The Black Flies (Diptera Simuliidae) of Vietnam-Monograph

Zootaxa 4261 (1): 001–165 ISSN 1175-5326 (print edition)
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Copyright © 2017 Magnolia Press
Monograph ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4261.1.1
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ZOOTAXA
4261
The black flies (Diptera: Simuliidae) of Vietnam
HIROYUKI TAKAOKA1,5, MOHD SOFIAN-AZIRUN1, ZUBAIDAH YA’COB1, CHEE DHANG

CHEN1, KOON WENG LAU1, VAN LUN LOW2, XUAN DA PHAM3 & PETER H. ADLER4
1
Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur, 50603 Malaysia
2
Tropical Infectious Diseases Research and Education Centre, University of Malaya, Kuala Lumpur, 50603 Malaysia
3
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)
4
Entomology Program, Clemson University, Clemson, South Carolina, 29634-0310, USA
5
Corresponding author. E-mail address: takaoka@oita-u.ac.jp
Magnolia Press
Auckland, New Zealand
Accepted by L. Hernandez-Triana: 15 Feb. 2017; published: 5 May 2017

HIROYUKI TAKAOKA, MOHD SOFIAN-AZIRUN, ZUBAIDAH YA’COB, CHEE DHANG CHEN, KOON
WENG LAU, VAN LUN LOW, XUAN DA PHAM & PETER H. ADLER
The black flies (Diptera: Simuliidae) of Vietnam
(Zootaxa 4261)
165 pp.; 30 cm.
5 May 2017
ISBN 978-1-77670-134-6 (paperback)
ISBN 978-1-77670-135-3 (Online edition)
FIRST PUBLISHED IN 2017 BY

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ISSN 1175-5326 (Print edition)

ISSN 1175-5334 (Online edition)
2 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.

Female adults of three black fly species, which are vectors of filariae in the Oriental Region (From top, Simulium asakoae
Takaoka & Davies, Simulium nigrogilvum Summers and Simulium nodosum Puri)
THE BLACK FLIES FROM VIETNAM Zootaxa 4261 (1) © 2017 Magnolia Press · 3
Synopsis
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.
Key words: black fly, Simulium, new species, fauna, Vietnam

Table of contents
Frontispiece . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Synopsis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Systematics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Genus Simulium Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Keys to 10 subgenera of the genus Simulium in the Oriental Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
I. Subgenus Gomphostilbia Enderlein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Keys to 25 species of the subgenus Gomphostilbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
1) Simulium asakoae species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Simulium (Gomphostilbia) asakoae Takaoka & Davies, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Simulium (Gomphostilbia) chaudinhense Takaoka & Sofian-Azirun sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Simulium (Gomphostilbia) confertum Takaoka & Sofian-Azirun, 2015. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Simulium (Gomphostilbia) fuscidorsum Takaoka & Ya’cob, 2015. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Simulium (Gomphostilbia) hmongense Takaoka & Ya’cob sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Simulium (Gomphostilbia) hongthaii Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Simulium (Gomphostilbia) phulocense Takaoka & Chen, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Simulium (Gomphostilbia) quychauense Takaoka & Chen sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Simulium (Gomphostilbia) sanchayense Takaoka & Lau sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Simulium (Gomphostilbia) tamdaoense Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Simulium (Gomphostilbia) unii Takaoka & Pham sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Simulium (Gomphostilbia) thituyenae Takaoka & Pham, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Simulium (Gomphostilbia) vinhphucense Takaoka & Low sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
2) Simulium batoense species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Simulium (Gomphostilbia) dachaisense Takaoka & Lau, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Simulium (Gomphostilbia) duolongum Takaoka & Davies, 1995. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Simulium (Gomphostilbia) lamdongense Takaoka & Sofian-Azirun, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Simulium (Gomphostilbia) longlanhense Takaoka & Ya’cob, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Simulium (Gomphostilbia) parahiyangum Takaoka & Sigit, 1992. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Simulium (Gomphostilbia) siamense Takaoka & Suzuki, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
3) Simulium ceylonicum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Simulium (Gomphostilbia) inthanonense Takaoka & Suzuki, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
4) Simulium darjeelingense species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Simulium (Gomphostilbia) eshimai Takaoka & Adler sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
5) Simulium epistum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Simulium (Gomphostilbia) montiblense Takaoka, 1983 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
6) Simulium gombakense species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Simulium (Gomphostilbia) thuathienense Takaoka & Sofian-Azirun, 2015. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
7) Simulium varicorne species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Simulium (Gomphostilbia) breviflagellum Takaoka & Sofian-Azirun, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
II. Subgenus Montisimulium Rubtsov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Simulium (Montisimulium) nigrofilum Takaoka & Sofian-Azirun sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
III. Subgenus Nevermannia Enderlein . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Keys to seven species of the subgenus Nevermannia from Vietnam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
1) Simulium feuerborni species-group. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Simulium (Nevermannia) bachmaense Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Simulium (Nevermannia) langbiangense Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Simulium (Nevermannia) maeaiense Takaoka & Srisuka, 2011 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Simulium (Nevermannia) phami Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
2) Simulium ruficorne species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Simulium (Nevermannia) aureohirtum Brunetti, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
3) Simulium vernum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Simulium (Nevermannia) laichauense Takaoka & Sofian-Azirun sp. nov.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Simulium (Nevermannia) tayense Takaoka & Ya’cob sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
IV. Subgenus Simulium Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Keys to 37 species of the subgenus Simulium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
1) Simulium argentipes species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Simulium (Simulium) sansahoense Takaoka & Chen sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
2) Simulium christophersi species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Simulium (Simulium) atipornae Takaoka, Srisuka & Choochote, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
3) Simulium griseifrons species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Simulium (Simulium) chungi Takaoka & Huang, 2006 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Simulium (Simulium) grossifilum Takaoka & Davies, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Simulium (Simulium) laocaiense Takaoka & Lau sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Simulium (Simulium) maenoi Takaoka & Choochote, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Simulium (Simulium) nigrogilvum Summers, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Simulium (Simulium) turgidum Takaoka & Pham sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Simulium (Simulium) vietnamense Takaoka, Sofian-Azirun & Chen, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
4) Simulium malyschevi species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Simulium (Simulium) obliquum Takaoka & Low sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
5) Simulium multistriatum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Simulium (Simulium) daoense Takaoka & Adler sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
Simulium (Simulium) hirtinervis Edwards, 1928 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Simulium (Simulium) lacduongense Takaoka & Ya’cob, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Simulium (Simulium) laui Takaoka & Sofian-Azirun, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Simulium (Simulium) malayense Takaoka & Davies, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
6) Simulium nobile species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Simulium (Simulium) nodosum Puri, 1933 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
7) Simulium striatum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Simulium (Simulium) chiangmaiense Takaoka & Suzuki, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Simulium (Simulium) jeffreyi Takaoka & Davies, 1995 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Simulium (Simulium) nakhonense Takaoka & Suzuki, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Simulium (Simulium) quinquestriatum (Shiraki, 1935) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Simulium (Simulium) thailandicum Takaoka & Suzuki, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Simulium (Simulium) tavanense Takaoka & Sofian-Azirun sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Simulium (Simulium) taythienense Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Simulium (Simulium) xuandai Takaoka, Sofian-Azirun & Ya’cob, 2014 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
8) Simulium tuberosum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
Simulium (Simulium) congi Takaoka & Sofian-Azirun, 2015. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
Simulium (Simulium) doipuiense Takaoka & Choochote, 2005 (complex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 126
Simulium (Simulium) fuscicoxae Takaoka & Ya’cob sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Simulium (Simulium) rosliramlii Takaoka & Chen sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Simulium (Simulium) giayense Takaoka & Lau sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Simulium (Simulium) huense Takaoka & Ya’cob (substitute name) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Simulium (Simulium) sapaense Takaoka & Low sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Simulium (Simulium) lowi Takaoka & Adler sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Simulium (Simulium) tani Takaoka & Davies, 1995 (complex) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Simulium (Simulium) suoivangense Takaoka & Pham sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Simulium (Simulium) xuandei Takaoka & Pham, 2015 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
9) Simulium variegatum species-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Simulium (Simulium) chamlongi Takaoka & Suzuki, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Simulium (Simulium) phuluense Takaoka & Sofian-Azirun sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Introduction
Adult black flies (Simuliidae) are small (1.5 to 6.0 mm in length), usually brown to black, two-winged insects
(hence, belonging to the order Diptera). Black flies are closely associated with clean, running water in streams and
rivers, where their immature stages develop and the gravid female adults return for oviposition. There are four
distinct developmental stages: egg, larva, pupa and adult. Female adults bite mammals and birds to obtain a blood
for development of their eggs. Due to their biting habits, black flies are insects of medical and veterinary
importance. Females of certain simuliid species can transmit viruses, protozoans and filarial parasites. The most
notorious filarial parasite transmitted by black flies is Onchocerca volvulus, a causative agent of human
onchocerciasis (also called “river blindness”) in Africa, and Central and South America (Crosskey 1990), followed
by O. dewittei japonica, a pathogen of zoonotic onchocerciasis, recently discovered in Japan (Takaoka et al. 2012).
The fauna of black flies in the Oriental Region, which was poorly studied before 1970 and once ranked last

among the six zoogeographical regions in number of species, has dramatically emerged as the second largest in the
number of constituent species following that of the Palaearctic Region. It has 524 species (23.8 % of the total world
number of extant species, 2,204; Adler and Crosskey 2016). The high biodiversity of black flies has been well
documented in some countries such as Indonesia, Malaysia, Philippines and Thailand (Takaoka 1983, 2003;
Takaoka and Choochote 2004b; Takaoka and Davies 1995, 1996; Takaoka and Suzuki 1984), whereas some other
countries in the region remain to be fully explored.
In Vietnam, the simuliid fauna has been poorly studied until recently. Only 16 species were recorded in the
twentieth century (Crosskey and Howard 1997; Pham 1998, 1999). In 2013 and 2014, we conducted surveys of
pupae and larvae of black flies in Tam Dao, Vinh Phuc Province (northern Vietnam); in Bach Ma and other
localities, Thua Thien Hue Province (central Vietnam); and in Dalat, Lam Dong Province (southern Vietnam) (Fig.
1), and added 30 more species including 22 new species (Takaoka et al. 2014a, b, c, 2015a). Of these 46 species,
which are all in the genus Simulium Latreille, 17, 4 and 25 species were placed in three subgenera: Gomphostilbia
Enderlein, Nevermannia Enderlein, and Simulium Latreille s. str., respectively (Adler and Crosskey 2016; Takaoka
et al. 2014a, b, c, 2015a).
Three among the 46 species recorded from Vietnam—S. (G.) asakoae Takaoka & Davies, S. (S.) nigrogilvum
Summers and S. (S.) nodosum Puri—were reported to transmit unknown filariae in Thailand (Takaoka et al. 2003,
Fukuda et al. 2003, Ishii et al. 2008). One species, S. (N.) aureohirtum Brunetti, was found to be autogenous in the
Ryukyu Islands, the Philippines, Malaysia, Thailand and Indonesia (Takaoka and Noda 1979; Takaoka 1989), but
biting habits and other biological aspects of these and other species remain to be studied in Vietnam, though six
species—S. (G.) asakoae, S. (S.) chungi Takaoka & Huang, S. (S.) maenoi Takaoka & Choochote, S. (S.)
grossifilum Takaoka & Davies, S. (S.) doipuiense Takaoka & Choochote (complex) (under the species name S. (S.)
rufibasis Brunetti), and S. (S.) vietnamense Takaoka, Sofian-Azirun & Chen—were attracted to humans in Tam
Dao National Park (Takaoka et al. 2014b).
We carried out surveys of larvae and pupae of black flies in Sapa, Lao Cai Province (northern Vietnam) in
December, 2014, and in Nghe An Province (northern Vietnam) in December, 2015 (Fig. 1). The results of these
surveys, coupled with reexaminations of black flies previously collected from Vinh Phuc Province and Lam Dong
Province (Fig. 1), yielded 22 new species and three new records. All these new species are here described and
illustrated.
The substitute name S. (S.) huense is given for the species that originally was described under the name S. (S.)
cavum Takaoka & Ya’cob from southern Vietnam. A redescription of the female, male, pupa and larva of S. (G.)
asakoae is presented, and the female and larva of S. (G.) hongthaii Takaoka, Sofian-Azirun & Ya’cob are described
for the first time.
The total number of species of black flies recorded from Vietnam is now 70, all of which are placed in four
subgenera of the genus Simulium (25 in Gomphostilbia; one in Montisimulium Rubtsov (a newly recorded
subgenus in Vietnam); seven in Nevermannia; and 37 in Simulium). Keys to identify the 10 subgenera in the
Oriental Region and all 70 species recorded from Vietnam are provided for females, males, pupae and mature
larvae.
The methods of collection, description and illustration, and terms for morphological features (Figs. 2 & 3)
follow those of Takaoka (2003) and partially those of Adler et al. (2004). Adult black flies examined
morphologically in this study were reared from pupae, and placed in 80% ethanol, together with their associated
pupal exuviae and cocoons, and larvae were fixed in Carnoy’s solution (one part glacial acetic acid and three parts
95% ethanol). Thoracic tissues of several adults fixed in 80% ethanol were used for a DNA sequence-based
analysis. The results of DNA analysis of the Simulium tani Takaoka & Davies complex was previously published
(Low et al. 2016), demonstrating the rich biodiversity of the Simulium tani complex and the discovery of more
cryptic taxa in Vietnam, and delineating taxonomic boundaries for each taxon within this species complex. Larvae
of the Simulium tuberosum species-group fixed in Carnoy’s solutions were chromosomally analyzed, by which we
also showed the wealth of biodiversity of the S. tuberosum species-group in Vietnam (Adler et al. 2016).
Taxonomic treatment of each taxon was mainly based on morphological characters, referring to molecular and
chromosomal data when available.
Holotypes and paratypes of the new species are deposited in the Institute of Biological Sciences, Faculty of
Science, University of Malaya, Kuala Lumpur, Malaysia. The register numbers of University Malaya Simuliidae
Research Project (abbreviated as UMSRP) are given for holotypes and paratypes.
FIGURE 1. Map of Vietnam showing the provinces where the current and previous surveys of black flies were carried out.
Nos. 1–5 show the five provinces where the current surveys were conducted (1, Lao Cai; 2, Vinh Phuc; 3, Nghe An; 4, Thua
Thien Hue; 5, Lam Dong), and nos. 6–11 show the provinces where Dr. X.D. Pham collected and recorded black flies (6, Lang
Son; 7, Bac Kan (a part of formerly Bac Thai); 8, Bac Giang; 9, Yen Bai; 10, Ha Tay; 11, Hoa Binh).

FIGURE 2. Morphological features and their terms of adults of black flies. A, Female head (front view); B, Thorax (lateral
view); C, Female legs (fore, mid and hind legs from left); D; Wing; E, Male genitalia (ventral view); F, Female terminalia
(ventral view). (Reproduced from Takaoka 2003).
FIGURE 3. Morphological features and their terms of the pupa and larva of black flies. A, Pupa in cocoon; B, Head
integument of pupa spread flattened (front view); C & D, Pupal abdomens (C, dorsal view; D, ventral view); E, Mature larva
(lateral view); F & G, Head capsules of larva (F, dorsal view; G, ventral view); H, Hypostoma; I, Posterior tip of larval abdomen
(posterodorsal view). (Reproduced from Takaoka 2003).
Systematics
Genus Simulium Latreille
The family Simuliidae is classified in 26 genera, of which Simulium is the largest and most widely distributed
genus, with 37 subgenera (Adler and Crosskey 2016). Simulium is the only genus recognized in the Oriental

Region, where a total of 524 species are recorded (Adler and Crosskey 2016), and they are further classified in 10
subgenera, i.e., three endemic subgenera (Asiosimulium Takaoka & Choochote, Daviesellum Takaoka & Adler, and
Wallacellum Takaoka), one semi-endemic subgenus (Gomphostilbia), four cosmopolitan subgenera (Byssodon
Enderlein, Eusimulium Roubaud, Nevermannia and Simulium), and two subgenera with a Palaearctic distribution
(Montisimulium and Wilhelmia Enderlein). In the Oriental Region, the subgenus Simulium is dominant, having 232
species (44.3%), followed by Gomphostilbia with 193 component species (36.8%) and Nevermannia with 55
species (10.5%), while seven other subgenera consist of 1 to 16 species.
Four subgenera (Gomphostilbia, Montisimulium, Nevermannia and Simulium) have been reported in Vietnam.
Definitions of the genus Simulium and 10 subgenera follow those given by Crosskey (1969), Takaoka (2003, 2012),
Takaoka and Adler (1997) and Takaoka and Choochote (2005b).
The keys to all these subgenera of the genus Simulium are herein updated. The subgenera and species-groups
known from Vietnam are shown in bold face type.
Keys to 10 subgenera of the genus Simulium in the Oriental Region
Females
1. Katepisternum haired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Katepisternum bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Calcipala elongate, reaching apex of second tarsomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wallacellum
Calcipala medium-long, not reaching apex of second tarsomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gomphostilbia
3. Pleural membrane haired. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Pleural membrane bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4. Claw without tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Wilhelmia
Claw with small subbasal tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simulium (S. ornatum species-group)
5. Postnotum with hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eusimulium
Postnotum without hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..6
6. Claw simple or with small subbasal tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Claw with large or medium-sized basal tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
7. Paraproct with cluster of dark spines anteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Daviesellum
Paraproct without cluster of dark spines anteriorly. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simulium (most species)
8. Basal section of radial vein bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Byssodon
Basal section of radial vein haired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Claw with medium-sized basal tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Montisimulium
Claw with large basal tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Cibarium with numerous spinous processes medially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiosimulium
Cibarium bare or with minute processes near ventral margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Nevermannia
Males
1. Style longer than coxite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Style shorter than coxite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Coxite much longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Daviesellum
Coxite as long as, or slightly shorter than, wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simulium
3. Katepisternum haired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Katepisternum bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
4. Calcipala elongate, reaching apex of second tarsomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wallacellum
Calcipala medium-long, not reaching apex of second tarsomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gomphostilbia
5. Pleural membrane haired. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Wilhelmia
Pleural membrane bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Basal section of radial vein bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Byssodon
Basal section of radial vein haired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Paramere without hooks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiosimulium
Paramere with parameral hook(s) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8. Hind basitarsus slender; ventral plate with median keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Hind basitarsus enlarged; ventral plate without median keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9. Ventral plate narrow, with divergent long arms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Eusimulium
Ventral plate broad, with parallel-sided short arms. . . . . . . . . . . . . . . . . . . . . . . . . . Nevermannia (S. ruficorne species-group)
10. Paramere with one long hook; median sclerite forked apically . . . . . . . . . . . . . . . . . Nevermannia (S. vernum species-group)
Paramere with three or more hooks; median sclerite club-shaped, not forked apically. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Montisimulium & Nevermannia (S. feuerborni species-group)
Pupae
1. Gill filaments with numerous minute black spots (Fig. 27H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Montisimulium
Gill filaments without minute black spots. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Abdominal segments 5–9 without spine-combs dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Abdominal segments with spine-combs dorsally (at least on segment 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3. Gill with 19–33 filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiosimulium
Gill with three, four, six or eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Gill with three filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Byssodon
Gill with four, six or eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Gill with four filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wallacellum
Gill with six or eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Wilhelmia
Gill with six filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Cocoon boot-shaped, with high neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Daviesellum
Cocoon shoe-shaped, with low neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Simulium (two species)
8. Frons with three pairs of trichomes; grapnel-shaped hooklets present on last abdominal segment (except a few species) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Gomphostilbia
Frons with two pairs of trichomes; grapnel-shaped hooklets absent on last abdominal segment. . . . . . . . . . . . . . . . . . . . . . . . .9
9. Abdominal segments 7 and 8 (or only segment 8) with spine-combs dorsally; terminal hooks absent or small, cone-like, if pres-
ent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simulium
Abdominal segments 6–9 (or 5–9 or 6–8) with spine-combs dorsally; terminal hooks large, cone-like . . . . . . . . . . . . . . . . . .10
10. Gill with inflated structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nevermannia (one species of S. vernum species-group)
Gill with four, five, six or eight thread-like filaments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Gill with four filaments . . . . . . . . . . . . . Eusimulium & Nevermannia (most species of S. vernum species-group, one species
of S. feuerborni species-group & one species of S. ruficorne species-group)
Gill with five, six or eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nevermannia (one species of S. ruficorne species-group)
Gill with five or six filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Gill with five filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nevermannia (one species of S. feuerborni species-group)
Gill with six filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nevermannia (most species of S. feuerborni species-group
& one species of S. ruficorne species-group)
Mature larvae
1. Hypostoma wide, with prominent median tooth accompanied by six small teeth on each side; posterior circlet with over 400
rows of hooklets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Daviesellum
Hypostoma with median tooth accompanied by one corner tooth and three intermediate teeth on each side; posterior circlet
with fewer than 250 rows of hooklets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Lateral margins of hypostoma smooth (Fig. 7E) (except a few species); main tooth of mandibular serrations at an acute angle
to mandible apically (Fig. 7D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Gomphostilbia
Lateral margins of hypostoma serrated; main tooth of mandibular serrations at right or obtuse angle to mandible apically . . .3
3. Mandibular serrations composed of two large teeth . . . . . . . . . . . . .Eusimulium & Nevermannia (S. ruficorne species-group)
Mandibular serrations composed of one large tooth and one small tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4. Postgenal cleft absent or vestigial (Fig. 27R) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Montisimulium
Postgenal cleft small to large. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
5. Antenna as long as stem of labral fan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Asiosimulium
Antenna longer than stem of labral fan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Pectination of outermost primary ray of labral fan different from those of other primary rays . . . . . . . . . . . . . . . . Wallacellum
Pectination of outermost primary ray of labral fan similar to those of other primary rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
7. Ventral papillae large. . . . . . . . . . Nevermannia (most species of S. feuerborni species-group & S. ruficorne species-group)
Ventral papillae absent or small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8. Rectal organ compound, each of three lobes with secondary lobules. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Simulium
Rectal organ simple, each of three lobes without secondary lobules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Histoblast of pupal gill with three filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Byssodon
Histoblast of pupal gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Wilhelmia
* The characters of the subgenus Byssodon used above are those of S. (B.) languidum, the only species of the subgenus in
the Oriental Region, and are not necessarily shared with other species of the subgenus in other regions.
I. Subgenus Gomphostilbia Enderlein
Twenty-five species are classified in the subgenus Gomphostilbia, and are further placed in seven species-groups
(S. asakoae, S. batoense, S. ceylonicum, S. darjeelingense, S. epistum, S. gombakense and S. varicorne species-

groups), defined or redefined by Takaoka (2012). The S. asakoae species-group containing 13 species is dominant,
followed by the S. batoense species-group accommodating seven species, while the other five species-groups are
each represented by one species.
Keys to 25 species of the subgenus Gomphostilbia
Females*
1. Claw with small subbasal tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dachaisense
Claw with large basal tooth (Fig. 4G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Hind tibia darkened except base yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Hind tibia darkened on apical one-third to one-half and yellow on rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
3. Sensory vesicle elongate, longer than 0.4 times length of third maxillary segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Sensory vesicle medium-long, shorter than 0.4 times length of third maxillary segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
4. Mid and hind femora yellow except apical cap darkened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. parahiyangum
Mid and hind femora light to medium brown except base yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Sensory vesicle elongate, 0.4 times length of third maxillary segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lamdongense
Sensory vesicle elongate, 0.5 times length of third maxillary segment (Fig. 22A) . . . . . . . . . . . . . . . . . . . . .S. eshimai sp. nov.
6. Fore coxa light brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. longlanhense
Fore coxa yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
7. Sternite 8 with about 24–28 stout hairs on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. montiblense
Sternite 8 with about 10 stout hairs on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Frons:head ratio 1.0:5.0. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. duolongum
Frons:head ratio 1.0:4.5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. siamense
9. Sensory vesicle elongate, 0.6 times length of third maxillary palpal segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
Sensory vesicle short to medium-long, 0.4 or less times length of third maxillary palpal segment . . . . . . . . . . . . . . . . . . . . . . 11
10. Frons:head ratio 1.0:5.2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. fuscidorsum
Frons:head ratio 1.0:4.3. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. thituyenae
11. Hind tibia darkened on apical one-third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
Hind tibia darkened on apical one-half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
12. Mandible with 15 outer teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. thuathienense
Mandible lacking outer teeth or with 1–6 outer teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
13. Outer margin of mandible without tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hongthaii
Outer margin of mandible toothed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Outer margin of mandible with two teeth (Fig. 17B).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. unii sp. nov.
Outer margin of mandible with five or six teeth (Fig. 4C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Sensory vesicle 0.3–0.4 times length of third maxillary palpal segment (Fig. 4A, B) . . . . . . . . . . . . . . . . . . . . . . . . . S. asakoae
Sensory vesicle 0.2–0.3 times length of third maxillary palpal segment (Fig. 8A) . . . . . . . . . . . . . . . . S. chaudinhense sp. nov.
16. Tuft hairs of base of radius brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. inthanonense
Tuft hairs of base of radius yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. confertum & S. phulocense
Males**
1. Antenna with eight flagellomeres . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. breviflagellum
Antenna with nine flagellomeres. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Hind basitarsus enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Hind basitarsus slender, parallel-sided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
3. Fore coxa darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. eshimai sp. nov.
Fore coxa yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Upper-eye facets in 16 vertical columns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Upper-eye facets in 9–14 vertical columns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5. Hair tuft of base of radius dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. inthanonense
Hair tuft of base of radius yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hongthaii
6. Antenna yellow except apical four or five or six flagellomeres darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Antenna darkened except scape, pedicel and base of first flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
7. Upper-eye facets in 9 or 10 vertical columns and 11 or 12 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Upper-eye facets in 11–13 vertical columns and 13 or 14 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
8. Upper-eye facets in 9 vertical columns and 11 or 12 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . S. vinhphucense sp. nov.
Upper-eye facets in 10 vertical columns and 12 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chaudinhense sp. nov.
9. Upper-eye facets in 11 vertical columns and 13 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. asakoae
Upper-eye facets in 13 or 14 vertical columns and 14 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. unii sp. nov.
10. Upper-eye facets in 14 vertical columns and 14 or 15 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Upper-eye facets in 11–13 vertical columns and 13 or 14 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
11. Paramere with setae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tamdaoense
Paramere without setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Hind basitarsus as wide as hind femur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. confertum
Hind basitarsus 0.8 times as wide as hind femur . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. quychauense sp. nov.
13. Hind basitarsus yellowish on basal two-fifths and dark on rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sanchayense sp. nov.
Hind basitarsus white on little more than basal half and dark on rest. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. phulocense
14. Abdominal segments 2, 6 and 7 each with pair of shiny dorsolateral patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Abdominal segments 2, 5, 6 and 7 each with pair of shiny dorsolateral patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
15. Style without terminal spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. parahiyangum
Style with terminal spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Ventral plate in ventral view with body tapered from base to middle, abruptly narrowed, then tapered posteriorly . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lamdongense
Ventral plate in ventral view with body nearly parallel-sided, though posterolateral corners rounded . . . . . . . . S. longlanhense
17. Upper-eye facets in 10 vertical columns and 10 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. montiblense
Upper-eye facets in 15–18 vertical columns and 15–18 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18. Upper-eye facets in 18 vertical columns and 18 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dachaisense
Upper-eye facets in 15 or 16 vertical columns and 15 or 16 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Scutum with golden-yellow short hairs. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. siamense
Scutum with brassy short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. duolongum
Pupae
1. Gill composed of inflated structure with eight slender filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. thuathienense
Gill composed of eight or ten slender filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Gill with ten filaments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lamdongense
Gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
3. All gill filaments short (about 1.0 mm). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
All or some gill filaments medium-long to long (more than 1.5 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
4. Antennal sheath with tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Antennal sheath without tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5. Gill filaments arranged as 3+3+2 from dorsal to ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. parahiyangum
Gill filaments arranged as 2+1+3+2 from dorsal to ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. longlanhense
6. Face with additional trichome mediolaterally on each side (Fig. 24A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. eshimai sp. nov.
Face without additional trichome mediolaterally on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. montiblense
7. Common basal stalk longer than interspiracular trunk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. fuscidorsum
Common basal stalk shorter than interspiracular trunk. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Dorsal and middle triplets of gill filaments not sharing common stalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Dorsal and middle triplets of gill filaments sharing common stalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9. Gill filaments with indistinct annular ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. duolongum
Gill filaments with marked annular ridges . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Dorsal surface of abdominal segments 1 and 2 grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. dachaisense
Dorsal surface of abdominal segments 1 and 2 pale yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. siamense
11. Dorsal surface of abdominal segments 1 and 2 (or also 3 and 4) darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Dorsal surface of abdominal segments 1 and 2 unpigmented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
12. Cocoon with long anterodorsal projection (Fig. 9L). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hmongense sp. nov.
Cocoon without anterodorsal projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
13. Dorsal surface of abdominal segments 1 and 2 with minute tubercles (Fig. 8C) . . . . . . . . . . . . . . . . . S. chaudinhense sp. nov.
Dorsal surface of abdominal segments 1 and 2 lacking tubercles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. asakoae
14. Cocoon with anterodorsal margin produced anteromedially, appearing as letter W or with short projection (Fig. 19E). . . . . . 15
Cocoon with anterodorsal margin somewhat produced anteromedially or not. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
15. Stalk of dorsal triplet against that of ventral pair of filaments at angle of 90 degrees or greater . . . . . . . . . . . . . . . . . . . . . . . 16
Stalk of dorsal triplet against that of ventral pair of filaments at angle of less than 90 degrees . . . . . . . . . . . . . . . . . . . . . . . . 17
16. Dorsal triplet of filaments with secondary stalk longer than primary stalk; terminal hooks broad, plate-like, with outer serrated
margin and three times length of inner margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. breviflagellum
Dorsal triplet of filaments with secondary stalk much shorter than primary stalk; terminal hooks narrow, plate-like, with outer
margin 1.8 times length of inner margin (Fig. 19D). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. unii sp. nov.
17. Stalk of ventral pair of filaments twice length of primary and secondary stalks of middle triplet of filaments combined . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. phulocense
Stalk of ventral pair of filaments as long as or shorter than length of primary and secondary stalks of middle triplet of filaments
combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. inthanonense
18. Abdominal segment 9 without spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. quychauense sp. nov.
Abdominal segment 9 with spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Dorsal and middle triplets with short common stalk shorter than their primary stalks . . . . . . . . . . . . . . . . . . . . . . .S. confertum
Dorsal and middle triplets with short common stalk nearly as long as or longer than their primary stalks . . . . . . . . . . . . . . . . 20
20. Terminal hooks flat and narrow, longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. thituyenae

