azirun, M. & Belabut, D.M. (2012b) Descriptions of two new species of Simulium (Simulium) (Diptera: Simuliidae) from Tioman Island, Peninsular Malaysia. Raffle’s Bulletin of Zoology, 60, 399–409.

    Takaoka, H., Sofian-Azirun, M., Ya’cob, Z., Chen, C.D., Lau, K.W. & Pham, H.T. (2014a) New species and records of black flies (Diptera: Simuliidae) from Vinh Phuc Province, Vietnam. Zootaxa, 3838 (3), 347–366.
    https://doi.org/10.11646/zootaxa.3838.3.6

    Takaoka, H., Sofian-Azirun, M., Ya’cob, Z., Chen, C.D., Lau, K.W. & Pham, H.T. (2014b) Female black flies of Simulium (Diptera: Simuliidae) collected on humans in Tam Dao National Park, Vietnam: Description of a new species and notes on four species newly recorded from Vietnam. Tropical Biomedicne, 31 (4), 742–748.

    Takaoka, H., Sofian-Azirun, M., Ya’cob, Z., Chen, C.D., Lau, K.W. & Pham, X.D. (2014c) Three new species of Simulium (Nevermannia) (Diptera: Simuliidae) from Vietnam. Zootaxa, 3866 (4), 555–571.
    https://doi.org/10.11646/zootaxa.3866.4.6

    Takaoka, H., Sofian-Azirun, M., Ya’cob, Z., Chen, C.D., Lau, K.W. & Pham, X.D. (2015a) The black flies (Diptera: Simuliidae) from Thua Thien Hue and Lam Dong Provinces, Vietnam. Zootaxa, 3961 (1), 1–96.

    Takaoka, H., Sofian-Azirun, M., Ya’cob, Z. & Hashim, R. (2014d) Two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Cameron’s Highlands, Peninsular Malaysia, with keys to 21 species of the Simulium asakoae species-group. Zootaxa, 3765 (1), 54–68.
    https://doi.org/10.11646/zootaxa.3765.1.3

    Takaoka, H. & Somboon, P. (2008) Eleven new species and one new record of black flies (Diptera: Simuliidae) from Bhutan. Medical Entomology and Zoology, 59 (3), 213–262.
    https://doi.org/10.7601/mez.59.213

    Takaoka, H. & Srisuka, W. (2009) Simulium (Gomphostilbia) chiangdaoense sp. nov. (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 60 (4), 269–276.
    https://doi.org/10.7601/mez.60.269

    Takaoka, H. & Srisuka, W. (2010) Simulium (Gomphostilbia) chayamaritae, a new species of black fly (Diptera: Simuliidae) from northern Thailand. Medical Entomology and Zoology, 61 (2), 265–271.
    https://doi.org/10.7601/mez.61.265

    Takaoka, H. & Srisuka, W. (2011) A new species of Simulium (Nevermannia) (Diptera; Simuliidae) from Thailand, with keys to members of the Simulium feuerborni species-group in Thailand. ZooKeys, 89, 57–70.
    https://doi.org/10.3897/zookeys.89.761

    Takaoka, H., Srisuka, W., Saeung, A., Otsuka, Y. & Choochote, W. (2012c) Simulium (Nevermannia) chomthongense, a new species of black fly (Diptera: Simuliidae) from Chiang Mai, Thailand. Tropical Biomedicine, 29, 381–390.

    Takaoka, H., Srisuka, W., Saeung, A., Otsuka, Y. & Choochote, W. (2013) Simulium (Nevermannia) khunklangense, a new species of black fly (Diptera: Simuliidae) from Chiang Mai, Thailand. Zootaxa, 3694 (3), 280–288.
    https://doi.org/10.11646/zootaxa.3694.3.8

    Takaoka, H., Srisuka, W., Otsuka, Y. & Choochote, W. (2014e) A new species and species-group of Simulium (Simulium) (Diptera: Simuliidae) from Thailand. Journal of Medical Entomology, 51, 725–732.
    https://doi.org/10.1603/ME13241

    Takaoka, H. & Suzuki, H. (1984) The blackflies (Diptera: Simuliidae) from Thailand. Japanese Journal of Sanitary Zoology, 35 (1), 7–45.
    https://doi.org/10.7601/mez.35.7

    Tangkawanit, U., Kuvangkadilok, C., Baimai, V. & Adler, P.H. (2009) Cytosystematics of the Simulium tani group (Diptera: Simuliidae) in Thailand. Zoological Journal of Linnean Society, 155, 289–315.
    https://doi.org/10.1111/j.1096-3642.2008.00433.x

    Xue, H.T. (1992) A new species of blackfly Simulium from Yunnan Province, China (Diptera: Simuliidae). Acta Zootaxonomica Sinica, 17 (2), 93–96.

    Zhang C.L. & Chen, H.B. (2001) A new black fly of the genus Simulium Latreille from Guizhou Province, China (Diptera: Simuliidae). Acta Zootaxonomica Sinica, 26 (2), 216–218.

    Zhang, C.L., Yang, M. & Chen, H.B. (2003) Two new species of the genus Simulium breeding adjacent on a waterfall from Wuzhishan of Hainan Island, China (Diptera, Simuliidae). Acta Zootaxonomica Sinica, 28 (4), 741–744.

    Zhu, J.Z. & Wang, S.P. (1995) A new species of Simulium from China (Diptera: Simuliidae). Sichuan Journal of Zoology, 14, 13–15.