Terminal hooks flat and wide, shorter than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
21. Stalk of ventral paired filaments as long as or shorter than common basal stalk (Fig. 21K). . . . . . . . . S. vinhphucense sp. nov.
Stalk of ventral paired filaments longer than common basal stalk . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
22. Terminal hooks with outer margin 1.8–1.9 times length of inner margin; stalk of ventral paired filaments shorter than primary
and secondary stalks of middle triplet combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. tamdaoense
Terminal hooks with outer margin 2.6–3.6 times length of inner margin; stalk of ventral paired filaments longer than primary
and secondary stalks of middle triplet combined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23. Two filaments of ventral pair 2.6 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hongthaii
Two filaments of ventral pair 3.0 mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. sanchayense sp. nov.
Mature larvae***
1. Postgenal cleft reaching posterior margin of hypostoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Postgenal cleft not reaching posterior margin of hypostoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2. Abdominal segments 1–5 each with two dorsal and two dorsolateral protuberances. . . . . . . . . . . . . . . . . . . . .S. parahiyangum
Abdominal segments 1–5 without such protuberances . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Anterolateral surfaces of abdominal segment 5 covered with conspicuous dark spinous setae longer than those on dorsal sur-
face; histoblast of gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. longlanhense
Anterolateral surfaces of abdominal segment 5 lacking conspicuous dark spinous setae; histoblast of gill with 10 filaments . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lamdongense
4. Postgenal cleft shorter than postgenal bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
Postgenal cleft slightly to greatly longer than postgenal bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5. Posterior circlet with 88 rows of hooklets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. inthanonense
Posterior circlet with 70 rows of hooklets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. hmongense sp. nov.
6. Postgenal cleft medium-long, 1.1–3.0 times as long as postgenal bridge. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Postgenal cleft long, more than 4.0 times as long as postgenal bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
7. Pharate pupal gill with inflated structure having eight slender filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. thuathienense
Pharate pupal gill with eight slender filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Abdominal segments 1–4 ochreous, without grayish markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. asakoae
Abdominal segments 1 and 2, or 1–3, or 1–4 grayish, or grayish green. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
9. Abdominal segments 1–4 grayish green . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. hongthaii
Abdominal segments 1 and 2, or 1–3 grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Abdominal segments 1–3 grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. fuscidorsum
Abdominal segments 1 and 2 grayish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Abdominal segment 4 with grayish markings ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chaudinhense sp. nov.
Abdominal segment 4 without grayish markings ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. unii sp. nov.
12. Posterior circlet with 80–88 rows of hooklets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Posterior circlet with about 70 rows of hooklets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13. Posterior circlet with 80 rows of hooklets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. montiblense
Posterior circlet with 85–88 rows of hooklets. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. eshimai sp. nov.
14. Head spots indistinct or faintly negative . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. dachaisense
Head spots faintly positive . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Posterior abdominal segments with dark spinous setae with distinct dark branches arising basally, medially and apically . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. siamense
Posterior abdominal segments with dark flat setae with fine transparent branches only apically . . . . . . . . . . . . . . S. duolongum
* The females of S. breviflagellum, S. hmongense sp. nov., S. sanchayense sp. nov., S. tamdaoense and S. vinhphucense
sp. nov. are unknown.
** The males of S. fuscidorsum, S. hmongense sp. nov., S. thituyenae and S. thuathienense are unknown.
*** The larvae of S. breviflagellum, S. confertum, S. phulocense, S. sanchayense sp. nov., S. tamdaoense, S. thituyenae and
S. vinhphucense sp. nov. are unknown.
1) Simulium asakoae species-group
Simulium (Gomphostilbia) asakoae Takaoka & Davies, 1995
Simulium (Gomphostilbia) asakoae Takaoka & Davies, 1995: 55–60 (Female, male, pupa and larva); Takaoka et al., 2014b:
743; Pham, 1998: 121.
Simulium (G.) asakoae was originally described from Peninsular Malaysia (Takaoka and Davies 1995) and
recorded from Vietnam (Pham 1998). A reexaminantion shows that the following characters differ from the
original description of this species: male antenna yellowish except apical five or six flagellomeres darkened, pupal
abdominal segments 1 and 2 without minute tubercles, and larval labral fan with 42–48 primary rays, and length
ratio of the antennal articles 1–3 (1.0:0.9–1.0:0.9–1.1). An analysis of DNA sequences shows only a slight
difference between S. (G.) asakoae populations from Vietnam and those from Peninsular Malaysia (Low et al.
unpublished data).
Female. Body length 2.4–2.5 mm. Head. Slightly narrower than width of thorax. Frons brownish black,
densely covered with yellowish-white scale-like recumbent short hairs interspersed with few dark longer hairs near
vertex; frontal ratio 1.7–1.8:1.0:2.3–2.5; frons:head ratio 1.0:4.6–5.0. Fronto-ocular area well developed, narrow,
directed dorsolaterally. Clypeus brownish black, densely covered with yellowish-white scale-like hairs interspersed
with several dark longer hairs on each side. Labrum 0.54–0.56 times length of clypeus. Antenna composed of
scape, pedicel and nine flagellomeres, light to dark brown except scape and pedicel yellow, and base of first
flagellomere yellowish white. Maxillary palp composed of five segments, light to medium brown, proportional
lengths of third, fourth, and fifth segments 1.0:1.1–1.2:2.8–2.9; third segment (Fig. 4A, B) widened apically;
sensory vesicle (Fig. 4A, B) small or medium sized, ellipsoidal (0.3–0.4 times length of third segment), with
medium or large opening. Maxillary lacinia with 9–12 inner and 13 or 14 outer teeth. Mandible (Fig. 4C) with 22 or
23 inner teeth and five or six outer teeth at some distance from tip. Cibarium (Fig. 4D) medially forming
sclerotized plate folded forward from posterior margin, with moderately sclerotized mediolongitudinal ridge with
bifid apex. Thorax. Scutum dark brown except anterolateral calli ochreous, and three blackish longitudinal vittae
(one median, two submedian), thinly pruinose and shiny when illuminated at certain angles, densely covered with
whitish-yellow to yellow scale-like recumbent short hairs except three vittae with dark recumbent short hairs.
Scutellum medium brown, covered with yellow short hairs and dark-brown long upright hairs along posterior
margin. Postnotum dark brown, slightly shiny when illuminated at certain angles, and bare. Pleural membrane bare.
Katepisternum longer than deep, medium to dark brown, shiny when illuminated at certain angles, moderately
covered with fine short hairs. Legs. Foreleg: coxa and trochanter yellowish white; femur dark yellow with apical
cap light brown (though extreme tip yellowish, and basal two-thirds of inner surface yellowish white); tibia white
except apical three-tenths dark brown, and little more than basal one-fourth grayish on inner surface, and covered
with white fine hairs on basal four-fifths; tarsus brownish black, with moderate dorsal hair crest; basitarsus
moderately dilated, 6.0–6.4 times as long as its greatest width. Midleg: coxa medium brown except posterolateral
surface dark brown; trochanter whitish yellow; femur light brown with basal one-fifth whitish yellow and apical
cap medium brown (though extreme tip yellowish); tibia yellowish white on basal three-fourths to four-fifths
(though apical two-thirds of anterior surface appearing darkened due to dark hair covering) and apical cap
brownish black, covered with yellowish fine hairs on posterior and inner surfaces of basal two-thirds; tarsus dark
brown to brownish black though basal half of basitarsus yellowish white. Hind leg: coxa light brown except apical
one-third yellowish; trochanter whitish yellow; femur light brown with base whitish yellow and apical cap dark
brown (though extreme tip yellowish white); tibia (Fig. 4E) yellowish white on basal two-thirds and brownish
black on rest, covered with whitish fine hairs on outer and posterior surfaces of little more than basal three-fourths;
tarsus brownish black except basal two-thirds (though base light brown) and basal half of second tarsomere
yellowish white; basitarsus (Fig. 4F) narrow, nearly parallel-sided though slightly narrowed apically, 5.5–6.2 times
as long as wide, and 0.7 and 0.6–0.7 times as wide as greatest widths of tibia and femur, respectively; calcipala
(Fig. 4F) nearly as long as width at base, and 0.6 times as wide as greatest width of basitarsus; pedisulcus (Fig. 4F)
well developed; claw (Fig. 4G) with large basal tooth 0.46–0.50 times length of claw. Wing. Length 2.0–2.1 mm.
Costa with dark spinules and hairs except basal patch of hairs yellow. Subcosta with dark hairs except near apex
bare. Hair tuft on base of radius yellow. Basal portion of radius fully haired; R1 with dark spinules and hairs; R2
with hairs only. Basal cell absent. Halter. White except basal portion darkened. Abdomen. Basal scale ochreous,
with fringe of whitish-yellow hairs. Dorsal surface of abdomen medium to dark brown except anterior half or little
more of segment 2 whitish, moderately covered with dark short to long hairs; tergites of segments 2 and 6–9 shiny
when illuminated at certain angles. Ventral surface of segment 2 white, those of segments 3 and 4 dark yellow to
light brown and those of other segments medium brown; sternal plate on segment 7 undeveloped. Terminalia.
Sternite 8 (Fig. 4H) bare medially, with 18–22 medium-long to long hairs together with four or five slender short
hairs on each side. Ovipositor valve (Fig. 4H) triangular (though posteromedial corner rounded), thin,
membranous, moderately covered with microsetae interspersed with one or two short hairs; inner margins slightly
sinuous, somewhat sclerotized, and moderately separated from each other. Genital fork (Fig. 4I) of usual inverted-

FIGURE 4. Female of Simulium (Gomphostilbia) asakoae. A & B, Third segments of maxillary palps with sensory vesicles
each having opening of different size (A, with opening of moderate size; B, with large opening; both right side and front view);
C, Tip of mandible; D, Cibarium (front view); E, Hind tibia (left side; outer view); F, Hind basitarsus and second tarsomere (left
side; outer view); G, Claw; H, Sternite 8 and ovipositor valve (right side only; ventral view); I, Genital fork (ventral view); J &
K, Paraprocts and cerci (J, ventral view; K, lateral view); L, Spermatheca. Scale bars. 0.1 mm for E and F; 0.02 mm for A, B, D
and H–L; 0.01 mm for C and G.
Y form, with slender stem; arms of moderate width, moderately folded medially, without lobe directed
posteromedially. Paraproct in ventral view (Fig. 4J) somewhat concave anterolaterally, with six sensilla on
anteromedial surface; paraproct in lateral view (Fig. 4K) slightly produced ventrally beyond ventral tip of cercus,
0.6–0.7 times as long as wide, with 22–27 medium-long to long hairs on ventral and lateral surfaces. Cercus in
lateral view (Fig. 4K) short, rounded posteriorly, 0.4 times as long as wide. Spermatheca (Fig. 4L) ellipsoidal, 1.4–
1.5 times as long as its greatest width, well sclerotized except duct and small area near juncture with duct
unsclerotized, and with many fissures on outer surface; internal setae absent; both accessory ducts slender,
subequal in diameter to major one.
Male. Body length 2.5–2.6 mm. Head. Somewhat wider than thorax. Upper eye bright medium brown,
consisting of 11 (rarely 12) vertical columns and 13 (rarely 14) horizontal rows of large facets. Face brownish
black, white pruinose. Clypeus brownish black, whitish pruinose, densely covered with golden-yellow scale-like
medium-long hairs (mostly directed upward) interspersed with several dark-brown simple longer hairs. Antenna
composed of scape, pedicel and nine flagellomeres, yellowish except flagellomeres 5–9 light to medium brown (or
yellow to dark yellow except base of flagellomere 1 yellowish white and flagellomeres 8 and 9 light to medium
brown in one male); first flagellomere elongate, 1.7 times length of second one. Maxillary palp light to medium
brown, with five segments, proportional lengths of third, fourth, and fifth segments 1.0:1.2–1.3:3.0–3.2; third
segment (Fig. 5A, B) slender; sensory vesicle (Fig. 5A, B) small, globular or ellipsoidal (0.2 times length of third
segment), and with small opening. Thorax. Scutum dark brown, with longitudinal vittae (one median, two
submedian) invisible or faintly visible, shiny and thinly gray pruinose when illuminated at certain angles, and
densely covered with whitish-yellow scale-like recumbent short hairs. Scutellum medium brown, covered with
yellow short hairs and dark-brown long upright hairs along posterior margin. Postnotum dark brown, slightly shiny
and white pruinose when illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum medium to
dark brown, longer than deep, shiny when illuminated at certain angles, moderately covered with fine short hairs.
Legs. Foreleg: coxa whitish yellow; trochanter yellow; femur dark yellow to light brown with apical cap medium
brown (though apical tip yellowish); tibia white except apical three-tenths dark brown, and covered with white
hairs on white portion; tarsus brownish black; basitarsus moderately dilated, 7.1–7.4 times as long as its greatest
width. Midleg: coxa light brown except posterior surface dark brown; trochanter yellowish; femur light brown with
base yellowish and apical cap medium brown (though apical tip yellow); tibia medium to dark brown except basal
one-third or little more whitish; tarsus dark brown except basal one-third or less of basitarsus yellow (border not
well defined). Hind leg: coxa light brown except apical portion yellow; trochanter yellowish; femur medium brown
with base yellowish white and apical cap dark brown (though apical tip yellow); tibia (Fig. 5C) dark brown to
brownish black except basal three-sevenths yellowish white; tarsus (Fig. 5D) medium to dark brown except little
less than basal half of basitarsus and basal half of second tarsomere whitish yellow; basitarsus (Fig. 5D) enlarged,
wedge-shaped, 3.3–3.5 times as long as wide, and 1.0–1.1 and 1.1–1.3 times as wide as greatest width of tibia and
femur, respectively; calcipala (Fig. 5D) slightly shorter than basal width, and 0.3 times as wide as greatest width of
basitarsus. Pedisulcus (Fig. 5D) well developed. Wing. Length 2.0 mm. Other characters as in female except
subcosta with one to three hairs (rarely no hairs). Halter. Dull white except basal stem darkened. Abdomen. Basal
scale dark brown, with fringe of light-brown hairs. Dorsal surface of abdomen dark brown to brownish black
except segment 2 light brown, covered with dark-brown short to long hairs except segment 2 with yellowish hairs;
segments 2 and 5–8 each with pair of shiny dorsolateral or lateral patches; ventral surface of segment 2 whitish,
those of segments 3 and 4 grayish white though sternal plates light brown, and those of other segments medium to
dark brown. Genitalia. Coxite in ventral view (Fig. 5E) nearly rectangular, 1.7 times as long as its greatest width.
Style in ventral view (Fig. 5E) bent inward, with triangular apex having single spine; style in ventrolateral view
(Fig. 5F) slightly tapered toward apex, with truncated apex. Ventral plate in ventral view (Fig. 5E) with body
transverse, 0.6 times as long as wide, with anterior margin produced anteromedially, posterior margin somewhat
concave medially, and lateral margin emarginated medially, and densely covered with microsetae on ventral
surface; basal arms of moderate length, slightly divergent, then convergent apically; ventral plate in lateral view
(Fig. 5G) moderately produced ventrally; ventral plate in caudal view (Fig. 5H) rounded ventrally, though ventral
margin nearly straight medially, densely covered with microsetae on posterior surface. Median sclerite (Fig. 5I)
plate-like, wide. Parameres (Fig. 5J) of moderate size, each with four distinct long and medium-long stout hooks,
and without minute setae on outer surface of basal arm. Aedeagal membrane (Fig. 5J) moderately setose; dorsal
plate not defined. Ventral surface of abdominal segment 10 (Fig. 5K, L) slightly sclerotized along anterior margin
and without distinct hairs near posterolateral corners. Cercus (Fig. 5K, L) small, rounded, with 12–17 hairs.

FIGURE 5. Male of Simulium (Gomphostilbia) asakoae. A & B, Third segments of maxillary palps with different shaped
sensory vesicles (A, globular; B, ellipsoidal; A, right side; B, left side; both front view); C, Hind tibia (left side; outer view); D,
Hind basitarsus and second tarsomere (left side; outer view); E, Coxites, styles and ventral plate (ventral view); F, style (right
side; ventrolateral view); G, Ventral plate and median sclerite (lateral view); H, Ventral plate (caudal view); I, Median sclerite
(caudal view); J, Paramere and aedeagal membrane (left side; caudal view); K & L, Abdominal segment 10 and cerci (right
side; K, lateral view; L, caudal view). Scale bars. 0.1 mm for C and D; 0.02 mm for A, B and E–L.
Pupa. Body length 2.5–2.7 mm. Head. Integument yellow, moderately covered with small round tubercles
except antennal sheaths and ventral surface almost bare; antennal sheath without any protuberances; frons with
three pairs of unbranched long trichomes with or without coiled apices (Fig. 6A); face with pair of unbranched
(rarely bifid) long trichomes with straight apices (Fig. 6A); three frontal trichomes on each side arising close
together, subequal in length to one another and slightly longer than facial one. Thorax. Integument yellow,
moderately covered with round tubercles, and with three long dorsomedial trichomes of different lengths (anterior
trichome longest, posterior one shortest) with coiled apices (Fig. 6A), two long anterolateral trichomes (anterior
trichome more slender and shorter with straight apex, posterior one with coiled apex) (Fig. 6A), one medium-long
mediolateral trichome with straight apex (Fig. 6A), and three ventrolateral trichomes (one medium-long, two short)
with straight apices (Fig. 6A) on each side; all trichomes unbranched except dorsal trichomes rarely bifid. Gill (Fig.
6B) composed of eight slender thread-like filaments, arranged as [3+(1+2)]+2 or [(2+1)+(1+2)]+2 from dorsal to
ventral, with short common basal stalk having somewhat swollen transparent basal fenestra at base; common basal
stalk 0.6 times length of interspiracular trunk; dorsal and middle triplets sharing short stalk, and dorsal triplet
composed of three individual filaments arising at same level, with short stalk, or composed of one individual and
two paired filaments with extremely short stalk, middle triplet composed of one individual and two paired
filaments and bearing short primary and secondary stalks; stalk of ventral pair of filaments medium-long, 1.4–1.7
times length of common basal stalk, and 0.8–1.0 times length of interspiracular trunk, and 0.9 times as thick as
common stalk of middle and dorsal triplets; common stalk of dorsal and middle triplets shorter than primary stalk
of middle triplet, and shorter or longer than stalk of dorsal triplet; middle triplet primary stalk longer than
secondary stalk; primary stalk of dorsal triplet lying against that of lower pair at angle of 70–90 degrees when
viewed laterally; filaments of dorsal triplet subequal in length (1.4–2.0 mm) and thickness to one another; filaments
of middle triplet subequal in length (1.8–2.7 mm) and thickness to one another; two filaments of ventral pair
subequal in length (2.6–3.2 mm) and thickness to each other and 1.4 and 1.3 times as thick as dorsal and middle
triplets of filaments when compared basally; all filaments medium to dark brown, gradually tapered toward apex;
cuticle of all filaments with well-defined annular ridges and furrows though becoming less marked apically,
densely covered with minute tubercles. Abdomen (Fig. 6C, E). Dorsally, segments 1 and 2 and basal half or more
of segment 3 (and rarely segment 4) light grayish, and other segments yellowish; all segments without tubercles;
segment 1 with one unbranched slender medium-long hair-like seta on each side; segment 2 with one unbranched
slender medium-long hair-like seta and five somewhat spinous minute setae submedially on each side; segments 3
and 4 each with four hooked spines and one somewhat spinous minute seta on each side; segment 5 lacking spine-
combs and comb-like groups of minute spines on each side; segments 6–9 each with spine-combs in transverse row
and comb-like groups of minute spines on each side (though spine-combs rarely absent on segment 9); segments 5–
8 each with few minute setae near posterior margin on each side; segment 9 with pair of wide flat terminal hooks,
of which outer margin is 3.5–4.1 times length of inner margin and crenulated (Fig. 6D) when viewed caudally.
Ventrally, segment 4 with one unbranched hook (subequal in size to those on segments 5–7) and few slender short
setae on each side; segment 5 with pair of bifid hooks submedially and few short slender setae on each side;
segments 6 and 7 each with pair of bifid (rarely trifid) inner and unbranched outer hooks somewhat spaced from
each other and few short slender setae on each side; segments 4–8 each with comb-like groups of minute spines.
Each side of segment 9 with three or four grapnel-shaped hooklets (Fig. 6F). Cocoon (Fig. 6G). Wall-pocket-
shaped, roughly to moderately woven, widely extended ventrolaterally; anterior margin thickly woven medially,
often with buldge; posterior three-fifths with floor roughly woven; individual threads visible; 3.4–4.0 mm long by
2.0–3.0 mm wide.
Mature larva. Body length 5.0–5.6 mm. Body light ochreous except ventral surface of abdominal segments 5–
9 whitish, with following color markings: thoracic segment 1 encircled with reddish-brown band (though
disconnected ventromedially) narrowly along anterior margin, thoracic segments 2 and 3 reddish brown on ventral
surface; abdominal segment 3 with or without pair of narrow reddish-brown lateral spots; abdominal segment 4
with pair of reddish-brown dorsolateral spots rarely faintly connected to each other ventrally, abdominal segments
5 and 6 each encircled by reddish-brown transverse band though disconnected ventromedially (often faded out
leaving round dorsomedian spot) and three similar colored spots on ventral surface, though those on segment 6
much lighter, abdominal segments 7–9 each covered with similar colored pigments to varying extent (as transverse
band similar to those on segments 5 and 6 in some larvae, but completely faded out in other larvae) on dorsal and
dorsolateral surface; abdominal segment 7 with similar colored transverse band ventrally (though usually faded out

FIGURE 6. Pupa of Simulium (Gomphostilbia) asakoae. A, Head and thorax (lateral view); B, Gill filaments (right side; outer
view); C, Abdomen (dorsal view); D, Terminal hooks (caudal view); E, Abdomen (ventral view); F, Grapnel-shaped hooklet; G,
Cocoon (dorsal view). Scale bars. 1.0 mm for G; 0.2 mm for A–C and E; 0.02 mm for D; 0.01 mm for F.
in most larvae). Head. Head capsule (Fig. 7A–C) yellow to dark yellow except eye-spot region whitish, sparsely
covered with minute setae (though moderately to densely on dorsal surface); head spots moderately positive though
anterior spots of posterolateral spots usually faint or indistinct (Fig. 7A); eyebrow distinct with small round dark
spot above it (Fig. 7B). Antenna composed of three articles and apical sensillum, longer than stem of labral fan;
proportional lengths of first, second, and third articles 1.0:0.9–1.0:0.9–1.1. Labral fan with 42–48 primary rays.
Mandible (Fig. 7D) with three comb-teeth decreasing in length from first tooth to third; mandibular serration
composed of two teeth (one medium-sized, one small); major tooth at angle of little less than 90 degrees against
mandible on apical side; supernumerary serrations absent. Hypostoma (Fig. 7E) with row of nine apical teeth, of
which median tooth is longer than each corner tooth; lateral margin smooth; five or six hypostomal bristles per side
lying nearly parallel to lateral margin. Postgenal cleft (Fig. 7C) arrow-head-shaped, medium-long, 2.2–2.7 times
length of postgenal bridge. Cervical sclerites composed of pair of small yellow rod-like pieces. Thorax and
Abdomen. Thoracic proleg without hairs; thoracic cuticle and abdominal cuticle of segments 1 and 2 sparsely
covered with unbranched unpigmented minute setae on dorsal surface; abdominal cuticle of segments 3–5 sparsely
covered with unbranched dark minute setae and unpigmented minute setae on dorsal surface, that of segments 6–9
moderately covered with slightly darkened unbranched minute setae mixed with unpigmented shorter minute setae
(Fig. 7F) on dorsal and dorsolateral surfaces; last abdominal segment densely covered with unbranched colorless
minute setae on dorsolateral and lateral surfaces of each side of anal sclerite and on each lateral surface even down
to base of ventral papilla; thorax and abdomen without dorsal protuberances. Rectal scales absent. Rectal organ
compound, each of three lobes with 9–11 finger-like secondary lobules. Anal sclerite of usual X-form, with anterior
arms 1.1 times as long as posterior ones, broadly sclerotized at base; no sensilla on broad base and posterior to
posterior arms; accessory sclerite absent. Last abdominal segment with pair of large conical ventral papillae.
Posterior circlet with 82–85 rows of hooklets with up to 13–15 hooklets per row.
FIGURE 7. Larva of Simulium (Gomphostilbia) asakoae. A, B & C, Head capsules (A, dorsal view; B, lateral view; C, ventral
view); D, Mandible; E, Hypostoma; F, Setae on dorsal surface of abdominal segment 8. Scale bars. 0.1 mm for A–C; 0.02 mm
for D–F.

Specimens examined: Five females, five males (together with their associated pupal exuviae), reared from
pupae and 10 mature larvae, collected from a slow-flowing stream (width 0.5–1.5 m, depth 10–20 cm, bottom
rocky, water temperature 18.0˚C, exposed to the sun, elevation 920 m, 21˚27’501’’N/105˚38’941’’E), Tam Dao,
Vinh Phuc Province, northern Vietnam, 8-XI-2013, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen &
K.W. Lau; one female and one male, one pupal exuviae and one mature larva, collected from a stream (width 30
cm, depth 5 cm, bottom sandy, water temperature 18.0˚C, exposed to the sun, elevation 80 m, 19˚20’12.147”N/
105˚09’15.908”E) slow flowing near a forest, Chau Dinh, Quy Chau, Nghe An Province, northern Vietnam, 11-
XII-2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W. Lau.
Distribution. Peninsular Malaysia, Hong Kong, Thailand, Vietnam (Lam Dong, Thua Thien Hue, Nghe An
and Vinh Phuc)
Remarks. Simulium (G.) asakoae is characterized in the female by the mandible with several teeth on the outer
margin (Fig. 4C) and hind tibia yellowish on the basal two-thirds (Fig. 4E), in the male by the presence of hairs on
the subcosta and much widened hind basitarsus as wide as or slightly wider than the hind tibia (Fig. 5D), in the
pupa by the dorsal surface of abdominal segments 1–3 (rarely 1–4) darkened (Fig. 6C), and in the larva by the
medium-sized postgenal cleft (Fig. 7C) and unicolored abdominal segments without dark transverse bands, though
reddish-brown pigment present usually on the dorsal surface of segments 5 and 6. A combination of these
characters separates this species from most other members of the S. asakoae species-group.
Females of S. (G.) asakoae were reported to bite humans (Choochote et al. 2005) and to serve as a vector of
unknown filarial species in Thailand (Fukuda et al. 2003; Ishii et al. 2008).
Simulium (Gomphostilbia) chaudinhense Takaoka & Sofian-Azirun sp. nov.
Female. Nearly as in female of S. (G.) asakoae except following characters. Body length 2.0–2.2 mm. Head.
Frontal ratio 1.6–1.8:1.0:2.2–2.4; frons:head ratio 1.0:4.0–4.8. Labrum 0.5 times length of clypeus. Maxillary palp:
proportional lengths of third, fourth, and fifth segments 1.0:1.2:2.7–2.9; sensory vesicle (Fig. 8A) small, ellipsoidal
(0.2–0.3 times length of third segment), with small opening. Maxillary lacinia with 9–11 inner and 11–15 outer
teeth. Mandible with 23 or 24 inner teeth and three or four outer teeth at some distance from tip. Legs. Foreleg:
coxa and trochanter yellowish white; femur light brown with apical cap medium brown (though extreme tip
yellowish); tibia white except little more than apical one-fourth dark brown, covered with white fine hairs on basal
four-fifths; basitarsus moderately dilated, 6.2–6.6 times as long as its greatest width. Midleg: femur light brown
with base yellowish white (though inner surface of basal half yellowish white) and apical cap medium brown
(though extreme tip yellowish); tibia light brown except basal one-third yellowish white (though inner surface of
basal two-thirds yellowish white) and apical cap brownish black, covered with yellowish fine hairs on posterior and
inner surfaces of basal two-thirds; tarsus medium to dark brown though basal half or little more of basitarsus
yellow. Hind leg: basitarsus 5.7–5.8 times as long as wide, and 0.7 and 0.6–0.7 times as wide as greatest widths of
tibia and femur, respectively; calcipala 0.5 times as wide as greatest width of basitarsus; claw with large basal tooth
0.53 times length of claw. Wing. Length 2.0–2.2 mm. Abdomen. Basal scale light brown, with fringe of whitish-
yellow hairs. Terminalia. Sternite 8 bare medially, with 14–22 medium-long to long hairs together with one or two
slender short hairs on each side. Ovipositor valves moderately covered with microsetae interspersed with two short
hairs; inner margins nearly straight or slightly sinuous, somewhat sclerotized. Paraproct in ventral view with three
to six sensilla on anteromedial surface; paraproct in lateral view 0.7 times as long as wide, with 19–32 medium-
long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.33 times as long as wide. Spermatheca
ellipsoidal, 1.5–1.6 times as long as its greatest width.
Male. Nearly as in male of S. (G.) asakoae except following characters. Body length 2.0–2.5 mm. Head.
Upper-eye facets in 10 (rarely 9) vertical columns and 12 horizontal rows. Antenna: first flagellomere 1.8 times
length of second one. Maxillary palp: proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.5; sensory
vesicle small, ellipsoidal (0.2 times length of third segment), and with small opening. Legs. Foreleg: basitarsus
moderately dilated, 7.6–7.9 times as long as its greatest width. Hind leg: basitarsus enlarged, wedge-shaped, 3.4
times as long as wide, and 1.0 and 1.2–1.3 times as wide as greatest width of tibia and femur, respectively;
calcipala slightly shorter than basal width, and 0.3 times as wide as greatest width of basitarsus. Wing. Length 2.0–
2.1 mm. Subcosta bare or with one to five hairs. Genitalia. Coxite in ventral view 1.9 times as long as its greatest
width. Style in ventrolateral view slightly tapered toward apex, with truncated apex, 0.8 times length of coxite.
Ventral plate in ventral view with body transverse, 0.5 times as long as wide, and posterior margin somewhat
concave medially (though posterior margin slightly convex medially when ventral plate is slightly tilted). Cercus
small, rounded, with 17 or 18 hairs.
Pupa. Body length 2.4–2.5 mm. As in pupa of S. (G.) asakoae except following characters. Thorax. Gill (Fig.
8B) composed of eight slender thread-like filaments, arranged as [(2+1)+(1+2)]+2 from dorsal to ventral; common
basal stalk 0.6–0.7 times length of interspiracular trunk; dorsal and middle triplets sharing short stalk 0.4–0.6 times
length of common basal stalk; dorsal triplet composed of one individual filament and two paired filaments with
short primary and secondary stalks (primary stalk usually much longer than secondary stalk) or three filaments
rarely arising at same level from primary stalk); middle triplet composed of one individual filament and two paired
filaments with short primary and secondary stalks (primary stalk usually much longer than secondary stalk, though
primary stalk slightly shorter than secondary stalk in one pupa); ventral paired filaments with medium-long stalk
1.1–1.5 times length of common basal stalk; primary stalk of dorsal triplet lying against that of lower pair at angle
of 75–90 degrees when viewed laterally; three filaments of dorsal triplet subequal in length (2.0–2.5 mm long
including their own stalks and common basal stalk) and thickness to one another; three filaments of middle triplet
subequal in length (2.7–2.9 mm) and thickness; two filaments of ventral pair subequal in length (3.0–3.1 mm) and
thickness to each other. Abdomen. Dorsally, segments 1 and 2 sparsely covered with minute tubercles (Fig. 8C);
segment 5 bare (though one spine-comb on each side in two pupae); segment 9 with pair of flat triangular terminal
hooks (Fig. 8D), of which outer margin 2.4–2.9 times length of inner margin and crenulated. Last segment with
three grapnel-shaped hooklets on each side. Cocoon. 3.3–4.0 mm long by 2.2–3.0 mm wide.
Mature larva. Body length 4.6–5.1 mm. Body whitish with following grayish and ochreous markings.
Grayish markings as in Fig. 8E: thoracic segment 1 encircled with grayish band (though disconnected
ventromedially) narrowly along anterior margin and grayish area on anterior surface of proleg, thoracic segment 2
grayish ventrally and faintly grayish dorsally; all these grayish markings on thorax faded out in some larvae;
abdominal segments 1 and 2 each always encircled with dark gray band; abdominal segment 3 rarely grayish
ventrally; abdominal segment 4 encircled with light grayish band, though often faded dorsally; abdominal segment
5 encircled with light grayish band, though often faded ventrally or dorsally or entirely; abdominal segment 6
rarely with gray areas laterally and ventrolaterally; abdominal segment 7 rarely with gray areas laterally and
ventrally. Ochreous markings as in Fig. 8F: thoracic segment 1 encircled with ochreous band; thoracic segments 2
and 3 faintly light ochreous dorsally and ventrally; abdominal segment 3 rarely light ochreous dorsally, abdominal
segments 4–9 each with distinct ochreous transverse band dorsally, of which those on segments 4–6 often extended
laterally and even ventrolaterally, and those of segments 6–8 usually fused to each other laterally, and abdominal
segment 7 often with ochreous transverse band ventrally. Head. Head capsule whitish yellow to yellow except
dorsomedial area along posterior margin sometimes darkened, eyes-pot region whitish, and lateral area along
posterior margin often darkened widely so that two large dark spots and two small dark spots in front of posterior
margin appearing to be obscure (Fig. 8G), sparsely covered with minute setae (though moderately on dorsal
surface); head spots distinctively positive though anterior spots of posterolateral spots usually faint or indistinct;
eyebrow not darkened, with small round dark spot above it (Fig. 8G). Antenna composed of three articles and
apical sensillum, longer than stem of labral fan; proportional lengths of first, second, and third articles 1.0:0.7:0.8–
0.9. Labral fan with 38–42 primary rays. Mandible (Fig. 8H) with three comb-teeth decreasing in length from first
tooth to third; mandibular serration composed of two teeth (one medium-sized, one small); major tooth at angle of
little less than 90 degree against mandible on apical side; supernumerary serrations absent. Hypostoma (Fig. 8I)
with row of nine apical teeth, of which median tooth is longer than each corner tooth; lateral margin smooth; four
or five hypostomal bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig. 8J) arrow-head-
shaped (though not pointed apically), medium-long, 2.1–2.6 times length of postgenal bridge. Cervical sclerites
composed of pair of small yellow rod-like pieces. Thorax and Abdomen. Thoracic proleg without hairs; thoracic
cuticle and abdominal cuticle of segments 1 and 2 almost bare; abdominal cuticle of segments 3 and 4 sparsely
covered with minute seate dorsally, that of segments 5–9 moderately covered with slightly darkened unbranched
minute setae mixed with unpigmented shorter minute setae (Fig. 8K) on dorsal and dorsolateral surfaces; last
abdominal segment densely covered with unbranched colorless minute setae on dorsolateral and lateral surfaces of
each side of anal sclerite and on each lateral surface even down to base of ventral papilla; thorax and abdomen
without dorsal protuberances. Rectal scales absent. Rectal organ compound,

FIGURE 8. Female, pupa and larva of Simulium (Gomphostilbia) chaudinhense sp. nov. A, Third segment of female maxillary
palp with sensory vesicle (right side, front view); B, Anterior half of pupal thorax and basal portion of gill filaments (right side;
outer view); C, Tubercles on pupal abdominal segment 1; D, Terminal hooks of pupal abdominal segment 9 (caudal view); E &
F, Entire body of mature larvae (both lateral view; E, grayish marking pattern; F, ochreous marking pattern); G, Larval head
capsule (lateral view); H, Larval mandible; I, Larval hypostoma; J, Larval head capsule (ventral view); K, Setae on dorsal
surface of larval abdominal segment 8. Scale bars. 0.5 mm for E and F; 0.1 mm for B, G and J; 0.02 mm for A, C, D, H, I and
K.
each of three lobes with 8–11 finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms 1.1
times as long as posterior ones, broadly sclerotized at base; no sensilla on broad base and posterior to posterior
arms; accessory sclerite absent. Last abdominal segment with pair of large conical ventral papillae. Posterior circlet
with 74 or 75 rows of hooklets with up to 13 or 14 hooklets per row.
Type material. HOLOTYPE: Male (with its associated pupal exuviae and cocoon) in 80% ethanol) labeled in
a vial as [UMSRP: Vietnam 024, Holotype, Simulium (G.) chaudinhense, Male, coll. Vietnam, 11-XII-2015,
Takaoka et al.], reared from a pupa collected from a slow-flowing stream (width 1.0 m, depth 5–10 cm, bottom
sandy, water temperature 18.0˚C, shaded, elevation 80 m, 19˚30’46.589”N/105˚09’10.470”E), Chau Dinh, Quy
Chau, Nghe An Province, northern Vietnam, 11-XII-2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W.
Lau. PARATYPES: Six females, two males and three mature larvae, labeled in vials as [UMSRP: Vietnam 024,
Paratype, Simulium (G.) chaudinhense, coll. Vietnam, 11-XII-2015, Takaoka et al.], same data as those of holotype.
Ecological notes. The pupae and larvae of this new species were attached to grass leaves trailing in the current.
Associated species were S. (G.) parahiyangum and S. (G.) unii sp. nov.
Distribution. Vietnam (Nghe An).
Etymology. The species name chaudinhense refers to the locality name, Chau Dinh, where this new species
was collected.
Remarks. This new species is similar to S. (G.) asakoae but is distinguished in the female by the shorter
sensory vesicle (Fig. 8A) and longer claw tooth, in the male by the smaller number of upper-eye facets, in the pupa
by the presence of minute tubercles on abdominal segments 1 and 2 (Fig. 8C), and in the larva by abdominal
segments 1 and 2 each encircled with a grayish band (Fig. 8E).
Simulium (Gomphostilbia) confertum Takaoka & Sofian-Azirun, 2015
Simulium (Gomphostilbia) confertum Takaoka & Sofian-Azirun (in Takaoka et al., 2015a): 7–13 (Female, male and pupa).
Specimens examined. Four males, reared from pupae collected from a moderately flowing stream (width 1.0 m,
depth 7 cm, bottom muddy, water temperature 13.0˚C, exposed to the sun, elevation 1,223 m, 22˚19’04.333”N/
103˚52’04.265”E), Lao Chai, Sapa, Lao Cai Province, 21-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob,
C.D. Chen & K.W. Lau.
Distribution. Vietnam (Lam Dong and Lao Cai).
Remarks. This species was described from Lam Dong Province in southern Vietnam (Takaoka et al. 2015a).
The males of this species collected from Sapa, Lao Cai Province in northern Vietnam have the subcosta with many
hairs except near the apex which is bare, differing from the male from Lam Dong (Takaoka et al. 2015a).
Simulium (Gomphostilbia) fuscidorsum Takaoka & Ya’cob, 2015
Simulium (Gomphostilbia) fuscidorsum Takaoka & Ya’cob (in Takaoka et al., 2015a): 13–17 (Female, pupa and larva).
Distribution. Vietnam (Lam Dong).

Remark. This species was described from Lam Dong Province in southern Vietnam (Takaoka et al. 2015a).
Simulium (Gomphostilbia) hmongense Takaoka & Ya’cob sp. nov.
Pharate Male. Only following characters observed. Upper-eye facets in 17 vertical columns and 17 horizontal
rows. Antenna composed of scape, pedicel and nine flagellomeres. Sensory vesicle small and rounded. Hind
basitarsus (Fig. 9A) enlarged. Costa with spinules and hairs. Basal portion of radius haired.
Pupa. Body length 3.4 mm. Head. Integument yellow, moderately covered with small round tubercles;
antennal sheath without any protuberances; frons with three pairs of unbranched long trichomes with coiled or
straight apices (Fig. 9B); face with pair of unbranched medium-long trichomes with straight apices (Fig. 9C); three
frontal trichomes on each side arising close together, subequal in length to one another and somewhat longer than

facial one. Thorax. Integument yellow except narrow mediolongitudinal portion of dorsal surface darkened,
sparsely covered with small round tubercles, with three long anterodorsal trichomes with coiled apices (Fig. 9D),
two long anterolateral trichomes (anterior trichome somewhat shorter than posterior one and with straight apex,
posterior trichome with coiled apex) (Fig. 9E), mediolateral trichome lost, and three ventrolateral trichomes with
straight apices (one medium-long, two short) (Fig. 9F), on each side; all trichomes unbranched. Gill (Fig. 9G)
composed of eight slender thread-like filaments, arranged as [(2+1)+(2+1)]+2 from dorsal to ventral; common
basal stalk medium-long, 0.8 times length of interspiracular trunk, having somewhat swollen basal fenestra at base;
dorsal and middle triplets sharing medium-long common stalk 0.9 times length of common basal stalk; dorsal
triplet composed of one individual and two paired filaments with short primary and secondary stalk in left gill or
composed of three individual filaments arising at same level from short stalk in right gill; primary stalk directed
inward and slightly upward, thus appearing to be hidden behind middle triplet; middle triplet directed forward,
composed of one individual filament and two paired filaments with long primary and medium-long secondary
stalks; stalk of ventral pair directed downward, then forward, 0.9 times as long as primary stalk of middle triplet
and 1.3 times length of interspiracular trunk; stalk of ventral pair 0.8 times as thick as common stalk of dorsal and
middle triplets, 1.1 and 1.3 times as thick as primary stalks of middle and dorsal triplets respectively; all filaments
grayish light-brown, gradually tapered toward apices; lengths of filaments probably little longer than 3.0 mm
(exact length of filaments not measurable due to loss of apical portions of all filaments); filaments of dorsal and
middle triplets subequal in length and thickness to one another, and appearing to be slightly shorter and thinner
than ventral paired filaments; cuticle of all filaments with well-marked annular ridges and furrows, which are,
though, becoming less distinct apically, and densely covered with minute tubercles, relatively larger ones on ridges.
Abdomen. Dorsally, segments 1–4 grayish light-brown, segment 9 yellowish, other segments unpigmented;
segment 1 with one unbranched slender medium-long hair-like seta (Fig. 9H) on each side; segment 2 with one
unbranched slender short hair-like seta (Fig. 9I) and four short somewhat spinous setae submedially near posterior
margin on each side; segments 3 and 4 each with four hooked spines and one short somewhat spinous seta near
posterior margin on each side; segments 5–8 each with two unbranched short setae near posterior margin on each
side; segments 5–9 each comb-like groups of minute spines near anterior margin; segments 6–9 each with spine-
combs in transverse row on each side; segment 9 with pair of small narrow blunt terminal hooks with rugged
surface apically (Fig. 9J, K). Ventrally, segment 4 with one unbranched hook and few unbranched short setae on
each side; segment 5 with pair of bifid hooks submedially and few unbranched short slender setae on each side;
segments 6 and 7 each with pair of bifid inner and unbranched outer hooks somewhat spaced from each other and
few unbranched short slender setae on each side; segments 4–8 each with comb-like groups of minute spines. Each
side of segment 9 with two grapnel-shaped hooklets. Cocoon (Fig. 9L). Wall-pocket-shaped, moderately and thinly
woven with no spaces in webs, widely extended ventrolaterally; anterior margin somewhat thickly woven, with
long anterodorsal projection directed forward; posterior half with floor roughly woven; individual threads partially
visible; 4.0 mm long by 3.6 mm wide.
Mature larva. Body length 5.0–5.5 mm. Body creamy except abdominal segments 1–3 greenish and dorsal
and dorsolateral surface of abdominal segments 4–9 light ochreous, with color markings as follows: thoracic
segment 1 with reddish-brown transverse band (though disconnected ventromedially), dorsal surface of thorax
thinly pinkish, ventral surface of thoracic segments 2 and 3 reddish brown, abdominal segment 3 thinly reddish
brown entirely or partially or not, abdominal segments 4–8 entirely or partially reddish brown dorsally,
dorsolaterally and ventrally (in particular markings on dorsal surface of segments 5 and 6 distinct). Head. Head
capsule whitish yellow to yellow, with faint positive head spots, and sparsely covered with unpigmented minute
setae (though moderately on dorsal surface). Antenna composed of three articles and apical sensillum, longer than
stem of labral fan; proportional lengths of first, second, and third articles 1.0:0.9–1.0:0.9. Labral fan with 42–44
primary rays. Mandible (Fig. 10A) with three comb-teeth decreasing in length from first tooth to third; mandibular
serration composed of two teeth (one medium-sized, one small) (left mandible in one larva with additional small
tooth at base of major tooth); major tooth at acute angle against mandible on apical side; supernumerary serrations
absent. Hypostoma (Fig. 10B) with row of nine apical teeth, of which median tooth is slightly longer than each
corner tooth; lateral margin smooth; four hypostomal bristles per side lying slightly divergent posteriorly from
lateral margin. Postgenal cleft (Fig. 10C) short, nearly quadrate, 0.9 times as long as wide, and 0.7 times length of
postgenal bridge. Cervical sclerites composed of pair of yellow small rod-like pieces. Thorax and Abdomen.
Thoracic proleg without hairs; thoracic cuticle bare; abdominal cuticle bare except last abdominal segment
moderately covered with colorless unbranched setae on each side of anal sclerite (even down to base of ventral
FIGURE 9. Male and pupa of Simulium (Gomphostilbia) hmongense sp. nov. A, Hind basitarsus and second tarsomere of male
leg (left side; outer view); B & C, Pupal cephalic trichomes (B, frontal; C, facial); D–F, Pupal thoracic trichomes (D,
anterodorsal; E, anterolateral; F, ventolateral); G, Anterior part of pupal thorax and gill filaments (left side; outer view); H & I,
Hair-like setae on dorsal surface of pupal abdomen ( H, on segment 1; I, on segment 2); J & K, Pupal terminal hooks (J, lateral
view; K, caudal view); L, cocoon (dorsal view). Scale bars. 1.0 mm for L; 0.2 mm for G; 0.1 mm for A; 0.04 mm for B–F, H
and I; 0.02 mm for J and K.

FIGURE 10. Larva of Simulium (Gomphostilbia) hmongense sp. nov. A, Mandible; B, Hypostoma; C, Head capsule (ventral
view). Scale bars. 0.1 mm for C; 0.02 mm for A and B.
papilla on each side); thorax and abdomen without dorsal protuberances. Rectal scales present, unpigmented.
Rectal organ compound, each of three lobes with several finger-like secondary lobules (exact number of secondary
lobules uncountable). Anal sclerite of usual X-form, with anterior arms 1.2 times length of posterior ones, broadly
sclerotized at base; no sensilla on broad base and posterior to posterior arms; accessory sclerite absent. Last
abdominal segment with pair of large conical ventral papillae. Posterior circlet with 70 rows of hooklets with up to
12 hooklets per row.
Type material. HOLOTYPE: Pupa (pharate male) in 80% ethanol, labeled in a vial as [UMSRP: Vietnam 025,
Holotype, Simulium (G.) hmongense, Pharate male, coll. Vietnam, 20-XII-2014, Takaoka et al.], collected from a
small stream (width 0.2 m, depth 2 cm, bottom rocky, water temperature 7.0˚C, partially shaded, elevation 1,893 m,
22˚21’28.378”N/103˚45’52.084”E) slowly flowing in shrub, Lai Chau, Sapa, Lao Cai Province, northern Vietnam,
20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: Two mature
larvae in 80% ethanol, labeled in a vial as [UMSRP: Vietnam 025, Paratype, Simulium (G.) hmongense, Larvae,
coll. Vietnam, 20-XII-2014, Takaoka et al.], collected from a stream (width 0.2 m, depth 5 cm, bottom sandy and
muddy, water temperature 8.0˚C, exposed to the sun, elevation 1,925 m, 22˚21’11.767”N/103˚46’28.309”E) slowly
flowing in shrub in grassland, Lai Chau, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka,
M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
Biological notes. The pupa and larvae of this new species were collected from plastic sheets in the current.
Associated species were S. (M.) nigrofilum sp. nov., S. (N.) maeaiense and S. (S.) daoense sp. nov.
Etymology. The species name hmongense refers to the land in which the Hmong tribe lives.
Remarks. This new species may be assigned to one of the three species-groups (S. asakoae, S. ceylonicum and
S. darjeelingense species-groups) of the subgenus Gomphostilbia, redefined by Takaoka (2012), by having the
enlarged male hind basitarsus (Fig. 9A). This new species is placed in the S. asakoae species-group due to the
similarity of the unique pupal terminal hooks (Fig. 9J, K) to those of S. (G.) lurauense Takaoka, Sofian-Azirun &
Hashim from Peninsular Malaysia and S. (G.) thituyenae Takaoka & Pham from Vietnam, both of which are
members of the S. asakoae species-group (Takaoka et al. 2011, 2015a).
This new species is characterized by the greater number of male upper-eye facets in 17 vertical columns and
17 horizontal rows, pupal thoracic integument sparsely covered with tubercles, middle triplet of the gill with a long
primary stalk (Fig. 9G), cocoon with a long anterodorsal projection (Fig. 9L), and small larval postgenal cleft (Fig.
10C). Among the species of the S. asakoae species-group, only two species, S. (G.) chiangdaoense Takaoka &
Srisuka from Thailand and S. (G.) fuscidorsum Takaoka & Ya’cob from Vietnam, have the cocoon with a long
anterodorsal projection (Takaoka and Srisuka 2009; Takaoka et al. 2015a). However, S. (G.) chiangdaoense differs
by having the number of male upper-eye facets in 13 or 14 vertical columns and 15 or 16 horizontal rows, pupal
thoracic integument moderately covered with tubercles and middle triplet of the pupal gill with a short primary
stalk, and S. (G.) fuscidorsum differs by having the pupal thoracic integument moderately covered with tubercles,
long common basal stalk of the pupal gill, and shape of the terminal hooks, of which the outer margin is longer than
the inner one and crenulated.
Out of seven species among the S. ceylonicum species-group, which have the cocoon with a long anterodorsal
projection, S. (G.) inthanonense Takaoka & Suzuki described from Thailand (Takaoka and Suzuki 1984) has a
similar number of male upper-eye facets (16 vertical columns and 16 horizontal rows) and small larval postgenal
cleft, but is distinguished from this new species by the ellipsoidal male sensory vesicle, arrangement of the pupal
gill filaments (length ratio of the primary stalk of the middle triplet against the stalk of the ventral pair is 0.6–0.7,
whereas it is 1.1 in this new species) and shape of the terminal hooks, of which the outer margin is much longer
than the inner one. Another species, S. (G.) ogatai (Rubtsov) described from Japan (Rubtsov 1959–1964), shares a
similar number of male upper-eye facets in 14 or 15 vertical columns and 16 horizontal rows, and similar small
pupal terminal hooks, but differs by the arrangement of the pupal gill filaments (length ratio of the primary stalk of
the middle triplet against the stalk of the ventral pair is 0.4–0.5, whereas it is 1.1 in this new species), and dorsal
surface of pupal abdominal segments 1–4 unpigmented. (Characters of S. (G.) ogatai used here are based on
specimens collected in Gifu, Japan, in July 17, 2011 by Dr. Y. Otsuka).
All five species of the S. darjeelingense species-group have simple wall-pocket-shaped cocoons, thus differing
from this new species.
Simulium (Gomphostilbia) hongthaii Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Gomphostilbia) hongthaii Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014a): 348– 351 (Male and pupa).
This species was described from males and pupae collected from Tam Dao National Park in northern Vietnam
(Takaoka et al. 2014a). The female and larva of this species are here described for the first time.
Female. Body length 2.0–2.3 mm. Head. Slightly narrower than width of thorax. Frons dark brown, densely
covered with yellowish-white scale-like recumbent short hairs interspersed with few dark longer hairs near vertex;
frontal ratio 1.6–1.7:1.0:1.9–2.2; frons:head ratio 1.0:4.1–4.2. Fronto-ocular area well developed, narrow, directed
dorsolaterally. Clypeus dark brown, densely covered with yellowish-white short hairs interspersed with several
dark longer hairs near ventral margin on each side. Labrum 0.65–0.69 times length of clypeus. Antenna composed
of scape, pedicel and nine flagellomeres, dark brown except scape, pedicel, and base of first flagellomere yellow.
Maxillary palp composed of five segments, light to medium brown except segments 1 and 2 dark yellow,
proportional lengths of third, fourth, and fifth segments 1.0:1.1–1.2:2.5–2.6; third segment (Fig. 11A) widened;
sensory vesicle (Fig. 11A) ellipsoidal (0.3–0.4 times length of third segment), with medium-sized opening.
Maxillary lacinia with 8 inner and 12–14 outer teeth. Mandible (Fig. 11B) with 17–21 inner teeth and lacking outer
teeth though outer margin undulate, appearing to have four to six rudimentary teeth. Cibarium (Fig. 11C) medially
forming sclerotized plate folded forward from posterior margin, with moderately sclerotized mediolongitudinal
ridge with bifid apex. Thorax. Scutum dark brown to brownish black except anterolateral calli ochreous, and three
blackish longitudinal vittae (one median, two submedian) though not clearly demarcated, thinly pruinose and shiny
when illuminated at certain angles, densely covered with whitish-yellow to yellow scale-like recumbent short hairs
except three longitudinal vittae covered with dark short hairs. Scutellum ochreous, covered with yellow short hairs
and dark-brown longer upright hairs along posterior margin. Postnotum dark brown, whitish-gray pruinose and
shiny when illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum longer than deep,
medium to dark brown, shiny when illuminated at certain angles, moderately covered with fine short hairs. Legs.
Foreleg: coxa and trochanter whitish yellow; femur light brown with apical cap medium brown (though extreme tip
yellowish); tibia white except apical one-fourth dark brown, covered with white fine hairs on basal four-fifths;
tarsus brownish black, with moderate dorsal hair crest; basitarsus moderately dilated, 6.6 times as long as its
greatest width. Midleg: coxa medium brown except posterolateral surface dark brown; trochanter dark yellow with

FIGURE 11. Female of Simulium (Gomphostilbia) hongthaii. A, Third segment of maxillary palp with sensory vesicle (right
side; front view); B, Tip of mandible; C, Cibarium (front view); D, Hind basitarsus and second tarsomere (left side; outer view).
E, Claw; F, Sternite 8 and ovipositor valve (right side only; ventral view); G, Genital fork (ventral view); H & I, Paraprocts and
cerci (H, ventral view; I, lateral view); J, Spermatheca. Scale bars. 0.1 mm for D; 0.02 mm for A, C and E–J; 0.01 mm for B.
base whitish yellow; femur dark yellow to light brown with basal half of inner surface yellow and apical cap
medium brown (though extreme tip yellowish); tibia light brown except basal two-fifths whitish yellow and apical
cap dark brown (and apical three-fifths of posterior surface dark brown), covered with whitish and yellowish fine
hairs on posterior and inner surfaces of little more than basal half; tarsus dark brown though basal half or little less
of basitarsus yellow. Hind leg: coxa light brown; trochanter whitish yellow; femur light brown with base whitish
yellow and apical cap medium brown (though extreme tip yellowish); tibia white or yellowish white on basal two-
thirds and light brown on rest though apical cap brownish black, covered with whitish fine hairs on outer and
posterior surfaces of little more than basal three-fourths; tarsus brownish black except basal two-thirds or little
more of basitarsus (though base light brown) and basal half of second tarsomere yellowish white; basitarsus (Fig.
11D) narrow, nearly parallel-sided though slightly narrowed apically, 7.0 times as long as wide, and 0.6 and 0.6
times as wide as greatest widths of tibia and femur, respectively; calcipala (Fig. 11D) nearly as long as width at
base, and 0.6 times as wide as greatest width of basitarsus; pedisulcus (Fig. 11D) well developed; claw (Fig. 11E)
with large basal tooth 0.46 times length of claw. Wing. Length 2.2 mm. Costa with dark spinules and hairs except
basal patch of hairs yellow. Subcosta with dark hairs except near apex bare. Hair tuft on base of radius yellow.
Basal portion of radius fully haired; R1 with dark spinules and hairs; R2 with hairs only. Basal cell absent. Halter.
White except basal portion darkened. Abdomen. Basal scale light ochreous, with fringe of whitish-yellow hairs.
Dorsal surface of abdomen light to dark brown except segment 2 yellow to light ochreous, moderately covered with
dark short to long hairs; tergites of segments 2 and 6–9 shiny when illuminated at certain angles. Ventral surface of
segment 2 yellowish white, those of segments 3–5 ochreous and those of other segments medium brown; sternal
plate on segment 7 undeveloped. Terminalia. Sternite 8 (Fig. 11F) bare medially, with 17–22 medium-long to long
hairs together with few slender short hairs on each side. Ovipositor valve (Fig. 11F) triangular (though
posteromedial corner rounded), thin, membranous, moderately covered with microsetae interspersed with one to
three short hairs; inner margins nearly straight or slightly sinuous, somewhat sclerotized, and moderately separated
from each other. Genital fork (Fig. 11G) of usual inverted-Y form, with slender stem; arms of moderate width,
moderately folded medially, without lobe directed posteromedially. Paraproct in ventral view (Fig. 11H) somewhat
concave anterolaterally, with three or four sensilla on anteromedial surface; paraproct in lateral view (Fig. 11I)
slightly produced ventrally beyond ventral tip of cercus, 0.7 times as long as wide, with 17–24 medium-long to
long hairs on ventral and lateral surfaces. Cercus in lateral view (Fig. 11I) short, rounded posteriorly, 0.5 times as
long as wide. Spermatheca (Fig. 11J) ellipsoidal, 1.4 times as long as its greatest width, well sclerotized except duct
and small area near juncture with duct unsclerotized, and with many fissures on outer surface; internal setae absent;
both accessory ducts slender, subequal in diameter to major one.
Mature larva. Body length 5.0 mm. Thorax whitish to light ochreous except anterior portion of first segment,
proleg and ventral surface of segments 2 and 3 light gray, with reddish-brown transverse band on dorsal surface of
segment 1 (though often disconnected medially); abdominal segments 1–4 entirely grayish, abdominal segments 5–
9 light gray on dorsal and dorsolateral surfaces, abdominal segments 5 and 6 each with reddish-brown transverse
band dorsally, and abdominal segments 7–9 faintly and irregularly mottled with reddish-brown markings dorsally.
Head. Head capsule (Fig. 12A) whitish yellow except surrounding areas of posterior spots of posterolateral spots
darkened, and sparsely covered with unpigmented minute setae (though moderately on dorsal surface); all head
spots on dorsal, lateral and ventral surfaces distinctively positive except anterior spots of posterolateral spots
indistinct and posterior spots of posterolateral spots merged into darkened surrounding areas on cephalic apotome;
eyebrow faintly visible. Antenna composed of three articles and apical sensillum, longer than stem of labral fan;
proportional lengths of first, second, and third articles 1.0:0.7:0.9. Labral fan with 41 primary rays. Mandible (Fig.
12C) with three comb-teeth decreasing in length from first tooth to third; mandibular serration composed of two
teeth (one medium-sized, one small); major tooth at acute angle against mandible on apical side; supernumerary
serrations absent. Hypostoma (Fig. 12D) with row of nine apical teeth, of which median tooth is longer than each
corner tooth; lateral margin smooth; five hypostomal bristles per side lying slightly divergent posteriorly from
lateral margin. Postgenal cleft (Fig. 12B) rounded, medium-long, 1.3 times length of postgenal bridge. Cervical
sclerites composed of pair of small light-brown rod-like pieces. Thorax and Abdomen. Thoracic proleg without
hairs; thoracic cuticle bare; abdominal cuticle of segments 1–6 sparsely covered with unpigmented minute setae on
dorsal and dorsolateral surface, and that of segments 7–9 sparsely to moderately covered with slightly darkened
unbranched minute setae interspersed with unbranched colorless setae (Fig. 12E) on dorsal and dorsolateral
surfaces; last abdominal segment densely covered with unbranched colorless longer setae on dorsolateral and
lateral surfaces of each side of anal sclerite and on each lateral surface even down to base of ventral papilla; thorax
and abdomen without dorsal protuberances. Rectal scales absent. Rectal organ compound, each of three lobes with
11–14 finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms 1.1 times length of posterior
ones, broadly sclerotized at base; no sensilla on broad base and posterior to posterior arms; accessory sclerite
absent. Last abdominal segment with pair of large conical ventral papillae. Posterior circlet with 83 rows of
hooklets with up to 13 hooklets per row.

FIGURE 12. Larva of Simulium (Gomphostilbia) hongthaii. A & B, Head capsules (A, dorsal view; B, ventral view); C,
Mandible; D, Hypostoma; E, Setae on dorsal surface of abdominal segment 8. Scale bars. 0.1 mm for A and B; 0.02 mm for C–
E.
Specimens examined. Four females, four males and one mature larva collected from a small moderately-
flowing stream (width 1.5 m, depth 10 cm, water temperature 13.0˚C, partially shaded, elevation 1,105 m,
22˚18’48.273”N/103˚53’10.384”E), Sapa, Lao Cai Province, northern Vietnam, 21-XII-2014, by H. Takaoka, M.
Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
Distribution. Vietnam (Lam Dong, Lao Cai, Thua Thien Hue and Vinh Phuc).
Remarks. The female of S. (G.) hongthaii is similar to that of S. (G.) asakoae originally described from
Peninsular Malaysia (Takaoka and Davies 1995) in many characters including the size of the sensory vesicle (Fig.
11A) and leg colors but is distinguished from the latter by the mandible lacking distinct outer teeth (Fig. 11B). The
larva of this species is distinguished from that of S. (G.) asakoae by the grayish abdominal segments 1–4.
Simulium (Gomphostilbia) phulocense Takaoka & Chen, 2015
Simulium (Gomphostilbia) phulocense Takaoka & Chen (inTakaoka et al., 2015a):18–24 (Female, male and pupa).
Distribution. Vietnam (Thua Thien Hue).
Remark. This species was described from Thua Thien Hue Province in central Vietnam (Takaoka et al.
2015a).
Simulium (Gomphostilbia) quychauense Takaoka & Chen sp. nov.
Male. Body length 2.3 mm. Head. Wider than thorax. Upper eye medium brown, consisting of large facets in 14
vertical columns and 14 horizontal rows. Face brownish black, white pruinose. Clypeus brownish black, whitish
pruinose, densely covered with golden-yellow scale-like medium-long hairs (mostly directed upward) interspersed
with several dark-brown longer hairs. Antenna composed of scape, pedicel and nine flagellomeres, dark brown
except scape and pedicel yellow and base of first flagellomere whitish yellow; first flagellomere elongate, 1.8 times
length of second one. Maxillary palp light to medium brown, with five segments, proportional lengths of third,
fourth, and fifth segments 1.0:1.2:2.5; third segment (Fig. 13A) slender; sensory vesicle (Fig. 13A) ellipsoidal,
small (0.2 times length of third segment), and with small opening. Thorax. Scutum brownish black, shiny and
thinly gray pruinose on prescutellar area when illuminated at certain angles, densely covered with golden-yellow
scale-like recumbent short hairs. Scutellum dark brown, covered with yellow short hairs and dark-brown longer
upright hairs. Postnotum brownish black, and white pruinose when illuminated at certain angles, and bare. Pleural
membrane bare. Katepisternum dark brown, longer than deep, moderately covered with fine short hairs. Legs.
Foreleg: coxa whitish yellow; trochanter light brown except base yellow; femur medium brown with apical cap
dark brown (though apical tip yellowish); tibia light brown except little less than apical one-third brownish black,
and outer surface paler medially, and covered with yellow hairs on basal two-thirds of outer surface; tarsus
brownish black; basitarsus moderately dilated, 6.9 times as long as its greatest width. Midleg: coxa dark brown;
trochanter light brown except base whitish yellow; femur light to medium brown with apical cap dark brown
(though apical tip yellow); tibia brownish black except basal one-third whitish yellow; tarsus brownish black
except base of basitarsus dark yellow (border not well defined). Hind leg: coxa medium brown; trochanter whitish
yellow; femur medium brown with base narrowly whitish yellow and apical cap dark brown (though apical tip
whitish yellow); tibia (Fig. 13B) medium to dark brown except basal two-fifths yellow, and apical cap brownish
black; tarsus (Fig. 13C) medium brown except basal half of basitarsus grayish white (widely covered with dark
hairs, thus border not well defined) and basal two-fifths of second tarsomere whitish; basitarsus (Fig. 13C)
enlarged, 4.4 times as long as wide, and 0.9 and 0.8 times as wide as greatest width of tibia and femur, respectively;
calcipala (Fig. 13C) as long as basal width, and 0.4 times as wide as greatest width of basitarsus. Pedisulcus (Fig.
13C) well developed. Wing. Length 1.9 mm. Costa with dark spinules and hairs except hairs on basal portion
yellow. Subcosta without hairs. Hair tuft on base of radius yellow. Basal portion of radius fully haired; R1 with dark
spinules and hairs; R2 with hairs only. Basal cell absent. Halter. White except outer surface and basal stem
darkened. Abdomen. Basal scale dark brown, with fringe of dark hairs. Dorsal surface of abdomen medium to
brownish black, covered with dark-brown short to long hairs; segments 2 and 5–7 each with pair of shiny
dorsolateral or lateral patches; ventral surface of abdomen light to dark brown though segments 2–4 paler.
Genitalia. Coxites, styles, ventral plate and median sclerite in ventral view as in Fig. 13D. Coxite in ventral view
(Fig. 13D) nearly rectangular, 1.8 times as long as its greatest width. Style in ventral view (Fig. 13D) bent inward,
nearly parallel-sided, bluntly pointed apically and with apical spine; style in ventrolateral view (Fig. 13E) shorter
than coxite (0.8 times length of coxite), 2.3 times as long as its greatest width at base, tapered to middle, then
nearly parallel-sided and with round apex. Ventral plate in ventral view (Fig. 13D) with body transverse, 0.5 times
as long as wide, with anterior margin produced anteromedially, and posterior margin appearing as flattened letter
W, lateral margins somewhat emarginated basally, then slightly divergent posteriorly, and densely covered with
microsetae on ventral surface; basal arms of moderate length, directed forward, then convergent apically; ventral
plate in lateral view (Fig. 13F) moderately produced ventrally; ventral plate in caudal view (Fig. 13G) trapezoidal,
with ventral margin nearly straight though slightly produced medially, densely covered with microsetae on
posterior surface except portion near each lateral tip narrowly bare. Median sclerite (Fig. 13D, F, H) plate-like and
wide. Parameres (Fig. 13I) of moderate size, each with four or five distinct long to short stout hooks. Aedeagal
membrane (Fig. 13I) moderately setose; dorsal plate not defined. Ventral surface of abdominal segment 10 (Fig.
13J, K) weakly sclerotized anterolaterally and without distinct hairs near posterior margin. Cercus (Fig. 13J, K)

FIGURE 13. Male of Simulium (Gomphostilbia) quychauense sp. nov. A, Third segment of maxillary palp with sensory
vesicle; B, Hind tibia (left side; outer view); C, Hind basitarsus and second tarsomere (left side; outer view); D, Coxites, styles
and ventral plate (ventral view); E, Style (right side; ventrolateral view); F, Ventral plate and median sclerite (lateral view); G,
Ventral plate (caudal view); H, Median sclerite (caudal view); I, Paramere and aedeagal membrane (left side; caudal view); J &
K, Abdominal segment 10 and cerci (left side; J, lateral view; K, caudal view). Scale bars. 0.1 mm for B and C; 0.02 mm for A
and D–K.
Pupa. Body length 2.4 mm. Head. Integument yellow, moderately and neatly covered with small round
tubercles except antennal sheaths and ventral surface almost bare; antennal sheath without any protuberances; frons
with three pairs of unbranched long trichomes with coiled or straight apices (Fig. 14A); face with pair of
unbranched long trichomes with straight apices (Fig. 14B); three frontal trichomes on each side arising close
together, subequal in length to one another and somewhat longer than facial one. Thorax. Integument yellow,
moderately covered with round tubercles except dorsolateral portion of posterior half sparsely covered, and with
three long dorsomedial trichomes of different lengths (anterior trichome longest, posterior one shortest) with coiled
apices (Fig. 14C), two long anterolateral trichomes with straight apices (anterior trichome more slender and
slightly shorter than posterior one) (Fig. 14D), one medium-long mediolateral trichome with straight apex (Fig.
14E), and three ventrolateral trichomes with straight apices (one long, two short) (Fig. 14F) on each side; all
trichomes unbranched. Gill (Fig. 14G) composed of eight slender thread-like filaments, arranged as
[(2+1)+(1+2)]+2 from dorsal to ventral, with medium-long common basal stalk having somewhat swollen
transparent basal fenestra at base; common basal stalk 0.8–0.9 times length of interspiracular trunk; dorsal and
middle triplets sharing short stalk, and dorsal and middle triplets each composed of one individual and two paired
filaments and bearing short to medium-long primary and short secondary stalks; stalk of ventral pair of filaments
medium-long, 0.9–1.0 times length of common basal stalk, and 0.7–0.8 times length of interspiracular trunk, and
0.9 times as thick as common stalk of middle and dorsal triplets, 1.3 and 1.1 times as thick as stalks of dorsal and
middle triplets, respectively; common stalk of dorsal and middle triplets as long as or shorter than primary stalks of
dorsal and middle triplets; primary stalks of dorsal and middle triplets each much longer than secondary stalk;
primary stalk of dorsal triplet lying against that of lower pair at angle of about 70 degrees when viewed laterally; all
filaments damaged losing apical portion, thus no exact lengths known except one of middle triplet, which was
intact and 1.9 mm long from base of common basal stalk to apical tip); six filaments of dorsal and middle triplets
probably subequal in length and thickness to one another; two filaments of ventral pair probably subequal in length
and thickness to each other (outer filament 2.5 mm from base of common basal stalk to broken tip), and 1.7 times
as thick as six other filaments when compared basally; all filaments light brown, gradually tapered toward apex;
cuticle of all filaments with moderately-defined annular ridges and furrows though becoming less marked apically,
densely covered with minute tubercles. Abdomen. Dorsally, all segments light yellow and without tubercles;
segment 1 with one unbranched slender medium-long hair-like seta (Fig. 14H) on each side; segment 2 with one
unbranched slender medium-long hair-like seta and five minute setae (Fig. 14I) submedially on each side;
segments 3 and 4 each with four hooked spines and one minute seta on each side; segment 5 with one spine (Fig.
14J) on right side and lacking spine-combs on left side; segments 6–8 each with spine-combs in transverse row and
comb-like groups of minute spines on each side; segments 5–8 each with few minute setae near posterior margin on
each side; segment 9 with comb-like groups of minute spines and pair of small terminal hooks (Fig. 14K, L).
Ventrally, segment 4 with one unbranched hook (much smaller in size than those on segments 5–7) and few minute
setae (Fig. 14M) on each side; segment 5 with pair of bifid hooks submedially and few short slender setae on each
side; segments 6 and 7 each with pair of bifid inner and unbranched outer hooks somewhat spaced from each other
and few short slender setae on each side; segments 4–8 with comb-like groups of minute spines. Each side of
segment 9 with three grapnel-shaped hooklets. Cocoon (Fig. 14N). Wall-pocket-shaped, thinly and moderately
woven except anterolateral and peripheral portions roughly woven, widely extended ventrolaterally; dorsomedial
portion of anterior margin slightly produced anteriorly; 3.0 mm long by 2.1 mm wide.
Female and larva. Unknown.
Type material. HOLOTYPE: Male (with its associated pupal exuviae) in 80% ethanol, labeled in a vial as
[UMSRP: Vietnam 026, Holotype, Simulium (G.) quychauense, Male, coll. Vietnam, 11-XII-2015, Takaoka et al.],
reared from pupa collected from a small stream (width 1.5–3.0 m, bottom of pebbles and sands, water temperature
18 ˚C, exposed to the sun, elevation 73 m, 19˚30’45.487’’N/105˚09’09.261’’E), moderately flowing in a forest,
Chau Dinh, Quy Chau, Nghe An Province, northern Vietnam, 11-XII-2015, by H. Takaoka, M. Sofian-Azirun, C.
D. Chen & K.W. Lau.
Biological notes. The pupa of this new species was collected from a dead tree leaf. Associated species were S.
(G.) chaudinhense sp. nov., S. (S.) nodosum and S. (S.) tani (complex).
Etymology. The species name quychauense refers to the district name, Quy Chau, where this new species was
collected.
Remarks. This new species is assigned to the S. asakoae species-group of the subgenus Gomphostilbia by
having the yellow hair tuft on the base of the radius, enlarged male hind basitarsi (Fig. 13C) and ventral plate with
lateral margins emarginated basally (Fig. 13D). This new species is similar to S. (G.) tamdaoense described from a
male reared from a pupa in northern Vietnam (Takaoka et al. 2014a) in having the same number of male upper-eye
(large) facets in 14 vertical columns and 14 horizontal rows, and relative size of the fore and hind basitarsi. It also
is similar to S. (G.) thituyenae described from a female reared from a pupa in southern Vietnam (Takaoka et al.
2015a) in having the small narrow pupal terminal hooks (Fig. 14L). However, this new species is distinguished

FIGURE 14. Pupa of Simulium (Gomphostilbia) quychauense sp. nov. A, Frontal trichome; B, Facial trichome; C–F, Thoracic
trichomes (C, anterodorsal; D, anterolateral; E, mediolateral; F, ventrolateral); G, Anterior part of thorax and gill filaments (left
side; outer view); H, Hair-like seta on dorsal surface of abdominal segment 1; I, Hair-like seta and minute seta on dorsal surface
of abdominal segment 2; J, Spine-comb on dorsal surface of abdominal segment 5; K & L, Terminal hooks (K, lateral view; L,
caudal view); M, Hooklet and minute seta on ventral surface of abdominal segment 4; N, Cocoon (dorsal view). Scale bars. 1.0
mm for N; 0.1 mm for G; 0.02 mm for A–F, H, I, K and L; 0.01 mm for J and M.
from both species by the absence of the spine-combs on abdominal segment 9, and also from S. (G.) tamdaoense by
the absence of setae on the basal portion of the paramere (Fig. 13I) and small narrow pupal terminal hooks (Fig.
14L).
Simulium (Gomphostilbia) sanchayense Takaoka & Lau sp. nov.
Simulium (Gomphostilbia) brinchangense: not Takaoka, Sofian-Azirun & Hashim (in Takaoka et al., 2014a): 364–365.
This species was previously reported from Vietnam as S. (G.) brinchangense Takaoka, Sofian-Azirun & Hashim,
originally described from Peninsular Malaysia (Takaoka et al. 2014a, d). A DNA sequence-based analysis using
the COI gene shows that this species is distinct from S. (G.) brinchangense (Low et al. unpublished data). This
species is here described as a new species although our morphological reexamination of the males and their
associated pupal exuviae collected from Tam Dao, Vinh Phuc Province, reveals that they are almost identical to
those of S. (G.) brinchangense except for a few morphological features as noted later.
Male. Body length 2.9 mm. Head. Nearly as wide as thorax. Upper eye medium brown, consisting of large
facets in 12 vertical columns and 14 horizontal rows. Face brownish black, white pruinose. Clypeus brownish
black, whitish pruinose, moderately covered with golden-yellow scale-like medium-long hairs (mostly directed
upward) interspersed with several dark-brown longer hairs. Antenna composed of scape, pedicel and nine
flagellomeres, medium brown except scape and pedicel yellow and base of first flagellomere whitish yellow; first
flagellomere elongate, 1.75 times length of second one. Maxillary palp light to medium brown, with five segments,
proportional lengths of third, fourth, and fifth segments 1.0:1.2:2.9; third segment (Fig. 15A) slender, somewhat
produced posteromedially; sensory vesicle (Fig. 15A) ellipsoidal, small (0.2 times length of third segment), and
with opening of medium-size. Thorax. Scutum brownish black except anterolateral calli medium brown, shiny and
thinly gray pruinose on shoulders, on wide area along each lateral margin and on prescutellar area when
illuminated at certain angles, densely covered with golden-yellow scale-like recumbent short hairs. Scutellum light
brown, covered with yellow short hairs and dark-brown longer upright hairs along posterior margin. Postnotum
brownish black, shiny and white pruinose when illuminated at certain angles, and bare. Pleural membrane bare.
Katepisternum dark brown, longer than deep, shiny when illuminated at certain angles, moderately covered with
fine short hairs. Legs. Foreleg: coxa whitish yellow; trochanter light brown except base yellow; femur light brown
with apical cap medium brown (though apical tip yellowish); tibia white to grayish except little less than apical
one-third brownish black, and covered with white and yellow hairs on basal two-thirds of outer surface and white
sheen on basal two-thirds of outer surface when illuminated at certain angles; tarsus brownish black; basitarsus
moderately dilated, 8.0 times as long as its greatest width. Midleg: coxa medium brown except posterior surface
dark brown; trochanter light brown except base whitish yellow; femur light brown with apical cap medium brown
(though apical tip yellow); tibia medium brown except basal one-third whitish yellow; tarsus dark brown except
basal half of basitarsus dark yellow to light brown (border in middle not well defined). Hind leg: coxa light brown
though apical portion whitish yellow; trochanter whitish yellow; femur light brown with base narrowly whitish
yellow and apical cap dark brown (though apical tip whitish yellow); tibia (Fig. 15B) dark brown to brownish black
except little less than basal half whitish yellow; tarsus (Fig. 15C) medium to dark brown except little less than basal
half of basitarsus and basal half of second tarsomere yellowish; basitarsus (Fig. 15C) enlarged, 3.5 times as long as
wide, and 1.0 and 1.2 times as wide as greatest width of tibia and femur, respectively; calcipala (Fig. 15C) slightly
shorter than basal width, and 0.3 times as wide as greatest width of basitarsus. Pedisulcus (Fig. 15C) well
developed. Wing. Length 2.1 mm. Costa with dark spinules and hairs except hairs on basal portion yellow.
Subcosta with two to four hairs. Hair tuft on base of radius yellow. Basal portion of radius fully haired; R1 with
dark spinules and hairs; R2 with hairs only. Basal cell absent. Halter. White except basal stem darkened. Abdomen.
Basal scale dark brown, with fringe of dark hairs. Dorsal surface of abdomen medium to dark brown, covered with
dark-brown short to long hairs; segments 2 and 5–7 each with pair of shiny dorsolateral or lateral patches; ventral
surface of abdomen light to dark brown except segments 2 and 3 pale. Genitalia. Coxites, styles and ventral plate in
ventral view as in Fig. 15D. Coxite in ventral view (Fig. 15D) nearly rectangular, 1.91 times as long as its greatest
width. Style in ventral view (Fig. 15D) bent inward, nearly parallel-sided, bluntly pointed apically and with apical
spine; style in ventrolateral view (Fig. 15E) shorter than coxite (0.8 times length of coxite), 2.1 times as long as its

FIGURE 15. Male of Simulium (Gomphostilbia) sanchayense sp. nov. A, Third segment of maxillary palp with sensory
vesicle (left side; front view); B, Hind tibia (left side; outer view); C, Hind basitarsus and second tarsomere (left side; outer
view); D, Coxites, styles and ventral plate (ventral view); E, Style (right side; ventrolateral view); F, Ventral plate and median
sclerite (lateral view); G, Ventral plate (caudal view); H, Median sclerite (caudal view); I, Paramere and aedeagal membrane
(left side; caudal view); J & K, Abdominal segment 10 and cerci (right side; J, lateral view; K, caudal view). Scale bars. 0.1 mm
for B and C; 0.02 mm for A and D–K.
greatest width at base, slightly tapered to apex, and with truncated apex. Ventral plate in ventral view (Fig. 15D)
with body transverse, 0.6 times as long as wide, with anterior margin produced anteromedially, and posterior
margin somewhat concave medially (though posterior margin slightly convex medially), lateral margins slightly
emarginated near base, and densely covered with microsetae on ventral surface; basal arms of moderate length,
slightly divergent, then convergent apically; ventral plate in lateral view (Fig. 15F) moderately produced ventrally;
ventral plate in caudal view (Fig. 15G) trapezoidal, with ventral margin nearly straight though slightly produced
medially, densely covered with microsetae on posterior surface except portion near each lateral tip narrowly bare.
Median sclerite (Fig. 15H) plate-like and wide. Parameres (Fig. 15I) of moderate size, each with four distinct long
and medium-long stout hooks and several smaller ones including smallest ones on middle part of basal arm.
Aedeagal membrane (Fig. 15I) moderately setose; dorsal plate not defined. Ventral surface of abdominal segment
10 (Fig. 15J, K) weakly sclerotized anteriorly and without distinct hairs near posterior margin. Cercus (Fig. 15J, K)
Pupa. Body length 2.4 mm. Head. Integument yellow, moderately covered with small round tubercles except
antennal sheaths and ventral surface almost bare; antennal sheath without any protuberances; frons with three pairs
of unbranched long trichomes with coiled apices (Fig. 16A); face with pair of unbranched long trichomes with
coiled apices (Fig. 16B); three frontal trichomes on each side arising close together, subequal in length to one
another and slightly longer than facial one. Thorax. Integument yellow, moderately covered with round tubercles,
and with three long dorsomedial trichomes of different lengths (anterior trichome longest, posterior one shortest)
with coiled apices (Fig. 16C), two long anterolateral trichomes (anterior trichome shorter and more slender than
posterior one and with straight apex, and posterior one with coiled apex) (Fig. 16D), one medium-long mediolateral
trichome with straight apex (Fig. 16E), and three ventrolateral trichomes (one medium-long and two short) with
straight apices (Fig. 16F) on each side; all trichomes unbranched. Gill (Fig. 16G) composed of eight slender thread-
like filaments, arranged as [3+(1+2)]+2 from dorsal to ventral, with medium-long common basal stalk having
somewhat swollen transparent basal fenestra at base; common basal stalk 0.65–0.76 times length of interspiracular
trunk; dorsal and middle triplets sharing short stalk, and dorsal triplet composed of three individual filaments
arising at same level, with short stalk, middle triplet composed of one individual and two paired filaments and
bearing short primary and secondary stalks; stalk of ventral pair of filaments medium-long, 1.2–1.6 times length of
common basal stalk, and 0.9 times length of interspiracular trunk, and 0.9–1.0 times as thick as common stalk of
middle and dorsal triplets; common stalk of dorsal and middle triplets 0.7–1.3 and 0.5–1.0 times lengths of primary
stalks of dorsal and middle triplets, respectively; middle triplet primary stalk 1.3–9.3 times length of secondary
stalk; primary stalk of dorsal triplet lying against that of lower pair at angle of 60–80 degrees when viewed
laterally; six filaments of dorsal and middle triplets subequal in length (1.9–2.2 mm) and thickness to one another;
two filaments of ventral pair subequal in length (3.0 mm) and thickness to each other, and 1.4 times as thick as six
other filaments when compared basally; all filaments light brown, gradually tapered toward apex; cuticle of all
filaments with well-defined annular ridges and furrows though becoming less marked apically, densely covered
with minute tubercles. Abdomen. Dorsally, all segments unpigmented and without tubercles; segment 1 with one
unbranched slender short hair-like seta (similar to Fig. 16H) on each side; segment 2 with one unbranched slender
short hair-like seta and five somewhat spinous minute setae (Fig. 16H) submedially on each side; segments 3 and 4
each with four hooked spines and one somewhat spinous minute seta on each side; segment 5 lacking spine-combs
and comb-like groups of minute spines on each side; segments 6–9 each with spine-combs in transverse row and
comb-like groups of minute spines on each side; segments 5–8 each with few minute setae near posterior margin on
each side; segment 9 with pair of wide flat terminal hooks, of which outer margin is 2.6–3.2 times length of inner
margin and crenulated (Fig. 16I) when viewed caudally. Ventrally, segment 4 with one unbranched hook (subequal
in size to those on segments 5–7) and few slender short setae on each side; segment 5 with pair of bifid hooks
submedially and few short slender setae on each side; segments 6 and 7 each with pair of bifid inner and
unbranched outer hooks somewhat spaced from each other and few short slender setae on each side; segments 4–8
each with comb-like groups of minute spines. Each side of segment 9 with three grapnel-shaped hooklets. Cocoon
(Fig. 16J, K). Wall-pocket-shaped, thinly and moderately woven, widely extended ventrolaterally; dorsomedial
portion of anterior margin slightly to somewhat produced anteriorly; 3.0–3.1 mm long by 2.0–2.2 mm wide.
Type material. HOLOTYPE: Male (with its associated pupal exuviae and cocoon) in 80% ethanol, labeled in
a vial as [UMSRP: Vietnam 027, Holotype, Simulium (G.) sanchayense, Male, coll. Vietnam, 8-XI-2013, Takaoka
et al.], reared from a pupa collected from a small stream (width 0.5 m, bottom rocky, water temperature 17˚C,
shaded, elevation 975 m, 21˚27’501’’N/105˚38’941’’E), slow-flowing in bushes along the road near the Tam Dao
National Park, Vinh Phuc Province, northern Vietnam, 8-XI-2013, by H. Takaoka, M. Sofian-Azirun & Z. Ya’cob.
PARATYPES: Two males (with their associated pupal exuviae and cocoons) in 80% ethanol, same data as those of
the holotype, labeled in each vial as [UMSRP: Vietnam 027, Paratype, Simulium (G.) sanchayense, Male, coll.
Vietnam, 8-XI-2013, Takaoka et al.].

FIGURE 16. Pupa of Simulium (Gomphostilbia) sanchayense sp. nov. A, Frontal trichome; B, Facial trichome; C–F, Thoracic
side; outer view); H, Hair-like seta and minute seta on dorsal surface of abdominal segment 2; I, Terminal hooks (caudal view);
J & K, Cocoons with slightly different anterodorsal margins (dorsal view). Scale bars. 1.0 mm for J and K; 0.1 mm for G; 0.02
mm for A–F, H and I.
Biological notes. The pupae of this new species were collected from dead tree leaves. The associated species
were S. (G.) hongthaii, S. (G.) vinhphucense sp. nov., S. (N.) sp. (Simulium feuerborni species-group) and S. (S.)
lowi sp. nov.
Distribution. Vietnam (Vinh Phuc).
Etymology. The species name sanchayense refers to the land in which the San Chay tribe lives.
having the yellow hair tuft on the base of the radius, enlarged male hind basitarsi (Fig. 15C) and ventral plate with
lateral margins emarginated (Fig. 15D). The male of this new species is almost identical to that of S. (G.)
brinchangense, including the moderate number of upper-eye (large) facets in 12 vertical columns and 14 horizontal
rows and color of the legs (Takaoka et al. 2014d). There are slight differences in some male features between this
new species and S. (G.) brinchangense (characters of S. (G.) brinchangense in parentheses): length ratio of the first
flagellomere of the antenna against the second one is 1.8 (1.9), length ratio of the hind basitarsus against its greatest
width is 3.5 (3.7–3.8), ventral margin of the ventral plate is nearly straight when viewed posteriorly (concave).
The pupa of this new species is almost identical to that of S. (G.) brinchangense except that the middle triplet
of the gill is composed of one individual and two paired filaments (Fig. 16G) (three filaments arising at the same
level in S. (G.) brinchangense) and the dorsal triplet of the gill is composed of three filaments arising at the same
level (Fig. 16G) (one individual and two paired filaments in S. (G.) brinchangense).
Simulium (Gomphostilbia) tamdaoense Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Gomphostilbia) tamdaoense Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014a): 351–353 (Male and
pupa).

Remarks. This species was described from Vinh Phuc Province in northern Vietnam (Takaoka et al. 2014a).
The female and larva of this species remain to be collected.
Simulium (Gomphostilbia) unii Takaoka & Pham sp. nov.
Female. Body length 2.0–2.5 mm. Head. Slightly narrower than width of thorax. Frons brownish black, densely
covered with yellowish-white scale-like recumbent short hairs interspersed with few dark longer hairs near vertex;
frontal ratio 1.8:1.0:2.3–2.5; frons:head ratio 1.0:4.6. Fronto-ocular area well developed, narrow, directed
dorsolaterally. Clypeus brownish black, densely covered with yellowish-white scale-like hairs interspersed with
10–12 dark longer hairs on each side. Labrum 0.6 times length of clypeus. Antenna composed of scape, pedicel and
nine flagellomeres, light to medium brown except scape, pedicel, and basal half of first flagellomere yellow.
Maxillary palp composed of five segments, light to medium brown, proportional lengths of third, fourth, and fifth
segments 1.0:1.1:2.4–2.5; third segment (Fig. 17A) widened apically; sensory vesicle (Fig. 17A) small, ellipsoidal
(0.2 times length of third segment), with small opening. Maxillary lacinia with 10–12 inner and 15–17 outer teeth.
Mandible (Fig. 17B) with 28 or 29 inner teeth and one or two outer teeth at some distance from tip. Cibarium (Fig.
17C) medially forming sclerotized plate folded forward from posterior margin, with moderately sclerotized
mediolongitudinal ridge with bifid apex. Thorax. Scutum dark brown except anterolateral calli ochreous, and three
blackish longitudinal vittae (one median, two submedian), thinly pruinose and shiny when illuminated at certain
angles, densely covered with whitish-yellow to yellow scale-like recumbent short hairs except three longitudinal
vittae covered with dark short hairs. Scutellum ochreous, covered with yellow short hairs and dark-brown longer
upright hairs along posterior margin. Postnotum dark brown, slightly shiny when illuminated at certain angles, and
bare. Pleural membrane bare. Katepisternum longer than deep, medium to dark brown, shiny when illuminated at
certain angles, moderately covered with fine short hairs. Legs. Foreleg: coxa whitish yellow; trochanter light brown
except base whitish yellow; femur light brown with apical cap medium brown (though extreme tip yellowish); tibia
white except apical one-fourth dark brown, covered with white fine hairs on basal four-fifths; tarsus brownish
black, with moderate dorsal hair crest; basitarsus moderately dilated, 6.4–6.8 times as long as its greatest width.
Midleg: coxa yellow with light brown basal portion except posterolateral surface dark brown; trochanter whitish
yellow; femur light brown with base yellow (though inner surface of basal half yellow) and apical cap medium
brown (though extreme tip yellowish); tibia medium to dark brown except basal one-third whitish yellow (though

FIGURE 17. Female of Simulium (Gomphostilbia) unii sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
side; front view); B, Tip of mandible; C, Cibarium (front view); D, Hind tibia (left side; outer view); E, Hind basitarsus and
second tarsomere (left side; outer view). F, Claw; G, Sternite 8 and ovipositor valve (right side only; ventral view); H, Genital
fork (ventral view); I & J, Paraprocts and cerci (I, ventral view; J, lateral view). K, Spermatheca. Scale bars. 0.1 mm for D and
E; 0.02 mm for A, C and G–K; 0.01 mm for B.
inner surface of basal two-thirds yellow), covered with whitish and yellowish fine hairs on posterior and inner
surfaces of basal two-thirds; tarsus dark brown though basal half or little less of basitarsus yellow. Hind leg: coxa
light brown except anterior two-fifths yellow; trochanter whitish yellow; femur light to medium brown with base
whitish yellow and apical cap dark brown (though extreme tip yellowish); tibia (Fig. 17D) yellowish white on basal
two-thirds and brownish black on rest, covered with whitish fine hairs on outer and posterior surfaces of little more
than basal three-fourths; tarsus brownish black except basal two-thirds or little more of basitarsus (though base
light brown) and basal half of second tarsomere yellowish white; basitarsus (Fig. 17E) narrow, nearly parallel-sided
though slightly narrowed apically, 6.0–6.1 times as long as wide, and 0.7 and 0.6 times as wide as greatest widths
of tibia and femur, respectively; calcipala (Fig. 17E) nearly as long as width at base, and 0.5 times as wide as
greatest width of basitarsus; pedisulcus (Fig. 17E) well developed; claw (Fig. 17F) with large basal tooth 0.5 times
length of claw. Wing. Length 2.0–2.2 mm. Costa with dark spinules and hairs except basal patch of hairs yellow.
Subcosta with dark hairs except near apex bare. Hair tuft on base of radius yellow. Basal portion of radius fully
haired; R1 with dark spinules and hairs; R2 with hairs only. Basal cell absent. Halter. White except basal portion
darkened. Abdomen. Basal scale yellow, with fringe of whitish-yellow hairs. Dorsal surface of abdomen light to
dark brown except anterior two-thirds of segment 2 whitish, moderately covered with dark short to long hairs;
tergites of segments 2 and 6–9 shiny when illuminated at certain angles. Ventral surface of segment 2 yellowish
white, that of segment 3 dark yellow and those of other segments medium brown; sternal plate on segment 7
undeveloped. Terminalia. Sternite 8 (Fig. 17G) bare medially, with 17–28 medium-long to long hairs together with
two to four slender short hairs on each side. Ovipositor valve (Fig. 17G) triangular (though posteromedial corner
rounded), thin, membranous, moderately covered with microsetae interspersed with two short hairs; inner margins
nearly straight or slightly sinuous, somewhat sclerotized, and moderately separated from each other. Genital fork
(Fig. 17H) of usual inverted-Y form, with slender stem; arms of moderate width, moderately folded medially,
without lobe directed posteromedially. Paraproct in ventral view (Fig. 17I) somewhat concave anterolaterally, with
three to six sensilla on anteromedial surface; paraproct in lateral view (Fig. 17J) slightly produced ventrally beyond
ventral tip of cercus, 0.7 times as long as wide, with 25–30 medium-long to long hairs on ventral and lateral
surfaces. Cercus in lateral view (Fig. 17J) short, rounded posteriorly, 0.5 times as long as wide. Spermatheca (Fig.
17K) ellipsoidal, 1.2–1.5 times as long as its greatest width, well sclerotized except duct and small area near
juncture with duct unsclerotized, and with many fissures on outer surface; internal setae absent; both accessory
ducts slender, subequal in diameter to major one.
Male. Body length 2.1–2.4 mm. Head. Somewhat wider than thorax. Upper eye medium brown, consisting of
13 or 14 vertical columns and 14 horizontal rows of large facets. Face brownish black, white pruinose. Clypeus
brownish black, whitish pruinose, densely covered with golden-yellow scale-like medium-long hairs (mostly
directed upward) interspersed with several dark-brown unbranched longer hairs. Antenna composed of scape,
pedicel and nine flagellomeres, yellowish except flagellomeres 5–9 or 6–9 light to medium brown; first
flagellomere elongate, 1.8 times length of second one. Maxillary palp light to medium brown, with five segments,
proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.2; third segment (Fig. 18A, B) slender; sensory
vesicle (Fig. 18A, B) small, globular or ellipsoidal (0.1 times length of third segment), and with small opening.
Thorax. Scutum dark brown, without longitudinal vittae, shiny and thinly gray pruinose on shoulders, on wide area
along each lateral margin and on prescutellar area when illuminated at certain angles, and densely covered with
golden-yellow scale-like recumbent short hairs. Scutellum medium brown, covered with yellow short hairs and
dark-brown long upright hairs along posterior margin. Postnotum dark brown, slightly shiny and white pruinose
when illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum medium brown, longer than
deep, shiny when illuminated at certain angles, moderately covered with fine short hairs. Legs. Foreleg: coxa
whitish yellow; trochanter light brown with base whitish yellow; femur light brown with apical cap medium brown
(though apical tip yellowish); tibia yellow except basal one-third light brown and apical one-third medium brown,
and covered with yellow hairs on basal two-thirds of outer surface; tarsus brownish black; basitarsus moderately
dilated, 7.6–7.8 times as long as its greatest width. Midleg: coxa light brown except posterior surface dark brown;
trochanter yellowish; femur light brown with base yellowish and apical cap medium brown (though apical tip
yellow); tibia medium brown except basal one-third yellowish; tarsus dark brown except basal half of basitarsus
yellow (border not well defined). Hind leg: coxa light brown though apical portion yellow; trochanter yellow;
femur light to medium brown with base narrowly yellow and apical cap dark brown (though apical tip whitish
yellow); tibia (Fig. 18C) medium to dark brown except little less than basal half whitish yellow and apex brownish
black, covered with yellow hairs on basal three-fourths of outer and posterior surface; tarsus (Fig. 18D) medium to
dark brown except basal half or little more of basitarsus and basal half of second tarsomere whitish yellow;
basitarsus (Fig. 18D) enlarged, spindle-shaped, 4.3–4.6 times as long as wide, and 0.8–0.9 and 0.8 times as wide as
greatest width of tibia and femur, respectively; calcipala (Fig. 18D) nearly as long as basal width, and 0.4 times as

FIGURE 18. Male of Simulium (Gomphostilbia) unii sp. nov. A & B, Third segments of maxillary palps with sensory vesicles
of different shape (A, right side; with ellipsoidal sensory vesicle; B, left side; with globular sensory vesicle; both front view); C,
Hind tibia (left side; outer view); D, Hind basitarsus and second tarsomere (left side; outer view); E, Coxites, styles and ventral
plate (ventral view); F, Style (right side; ventrolateral view); G, Ventral plate and median sclerite (lateral view); H, Ventral plate
(caudal view); I, Median sclerite (caudal view); J, Paramere and aedeagal membrane (left side; caudal view); K & L,
Abdominal segment 10 and cerci (right side; K, lateral view; L, caudal view). Scale bars. 0.1 mm for C and D; 0.02 mm for A,
B and E–L.
wide as greatest width of basitarsus. Pedisulcus (Fig. 18D) well developed. Wing. Length 1.9–2.0 mm. Other
characters as in female except subcosta bare. Halter. Grayish white except basal stem darkened and apical half
grayish ochreous. Abdomen. Basal scale medium brown, with fringe of light-brown hairs. Dorsal surface of
abdomen dark brown to brownish black except anterior one-third of segment 2 yellowish, covered with dark-brown
short to long hairs; segments 2 and 5–8 each with pair of shiny dorsolateral or lateral patches; ventral surface of
segments 2 and 3 whitish though sternal plate of segment 3 darkened, and those of other segments medium to dark
brown. Genitalia. Coxite in ventral view (Fig. 18E) nearly rectangular, 1.9 times as long as its greatest width. Style
in ventral view (Fig. 18E) bent inward, with blunt apex having single spine; style in ventrolateral view (Fig. 18F)
slightly tapered toward apex, with truncated apex. Ventral plate in ventral view (Fig. 18E) with body transverse, 0.6
times as long as wide, with anterior margin produced anteromedially, and posterior margin somewhat concave
medially (though posterior margin slightly convex medially when ventral plate is slightly tilted), slightly narrowed
medially, then nearly parallel-sided, and densely covered with microsetae on ventral surface; basal arms of
moderate length, directed forward; ventral plate in lateral view (Fig. 18G) moderately produced ventrally; ventral
plate in caudal view (Fig. 18H) gently rounded ventrally, densely covered with microsetae on posterior surface.
Median sclerite (Fig. 18I) plate-like and wide. Parameres (Fig. 18J) of moderate size, each with four distinct long
and medium-long stout hooks, and without minute setae on outer surface of basal arm. Aedeagal membrane (Fig.
18J) moderately setose; dorsal plate not defined. Ventral surface of abdominal segment 10 (Fig. 18K, L) without
distinct hairs near posterior margin. Cercus (Fig. 18K, L) small, rounded, with 14–16 hairs.
Pupa. Body length 2.4–2.5 mm. As in S. (G.) sanchayense sp. nov. except following characters. Thorax. Gill
(Fig. 19A, B, C) composed of eight slender thread-like filaments, arranged as [(2+1)+(2+1)]+2, or [3+(2+1)]+2, or
[(2+1)+3]+2, or (3+3)+2 from dorsal to ventral, with short common basal stalk having somewhat swollen basal
fenestra ventrally at base; common basal stalk 0.6–0.8 times length of interspiracular trunk; dorsal and middle
triplets sharing short stalk 0.6–0.9 times length of common basal stalk; dorsal and middle triplets each composed of
one individual filament and two paired filaments with short primary and secondary stalks (secondary stalk usually
much shorter than primary stalk but in left gill of one pupa secondary stalk is exceptionally long, twice length of
primary stalk as shown in Fig. 19C), or three filaments arising at same level from primary stalk; ventral paired
filaments with medium-long stalk 0.9–1.5 times length of common basal stalk; primary stalk of dorsal triplet lying
against that of ventral pair at angle of 90 degrees or little greater when viewed laterally; all filaments medium
brown, gradually tapered toward apex; three filaments of dorsal triplet subequal in length (2.3–2.5 mm long
including their own stalks and common basal stalk) and thickness to one another; three filaments of middle triplet
subequal in length (2.6–3.0 mm) and thickness; inner filament of ventral pair longest (2.8–3.2 mm), slightly longer
than outer filament (2.7–3.1 mm); two filaments of ventral pair subequal in thickness to each other, and 1.2–1.5
times as thick as six other filaments when compared basally; cuticle of all filaments with well-defined annular
ridges and furrows, and densely covered with minute tubercles. Abdomen. Dorsally, segments 1, 2 and 9 yellow,
and other segments whitish yellow at least medially; segment 9 with pair of flat triangular terminal hooks, of which
outer margin 1.8 times length of inner margin and crenulated (Fig. 19D). Cocoon (Fig. 19E, F). Wall-pocket-
shaped, moderately woven, widely extended ventrolaterally; anterior margin thickly woven medially, with
dorsomedial projection, resembling letter W when viewed dorsally; posterior three-fifths with floor roughly
woven; individual threads visible; 3.0–3.9 mm long by 2.0–2.9 mm wide.
Mature larva. Body length 4.4–5.5 mm. Body whitish to light ochreous, with following color markings:
thoracic segment 1 encircled with reddish-brown band (though often disconnected ventromedially), proleg light
ochreous, thoracic segments 2 and 3 reddish brown on ventral surface, though lighter on segment 3; abdominal
segments 1 and 2 each encircled by grayish broad band; abdominal segments 3 and 4 each with or without pair of
narrow light reddish-brown dorsolateral spots; abdominal segment 5 encircled by distinct reddish-brown transverse
band though disconnected ventromedially, abdominal segment 6 with similar colored transverse band (though
much narrower) on dorsal and dorsolateral surfaces, though often faded to varying extent leaving reddish-brown
round spot dorsomedially, and pair of light reddish-brown spots ventrally; abdominal segments 7 and 8 widely
reddish brown on dorsal and dorsolateral surfaces, though faded to varying extent anteromedially; abdominal
segment 7 with reddish-brown narrow transverse band ventrally. Head. Head capsule whitish yellow, sparsely
covered with minute setae (though moderately on dorsal surface); head spots faintly positive though two relatively
large spots near posterior margin on lateral surface usually moderately positive, and two small spots below eye-
spot region often indistinct; eyebrow indistinct. Antenna composed of three articles and apical sensillum, longer

FIGURE 19. Pupa of Simulium (Gomphostilbia) unii sp. nov. A, Anterior part of thorax and gill filaments (left side; outer
view); B & C, Basal portions of gill filaments with different arrangements of dorsal and medial triplets (left side; outer view);
D, Terminal hooks (caudal view); E & F, Cocoons (E, dorsal view; F, lateral view). Scale bars. 1.0 mm for E and F; 0.1 mm for
A–C; 0.02 mm for D.
than stem of labral fan; proportional lengths of first, second, and third articles 1.0:0.6–0.8:0.9. Labral fan with 36–
41 primary rays. Mandible (Fig. 20A) with three comb-teeth decreasing in length from first tooth to third;
mandibular serration composed of two teeth (one medium-sized, one small); major tooth at acute angle against
mandible on apical side; supernumerary serrations absent. Hypostoma (Fig. 20B) with row of nine apical teeth, of
which median tooth is longer than each corner tooth; lateral margin smooth; four or five hypostomal bristles per
side lying slightly divergent posteriorly from lateral margin. Postgenal cleft (Fig. 20C) rounded, medium-long, 1.4
times length of postgenal bridge. Cervical sclerites composed of pair of small unpigmented rod-like pieces. Thorax
and Abdomen. Thoracic proleg without hairs; thoracic cuticle almost bare; abdominal cuticle of segments 1–6
almost bare, and that of segments 5–9 sparsely covered with slightly darkened unbranched or bifid or trifid minute
setae (Fig. 20D) on dorsal and dorsolateral surfaces; last abdominal segment densely covered with unbranched
colorless longer setae on dorsolateral and lateral surfaces of each side of anal sclerite and on each lateral surface
even down to base of ventral papilla; thorax and abdomen without dorsal protuberances. Rectal scales absent.
Rectal organ compound, each of three lobes with six to nine finger-like secondary lobules. Anal sclerite of usual X-
form, with anterior arms nearly as long as posterior ones, broadly sclerotized at base; no sensilla on broad base and
posterior to posterior arms; accessory sclerite absent. Last abdominal segment with pair of large conical ventral
papillae. Posterior circlet with 68 rows of hooklets with up to 12 hooklets per row.
FIGURE 20. Larva of Simulium (Gomphostilbia) unii sp. nov. A, Mandible; B, Hypostoma; C, Head capsule (ventral view);
D, Setae on dorsal surface of abdominal segment 8. Scale bars. 0.05 mm for C; 0.02 mm for B and D; 0.01 mm for A.
Type material. HOLOTYPE: Female (with its associated pupal exuviae and cocoon) in 80% ethanol, labeled
in a vial as [UMSRP: Vietnam 028, Holotype, Simulium (G.) unii, Female, coll. Vietnam, 11-XII-2015, Takaoka et
al.], reared from a pupa collected from a slow-flowing stream (width 1.0 m, depth 5–10 cm, bottom sandy, water
temperature 18.0˚C, shaded, elevation 80 m, 19˚30’46.589”N/105˚09’10.470”E), Chau Dinh, Quy Chau, Nghe An
Province, northern Vietnam, 11-XII-2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W. Lau.

PARATYPES: Six females, three males and two mature larvae, same data as those of holotype, labeled in each vial
as [UMSRP: Vietnam 028, Paratype, Simulium (G.) unii, coll. Vietnam, 11-XII-2015, Takaoka et al.].
Biological notes. The pupae and larvae of this new species were collected from fallen leaves and grasses
tailing in the current. Associated species were S. (G.) chaudinhense sp. nov., S. (G.) parahiyangum, S. (S.) nodosum
and S. (S.) tani (complex).
Etymology. The species name unii is in honor of Prof. Sigehiko Uni, Institute of Biological Sciences, Faculty
of Science, University of Malaya, for his great contribution to taxonomic studies of filariae of wild animals in
relation to zoonotic onchocerciasis in Japan.
Remarks. Simulium (G.) unii sp. nov. is placed in the S. asakoae species-group by having the yellow hair tuft
on the base of the radius and enlarged male hind basitarsus (Fig. 18D), although the ventral plate of this new
species is not clearly emarginated on each lateral margin (Fig. 18E), a character disagreeing with one of the key
characters of this species-group defined by Takaoka (2012).
This new species is characterized by the cocoon having an anterodorsal projection (Fig. 19E). Among the 25
species of the S. asakoae species-group (Takaoka 2012; Takaoka et al. 2014a, d, 2015a), S. (G.) gyorkosae Takaoka
& Davies from Java, Indonesia (Takaoka and Davies 1996); S. (G.) fanjingshanense Chen, Zhang & Wen from
Guizhou, south China (Chen et al. 2000); S. (G.) fuscidorsum Takaoka & Ya’cob from Vietnam (Takaoka et al.
2015a); S. (G.) phulocense Takaoka & Chen from Vietnam (Takaoka et al. 2015a); and S. (G.) yunnanense Chen &
Zhang from Yunnan, China (Chen and Zhang 2004) have a similar cocoon. However, S. (G.) unii sp. nov. is
distinguished from these five known species by the following characters (those of each related species in
parentheses): from (G.) gyorkosae by the number of male upper-eye facets in 13 or 14 vertical columns (in 10 or 11
vertical columns); from S. (G.) fanjingshanense by the female hind tibia which is yellowish on the basal two-thirds
(Fig. 17D) (yellowish on the basal two-fifths), and angle of the stalk of the dorsal triplet against that of the ventral
pair of pupal gill filaments when viewed laterally being 90 degrees or slightly greater (Fig. 19A) (60 degrees); from
S. (G.) fuscidorsum by the small female sensory vesicle (Fig. 17A) (enlarged sensory vesicle) and short common
basal stalk (Fig. 19A) (long common basal stalk); from S. (G.) phulocense by the female hind tibia which is
yellowish on the basal two-thirds (Fig. 17D) (yellowish on the basal half), and angle of the stalk of the dorsal triplet
against that of the ventral pair of pupal gill filaments when viewed laterally being 90 degrees or slightly greater
(Fig. 19A) (60 degrees); from S. (G.) yunnanense by the yellow hair tuft on the base of the radius (black hair tuft),
and number of male upper-eye facets in 13 or 14 vertical columns (in 10 vertical columns).
The larva of this new species is characterized by the abdomen having grayish bands only on segments 1 and 2,
faint head spots and medium-long postgenal cleft (Fig. 20C).
Simulium (Gomphostilbia) thituyenae Takaoka & Pham, 2015
Simulium (Gomphostilbia) thituyenae Takaoka & Pham (in Takaoka et al., 2015a): 24 –28 (Female and pupa)

Remarks. This species was described from Lam Dong Province in southern Vietnam (Takaoka et al. 2015a).
The male and larva of this species are still unknown.
Simulium (Gomphostilbia) vinhphucense Takaoka & Low sp. nov.
Male. Body length 2.5 mm. Nearly as in male of S. (G.) sanchayense sp. nov. except following characters. Head.
Upper eye medium brown, consisting of large facets in 9 vertical columns and 11 or 12 horizontal rows. Antenna
medium brown except scape and pedicel yellow, base of first flagellomere whitish yellow, and rest of first
flagellomere and second and third flagellomeres yellow to dark yellow; first flagellomere elongate, 1.7 times
length of second one. Maxillary palp: proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.4; third
segment (Fig. 21A) slender; sensory vesicle (Fig. 21A) ellipsoidal, small (0.2 times length of third segment), and
with small opening. Thorax. Scutum dark brown except anterolateral calli ochreous, with faint longitudinal vittae
(one median and two submedian), shiny and thinly gray pruinose on entire surface. Legs. Fore and mid trochanters
and base of mid femur yellow. Fore basitarsus moderately dilated, 8.2 times as long as its greatest width. Hind
basitarsus (Fig. 21B) enlarged, 3.3 times as long as wide, and 1.1 and 1.5 times as wide as greatest width of tibia
and femur, respectively; calcipala (Fig. 21B) slightly shorter than basal width, and 0.2 times as wide as greatest
width of basitarsus. Wing. Length 2.0 mm. Subcosta with two hairs. Abdomen. Dorsal surface of abdomen medium
to dark brown except segment 2 pale though narrow portion along posterior margin somewhat darkened; tergites 3–
6 each with markedly lighter medial portion; ventral surface of abdomen light to medium brown except segment 2–
4 pale though sternites 3 and 4 somewhat darkened. Genitalia. Coxite in ventral view (Fig. 21C) nearly rectangular,
twice as long as its greatest width. Style in ventral view (Fig. 21C) bent inward, bluntly rounded apically and with
apical spine; style in ventrolateral view (Fig. 21D) 0.8 times length of coxite, gradually tapered from base to apex,
and with truncated apex. Ventral plate in ventral view (Fig. 21C) with body transverse, 0.5 times as long as wide,
with anterior margin produced anteromedially, and posterior margin slightly concave medially, lateral margins
nearly straight or slightly emarginated basally, and densely covered with microsetae on ventral surface; basal arms
of moderate length, directed forward, then convergent apically; ventral plate in lateral view (Fig. 21E) moderately
produced ventrally; ventral plate in caudal view (Fig. 21F) rounded ventrally, densely covered with microsetae on
posterior surface except portion near each lateral tip narrowly bare. Median sclerite plate-like, wide (Fig. 21G).
Parameres (Fig. 21H) of moderate size, each with four distinct long and medium-long stout hooks and several
smaller ones. Aedeagal membrane (Fig. 21H) moderately setose; dorsal plate not defined. Ventral surface of
abdominal segment 10 (Fig. 21I, J) without distinct hairs near posterior margin. Cercus (Fig. 21I, J) small, rounded,
with 12 or 14 hairs.
Pupa. Body length 2.6 mm. Nearly as in pupa of S. (G.) sanchayense sp. nov. except following characters.
Head. Frons with three pairs of unbranched long trichomes with straight or coiled apices; face with pair of
unbranched long trichomes with straight or coiled apices. Thorax. Three unbranched ventrolateral trichomes (one
medium-long with coiled apex, two short with straight apices) on each side. Gill (Fig. 21K) composed of eight
slender thread-like filaments, arranged as [(1+2)+(1+2)]+2 from dorsal to ventral, with medium-long common
basal stalk having somewhat swollen transparent basal fenestra at base; common basal stalk 0.7 times length of
interspiracular trunk; dorsal and middle triplets sharing short stalk, and each triplet composed of one individual and
two paired filaments and bearing short primary and secondary stalks; stalk of ventral pair of filaments medium-
long, 0.8–1.0 times length of common basal stalk, and 0.6 times length of interspiracular trunk, and 0.8–0.9 times
as thick as common stalk of middle and dorsal triplets and 1.2–1.3 times as thick as primary stalks of dorsal and
middle triplets; common stalk of dorsal and middle triplets 1.3–2.1 and 0.7–1.1 times lengths of primary stalks of
dorsal and middle triplets, respectively; primary stalks of dorsal and middle triplets each much longer than
secondary stalk; primary stalk of dorsal triplet lying against that of ventral pair at angle of 60–70 degrees when
viewed laterally; six filaments of dorsal and middle triplets subequal in length (1.8–2.1 mm) and thickness to one
another; two filaments of ventral pair subequal in length (2.6–2.7 mm) and thickness to each other, and 1.7 times as
thick as six other filaments when compared basally Abdomen. Dorsally, segment 1 with one unbranched slender
medium-long hair-like seta on each side; segment 2 with one unbranched slender medium-long hair-like seta and
five somewhat spinous minute setae submedially on each side; segment 9 with pair of wide flat terminal hooks, of
which outer margin is 3.0 times length of inner margin and crenulated (Fig. 21L) when viewed caudally. Cocoon
(Fig. 21M). Wall-pocket-shaped, thinly and moderately woven, widely extended ventrolaterally; right submedial
portion of anterior margin slightly produced anteriorly; individual threads clearly visible; 3.5 mm long by 2.3 mm
wide.
a vial as [UMSRP: Vietnam 029, Holotype, Simulium (G.) vinhphucense, Male, coll. Vietnam, 8-XI-2013, Takaoka
et al.], reared from a pupa collected from a small stream (width 0.5 m, bottom rocky, water temperature 17˚C,
shaded, elevation 975 m, 21˚27’501’’N/105˚38’941’’E), slow-flowing in bushes along the road near the Tam Dao
National Park, Vinh Phuc Province, northern Vietnam, 8-XI-2013, by H. Takaoka, M. Sofian-Azirun & Z. Ya’cob.
Biological notes. The pupa of this new species was collected from a grass leaf trailing in the current.
Associated species were S. (G.) hongthaii, S. (G.) sanchayense sp. nov., S. (N.) sp. (S. feuerborni species-group)
and S. (S.) lowi sp. nov.

FIGURE 21. Male and pupa of Simulium (Gomphostilbia) vinhphucense sp. nov. A–J, Male and K–M, Pupa. A, Third segment
of maxillary palp with sensory vesicle (right side; front view); B, Hind basitarsus and second tarsomere (right side; outer view);
C, Coxites, styles and ventral plate (ventral view); D, style (right side; ventrolateral view); E, Ventral plate and median sclerite
(lateral view); F, Ventral plate (caudal view); G, Median sclerite (caudal view); H, Paramere and aedeagal membrane (right
side; caudal view); I & J, Abdominal segment 10 and cerci (right side; I, lateral view; J, caudal view); K, Anterior part of pupal
thorax and gill filaments (left side; outer view); L, Terminal hooks (caudal view); M, Cocoon (dorsal view). Scale bars. 1.0 mm
for M; 0.1 mm for B and K; 0.02 mm for A, C–J and L.
Etymology. The species name vinhphucense refers to the province name, Vinh Phuc, where this new species
was collected.
having the yellow hair tuft on the base of the radius and enlarged male hind basitarsi (Fig. 21B), although the
lateral margins of the ventral plate are not emarginated when viewed ventrally (Fig. 21C).
The male of this new species is characterized by the smaller number of upper-eye (large) facets in 9 vertical
columns and 11 or 12 horizontal rows. This smaller number of male upper-eye facets was recorded in S. (G.) mengi
Chen, Zhang & Wen, and S. (G.) miaolingense Wen & Chen, both described from Guizhou Province, China, which
have the male upper-eye facets in eight and nine horizontal rows, respectively (Chen and An 2003). This new
species is distinguished from S. (G.) mengi and S. (G.) miaolingense by the larger number of horizontal rows of the
male upper-eye facets, from the former species by the stalk of the ventral paired pupal gill filaments as long as or
shorter than the common basal stalk (Fig. 21K) (stalk of the ventral paired filaments longer than the common basal
stalk in S. (G.) mengi), and from the latter species by the shape of the ventral plate not angulated laterally (Fig. 21C)
(ventral plate basally angulated outward when viewed ventrally in S. (G.) miaolingense).
2) Simulium batoense species-group
Simulium (Gomphostilbia) dachaisense Takaoka & Lau, 2015
Simulium (Gomphostilbia) dachaisense Takaoka & Lau (in Takaoka et al., 2015a): 28–34 (Female, male, pupa and larva).

Simulium (Gomphostilbia) duolongum Takaoka & Davies, 1995
Simulium (Gomphostilbia) duolongum Takaoka & Davies, 1995: 19–24 (Female, male, pupa and larva).
Distribution. Peninsular Malaysia, Sumatra and Vietnam (Bac Thai).

Remarks. This species was originally described from Peninsular Malaysia (Takaoka and Davies 1995) and
recorded from Vietnam based on two pupae and two larvae by Pham (1999). The identity of the species regarded as
S. (G.) duolongum in Vietnam should be confirmed based on the adult males and females.
Simulium (Gomphostilbia) lamdongense Takaoka & Sofian-Azirun, 2015
Simulium (Gomphostilbia) lamdongense Takaoka & Sofian-Azirun (in Takaoka et al., 2015a): 35–41 (Female, male, pupa and
larva).

Simulium (Gomphostilbia) longlanhense Takaoka & Ya’cob, 2015
Simulium (Gomphostilbia) longlanhense Takaoka & Ya’cob (in Takaoka et al., 2015a): 41–48 (Female, male, pupa and larva).


Simulium (Gomphostilbia) parahiyangum Takaoka & Sigit, 1992
Simulium (Gomphostilbia) parahiyangum Takaoka & Sigit, 1992: 135–142 (Female, male, pupa and larva).
Specimens examined. Seven females, nine males (all reared from pupae), and six mature larvae, collected from a
slow-flowing stream (width 1.0 m, depth 5–10 cm, bottom sandy, water temperature 18.0˚C, shaded, elevation 80
m, 19˚30’46.589”N/105˚09’10.470”E), Chau Dinh, Quy Chau, Nghe An Province, northern Vietnam, 11-XII-2015,
by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W. Lau.
Distribution. Java, Sumatra, Malaysia, Thailand, Assam and Vietnam (Nghe An) (New record).
Remarks. The male of this species collected from Vietnam differs from that of S. (G.) parahiyangum from
Java by the number of upper-eye facets in 14 vertical columns and 14 (rarely 15) horizontal rows (in 17 vertical
columns and 15 horizontal rows in the original description) (Takaoka and Sigit 1992).
Simulium (Gomphostilbia) siamense Takaoka & Suzuki, 1984
Simulium (Gomphostilbia) siamense Takaoka & Suzuki, 1984: 14–18 (Female, male, pupa and larva).
Distribution. Thailand and Vietnam.

Remarks. This species was described from Thailand by Takaoka and Suzuki (1984), and later recorded from
Vietnam by Crosskey and Howard (1997). No detailed information of this record was given, leaving the locality in
Vietnam unknown. There is a possibility that the taxa regarded as S. (G.) siamense and S. (G.) duolongum are the
same species because the pupal gills of both species are morphologically indistinguishable from each other
(Takaoka and Suzuki 1984; Takaoka and Davies 1995).
3) Simulium ceylonicum species-group
Simulium (Gomphostilbia) inthanonense Takaoka & Suzuki, 1984
Simulium (Gomphostilbia) inthanonense Takaoka & Suzuki, 1984: 18–21 (Female, male, pupa and larva).
Distribution. South China, Thailand and Vietnam (Bac Thai and Ha Thai).
Remarks. This species was originally described from Thailand by Takaoka and Suzuki (1984), and later
recorded from Vietnam by Pham (1998, 1999). There remains a possibility that the species recorded as S. (G.)
inthanonense is either S. (G.) phulocense or S. (G.) unii sp. nov. or another related species, as its identification was
based on only two pupae and three larvae. These three species and a few other related species have the same shape
of the cocoon with a short anterodorsal projection, as shown in Fig. 19E.
4) Simulium darjeelingense species-group
Simulium (Gomphostilbia) eshimai Takaoka & Adler sp. nov.
Female. Body length 2.0 mm. Head. Slightly narrower than thorax. Frons brownish black, densely covered with
yellow scale-like recumbent short hairs and few dark hairs along each lateral margin; frontal ratio 1.8:1.0:1.8;
frons:head ratio 1.0:3.7. Fronto-ocular area well developed, narrow, directed dorsolaterally. Clypeus brownish
black, slightly white pruinose, and densely covered with yellowish-white recumbent hairs interspersed with several
dark longer hairs on each side. Labrum 0.8 times as long as clypeus. Antenna composed of scape, pedicel and nine
flagellomeres, dark brown, though scape, pedicel and base of first flagellomere slightly lighter. Maxillary palp
composed of five segments, light brown except third segment dark brown, proportional lengths of third, fourth, and
fifth segments 1.0:1.1:2.2; third segment (Fig. 22A) somewhat enlarged; sensory vesicle (Fig. 22A) elongate, 0.5
times length of third segment and with medium-sized opening. Maxillary lacinia with 13 or 14 inner and 20 outer
FIGURE 22. Female of Simulium (Gomphostilbia) eshimai sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Cibarium (front view); C, Hind basitarsus and second tarsomere (left side; outer view). D, Claw; E,
Sternite 8 and ovipositor valve (left side only; ventral view); F, Genital fork (ventral view); G & H, Paraprocts and cerci (G,
ventral view; H, lateral view). I, Spermatheca. Scale bars. 0.1 mm for C; 0.02 mm for A, B and D–I.
teeth. Mandible with 36 inner teeth and four or five outer teeth at some distance from apex. Cibarium (Fig. 22B)
with dorsal margin having short plate produced forward and downward, and with weakly-sclerotized
mediolongitudinal stripe bearing well-sclerotized U-shaped ridge apically. Thorax. Scutum brownish black,
slightly shiny when illuminated at certain angles, densely covered with yellow scale-like recumbent hairs mixed

with whitish similar hairs on peripheral portions. Scutellum dark brown, covered with yellow short hairs and dark-
brown long upright hairs. Postnotum brownish black, white pruinose and slightly shiny when illuminated at certain
angles, and bare. Pleural membrane bare. Katepisternum longer than deep, dark brown, white pruinose and shiny
when illuminated at certain angles, moderately covered with pale and dark short hairs. Legs. Foreleg: coxa yellow;
trochanter light brown except base yellow; femur medium brown with apical cap dark brown (though extreme tip
yellow); tibia medium brown except base yellow and apical cap dark brown, though median portion of outer
surface widely lighter; tarsus brownish black to black, with moderate dorsal hair crest; basitarsus somewhat
dilated, 7.0 times as long as its greatest width. Midleg: coxa medium brown except posterolateral surface dark
brown; trochanter light brown except base yellow; femur medium brown except basal one-fourth yellow and apical
cap medium brown (though extreme tip yellow); tibia medium to dark brown except extreme base yellow and
apical cap brownish black; tarsus dark brown to brownish black, though base of basitarsus somewhat lighter. Hind
leg: coxa medium brown; trochanter yellow; femur light to medium brown except base yellow and apical cap dark
brown (though extreme tip yellow); tibia light to medium brown except base yellow, apical cap dark brown and
subbasal portion medium to dark brown; tarsus (Fig. 22C) dark brown except basal two-thirds of basitarsus (though
base light brown) and little less than basal half of second tarsomere whitish; basitarsus (Fig. 22C) narrow, slightly
narrowed toward apex, 7.6 times as long as wide, and 0.64 and 0.54 times as wide as greatest widths of tibia and
femur, respectively; calcipala (Fig. 22C) well developed, slightly longer than wide, and 0.6 times as wide as
greatest width of basitarsus; pedisulcus (Fig. 22C) well developed; claw (Fig. 22D) with large basal tooth 0.5 times
length of claw. Wing. Length 1.5 mm. Costa with dark-brown spinules and light-brown hairs except basal patch of
yellow hairs. Subcosta haired except near apex bare. Hair tuft on base of radius dark brown intermixed with several
yellow hairs. Basal portion of radius fully haired. Basal cell absent. Halter. Creamy except basal stem darkened.
Abdomen. Basal scale grayish yellow, with fringe of yellowish-white hairs. Dorsal surface of abdomen medium
brown except segment 2 grayish yellow though tergal plate medium brown, and moderately covered with dark
short to long hairs; tergites of segments 2 and 6–9 shiny when illuminated at certain angles. Ventral surface of
segment 2 creamy, those of other segments ochreous except sternite 8 dark brown; sternal plate on segment 7
undeveloped. Terminalia. Sternite 8 (Fig. 22E) bare medially, with 12–14 medium-long to long hairs together with
few slender short hairs on each side. Ovipositor valve (Fig. 22E) triangular (though posteromedial corner appearing
to be truncated), thin, membranous, moderately covered with microsetae interspersed with three or four short hairs;
inner margins nearly straight or sinuous, with small depression subapically, moderately sclerotized, and somewhat
separated from each other. Genital fork (Fig. 22F) of usual inverted-Y form, with slender stem; arms of moderate
width, with lateral plate angulated anterodorsally. Paraproct in ventral view (Fig. 22G) with anteromedian margin
slightly raised ventrally, with unpigmented anteromedial surface having four or five sensilla; paraproct in lateral
view (Fig. 22H) much produced ventrally, 0.6 times as long as wide, with 18 or 19 medium-long to long hairs on
ventral and lateral surfaces. Cercus in lateral view (Fig. 22H) short, rounded posteriorly, 0.5 times as long as wide.
Spermatheca (Fig. 22I) ellipsoidal, 1.6 times as long as greatest width, well sclerotized except duct and small area
near juncture with duct unsclerotized, and with many fissures on surface; internal setae absent; both accessory
ducts slender, subequal in diameter to major one.
Male. Body length 2.2–2.5 mm. Head. As wide as thorax. Upper eye medium brown, with large facets in 16 or
17 vertical columns and 17 horizontal rows. Face brownish black, whitish-gray pruinose when illuminated at
certain angles. Clypeus brownish black, whitish-gray pruinose when illuminated at certain angles, and moderately
covered with yellow short hairs interspersed with 12–16 dark-brown longer hairs on each side. Antenna composed
of scape, pedicel and nine flagellomeres, brownish black except base of first flagellomere light brown; first
flagellomere elongate, twice length of second one. Maxillary palp with five segments, light brown except third
segment dark brown, proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.1; third segment (Fig. 23A)
somewhat widened apically; sensory vesicle (Fig. 23A) ellipsoidal, medium-long, 0.3 times length of third
segment, and with opening of medium-size. Thorax. Scutum black, slightly shiny when illuminated at certain
angles, densely covered with golden-yellow short hairs. Scutellum brownish back, with golden-yellow short hairs
and dark-brown long upright hairs. Postnotum brownish black and bare. Pleural membrane bare. Katepisternum
longer than deep, brownish black, thinly pruinose and slightly shiny when illuminated at certain angles, moderately
covered with dark hairs. Legs. Foreleg: coxa and trochanter medium brown; femur medium brown with apical cap
dark brown; tibia medium brown except apical cap dark brown and medial portion of outer surface light brown:
tarsus brownish black; basitarsus slightly flattened, 9.1 times as long as greatest width. Midleg: coxa dark brown
except posterolateral surface brownish black; trochanter medium brown; femur medium brown with apical cap
FIGURE 23. Male of Simulium (Gomphostilbia) eshimai sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Hind basitarsus and second tarsomere (left side; outer view); C, Coxites, styles and ventral plate
(ventral view); D, Style (right side; ventrolateral view); E, Ventral plate and median sclerite (lateral view); F, Ventral plate
(caudal view); G, Median sclerite (caudal view); H, Paramere and aedeagal membrane (left side; caudal view); I & J,
Abdominal segment 10 and cerci (right side; I, lateral view; J, caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–J.

dark brown (though extreme tip somewhat paler); tibia medium brown except base dark yellow and apical cap dark
brown, and with golden-yellow short hairs on posterior surface of basal one-third; tarsus dark brown to brownish
black. Hind leg: coxa dark brown; trochanter light brown; femur medium brown with apical cap dark brown
(though extreme tip yellow); tibia medium brown to brownish black except base dark yellow to light brown; tarsus
dark brown except basitarsus grayish to light brown and little less than basal half of second tarsomere whitish;
basitarsus (Fig. 23B) enlarged, spindle-shaped, 4.2 times as long as wide, and 0.9 and 1.0 times as wide as greatest
widths of tibia and femur, respectively; calcipala (Fig. 23B) well developed, slightly longer than width at base, and
0.4 times as wide as greatest width of basitarsus; pedisulcus (Fig. 23B) well developed. Wing. Length 2.2 mm.
Other characters as in female except costa without basal patch of yellow hairs, subcosta bare, and hair tuft on base
of radius dark brown. Halter. Ochreous with basal portion darkened. Abdomen. Basal scale brownish black, with
fringe of dark-brown long hairs. Dorsal surface of abdomen dark brown to brownish black, covered with light-
brown short to long hairs; segments 2, 6 and 7 each with pair of shiny dorsolateral patches, when illuminated at
certain angles; ventral surface of segment 2 grayish white, those of other segments medium to dark brown.
Genitalia. Coxite in ventral view (Fig. 23C) nearly rectangular, 1.7 times as long as its greatest width. Style in
ventral view (Fig. 23C) slender, curved inward, tapered toward middle, with apical spine; style in ventrolateral
view (Fig. 23D) 0.9 times as long as coxite, tapered from base to middle, then nearly parallel-sided toward apex,
with round apex. Ventral plate in ventral view (Fig. 23C) transverse, 0.7 times as long as greatest width at base,
with body narrowed posteriorly, with anterior margin produced anteromedially, posterior margin concave medially,
and densely covered with microsetae on ventral surface except anterolateral areas bare; basal arms of moderate
length, somewhat divergent, then convergent apically; ventral plate in lateral view (Fig. 23E) with posterior portion
of body not produced ventrally; ventral plate in caudal view (Fig. 23F) slightly rounded ventrally
(width:height=1.0:0.3), with dorsal margin concave medially, densely covered with microsetae on posterior
surface. Median sclerite (Fig. 23C, E, G) plate-like, arising from level near anteromedial tip of ventral plate, and
directed dorsally. Paramere (Fig. 23H) with four distinct hooks. Aedeagal membrane (Fig. 23H) moderately
covered with microsetae, and with no sclerotized dorsal plate. Ventral surface of abdominal segment 10 (Fig. 23I,
J) without distinct hair near posterior margin on each side. Cercus (Fig. 23I, J) rounded, slightly produced ventrally,
with 17 or 18 hairs.
Pupa. Body length 2.3–2.5 mm. Head. Integument yellow, moderately covered with small round tubercles
except most of ventral surface of face bare; antennal sheath without tubercles; frons with three pairs of unbranched
long trichomes with straight apices; face with two pairs of unbranched long trichomes with straight apices (Fig.
24A); three frontal trichomes on each side arising close together, subequal in length to one another and nearly as
long as or somewhat longer than facial ones. Thorax (Fig. 24B). Integument yellow, moderately covered with
round tubercles on anterior one-third, and almost bare on other portion except dorsal surface near posterior margin
sparsely covered with small tubercles; thorax with three long anterodorsal trichomes with coiled or straight apices,
two anterolateral trichomes with straight apices (anterior one medium-long, posterior one long), one long
mediolateral trichome with straight apex, and three ventrolateral trichomes with straight apices (anterior one
medium-long, two others long) on each side; all trichomes unbranched. Gill (Fig. 24B, C) composed of eight
slender thread-like filaments, arranged in three groups (one dorsal, one ventral inner and one ventral outer) arising
from short common basal stalk, which has somewhat swollen basal fenestra; dorsal group composed of three
filaments all arising upward at same level from short stalk, ventral inner group composed of one individual and two
paired filaments with short primary and secondary stalks, or all three filaments arising forward at same level from
short stalk, and ventral outer group composed of two paired filaments with short stalk; all filaments light brown,
subequal in length (0.7–1.0 mm) and thickness to one another except ventral filament of ventral pair somewhat
thicker than others when compared basally; cuticle of all filaments without annular ridges and furrows, and densely
covered with minute tubercles. Abdomen. Dorsally, all segments nearly unpigmented and without tubercles except
segments 1 and 2 pale yellowish, and segment 9 and basal portions of spine-combs on segments 6–9 yellowish;
segment 1 with one unbranched slender medium-long hair-like seta on each side; segment 2 with one unbranched
slender medium-long or short hair-like seta and five short somewhat spinous setae submedially near posterior
margin on each side; segments 3 and 4 each with four hooked spines and one short somewhat spinous seta near
posterior margin on each side; segments 5–8 each with two or three unbranched short setae near posterior margin
on each side; segments 6–9 each with spine-combs in transverse row and comb-like groups of minute spines near
FIGURE 24. Pupa of Simulium (Gomphostilbia) eshimai sp. nov. A, Head (left side; outer view); B, Thorax and gill filaments
(left side; outer view); C, Basal portions of gill filaments (left side; dorsal view); D, Terminal hooks (caudal view); E, Cocoon
(dorsal view). Scale bars. 1.0 mm for E; 0.1 mm for A and B; 0.05 mm for C; 0.02 mm for D.
anterior margin; segment 9 with pair of cone-like terminal hooks (Fig. 24D). Ventrally, segment 4 with one
unbranched hook and few unbranched short setae on each side; segment 5 with pair of bifid or trifid hooks
submedially and few unbranched short slender setae on each side; segments 6 and 7 each with pair of bifid or trifid
inner and unbranched outer hooks somewhat spaced from each other and few unbranched short slender setae on
each side; segments 4–8 with comb-like groups of minute spines. Each side of segment 9 with three grapnel-shaped
hooklets. Cocoon (Fig. 24E). Wall-pocket-shaped, moderately woven, widely extended ventrolaterally; anterior
margin thickly woven, without anterodorsal bulge or projection; posterior half with floor roughly or moderately
woven; individual threads visible; 3.0 mm long by 2.6–2.8 mm wide.
Mature larva. Body length 4.0–4.6 mm. Body light ochreous except anterior surface of thoracic proleg,
thoracic segment 3 (except ventral surface light ochreous) and abdominal segments 1–4 entirely greenish, and
ventral surface of abdominal segments 5–9 creamy (though light-ochreous transverse band on ventral surface of
segment 7). Head. Head capsule whitish yellow, with no distinct head spots, and sparsely covered with
unpigmented minute setae (though moderately on dorsal surface); ventral surface slightly darkened near lateral
margins of postgenal cleft, and with negative elongate spot on each side of postgenal cleft. Antenna composed of

FIGURE 25. Larva of Simulium (Gomphostilbia) eshimai sp. nov. A, Mandible; B, Hypostoma; C, Head capsule (ventral
view); D, Setae on dorsal surface of abdominal segment 8. Scale bars. 0.1 mm for C; 0.02 mm for A, B and D.
three articles and apical sensillum, longer than stem of labral fan; proportional lengths of first, second, and third
articles 1.0:0.8:0.7–0.8. Labral fan with 42 primary rays. Mandible (Fig. 25A) with three comb-teeth decreasing in
length from first tooth to third; mandibular serration composed of two teeth (one medium-sized, one small); major
tooth at acute angle against mandible on apical side; supernumerary serrations absent. Hypostoma (Fig. 25B) with
row of nine apical teeth, of which median tooth is longer than each corner tooth; lateral margin smooth; four or five
hypostomal bristles per side lying nearly parallel to or slightly divergent posteriorly from lateral margin. Postgenal
cleft (Fig. 25C) arrow-head-shaped, long, 4.2 times length of postgenal bridge. Cervical sclerites composed of pair
of small yellow rod-like pieces. Thorax and Abdomen. Thoracic proleg without hairs; thoracic cuticle moderately
covered with dark setae with two to five branches (majority of setae with three or four branches) interspersed with
unbranched colorless setae (similar to those in Fig. 25D) on dorsal surface, abdominal cuticle of segments 1–4
sparsely covered with similar branched dark minute setae on dorsal surface, and that of segments 5–9 densely
covered with dark minute setae each with two to eight branches (majority of setae with four to six branches)
interspersed with unbranched colorless setae (Fig. 25D) on dorsal and dorsolateral surfaces, even on dorsal surface
of each side of anal sclerite; last abdominal segment densely covered with unbranched colorless longer setae on
dorsolateral and lateral surfaces of each side of anal sclerite and on each lateral surface down to base of ventral
papilla; thorax and abdomen without dorsal protuberances. Rectal scales present, unpigmented. Rectal organ
compound, each of three lobes with several finger-like secondary lobules (exact number of secondary lobules
uncountable). Anal sclerite of usual X-form, with anterior arms 1.1 times length of posterior ones, broadly
sclerotized at base; no sensilla on broad base and posterior to posterior arms; accessory sclerite absent. Last
abdominal segment with pair of large conical ventral papillae. Posterior circlet with 85–88 rows of hooklets with
up to 13 or 14 hooklets per row.
in a vial as [UMSRP: Vietnam 030, Holotype, Simulium (G.) eshimai, Female, coll. Vietnam, 20-XII-2014,
Takaoka et al.], reared from a pupa collected from a river (width 6 m, depth 0.6 m, bottom rocky, water temperature
11.0˚C, exposed to the sun, elevation 1,728 m, 22˚22’22.719”N/103˚45’24.852”E) fast-flowing in forest, Lai Chau,
Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen
& K.W. Lau. PARATYPES: Two males, three pupal exuviae, one mature larva, same data as those of the holotype
in 80% ethanol, labeled in a vial as [UMSRP: Vietnam 030, Paratype, Simulium (G.) eshimai, coll. Vietnam, 20-
XII-2014, Takaoka et al.]; one mature larva in 80% ethanol, labeled in a vial as [UMSRP: Vietnam 030, Paratype,
Simulium (G.) eshimai, coll. Vietnam, 20-XII-2014, Takaoka et al.], collected from a small cascading stream (width
2.5 m, depth 3 cm, bottom rocky, water temperature 11.0˚C, exposed to the sun, elevation 1,853 m,
22˚21’18.609”N/103˚46’27.550”E) slow-flowing in forest, Oguy Ho, Sapa, Lao Cai Province, northern Vietnam,
20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
Biological notes. The pupae of this new species were collected from plastic sheets in the current. Associated
species were S. (M.) nigrofilum sp. nov., S. (N.) laichauense sp. nov., S. (N.) tayense sp. nov., S. (S.) rosliramlii sp.
nov. and S. (S.) turgidum sp. nov.
Distribution. Vietnam (Lao Cai).
Etymology. The species name eshimai is in honor of Prof. Nobuoki Eshima, Kyoto University, for his great
contribution to statistical analysis of infectious diseases and long-standing support of our studies of black flies.
Remarks. This new species is placed in the S. darjeelingense species-group (five species included) of the
subgenus Gomphostilbia, defined by Takaoka (2012), based on the fore coxae darkened, male hind basitarsus
enlarged (Fig. 23B), ventral plate not produced ventrally (Fig. 23E, F) (its ratio of the height against the greatest
width is 0.33), and pupal gill with eight short slender filaments (Fig. 24B).
This new species shows great similarities to S. (G.) chayamaritae Takaoka & Srisuka from Thailand (Takaoka
and Srisuka 2010) in many characters including the number of male upper-eye facets, male genitalia (Fig. 23C),
arrangement of pupal gill filaments (Fig. 24B) and pupal face with an additional trichome on each mediolateral
surface (Fig. 24A). However, this new species is distinguished from the latter species by the round tubercles on the
frons (frons with cone-shaped tubercles each with a sharply-pointed apex in S. (G.) chayamaritae). There is no
information on the additional trichomes on the pupal face of the four other species of the S. darjeelingense species-
group.
This new species is distinguished from S. (G.) darjeelingense Datta from India (Datta 1973), S. (G.)
guizhouense Chen, Zhang & Yang from China (Chen et al. 2003a), and S. (G.) guniki Takaoka from Sabah,
Malaysia (Takaoka 2001b) by the pupal gill filaments arranged as 3+3+2 or 3+(1+2)+2 (Fig. 24B) (2+(2+2)+2 in S.
(G.) darjeelingense and S. (G.) guizhouense, and all eight filaments arising at the same level from the common
basal stalk in S. (G.) guniki). Simulium (G.) darjeelingense also differs from this new species by the number of male
upper-eye facets (in 14 vertical columns and 14 horizontal rows), and absence of pupal terminal hooks.
The remaining species, S. (G.) xizangense An, Zhang & Deng from China, which was described from a male
(An et al. 1990), differs from this new species by the small round sensory vesicle and ventral plate slightly widened
posteriorly.
5) Simulium epistum species-group
Simulium (Gomphostilbia) montiblense Takaoka, 1983
Simulium (Gomphostilbia) montiblense Takaoka, 1983: 71–73 (Female, male, pupa and larva).
Distribution. Philippines (Palawan) and Vietnam.

Remarks. This species was described from Palawan, the Philippines by Takaoka (1983), and later recorded
from Vietnam by Crosskey and Howard (1997). No detailed information of this record was available; hence, the
material and locality in Vietnam are unknown. If its identification was based only on the pupal stage, there is a
possibility that the species recorded as S. (G.) montiblense is one of three species, i.e., S. (G.) eshimai sp. nov., S.
(G.) longlanhense or S. (G.) parahiyangum because the pupal gills of S. (G.) montiblense and the three other species
are all short, and their arrangements are almost indistinguishable from one another (Takaoka 1983; Takaoka and
Sigit 1992; Takaoka et al. 2015a).
6) Simulium gombakense species-group
Simulium (Gomphostilbia) thuathienense Takaoka & Sofian-Azirun, 2015

Simulium (Gomphostilbia) thuathienense Takaoka & Sofian-Azirun: (in Takaoka et al., 2015a): 48–52 (Female, pupa and
larva).

Remarks. This species was described from Thua Thien Hue Province in central Vietnam by Takaoka et al.
(2015a). The male of this species remains unknown.
7) Simulium varicorne species-group
Simulium (Gomphostilbia) breviflagellum Takaoka & Sofian-Azirun, 2015
Simulium (Gomphostilbia) breviflagellum Takaoka & Sofian-Azirun (in Takaoka et al., 2015a): 53–57 (Male and pupa)

Remarks. This species was described from Lam Dong Province in southern Vietnam by Takaoka et al.
(2015a). The female and larva of this species are unknown.
II. Subgenus Montisimulium Rubtsov
The subgenus Montisimulium is represented by only one species, S. (M.) nigrofilum sp. nov. in Vietnam. This
subgenus is newly recorded from Vietnam.
Simulium (Montisimulium) nigrofilum Takaoka & Sofian-Azirun sp. nov.
Female. Unknown.
Male. Body length 3.1 mm. Head. As wide as thorax. Upper eye with large facets in 17 vertical columns and
19 horizontal rows. Face brownish black. Clypeus brownish black, densely covered with yellow short hairs
interspersed with many dark-brown hairs. Antenna composed of scape, pedicel and nine flagellomeres, brownish
black except scape and pedicel medium brown and base of first flagellomere whitish yellow; first flagellomere
elongate, twice length of second one. Maxillary palp with five segments, light brown except third segment dark
brown and first and second segments yellow, proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.1;
third segment (Fig. 26A) slender; sensory vesicle (Fig. 26A) ellipsoidal, medium-long, 0.3 times length of third
segment, and with small opening. Thorax. Scutum brownish black, densely covered with golden-yellow short
hairs. Scutellum brownish black, with golden-yellow short hairs and dark-brown long upright hairs. Postnotum
brownish black and bare. Pleural membrane bare. Katepisternum longer than deep, brownish black, and bare. Legs
(coloration appearing incomplete, in particular all femora and tibia). Foreleg: coxa dark yellow; trochanter light
brown; femur whitish yellow with apical cap dark brown; tibia dark brown except medial portion of outer surface
paler: tarsus brownish black; basitarsus slightly flattened, 11.0 times as long as greatest width. Midleg: medium
brown; trochanter light brown; femur light grayish with apical cap dark brown; tibia dark brown except median
large area paler; tarsus dark brown to brownish black. Hind leg: coxa medium brown; trochanter dark yellow;
femur light grayish with apical cap dark brown; tibia dark brown to brownish black except median large area paler;
tarsus dark brown except basitarsus grayish to light brown; basitarsus (Fig. 26B) somewhat enlarged, spindle-
shaped, 4.3 times as long as wide, and 0.9 and 0.9 times as wide as greatest widths of tibia and femur, respectively;
calcipala (Fig. 26B) well developed, as long as width at base; pedisulcus well developed. Wing. Length 2.2 mm.
Costa with dark-brown spinules and light-brown hairs except basal patch of dark hairs mixed with yellow hairs.
Subcosta with one or five hairs. Hair tuft on base of radius dark brown. Basal portion of radius fully haired. Basal
cell absent. Halter. Light ochreous with basal portion darkened. Abdomen. Basal scale dark brown, with fringe of
yellow long hairs. Dorsal surface of abdomen dark brown to brownish black, covered with yellow short hairs
interspersed with dark hairs. Genitalia. Coxite in ventral view (Fig. 26C) nearly rectangular, 2.1 times as long as its
greatest width. Style in ventral view (Fig. 26C) slender, curved inward, tapered from middle to apex, with apical
FIGURE 26. Male of Simulium (Montisimulium) nigrofilum sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Hind basitarsus (left side; outer view); C, Coxites, styles and ventral plate (ventral view); D & E,
Styles (right side; D, ventrolateral view; E, medial view); F, Ventral plate and median sclerite (lateral view); G, Ventral plate
(caudal view); H, Median sclerite (caudal view); I, Paramere (left side; ventral view); J, Aedeagal membrane and dorsal plate
(caudal view); K & L, Abdominal segment 10 and cerci (right side; K, lateral view; L, caudal view). Scale bars. 0.1 mm for B;
0.02 mm for A and C–L.

spine; style in ventrolateral view (Fig. 26D) 0.6 times as long as coxite, nearly parallel-sided from base to middle,
then tapered toward apex; style in medial view (Fig. 26E) boot-shaped, with apical portion twisted dorsally in form
of triangular projection. Ventral plate in ventral view (Fig. 26C) transverse, 0.4 times as long as greatest width at
base, with body narrowed posteriorly, with anterior margin produced anteromedially, posterior margin greatly
concave medially, and densely covered with microsetae on medial one-third of ventral surface; basal arms short,
nearly parallel-sided, though slightly convergent apically; ventral plate in lateral view (Fig. 26F) with apical
portion of arms slightly curved dorsally; ventral plate in caudal view (Fig. 26G) slightly rounded ventrally, with
dorsal margin greatly concave medially, densely covered with microsetae medially on posterior surface. Median
sclerite (Fig. 26C, F, H), plate-like, narrowed toward middle, then widened toward apex, arising from level near
anteromedial tip of ventral plate, and directed posteriorly. Paramere (Fig. 26I) with four distinct hooks. Aedeagal
membrane (Fig. 26J) moderately covered with microsetae, and with weakly sclerotized dorsal plate in form of
broad band. Ventral surface of abdominal segment 10 (Fig. 26K, L) with two to five distinct hairs near posterior
margin on each side. Cercus (Fig. 26K, L) rounded, slightly produced ventrally, with eight to eleven hairs.
Pupa. Body length 3.2 mm. Head. Integument including antennal sheaths yellow, densely covered with
tubercles having few to several minute secondary projections; frons with two pairs of unbranched short trichomes
with straight apices (one stout, one slender) (Fig. 27A); face with pair of unbranched short stout trichomes with
coiled apices (Fig. 27B); two frontal trichomes on each side arising close together, and much shorter than facial
ones. Thorax. Integument yellow, densely covered with tubercles bearing few to several minute secondary
projections; thorax with three short anterodorsal trichomes (two stout, with coiled apices, one slender, with straight
apex) (Fig. 27C), two short slender anterolateral trichomes with straight apices (anterior one shorter than posterior
one) (Fig. 27D), one short stout mediolateral trichome with straight apex (Fig. 27E), and three short ventrolateral
trichomes with straight apices (anterior one slender, two others stout) (Fig. 27F) on each side; all trichomes
unbranched. Gill (Fig. 27G) composed of 12 slender thread-like filaments, arranged in four groups (one inner, one
middle dorsal, one middle ventral and one outer) arising from medium-long common basal stalk, which has
transparent basal fenestra and 0.8 times length of interspiracular trunk; inner group composed of two filaments with
short stalk, middle dorsal and ventral groups each composed of one individual and two paired filaments with short
primary and medium-long secondary stalks, and outer group composed of two pairs of four filaments with short
common stalk and medium-long secondary stalks; these two pairs lying vertically, one pair dorsal and another
ventral; all filaments light brown except common basal stalk dark brown, subequal in length and thickness (though
their exact length unmeasurable due to loss of apical portions) and tapered toward apices; cuticle of all filaments
with well-defined annular ridges and furrows, and densely covered with minute tubercles, and with numerous dark-
brown minute particles inside outer cuticular layer (Fig. 27H). Abdomen. Dorsally, all segments yellowish;
segments 1 and 2 with minute tubercles, segment 1 with one unbranched slender short hair-like seta (Fig. 27I) on
each side; segment 2 with one unbranched slender short hair-like seta and five short somewhat spinous setae
submedially (Fig. 27J) near posterior margin on each side; segments 3 and 4 each with four hooked spines and one
short somewhat spinous seta near posterior margin on each side; segments 5–8 each with two unbranched short
setae near posterior margin on each side; segments 6–8 each with spine-combs in transverse row; segments 5–9
each with comb-like groups of minute spines near anterior margin on each side; segment 9 with pair of cone-like
terminal hooks (Fig. 27K). Ventrally, segment 4 with few unbranched short setae on each side; segment 5 with pair
of bifid hooks submedially and few unbranched short slender setae on each side; segments 6 and 7 each with pair of
bifid inner and outer hooks somewhat spaced from each other and few unbranched short slender setae on each side;
segments 4–8 each with comb-like groups of minute spines; segment 9 yellowish, covered sparsely with round
small tubercles. Segment 9 with three grapnel-shaped hooklets (Fig. 27L) on each lateral side. Cocoon (Fig. 27M,
N). Wall-pocket-shaped, thickly woven, not extended ventrolaterally, anterodorsal margin slightly produced
anteriorly but without bulge or projection; posterior two-thirds with floor moderately woven; individual threads
invisible; 3.0 mm long by 1.6–1.8 mm wide.
Mature larva. Body length 6.2–6.6 mm. Body creamy except anterior and posterior surfaces of thoracic
proleg, ventral surface of thoracic segments 2 and 3, and abdominal segments 1–4 light grayish-green, and with
reddish-brown or ochreous markings as follows: thoracic segment 1 encircled with reddish-brown band though
disconnected ventrally, thoracic segments 2 and 3 light reddish-brown dorsally and reddish brown ventrally,
abdominal segment 1 encircled with light reddish-brown or ochreous band though disconnected dorsomedially,
abdominal segments 2–4 each with pair of light reddish-brown or ochreous spots dorsally (though colored spots
only on segments 3 and 4 in one larva and absent in one larva), abdominal segments 5–9 each with reddish-brown
FIGURE 27. Pupa and larva of Simulium (Montisimulium) nigrofilum sp. nov. A–N, Pupa and O–R, Larva. A, Frontal
trichomes; B, Facial trichome; C–F, Thoracic trichomes (C, anterodorsal; D, anterolateral; E, mediolateral; F, ventrolateral); G,
Gill filaments (left side; dorsal view); H, Enlargement of filament showing black pigments inside cuticle; I, Hair-like seta on
dorsal surface of abdominal segment 1; J, Hair-like seta and minute seta on dorsal surface of abdominal segment 2; K, Terminal
hooks (caudal view); L, Grapnel-shaped hooklet on lateral surface of abdominal segment 9; M & N, Cocoons (M, dorsal view;
N, lateral view); O, Antenna showing three unpigmented annulations of segment 2; P, Mandible; Q, Hypostoma; R, Head
capsule (ventral view). Scale bars. 1.0 mm for M and N; 0.1 mm for G and R; 0.05 mm for O; 0.02 mm for H, K, P and Q; 0.01
mm for A-F, I, J and L.

or ochreous markings entirely or partially covering dorsal surface, and those on segments 5 and 6 extended laterally
and ventrally, though becoming lighter. Head. Head capsule yellow to dark yellow except areas in front of posterior
margin on dorsal and lateral surfaces, and area above eyebrow on lateral surface much darkened, and median
portion of ventral surface somewhat darkened, and with distinct head spots though anterior spots of posterolateral
spots faint or absent, and sparsely covered with unpigmented minute setae (though moderately on dorsal surface);
eyebrow distinct. Antenna composed of three articles and apical sensillum, longer than stem of labral fan;
proportional lengths of first, second, and third articles 1.0:0.9:0.7–0.8; antennal article 2 light yellow, with three
unpigmented annulations (Fig. 27O). Labral fan with 35 primary rays. Mandible (Fig. 27P) with three comb-teeth
decreasing in length from first tooth to third, or second and third teeth subequal; mandibular serration composed of
two teeth (one medium-sized, one small); major tooth at right angle against mandible on apical side;
supernumerary serrations absent. Hypostoma (Fig. 27Q) with row of nine apical teeth, of which median tooth as
long as or slightly longer than each corner tooth; lateral margin serrate apically; five or six hypostomal bristles per
side lying slightly divergent posteriorly from lateral margin. Postgenal cleft (Fig. 27R) absent (or vestigial if any).
Cervical sclerites composed of pair of small dark rod-like pieces. Thorax and Abdomen. Thoracic proleg without
hairs; thoracic and abdominal cuticle almost bare except last abdominal segment densely covered with unbranched
colorless setae on dorsolateral and lateral surfaces of each side of anal sclerite and on each lateral surface down to
base of ventral papilla. Rectal scales present, unpigmented. Rectal organ compound, each of three lobes with
several finger-like secondary lobules (exact number of secondary lobules uncountable). Anal sclerite of usual X-
form, with anterior arms as long as or slightly longer than posterior ones, broadly sclerotized at base; no sensilla on
broad base and posterior to posterior arms; accessory sclerite absent. Last abdominal segment with pair of large
conical ventral papillae. Posterior circlet with 78 or 79 rows of hooklets with up to 16 or 17 hooklets per row.
Type material. HOLOTYPE: Pharate male (with its associated pupal exuviae and cocoon) in 80% ethanol,
labeled in a vial as [UMSRP: Vietnam 031, Holotype, Simulium (M.) nigrofilum, Pharate male, coll. Vietnam, 20-
XII-2014, Takaoka et al.], dissected out from a pupa collected from a small stream (width 0.5 m, bottom rocky,
water temperature 11.0˚C, exposed to sun, elevation 1,750 m, 22˚21’43.110”N/103˚47’19.221”E) moderately
flowing in a secondary forest, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-
Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: One pupal exuviae and three mature larvae, same data
as those of the holotype. in 80% ethanol, labeled in each vial as [UMSRP: Vietnam 031, Paratype, Simulium (M.)
nigrofilum, coll. Vietnam, 20-XII-2014, Takaoka et al.]
species were S. (N.) tayense sp. nov., S. (S.) laocaiense sp. nov., S. (S.) sapaense sp. nov. and S. (S.) turgidum sp.
nov.
Etymology. The species name nigrofilum refers to the apparently black pupal gill filaments due to numerous
minute black spots (Fig. 27H).
Remarks. This new species is characterized by the pupal gill with 12 filaments arranged in four groups (Fig.
27G), and wall-pocket-shaped cocoon (Fig. 27M, N). In the arrangement of the pupal gill, this species is most
similar to S. (M.) angkaense Takaoka & Choochote from Thailand (Takaoka and Choochote 2005a, 2009) and S.
(M.) nemorivagum Datta from India (Datta 1973). This new species is distinguished from S. (M.) angkaense and S.
(M.) nemorivagum by the male upper-eye facets in 17 vertical columns and 19 horizontal rows (20 vertical columns
and 22 horizontal rows in S. (M.) angkaense, and 12 vertical columns and 14 horizontal rows in S. (M.)
nemorivagum), and also from S. (M.) angkaense in the pupa by the presence of grapnel-shaped hooklets on each
side of abdominal segment 9 (Fig. 27L).
III. Subgenus Nevermannia Enderlein
The subgenus Nevermannia is represented by seven species in Vietnam, of which four are placed in the S.
feuerborni species-group, one is in the S. ruficorne species-group and the remaining two are in the S. vernum
species-group, which is newly recorded from Vietnam. The definition of the S. feuerborni species-group follows
Datta (1973) and those of the other two species-groups follow Crosskey (1969).
Keys to seven species of the subgenus Nevermannia from Vietnam
Females
1. Fore coxa darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Fore coxa yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Relative length of labrum against clypeus 0.7; relative length of sensory vesicle against third maxillary palpal segment 0.5
(Fig. 28A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. laichauense sp. nov.
Relative length of labrum against clypeus 0.9; relative length of sensory vesicle against third maxillary palpal segment 0.6–0.7
(Fig. 30A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tayense sp. nov.
3. Scutum without dark longitudinal vittae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. aureohirtum
Scutum with three dark longitudinal vittae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Sensory vesicle with large opening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. langbiangense
Sensory vesicle with opening of moderate size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Genital fork with base of anteriorly directed projection with strongly sclerotized horizontal bar . . . . . . . . . . . . . . S. maeaiense
Genital fork with base of anteriorly directed projection without horizontal bar. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..6
6. Genital fork with rounded posteromedial projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. phami
Genital fork with pointed posteromedial projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bachmaense
Males*
1. Ventral plate with median keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aureohirtum
Ventral plate without median keel. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Fore coxa darkened; paramere with one hook . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tayense sp. nov.
Fore coxa yellow; paramere with several hooks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Antenna yellow except pedicel and little less than apical half of first flagellomere somewhat dark yellow to light brown, and
base of first flagellomere whitish yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bachmaense
Antenna light brown except scape and pedicel medium brown, and base of first flagellomere yellowish white. . . . . . . . . . . . .4
4. Upper-eye facets in 19 vertical columns and 20 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maeaiense
Upper-eye facets in 17 or 18 vertical columns and 18 or 19 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Relative length of maxillary palpal segment 5 against segment 3 equal to 1.7 . . . . . . . . . . . . . . . . . . . . . . . . . S. langbiangense
Relative length of maxillary palpal segment 5 against segment 3 equal to 1.9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. phami
Pupae
1. Gill with four filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Gill with five or six filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2. Cocoon with anterodorsal projection (Fig. 32K); frons and thorax light ochreous . . . . . . . . . . . . . . . . . . . . . S. tayense sp. nov.
Cocoon without anterodorsal projection (Fig. 29G); frons and anterolateral surface of thorax dark brown (Fig. 29A, B) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. laichauense sp. nov.
3. Gill with five filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. phami
Gill with six filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4. Six gill filaments widely divergent basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. aureohirtum
Six gill filaments not divergent, directed forward. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Cocoon with anterodorsal projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. maeaiense
Cocoon without anterodorsal projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6. Gill filaments arranged as 2+2+2 in three pairs (inner dorsal, outer dorsal, and ventral). . . . . . . . . . . . . . . . . ..S. langbiangense
Gill filaments arranged as four in dorsal group and two in ventral group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. bachmaense
Mature larvae**
1. Accessory sclerite (Fig. 33D) present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tayense sp. nov.
Accessory sclerite absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Mandibular serrations composed of two large teeth; body without reddish-brown markings . . . . . . . . . . . . . . . . S. aureohirtum
Mandibular serrations composed of one large tooth and one small tooth; body with reddish-brown markings . . . . . . . . . . . . . 3
3. Labral fan with 23–25 primary rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maeaiense
Labral fan with 34–44 primary rays… . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Postgenal cleft 0.6–0.7 length of postgenal bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. langbiangense
Postgenal cleft 0.4–0.5 length of postgenal bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
5. Length ratio of second antennal article against first article 1.1; posterior circlet with 74–76 rows of hooklets . . . . . . . S. phami
Length ratio of second antennal article against first article 0.6; posterior circlet with 88 rows of hooklets . . . . . S. bachmaense
* The male of S. laichauense sp. nov. is unknown.
** The larva of S. laichauense sp. nov. is unknown.

1) Simulium feuerborni species-group
Simulium (Nevermannia) bachmaense Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Nevermannia) bachmaense Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014c): 566–570 (Female, male,
pupa and larva).

Remarks. This species was described from Bach Ma National Park, central Vietnam (Takaoka et al. 2014c). It
is characterized by the female genital fork with a pointed posteromedial projection on each arm.
Simulium (Nevermannia) langbiangense Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Nevermannia) langbiangense Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014c): 556–563 (Female, male,
pupa and larva).

Remarks. This species was described from Lam Dong Province, southern Vietnam (Takaoka et al. 2014c). It is
characterized by the female sensory vesicle with an enlarged opening.
Simulium (Nevermannia) maeaiense Takaoka & Srisuka, 2011
Simulium (Nevermannia) maeaiense Takaoka & Srisuka, 2011: 57–70 (Female, male, pupa and larva).
Specimens examined. One female and one male, reared from pupae collected from a small stream (width 20 cm,
depth 2 cm, bottom rocky, water temperature 7.0˚C, partially shaded, elevation 1,893 m, 22˚21’28.378”N/
103˚45’52.084”E) slowly flowing in shrubs, Lai Chau, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by
H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
Distribution. Thailand and Vietnam (Lao Cai) (New record).
Remarks. Simulium (N.) maeaiense was described from Thailand (Takaoka and Srisuka 2011). This is the first
record of this species from Vietnam. This species is characterized by the female genital fork with a sclerotized
horizontal bar at the base of the anterodorsal projection on each lateral arm, male upper-eye facets in 19 vertical
columns and 20 horizontal rows, paramere with five or six hooks, pupal gill with six long filaments arranged
horizontally as 2+1+2+1 from inside to outside, and cocoon with an elongate anterodorsal projection (Takaoka and
Srisuka 2011). There are differences in several morphological features between Vietnamese and Thai specimens
(characters of Thai specimens in parentheses): in the female, body length 3.2 mm (2.4–2.6 mm), wing length 3.1
mm (2.6–2.8 mm), cibarium with 33 processes (40–44 processes), fore basitarsus 8.5 times as long as its greatest
width (8.4 times), hind basitarsus 7.6 times as long as its width (6.2–6.8 times), 0.7 times as wide as the hind tibia
(0.8 times), 0.6 times as wide as the hind femur (0.6–0.7 times), claw with a large basal tooth 0.46 times as long as
the claw (0.49 times), spermatheca ovoidal and 1.3 times as long as its width (1.2 times); in the male, body length
3.0 mm (2.6–2.9 mm), wing length 2.9 mm (2.4–2.5 mm), scutum brownish black to black with no longitudinal
vittae (dark brown with three longitudinal vittae), subcosta with three to five hairs (without hairs), hind basitarsus
4.4 times as long as wide (4.6–4.9 times), 1.0 times as wide as the hind tibia (0.9 times), 1.1 times as wide as the
hind femur (0.9 times).
Simulium (Nevermannia) phami Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Nevermannia) phami Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014c): 563–566 (Female, male, pupa
and larva).
Remarks. This species was described from Lam Dong Province, southern Vietnam (Takaoka et al. 2014c). It is
characterized by the pupal gill with five filaments.
2) Simulium ruficorne species-group
Simulium (Nevermannia) aureohirtum Brunetti, 1911
Simulium aureohirtum Brunetti, 1911: 283–288 (Male).

Simulium (Nevermannia) aureohirtum: Ogata, 1956: 61–62; Ogata, 1966: 129; Takaoka & Roberts, 1988: 194–195; Takaoka,
2003: 37–45 (Female, male, pupa and larva).
Simulium (Eusimulium) aureohirtum: Puri, 1933c: 1–7 (Female, male, pupa and larva); Takaoka, 1979: 382–384 (Female,
male, pupa and larva).
Specimens examined. Two males (reared from pupae), one pupal exuviae and one mature larva, collected from a
stream (width 30 cm, depth 5 cm, bottom sandy, water temperature 18.0˚C, exposed to the sun, elevation 80 m,
19˚20’12.147”N/105˚09’15.908”E) slow flowing near a forest, Chau Dinh, Quy Chau, Nghe An Province, northern
Vietnam, 11-XII-2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W. Lau.
Distribution. India, Bhutan, China, Guam, Indonesia, Japan, Malaysia, Nepal, Pakistan, Philippines, Sri
Lanka, Taiwan, Thailand, Vietnam (Lam Dong, Nghe An and Vinh Phuc).
Remarks. This species is characterized by the unique female and male genitalia, pupal gill with six filaments
and larval head capsule with bold head spots (Takaoka 2003). This is the only autogenous species in the Oriental
Region, of which the female can develop the first batch of eggs without blood-meals (Takaoka and Noda 1979).
The pupae and larvae of this species appear to be highly adapted to relatively high water temperatures, slow flow
and even somewhat muddy or contaminated waters, where most other simuliid species do not occur (Takaoka
2003). As noted previously (Takaoka et al. 2014a), these capabilities, together with other factors, may explain why
this species is widely distributed in the Oriental Region and extends into the Palearctic and Australasian Regions.
Alternatively, given its wide grographical range, it might be a complex of species.
3) Simulium vernum species-group
Simulium (Nevermannia) laichauense Takaoka & Sofian-Azirun sp. nov.
Female. Body length 2.5 mm. Head. Slightly narrower than width of thorax. Frons brownish black, moderately
covered with whitish yellow (golden yellow when illuminated at certain angles) recumbent short hairs interspersed
with few dark-brown long hairs along each lateral margin; frontal ratio 1.8:1.0:2.6. Frons:head ratio 1.0:5.9.
Fronto-ocular area directed laterally and slightly upward, rounded apically. Clypeus dark brown, thinly whitish-
gray pruinose, slightly shiny when illuminated at certain angles, moderately covered with yellow short hairs
interspersed with one to three dark-brown long hairs though mediolongitudinal area widely bare. Labrum 0.7 times
as long as clypeus. Antenna composed of scape, pedicel and nine flagellomeres, entirely brownish black; first
flagellomere 1.6 times as long as second one. Maxillary palp composed of five segments, medium brown except
segments 1 and 2 ochreous and segment 3 dark brown, proportional lengths of third, fourth, and fifth segments
1.0:0.8:1.5; third segment (Fig. 28A) somewhat enlarged; sensory vesicle (Fig. 28A) enlarged, ellipsoidal, 0.5
times as long as third segment, and with large opening apically. Maxillary lacinia with 15 inner and 17 or 18 outer
teeth. Mandible with 35 inner and 17 or 18 outer teeth. Cibarium (Fig. 28B) with enlarged cornuae; no processes
near anterodorsal surface. Thorax. Scutum black except anterolateral calli dark brown, thinly whitish-gray
pruinose and slightly shiny when illuminated at certain angles, densely covered with whitish yellow (golden yellow
when illuminated at certain angles) recumbent short hairs. Scutellum dark brown, covered with whitish-yellow
short hairs and dark-brown long hairs. Postnotum brownish black, whitish-gray pruinose and slightly shiny when
illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum longer than deep, brownish black,
whitish-gray pruinose and slightly shiny when illuminated at certain angles, and bare. Legs. Medium to dark brown

FIGURE 28. Female of Simulium (Nevermannia) laichauense sp. nov. A, Third segment of maxillary palp with sensory
vesicle (right side; front view); B, Cibarium (front view); C, Hind basitarsus and second tarsomere (left side; outer view). D,
Claw; E, Sternite 8 and ovipositor valve (left side only; ventral view); F, Genital fork (ventral view); G & H, Paraprocts and
cerci (G, ventral view; H, lateral view). I, Spermatheca. Scale bars. 0.1 mm for C; 0.02 mm for A, B and E–I; 0.01 mm for D.
except basal extreme of hind tibia and narrow portion of pedisulcus of second tarsomere pale. Fore basitarsus
somewhat dilated, 8.7 times as long as its greatest width. Hind basitarsus (Fig. 28C) slender, nearly parallel-sided,
7.8 times as long as wide, and 0.7 and 0.6 times as wide as tibia and femur, respectively; calcipala (Fig. 28C) well
developed, slightly longer than basal width, and 0.5 times as wide as greatest width of basitarsus; pedisulcus (Fig.
28C) well developed; claw (Fig. 28D) with large basal tooth 0.5 times length of claw. Wing. Length 2.9 mm. Costa
with dark-brown spinules and hairs except basal patch of hairs yellowish white. Subcosta with dark-brown hairs
except apical one-fifth or one-fourth bare. Hair tuft on base of radius dark brown. Basal portion of radius fully
haired. R1 with dark-brown spinules and hairs. R2 with dark-brown hairs only. Basal cell and basal median cell
absent. Halter. White except basal stem darkened. Abdomen. Basal scale ochreous to light brown, with fringe of
yellowish-white long hairs; dorsal surface of abdomen medium brown except rest of tergal plate of segment 2
ochreous, and moderately covered with whitish-yellow short hairs as well as dark-brown short to medium-long
hairs; tergites 6–8 shiny when illuminated at certain angles; ventral surface of segments 2–7 light ochreous, and
segment 8 medium brown; segment 7 with weakly defined large sternal plate medially. Terminalia. Sternite 8 (Fig.
28E) bare medially, and with 10–14 short to long hairs on each side. Ovipositor valve (Fig. 28E) triangular,
rounded posteromedially, thin, membranous, densely covered with microsetae interspersed with four to six fine
short setae (though one seta on each side medium-long); inner margins moderately sclerotized, narrowly darkened,
slightly sinuous, and moderately separated from each other. Genital fork (Fig. 28F) of usual inverted-Y form, stem
slender and moderately sclerotized; arms each with broad lateral plate having strongly sclerotized lateral margin.
Paraproct in ventral view (Fig. 28G) ovoidal, with strongly sclerotized anterior surface, covered with 10 medium-
long and long hairs on ventral and lateral surfaces, with shallow depression along anteromedial margin, with four
or seven sensilla on anteromedial transparent surface; paraproct in lateral view (Fig. 28H) slightly produced
ventrally beyond ventral margin of cercus. Cercus in ventral view (Fig. 28G) narrow, with widely concave inner
surface; cercus in lateral view (Fig. 28H) rounded posteriorly, 0.6 times as long as wide, and with numerous
medium-long to long hairs. Spermatheca (Fig. 28I) small, ovoidal, 1.2 times as long as its greatest width, well
sclerotized except area at juncture with duct, with reticulate surface pattern; internal setae absent; main duct and
both accessory ducts subequal in diameter to one another.
Pupa. Body length (excluding gill filaments) 3.4 mm. Head. Integument including antennal sheath yellow
except frons dark brown (Fig. 29A), and bare; frons with three unbranched long trichomes with straight apices (Fig.
29A) arising close together with one another on each side; face with one unbranched long trichome with straight
apex (Fig. 29A) on each side. Thorax. Integument yellow except anterolateral portion widely dark brown, and bare
except posterodorsal surface sparsely covered with small tubercles (Fig. 29B); thorax with three long anterodorsal
trichomes with coiled or straight apices, two long anterolateral trichomes with straight apices (one slightly shorter
than other), one medium-long mediolateral trichome with straight apex, and three ventrolateral trichomes with
straight apices (two long, one medium-long) (Fig. 29B), on each side; all trichomes unbranched. Gill (Fig. 29C)
with four slender thread-like filaments, arranged in dorsal and ventral pairs, each with short stalk arising from short
common basal stalk with moderate basal fenestra at base; common basal stalk 0.54 times length of interspiracular
trunk; stalks of dorsal and ventral pair subequal in length and thickness to each other, 0.4–0.5 times length of
common basal stalk; common basal stalk directed forward and somewhat outward, stalk of dorsal pair directed
upward and forward, with upper filament directed upward and forward, then curved downward, and lower filament
directed forward; stalk of ventral pair directed somewhat outward and forward, with outer and inner filaments
directed forward; upper filament of dorsal pair and outer filament of ventral pair on each side subequal in length
(2.6 mm) to each other, lower filament of dorsal pair and inner filament of ventral pair on each side broken losing
apical half, thus not measurable; upper filament of dorsal pair as thick as or very slightly thicker than outer filament
of ventral pair; lower filament of dorsal pair and inner filament of ventral pair subequal in thickness to each other,
and 0.9 times as thick as counter filament of each pair when compared basally; upper filament of dorsal pair and
outer filament of ventral pair slightly tapered toward apices, each with relatively thick apical portion (0.6 times as
thick as basal portion); all filaments medium brown, with annular ridges and furrows forming reticulate patterns on
surface of basal half, and densely covered with minute tubercles on outer surface. Abdomen. Dorsally, segments 1
and 2 dark gray and with minute spine-like tubercles, basal half of segment 3 grayish, and other segments
unpigmented except basal portions of spine-combs on segments 6–9 yellowish and posterior half of segment 9 light
grayish; segment 1 with one medium-long hair-like seta on each side; segment 2 with one medium-long hair-like
seta and five spinous short setae on each side; segments 3 and 4 each with four dark stout hooked spines and one
short spinous seta along posterior margin on each side; segments 2–4 each with one minute seta near anterior

FIGURE 29. Pupa of Simulium (Nevrmannia) laichauense sp. nov. A, Frons (left half; front view); B, Thorax showing
darkened portions (left side; outer view); C, Anterior portion of thorax and gill filaments (right side; outer view); D & E,
Terminal hooks (D, caudal view; E, lateral view); F, Bifid hooklet and minute spines on ventral surface of abdominal segment
4; G, Cocoon (dorsal view). Scale bars. 1.0 mm for G; 0.1 mm for A and B; 0.05 mm for C; 0.02 mm for D; 0.01 mm for E and
F.
margin on each side; segments 5–9 each with comb-like groups of minute spines in transverse rows on each side;
segments 6–9 each with spine-combs in transverse row on each side (though those on segment 9 much smaller in
size than those on other segments); segment 9 with pair of short blunt terminal hooks (Fig. 29D, E). Ventrally, all
segments unpigmented except segment 9 yellowish gray; segment 4 with one bifid hook and few unbranched short
setae on each side; segment 5 with two bifid hooks and few unbranched short setae on each side; segments 6 and 7
each with one bifid or trifid inner hook and one bifid outer hook and few unbranched short setae on each side;
segments 4–8 each with comb-like groups of minute spines; all hooks on segments 4–8 each with several minute
spines near its base (Fig. 29F). Cocoon (Fig. 29G). Wall-pocket-shaped, moderately woven, anterior margin thickly
woven, and extended ventrolaterally, without projection; floor woven on posterior two-thirds; individual threads
visible; 3.8 mm long by 3.1 mm wide.
Male and Mature larva. Unknown.
in a vial as [UMSRP: Vietnam 032, Holotype, Simulium (N.) laichauense, Female, coll. Vietnam, 20-XII-2014,
11.0˚C, exposed to the sun, elevation 1,728 m, 22˚22’22.719”N/103˚45’24.852”E) fast-flowing near a forest, Lai
Chau, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D.
Chen & K.W. Lau.
Biological notes. The pupa of this new species was collected from a grass leaf trailing in the current.
Associated species were S. (N.) tayense sp. nov. and S. (S.) giayense sp. nov.
Etymology. The species name laichauense refers to the locality name, Lai Chau, where this new species was
collected.
Remarks. This new species is assigned to the S. vernum species-group of the subgenus Nevermannia, defined
by Crosskey (1969), judging from the female terminalia (Fig. 28E–H) and pupal gill with four long slender
filaments (Fig. 29C). The pupa of this new species is remarkable in having a darkened frons and anterolateral
surface of the thorax (Fig. 29A, B), by which it is distinguished from other members of the S. vernum species-
group.
Simulium (Nevermannia) tayense Takaoka & Ya’cob sp. nov.
Female. Body length 2.8 mm. Head. Slightly narrower than width of thorax. Frons brownish black, gray pruinose,
not shiny, densely covered with whitish-yellow scale-like recumbent short hairs interspersed with few dark longer
hairs along each lateral margin; frontal ratio 1.7:1.0:2.4; frons:head ratio 1.0:4.9. Fronto-ocular area well
developed, narrow, directed dorsolaterally. Clypeus dark brown, gray pruinose, moderately covered with whitish-
yellow hairs interspersed with few dark longer hairs on each side, though upper and lower portions bare. Labrum
0.9 times as long as clypeus. Antenna composed of scape, pedicel and nine flagellomeres, brownish black except
scape and pedicel medium brown. Maxillary palp composed of five segments, light to medium brown except
anterior surface of segments 1 and 2 yellowish and third segment dark brown, proportional lengths of third, fourth,
and fifth segments 1.0:0.8:1.5; third segment (Fig. 30A) swollen; sensory vesicle (Fig. 30A) elongate (0.6–0.7
times as long as third segment), with medium-sized opening. Maxillary lacinia with 13 or 14 inner and 15 or 16
outer teeth. Mandible with 32–34 inner and 14 or 15 outer teeth. Cibarium (Fig. 30B) without any processes.
Thorax. Scutum brownish black except anterolateral calli medium brown, densely covered with yellow scale-like
recumbent hairs interspersed with several dark-brown long upright hairs on prescutellar area. Scutellum medium
brown, moderately covered with yellow short hairs mixed with several dark-brown long upright hairs. Postnotum
dark brown, and bare. Pleural membrane bare. Katepisternum dark brown to brownish black, longer than deep,
whitish-gray pruinose, slightly shiny when illuminated at certain angles, and bare. Legs. Foreleg: coxa and
trochanter light brown; femur dark yellow to light brown except apical cap dark brown; tibia dark brown, with
median large portion on outer surface light brown; tarsus dark brown, with moderate dorsal hair crest; basitarsus
moderately dilated, 8.2 times as long as its greatest width. Midleg: as in foreleg except coxa medium brown on
anterolateral surface and dark brown on posterolateral surface, and trochanter medium brown. Hind leg: coxa and
trochanter medium brown; femur dark yellow to light brown with apical cap medium brown; tibia medium to dark
brown except medial large portion on outer surface light to medium brown; tarsus dark brown except basitarsus
light to medium brown (though base dark brown) and basal half of second tarsomere light brown; basitarsus (Fig.
30C) narrow, nearly parallel-sided, 7.1 times as long as wide, and 0.8 and 0.7 times as wide as greatest widths of
tibia and femur, respectively; calcipala (Fig. 30C) nearly as long as width at base, and 0.5 times as wide as greatest

FIGURE 30. Female of Simulium (Nevermannia) tayense sp. nov. A, Third segment of maxillary palp with sensory vesicle
Sternite 8 and ovipositor valve (left side only; ventral view); F, Genital fork (ventral view); G & H, Paraprocts and cerci (G,
ventral view; H, lateral view). I, Spermatheca. Scale bars. 0.1 mm for C; 0.02 mm for A, B and D–I.
width of basitarsus; pedisulcus (Fig. 30C) shallow; claw (Fig. 30D) with large basal tooth 0.48 times as long as
claw. Wing. Length 3.0 mm. Costa with dark-brown spinules and light-brown hairs though intermixed with
yellowish hairs basally. Subcosta with light-brown hairs except apical one-fourth bare. Hair tuft on base of radius
dark brown. Basal portion of radius fully haired; R1 with dark spinules and hairs; R2 with hairs only. Basal cell
absent. Halter. White except basal stem darkened. Abdomen. Basal scale light brown, with fringe of whitish-
yellow hairs. Dorsal surface of abdomen dark brown except segment 2 light brown, and tergites 7–9 medium
brown, moderately covered with dark short to long hairs intermixed with yellow short hairs; tergites of segments 2
and 7–9 wide and shiny when illuminated at certain angles, tergites 3–6 relatively narrow and dull. Ventral surface
of abdomen mostly ochreous to medium brown except segment 2 whitish; sternal plate on segment 7 developed
medially. Terminalia. Sternite 8 (Fig. 30E) bare medially, with 11 medium-long to long yellow hairs together with
few slender short yellow hairs on each side. Ovipositor valve (Fig. 30E) triangular (though medioposterior corner
rounded), thin, membranous, moderately covered with microsetae interspersed with four short slender yellow hairs;
inner margins slightly concave medially, somewhat sclerotized, and somewhat separated from each other. Genital
fork (Fig. 30F) of usual inverted-Y form, with slender stem; arms of moderate width medially; lateral plate of each
arm strongly sclerotized along dorsolateral margin, and with thin lobe directed medioposteriorly. Paraproct in
ventral view (Fig. 30G) nearly quadrate with triangular projection directed medially, with six sensilla on
unpigmented anteromedial surface; paraproct in lateral view (Fig. 30H) somewhat produced ventrally, 0.7 times as
long as wide, with 12 or 13 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view (Fig.
30H) short, with truncate posterior margin, 0.7 times as long as wide; cercus in end view much curved outward.
Spermatheca (Fig. 30I) ovoidal, 1.2 times as long as its greatest width, well sclerotized except duct and small area
near juncture with duct unsclerotized, and with hexagonal patterns (though not well defined) on surface; internal
setae absent; both accessory ducts slender, subequal in diameter to major one.
Male. Body length 3.2 mm. Head. Wider than thorax. Upper eye medium brown, with large facets in 17
vertical columns and 19 horizontal rows. Face dark brown, whitish-gray pruinose. Clypeus brownish black,
whitish-gray pruinose, moderately covered with golden-yellow medium-long hairs (mostly directed upward).
Antenna composed of scape, pedicel and nine flagellomeres, dark brown except extreme base of first flagellomere
light brown; first flagellomere elongate, 1.9 times as long as second one. Maxillary palp composed of five
segments, light brown except segments 1 and 2 yellow; proportional lengths of third, fourth, and fifth segments
1.0:0.9:1.8; third segment (Fig. 31A) not swollen; sensory vesicle (Fig. 31A) ellipsoidal, small, 0.2 times as long as
third segment, and with small opening. Thorax. Scutum black, densely covered with golden-yellow recumbent
short hairs interspersed with several medium-brown upright long hairs on prescutellar area; other features as in
female. Legs. Color nearly as in female except fore trochanter medium brown except inner surface yellow, hind
trochanter yellowish except anterior surface medium brown, hind femur yellowish on inner surface. Fore basitarsus
slightly dilated, 10.4 times as long as its greatest width. Hind basitarsus (Fig. 31B) enlarged, spindle-shaped, 3.8
times as long as wide, and 1.1 and 1.2 times as wide as greatest widths of hind tibia and femur, respectively;
calcipala (Fig. 31B) slightly shorter than basal width, and 0.3 times as wide as greatest width of basitarsus.
Pedisulcus (Fig. 31B) well developed. Wing. Length 2.9 mm; other features as in female except subcosta with few
hairs. Halter. White except basal stem darkened. Abdomen. Basal scale dark brown, with fringe of dark long hairs.
Dorsal surface of abdomen medium brown except little less than basal half of segment 2 ochreous, moderately
covered with yellow short hairs intermixed with dark brown short to medium-long hairs; segments 7 and 8 each
with pair of slightly shiny lateral patches when illuminated at certain angles; ventral surface of segment 2 white,
those of segments 3–6 ochreous except sternites medium brown, and those of other segments medium brown.
Genitalia. Coxite in ventral view (Fig. 31C) nearly rectangular, twice as long as its greatest width. Style in ventral
view (Fig. 31C) bent inward, nearly parallel-sided, with truncated apex having apical spine; style in ventrolateral
view (Fig. 31D) shorter than coxite (0.8 times as long as coxite), straight up to apical one-third, then curved
inwardly, widest at base, nearly parallel-sided up to apical one-third, then gradually narrowed toward apex and with
rounded apex; style in medial view (Fig. 31E) boot-shaped, with triangular apical lobe directed dorsomedially;
style in caudal view (Fig. 31F) widely depressed on posterodorsal surface of medially-directed apical lobe. Ventral
plate in ventral view (Fig. 31C) with subquadrate body, 0.7 times as long as wide, with anterior margin undulate
with median concavity, both sides nearly straight though posterolateral corners rounded, and posterior margin with
two small and shallow concavities submedially; body of ventral plate darkened along anterior margin, and nearly

FIGURE 31. Male of Simulium (Nevermannia) tayense sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
side; front view); B, Hind basitarsus and second tarsomere (left side; outer view); C, Coxites, styles and ventral plate (ventral
view); D–F, Styles (right side; D, ventrolateral view; E, medial view; F, caudal view); G, Ventral plate and median sclerite
(lateral view); H, Ventral plate (caudal view); I, Median sclerite (caudal view); J, Paramere (left side; dorsal view) ; K,
Aedeagal membrane and dorsal plate (caudal view); L & M, Abdominal segment 10 and cerci (right side; L, lateral view; M,
caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–M.
bare except posteromedian portion of ventral surface densely covered with microsetae; basal arms of moderate
length, well sclerotized, divergent from base as shown in Fig. 31C; ventral plate in lateral view (Fig. 31G) with
body slightly narrowed posteriorly; ventral plate in caudal view (Fig. 31H) with body appearing as shallow
inverted-V shape, having similar width, posteroventral margin roughly undulate, and densely covered with
microsetae medially on posterior surface. Median sclerite (Fig. 31C, G, I) club-shaped, narrow, with forked apex,
and with base located in front of anterior margin of ventral plate. Parameres (Fig. 31J) large, each with small apical
appendix bearing one distinct long and stout hook and one small short hook at its base. Aedeagal membrane (Fig.
31K) moderately setose, dorsal plate well defined, broadly produced ventrally with round apex. Abdominal
segment 10 (Fig. 31L, M) with one distinct hair on ventral surface and one distinct hair on lateral surface near each
posterolateral margin. Cercus (Fig. 31L, M) small, rounded, with 14 hairs.
Pupa. Body length 2.8–3.3 mm. Head. Integument light ochreous, bare; antennal sheath without any
protuberances; frons with three pairs of unbranched long trichomes with coiled apices (Fig. 32A), and face with
pair of unbranched long trichomes with coiled apices (Fig. 32B). Thorax. Integument light ochreous, bare except
dorsal and dorsolateral surfaces of posterior half sparsely covered with small round tubercles (though small area
near posterior margin almost bare); thorax with three long dorsomedial trichomes with coiled or straight apices
(anterior and middle ones (Fig. 32C) slightly longer than posterior one), two long anterolateral trichomes with
coiled apices (anterior trichome somewhat shorter than posterior one) (Fig. 32D), one medium-long with coiled or
straight apex (Fig. 32E), and three ventrolateral trichomes with straight apices (one medium-long, two short) (Fig.
32F), on each side; all trichomes unbranched. Gill (Fig. 32G) composed of four slender thread-like filaments,
arranged in pairs, with medium-long common basal stalk having somewhat swollen transparent basal fenestra
ventrally at base; common basal stalk 0.7–1.4 times length of interspiracular trunk; stalk of dorsal pair slightly
shorter and thicker than that of ventral pair, and 0.2–0.6 times length of common basal stalk; stalk of ventral pair
0.3–0.7 times length of common basal stalk; stalk of dorsal pair 1.1–1.3 and 0.7–0.8 times as thick as stalk of
ventral pair and common basal stalk, respectively; all filaments slightly different in length and thickness from one
another, though filaments of ventral pair subequal in length and thickness; upper filament of dorsal pair longest,
with its length including its own stalk and common basal stalk 4.0–4.8 mm, ventral filament of dorsal pair second
longest with its length 3.3–4.5 mm, two filaments of ventral pair shortest, with their length 3.0–4.3 mm; relative
thickness of filaments from dorsal to ventral when their basal portion measured 1.0:0.8:0.7:0.7; cuticle of all
filaments grayish brown except common basal stalk much lighter, with well-defined annular ridges and furrows,
and densely covered with minute tubercles. Abdomen. Dorsally, segment 1 entirely dark grayish, segment 2 dark
grayish on anterior three-fourths, segment 3 dark grayish on basal half and segment 4 dark grayish on anterior one-
third, and segments 1 and 2 weakly tuberculate; segment 1 with one long slender hair-like seta and one to three
minute setae submedially on each side (Fig. 32H); segment 2 with one short slender hair-like seta and five short
somewhat spinous setae submedially along posterior margin (Fig. 32I) and one minute seta (similar to those on
segment 1) submedially near anterior margin on each side; segments 3 and 4 each with four hooked spines and one
short somewhat spinous seta along posterior margin and one minute seta submedially near anterior margin on each
side; segments 5–8 each with spine-combs in transverse row and comb-like groups of minute spines along anterior
margin and two short setae near posterior margin (except segment 5 with four or five setae) on each side; all setae
unbranched; segment 9 with pair of cone-like terminal hooks (Fig. 32J) and comb-like groups of minute spines.
Ventrally, segment 4 with one short seta and few minute setae on each side; segment 5 with pair of bifid hooks
submedially and two short setae on each side; segments 6 and 7 each with pair of bifid inner and unbranched outer
hooks somewhat spaced from each other and one short setae on each side; segments 3–8 each with comb-like
groups of minute spines. Each side of segment 9 without grapnel-shaped hooklets. Cocoon (Fig. 32K). Wall-
pocket-shaped, light yellowish, thinly woven except anterior margin thickly woven, widely extended
ventrolaterally, appearing round when viewed dorsally; anterior margin with medium to long dorsal projection
(1.0–1.2 mm long) usually slightly curved downward; posterior half with floor roughly or moderately woven;
individual threads partially visible; 3.4–4.0 mm long by 3.0–4.6 mm wide.
Mature larva. Body length 6.4 mm. Body light ochreous. Head. Cephalic apotome whitish yellow, though
narrow area along posterior margin darkened, and moderately covered with unpigmented short setae; head spots
moderately positive except anterior spot of mediolongitudinal spots and anterior spots of mediolateral spots in one
larva and anterior spots of posterolateral spots in two larvae much lighter. Lateral surface of head capsule whitish
yellow except eye-spot region whitish; eyebrow distinct; two relatively large spots and two or three smaller spots in

FIGURE 32. Pupa of Simulium (Nevermannia) tayense sp. nov. A, Frontal trichome; B, Facial trichome; C–F, Thoracic
trichomes (C, anterodorsal; D, anterolateral; E, mediolateral; F, ventrolateral); G, Anterior part of thorax and gill filaments
(right side; outer view); H, Hair-like seta and minute seta on dorsal surface of abdominal segment 1; I, Hair-like seta and
minute seta on dorsal surface of abdominal segment 2; J, Terminal hooks (caudal view); K, Cocoon (dorsal view). Scale bars.
1.0 mm for K; 0.1 mm for G; 0.02 mm for A–F and H–J.
FIGURE 33. Larva of Simulium (Nevermannia) tayense sp. nov. A, Mandible; B, Hypostoma; C, Head capsule (ventral view);
D, Accessory sclerite. Scale bars. 0.1 mm for C; 0.04 mm for B; 0.02 mm for A and D.
front of posterior margin moderately positive; one small round spot below eye-spot region faintly positive. Ventral
surface of head capsule whitish yellow to yellow except darkened area near posterior margin on each side of
postgenal cleft; horizontal elongate spot and round spot on each side of postgenal cleft moderately positive.
Antenna composed of three articles and apical sensillum, somewhat longer than stem of labral fan; proportional
lengths of first, second, and third articles 1.0:1.0:0.7. Labral fan with 35 primary rays. Mandible (Fig. 33A) with
three comb-teeth, of which first is longest and second is as long as or little shorter than third; mandibular serration
composed of two teeth (one medium-sized and one small); major tooth at obtuse angle against mandible on apical
side; supernumerary serrations present though consisting of one minute slender tooth or two). Hypostoma (Fig.
33B) with row of nine apical teeth; median and each corner tooth prominent, subequal in length to one another and
much longer than three intermediate teeth on each side; lateral margin serrate anterirorly; four or five hypostomal
bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig. 33C) short, rounded apically, 0.94 times
length of postgenal bridge. Cervical sclerite composed of two medium-brown small oblong pieces, not fused to
occiput, widely separated medially from each other. Thorax and Abdomen. Thoracic cuticle bare. Abdominal
cuticle almost bare except few posterior segments sparsely or moderately covered with unpigmented short setae,
and last segment moderately covered with unpigmented short setae on each side of anal sclerite including ventral
papillae. Rectal scales present. Rectal organ compound, with finger-like secondary lobules (number of secondary
lobules uncountable due to retracted rectal organ). Anal sclerite of usual X-form, with anterior arms 0.9 times as
long as posterior ones, broadly sclerotized at base; accessory sclerite (Fig. 33D) composed of four to six sclerotized
spots on each side. Last abdominal segment with pair of large conical ventral papillae. Posterior circlet with 78–85
rows of up to 13–15 hooklets per row.
a vial as [UMSRP: Vietnam 033, Holotype, Simulium (N.) tayense, Male, coll. Vietnam, 20-XII-2014, Takaoka et
al.], reared a from pupa, collected from a stream (width 50 cm, depth 10 cm, bottom rocky, water temperature
11.0˚C, exposed to the sun, elevation 1,750 m, 22˚21’43.110”N/103˚47’19.221”E) slowly flowing from a
secondary forest, Sapa, Lao Cai, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob,
C.D. Chen & K.W. Lau. PARATYPES: One pupal exuviae and one mature larva, same data and date as those of the
holotype in 80% ethanol, labeled in each vial as [UMSRP: Vietnam 033, Paratype, Simulium (N.) tayense, coll.
Vietnam, 20-XII-2014, Takaoka et al.]; two females (with their associated pupal exuviae and cocoons) in 80%
ethanol, labeled in each vial as [UMSRP: Vietnam 033, Paratype, Simulium (N.) tayense, Female, coll. Vietnam,
20-XII-2014, Takaoka et al.], collected from a river (width 6 m, depth 60 cm, bottom rocky, exposed to the sun,
elevation 1,728 m, 22˚22’22.719”N/103˚45’24.852” E) fast-flowing in a forest, Sapa, Lao Cai, northern Vietnam,
20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
Biological notes. The pupae and larvae of this new species were collected from plastic sheets and dead grass
leaves in the current. Associated species were S. (M.) nigrofilum sp. nov., S. (N.) laichauense sp. nov., S. (S.)
laocaiense sp. nov., S. (S.) rosliramlii sp. nov., S. (S.) sapaense sp. nov. and S. (S.) turgidum sp. nov.

Etymology. The species name tayense refers to the land in which the Tay tribe lives.
Remarks. Simulium (N.) tayense sp. nov. is placed in the S. vernum species-group of the subgenus
Nevermannia, based on the shape of the male genitalia (Fig. 31C–M). This new species is characterized by the
presence of an accessory sclerite on each side of the last larval abdominal segment (Fig. 33D). Beside this new
species, the following seven species are known to bear accessory sclerites among species of the S. vernum species-
group: S. (N.) aberrans Delfinado from the Philippines (Takaoka 1983), S. (N.) caudisclerum Takaoka & Davies
from Peninsular Malaysia (Takaoka and Davies 1995), S. (N.) chomthongense Takaoka, Srisuka & Choochote and
S. (N.) khunklangense Takaoka & Srisuka, both from Thailand (Takaoka et al. 2012c, 2013), S. (N.) ludingense
Chen, Zhang & Huang from Sichuan, China (Chen et al. 2005), S. (N.) yushangense Takaoka from Taiwan
(Takaoka 1979), and S. (N.) zhangjiajiense Chen, Zhang & Bi from Hunan, China (Chen et al. 2004) . However,
this new species is easily distinguished from these seven species by the following characteristics: from S. (N.)
aberrans, S. (N.) ludingense and S. (N.) khunklangense by the cocoon with an elongate anterodorsal projection (the
cocoon is simple, without a projection in the former two species and the cocoon has a short bulge in S. (N.)
khunklangense), and from four other species by the greater number (19) of horizontal rows of male upper-eye
facets (13 in S. (N.) zhangjiajiense, 15 in S. (N.) caudisclerum, 17 in S. (N.) yushangense, 17 or 18 in S. (N.)
chomthongense). The number (19) of horizontal rows in the male upper-eye facets of this new species is close to
that (17 or 18) of S. (N.) chomthongense but both species are separated from each other by the ratio of the postgenal
cleft against the postgenal bridge (0.9 in this new species versus 0.7–0.8 in S. (N.) chomthongense).
IV. Subgenus Simulium Latreille
Thirty-seven species are treated under the subgenus Simulium, which are further placed in nine species-groups (11
in the S. tuberosum species-group, eight in the S. striatum species-group, seven in the S. griseifrons species-group,
five in the S. multistriatum species-group, two in the S. variegatum species-group, and one each in the S.
argentipes, S. christophersi, S. malyschevi and S. nobile species-groups). The S. argentipes and S. malyschevi
species-groups are newly recorded from Vietnam. The definition of these species groups follows Rubtsov (1956),
Takaoka and Davies (1996), Takaoka and Saito (2007) and Takaoka et al. (2014e).
Keys to 37 species of the subgenus Simulium
Females*
1. Claws simple, without basal or subbasal tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Claws with subbasal tooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2. Basal section of radial vein haired . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Basal section of radial vein bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
3. Hind tibia darkened except base whitish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . S. chiangmaiense, S. jeffreyi, S. nakhonense, S. quinquestriatum, S. tavanense sp. nov. & S. taythienense
Hind tibia white on basal two-thirds or more and dark on rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Scutum without longitudinal vittae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. vietnamense
Scutum with five longitudinal vittae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Ovipositor valve tapered posteromedially. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chungi
Ovipositor valve rounded posteromedially . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lacduongense & S. hirtinervis
6. Scutum with five longitudinal vittae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Scutum without longitudinal vittae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7. Ovipositor valve with elongate projection on inner surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. grossifilum
Ovipositor valve without projection on inner surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Inner margins of ovipositor valves nearly parallel-sided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maenoi
Inner margins of ovipositor valves divergent posteriorly (Fig. 48D) . . . . . . . . . . . . . . . . . . . . . . . . S. daoense sp. nov. & S. laui
9. Abdominal segment 7 with pair of hair tuft ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Abdominal segment 7 without hair tuft ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
10. Sensory vesicle with large opening (Fig. 56A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. fuscicoxae sp. nov.
Sensory vesicle with opening of moderate size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Antenna darkened except scape, pedicel and base of first flagellomere yellow . . . . . . . . . . . . . . . . . . . S. doipuiense (complex)
Antenna almost entirely darkened. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Labrum 0.6–0.7 times length of clypeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. giayense sp. nov.
Labrum 0.7–0.8 times length of clypeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rosliramlii sp. nov.
13. Ovipositor valves with inner margins widely concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. nodosum
Ovipositor valves with inner margins nearly straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
14. Hind tibia medium to dark brown except base yellow . . . . . . . . . . . . .S. suoivangense sp. nov., S. tani (complex) & S. xuandei
Hind tibia yellow except apical cap medium brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Frons:head ratio 1.0:3.6. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. laocaiense sp. nov.
Frons:head ratio 1.0:4.3. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. turgidum sp. nov.
16. Frons densely covered with yellow short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. nigrogilvum
Frons sparsely covered with dark hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
17. Subcosta bare or rarely with one or two hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. obliquum sp. nov.
Subcosta with many hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
18. Ovipositor valves with inner margins nearly straight or slightly sinuous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. atipornae
Ovipositor valves with inner margins widely concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
19. Ovipositor valves rounded posteromedially (Fig. 34E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sansahoense sp. nov.
Ovipositor valves pointed posteromedially (Fig. 69G). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20. Fore basitarsus greatly dilated, 5.6–5.8 times as long as its greatest width . . . . . . . . . . . . . . . . . . . . . . . . . S. phuluense sp. nov.
Fore basitarsus greatly dilated, 5.1 times as long as its greatest width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. chamlongi
Males**
1. Scutum white pruinose with inverted-V-shaped, non-pruinose black area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. nodosum
Scutum white pruinose on each shoulder, along each lateral margin and on prescutellar area . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Basal portion of radial vein with four or five hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tavanense sp. nov.
Basal portion of radial vein bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Fore basitarsus with thick hair crest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigrogilvum
Fore basitarsus with moderate hair crest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Style without terminal spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chungi
Style with terminal spine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Ventral plate without teeth posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Ventral plate with teeth posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
6. Style with long horn-like protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. grossifilum
Style with short basal protuberance. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Ventral plate rectangular, longer than wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maenoi
Ventral plate saddle-shaped (Fig. 53C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Scutum with yellowish short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Scutum with brassy or copper-colored short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9. Upper-eye facets in 16 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. quinquestriatum
Upper-eye facets in 20 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. xuandai
10. Upper-eye facets in 17 or 18 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. taythienense
Upper-eye facets in 16 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. chiangmaiense, S. nakhonense & S. thailandicum
11. Style with numerous short spines on dorsal surface of basal portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Style without short spines on dorsal surface of basal portion or with one or two on tip of long basal protuberance . . . . . . . . 18
12. Abdominal segments 2, 6 and 7 each with pair of shiny dorsolateral patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Abdominal segments 2, 5, 6 and 7 each with pair of shiny dorsolateral patches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
13. Fore coxa darkened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
Fore coxa yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
14. Scutum with yellow, short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. giayense sp. nov.
Scutum with brassy, short hairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. rosliramlii sp. nov.
15. Upper-eye facets in 21 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. huense
Upper-eye facets in 16–19 horozontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
16. Upper-eye facets in 16 or 17 horizontal rows; fore basitarsus 5.9–6.1 times as long as its greatest width . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. doipuiense (complex)
Upper-eye facets in 18 or 19 horizontal rows; fore basitarsus 4.5–5.2 times as long as its greatest width . . . . . . . . . . . . . . . .17
17. Fore basitarsus 4.5 times as long as its greatest width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lowi sp. nov.
Fore basitarsus 5.2 times as long as its greatest width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. sapaense sp. nov.
18. Upper-eye facets in 20 vertical columns and 21 horizontal rows; fore coxa darkened . . . . . . . . . . . . . . . . . . . . . . . . . . .S. congi
Upper-eye facets in 18 vertical columns and 18 or 19 horizontal rows; fore coxa yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. suoivangense sp. nov., S. tani (complex) & S. xuandei
19. Style without any basal protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Style with horn-like or round basal protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
20. Scutum with dark short hairs; hind basitarsus slender, nearly parallel-sided (Fig. 45D) . . . . . . . . . . . . . . . S. obliquum sp. nov.
Scutum with yellow short hairs; hind basitarsus enlarged, spindle-shaped (Fig. 35B). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chamlongi, S. phuluense sp. nov. & S. sansahoense sp. nov.
21. Style with round, short basal protuberance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. atipornae
Style with elongate, horn-like basal protuberance with pointed apex (Fig. 49E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
22. Upper-eye facets in 14 or 15 vertical columns and 15 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. hirtinervis

Upper-eye facets in 17–21 vertical columns and 17–21 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
23. Upper-eye facets in 20 or 21 vertical columns and 21 horizontal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. laui
Upper-eye facets in 17–19 vertical columns and 17–20 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24. Upper-eye facets in 20 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lacduongense
Upper-eye facets in 17–19 horizontal rows. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
25. Hind basitarsus whitish yellow on basal half or little less and dark on rest . . . . . . . . . . . . . . . . . . . . . . . . . . S. daoense sp. nov.
Hind basitarsus whitish yellow on basal two-fifths and dark on rest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. malayense
Pupae***
1. Gill with three filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nodosum
Gill with six, eight or 10 filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Gill with six filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Gill with eight or 10 filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
3. Cocoon shoe-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigrogilvum
Cocoon wall-pocket-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Cocoon with anterolateral window(s) on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Cocoon without anterolateral window on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
5. Gill with inflated filaments (Fig. 42H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
Gill with thread-like filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
6. Gill filaments pale, divergent, with minute fine projections on their surface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. grossifilum
Gill filaments dark brown, not divergent, without minute projection on their surface . . . . . . . . . . . . . . . . . .S. turgidum sp. nov.
7 Cocoon with small anterolateral window on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. maenoi
Cocoon with large anterolateral window on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Terminal hook cone-shaped. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chungi
Terminal hook absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Abdominal segment 9 with spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. obliquum sp. nov.
Abdominal segment 9 without spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..S. atipornae
10. Cocoon with short broad anterodorsal projection (Fig. 39L) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. laochauense sp. nov.
Cocoon without anterodorsal projection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Thoracic integument with pit-like organ at base of gill (Fig. 64D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Thoracic integument without pit-like organ at base of gill . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
12. Tubercles present near pit-like organ (Fig. 64D) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. sapaense sp. nov.
Tubercles absent near pit-like organ (Fig. 65G) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
13. Relatively large tubercles on frons with minute secondary projections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lowi sp. nov.
Relatively large tubercles on frons without minute secondary projections. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. huense
14. Basal fenestra of gill inflated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. congi
Basal fenestra of gill not inflated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15. Frons bare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Frons covered with tubercles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
16. Stalks of dorsal and ventral pairs of gill filaments at about right angle when viewed laterally (Fig. 60F) . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. rosliramlii sp. nov.
Stalks of dorsal and ventral pairs of gill filaments at angle of about 120 degrees when viewed laterally . . . . . . . . S. chamlongi
17. Gill filaments directed forward in bundle (Fig. 68A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Gill filaments lying in vertical plane, and stalks of dorsal and ventral pairs of gill filaments lying at angle of 60–120 degrees
when viewed laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18. Terminal hooks present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tani (complex)
Terminal hooks absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. suoivangense sp. nov.
19. Abdominal segment 9 without spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. phuluense sp. nov.
Abdominal segment 9 with spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
20. Frons densely covered with tubercles (Fig. 61B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. giayense sp. nov.
Frons moderately covered with tubercles (Fig. 57A) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
21. Terminal hooks present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. xuandei
Terminal hooks absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
22. Abdominal segment 7 with spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. doipuiense (complex)
Abdominal segment 7 without spine-combs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. fuscicoxae sp. nov.
23. Gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Gill with 10 filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
24. Cocoon shoe-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25
Cocoon wall-pocket-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26
25. All filaments of same thickness (Fig. 36G). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. sansahoense sp. nov.
Ventral two filaments more slender than other filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chiangmaiense
26. Cocoon without anterolateral window on each side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. malayense
Cocoon with anterolateral window on each side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
27. Stalks of dorsal and ventral pairs of gill filaments lying at angle of about 120 degrees when viewed laterally . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. lacduongense & S. laui
Stalks of dorsal and ventral pairs of gill filaments lying at angle of about 90 degrees when viewed laterally (Fig. 50G) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. daoense sp. nov. & S. hirtinervis
28. Ten gill filaments arranged as 2+3+3+2 from dorsal to ventral (Fig. 54H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29
Ten gill filaments arranged as 2+2+2+2+2 or 2+1+2+1+2+2 from dorsal to ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
29. Frons with relatively large tubercles without secondary projections . . . . . . . . . . . . . . . . . S. quinquestriatum & S. taythienense
Frons with relatively large tubercles having minute secondary projections (Fig. 54A) . . . . . S. tavanense sp. nov. & S. xuandai
30. Ten gill filaments arranged as 2+1+2+1+2+2 from dorsal to ventral. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. jeffreyi
Ten gill filaments arranged as 2+2+2+2+2 from dorsal to ventral.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31. Frons covered with tubercles of various shapes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. thailandicum
Frons covered with round tubercles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nakhonense
Mature larvae****
1. Accessory sclerite present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. nigrogilvum
Accessory sclerite absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Abdominal segments1–4 each with six pairs of protuberances (Fig. 47F) . . . . . . . . . . . . . . . . . . . . . . . . . . S. obliquum sp. nov.
Abdominal segments 1–4 each with or without paired dorsal protuberances. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Abdomen with paired dorsal protuberances (Fig. 55E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Abdomen without paired dorsal protuberances. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
4. Postgenal cleft deep, reaching posterior border of hypostoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. nodosum
Postgenal cleft deep but not reaching posterior border of hypostoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Histoblast of pupal gill with eight filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. hirtinervis
Histoblast of pupal gill with 10 filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6. Histoblast of pupal gill with 10 filaments arranged as 2+3+3+2 from dorsal to ventral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. quinquestriatum, S. taythienense & S. tavanense sp. nov.
Histoblast of pupal gill with 10 filaments arranged as 2+2+2+2+2 or 2+1+2+1+2+2 from dorsal to ventral . . . . . . . . . . . . . . 7
7. Histoblast of pupal gill with 10 filaments arranged as 2+1+2+1+2+2 from dorsal to ventral . . . . . . . . . . . . . . . . . . . . S. jeffreyi
Histoblast of pupal gill with 10 filaments arranged as 2+2+2+2+2 from dorsal to ventral . . . . . . . . . . . . . . . . . . S. nakhonense
8. Abdominal segments 1–4 each with transverse dark band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Abdominal segments 1–4 each without a dark band. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
9. Postgenal cleft rounded laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. tani (complex) & S. suoivangense sp. nov.
Postgenal cleft angulated laterally. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. xuandei
10. Postgenal cleft triangular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
Postgenal cleft spear-head-shaped or mitre-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
11. Pharate gill with enlarged basal fenestra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. congi
Pharate gill without small basal fenestra. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Body length 5.8–6.6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. atipornae
Body length 5.5 mm or shorter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. huense
13. Postgenal cleft longer than wide (Fig. 37B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. chamlongi & S. sansahoense sp. nov.
Postgenal cleft nearly as long as wide (Fig. 51B) . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. daoense sp. nov., S. laui & S. malayense
* The females of S. huense and S. congi remain unknown.
** The male of S. vietnamense remains unknown.
*** The pupa of S. vietnamense remains unknown.
**** The larvae of S. chiangmaiense, S. lacduongense, S. thailandicum, S. fuscicoxae sp. nov., S. lowi sp. nov. S. rosliramlii sp.
nov., S. giayense sp. nov., S. doipuiense (complex) and S. vietnamense remain unknown.
1) Simulium argentipes species-group
Simulium (Simulium) sansahoense Takaoka & Chen sp. nov.
Female. Body length 2.5–2.6 mm. Head. Slightly narrower than thorax. Frons black, shiny, with bluish reflection
when illuminated at certain angles, with several dark stout hairs along lateral margins and few to several yellow
fine hairs near antennal bases; frontal ratio 1.3:1.0:0.9–1.1; frons:head ratio 1.0:3.3–3.5. Fronto-ocular area well
developed, short, directed laterally, and rounded or pointed apically. Clypeus black, shiny and white pruinose when
illuminated at certain angles, moderately covered with dark-brown medium-long hairs mixed with yellow fine hairs
(though mediolongitudinal portion somewhat widely bare on upper half to three-fifths). Labrum 0.6 times length of
clypeus. Antenna composed of scape, pedicel and nine flagellomeres, dark brown, except scape, pedicel and first
flagellomere yellow when viewed ventrally, but dark brown except scape, pedicel and basal one-third of first
flagellomere yellow when viewed dorsally. Maxillary palp with five segments, medium brown except third
segment dark brown; proportional lengths of third, fourth, and fifth segments 1.0:1.2:2.7; third segment (Fig. 34A)

FIGURE 34. Female of Simulium (Simulium) sansahoense sp. nov. A, Third segment of maxillary palp with sensory vesicle
Sternite 8 and ovipositor valve (right side only; ventral view); F, Genital fork (ventral view); G & H, Paraprocts and cerci (G,
ventral view; H, lateral view). I, Spermatheca. Scale bars. 0.1 mm for C; 0.02 mm for A, B and E–I; 0.01 mm for D.
of moderate size, with medium-sized ellipsoidal sensory vesicle (0.3–0.4 times length of third segment) having
medium-sized opening. Maxillary lacinia with 11 or 12 inner and 13–15 outer teeth. Mandible with 29 inner and 12
outer teeth. Cibarium (Fig. 34B) with 30 or 31 minute processes near posterodorsal margin. Thorax. Scutum black,
shiny and white pruinose with three faint dark vittae (one median, two submedian) when illuminated at certain
angles, moderately covered with yellow short fine hairs interspersed with several dark-brown long upright hairs on
prescutellar area. Scutellum brownish black, covered with dark-brown upright long and yellow short hairs.
Postnotum brownish black, shiny and white pruinose when illuminated at certain angles, and bare. Pleural
membrane bare. Katepisternum longer than deep, brownish black, shiny and white pruinose when illuminated at
certain angles, and bare. Legs. Foreleg: coxa and trochanter whitish yellow; femur yellow with apical cap light
brown; tibia whitish yellow except apical cap dark brown, with median outer surface widely white and with bright
white sheen when illuminated at certain angles; tarsus black, with moderate dorsal hair crest; basitarsus greatly
dilated, 5.8–5.9 times as long as its greatest width. Midleg: coxa dark brown; trochanter whitish yellow with apical
portion partially light brown; femur yellow except apical cap light brown; tibia whitish yellow except apical tip
light brown, and with whitish sheen widely on posterior surface when illuminated at certain angles; tarsus medium
brown except basal four-fifths of basitarsus whitish yellow. Hind leg: coxa medium to dark brown; trochanter
yellowish white; femur yellow except apical cap light brown; tibia white on posterior surface of basal half and
yellow on rest except apical cap medium brown, and with whitish sheen widely on posterior surface when
illuminated at certain angles; tarsus medium brown except little more than basal two-thirds of basitarsus (though
base light brown) and basal half of second tarsomere yellowish white; basitarsus (Fig. 34C) nearly parallel-sided,
6.2–6.3 times as long as wide, and 0.8 and 0.7 times as wide as greatest widths of hind tibia and femur,
respectively; calcipala (Fig. 34C) moderately developed, nearly as long as wide, and 0.4 times as wide as greatest
width of basitarsus; pedisulcus (Fig. 34C) well developed; claw (Fig. 34D) with small subbasal tooth. Wing.
Length 2.5–2.7 mm. Costa with dark spinules and hairs; subcosta haired except apical one-fifth to half bare; basal
section of radial vein bare; R1 with dark-brown spinules and hairs; R2 with dark-brown hairs; hair tuft on base of
radius dark brown; basal cell absent. Halter. White except base darkened. Abdomen. Basal scale dark brown, with
fringe of yellow hairs. Dorsal surface of abdomen medium brown to brownish black, with dark short hairs; tergite 2
shiny and silvery iridescent when illuminated at certain angles and tergites 6–9 shiny when illuminated at certain
angles. Ventral surface light to medium brown; segment 7 without sternal plate. Terminalia. Sternite 8 (Fig. 34E)
well sclerotized and bare medially, covered with 6–10 long and medium-long stout hairs and three or four short fine
hairs on each side. Ovipositor valves (Fig. 34E) nearly tongue-like, rounded posteromedially, membranous, each
densely covered with microsetae together with five to eight short to medium-long hairs; inner margin deeply
concave medially. Genital fork (Fig. 34F) of inverted-Y form; stem slender and well sclerotized; arms of moderate
width, each with strongly sclerotized apical portion having distinct projection directed anterodorsally. Paraproct in
ventral view (Fig. 34G) nearly quadrate, with anteromedian portion depressed and moderately sclerotized having
eight or nine sensilla on its surface; paraproct in lateral view (Fig. 34H) somewhat produced ventrally, 0.7 times as
long as wide, covered with 15–19 short to medium-long hairs and numerous microsetae on lateral and ventral
surfaces. Cercus in lateral view (Fig. 34H) short, rounded posteriorly, 0.56 times as long as its greatest width, and
covered with medium-long and short hairs. Spermatheca (Fig. 34I) ovoidal, 1.2 times as long as wide, sclerotized
except duct and narrow area of juncture with duct unsclerotized, without discernible reticulate surface patterns;
minute internal setae present; accessory ducts subequal in thickness to each other and slightly thicker than main
duct.
Male. Body length 2.6–3.0 mm. Head. Slightly wider than thorax. Upper eye medium brown, with large facets
in 21 or 22 vertical columns and in 21–23 horizontal rows. Clypeus black, thickly white pruinose, silvery or bluish,
shiny when illuminated at certain angles, covered with dark-brown hairs along lateral margins and near ventral
margin (medial portion widely bare). Antenna composed of scape, pedicel and nine flagellomeres, medium brown
to brownish black except base of first flagellomere dark yellow; first flagellomere elongate, 1.7 times as long as
second one. Maxillary palp grayish to dark brown, composed of five segments with proportional lengths of third,
fourth, and fifth segments 1.0:1.3:2.9; third segment (Fig. 35A) of moderate size; sensory vesicle (Fig. 35A) small,
ellipsoidal (0.3 times as long as third segment), and with small opening. Thorax. Scutum black, with white
pruinose pattern, i.e., anterior pair of crescent spots on shoulders extending posteriorly along lateral margins
connected near base of wings to large transverse posterior spot entirely covering prescutellar area; these pruinose

FIGURE 35. Male of Simulium (Simulium) sansahoense sp. nov. A, Third segment of maxillary palp with sensory vesicle (left
view); D, coxite (right side; ventrolateral view); E & F, Styles (right side; E, ventrolateral view; F, medial view); G, Ventral
plate and median sclerite (lateral view); H, Ventral plate (caudal view); I, Median sclerite (caudal view); J, Paramere (right side;
caudal view) ; K, Dorsal plate (caudal view); L & M, Abdominal segment 10 and cerci (right side; L, lateral view; M, caudal
view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–M.
areas silvery or bluish iridescent when illuminated at certain angles; scutum uniformly and moderately covered
with yellow recumbent short hairs interspersed with dark-brown long upright hairs on prescutellar area. Scutellum
brownish black, with several dark-brown long upright hairs and yellow short hairs. Postnotum black, white
pruinose when illuminated at certain angles and bare. Pleural membrane bare. Katepisternum longer than deep,
black, white pruinose when illuminated at certain angles, and bare. Legs. Foreleg: coxa whitish yellow; trochanter
light brown except base whitish yellow; femur light brown with apical cap medium brown, though inner surface of
basal half yellowish; tibia medium brown except median large portion widely white on outer surface, and with
bright white sheen widely on outer surface when illuminated at certain angles; tarsus brownish-black to black, with
moderate dorsal hair crest; basitarsus greatly dilated, 6.6–6.8 times as long as its greatest width. Midleg: coxa
brownish black; trochanter medium brown except base yellowish white; femur light brown with apical cap medium
brown; tibia light brown except basal one-third yellow and apical cap medium brown; tibia with white sheen on
posterior surface of basal one-third when illuminated at certain angles; tarsus medium to dark brown except basal
half or little more of basitarsus whitish yellow. Hind leg: coxa dark brown; trochanter whitish yellow; femur
medium brown except base whitish yellow and apical cap dark brown; tibia medium brown except apical cap
brownish black and basal tip yellowish white; tarsus (Fig. 35B) medium to dark brown except basal half or little
more of basitarsus whitish yellow (though extreme base darkened) and basal half of second tarsomere yellowish
white; basitarsus (Fig. 35B) much enlarged, gradually widened from base to apical one-third, then slightly
narrowed to apex, 4.1–4.9 times as long as its greatest width, and 0.7–0.8 and 0.9 times as wide as greatest widths
of hind tibia and femur, respectively; calcipala (Fig. 35B) small, slightly shorter than wide, 0.3 times as wide as
greatest width of basitarsus; pedisulcus (Fig. 35B) well developed. Wing. Length 2.6–2.7 mm. Other features as in
female except subcosta bare. Halter. White except basal portion darkened. Abdomen. Basal scale brownish black,
with fringe of dark long hairs. Dorsal surface of abdomen dark brown to brownish black, with dark short hairs;
segments 2, 5, 6 and 7 each with pair of silvery or bluish iridescent spots dorsolaterally, those on segment 2 broadly
connected in middle to each other. Genitalia. Coxites, styles and ventral plate in ventral view as in Fig. 35C. Coxite
in ventrolateral view (Fig. 35D) 0.8 times as long as wide. Style in ventrolateral view (Fig. 35E) elongate, 2.9 times
as long as its greatest width near base, nearly parallel-sided from base to little more than basal one-third, then
tapered toward middle, and nearly parallel-sided or slightly widened to apex, and with distinct apical spine; style in
medial view (Fig. 35F) 1.4 times as long as coxite, widened from base to basal one-fourth, tapered to little less than
apical one-third, then nearly parallel-sided to apex. Ventral plate in ventral view (Fig. 35C) well sclerotized, with
narrow body having ventrally produced process posteriorly, which bears toothed posterolateral margins and is
covered with many minute setae on anterolateral surface; arms divergent outward, then convergent apically; ventral
plate in lateral view (Fig. 35G) with posterior margin serrated from base to apical tip and with anterolateral
surfaces covered with many microsetae; ventral plate in end view (Fig. 35H) elliptical, with both lateral margins
gently rounded, and each with several teeth. Median sclerite in lateral view (Fig. 35G) arising from anterior margin
of body of ventral plate, directed dorsally; median sclerite in caudal view (Fig. 35I) plate-like, nearly parallel-sided
to apex, with round apex, brown basally, but not so well sclerotized apically. Paramere in caudal view (Fig. 35J)
with enlarged basal portion and with several distinct hooks and several smaller ones. Aedeagal membrane densely
covered with minute setae, and with moderately sclerotized dorsal plate in form of horizontal bar (Fig. 35K).
Abdominal segment 10 (Fig. 35L, M) with three or five hairs on ventral surface and three to five hairs on lateral
surface on each side. Cercus (Fig. 35L, M) small, rounded, with 10 or 11 distinct hairs.
Pupa. Body length 3.0–3.2 mm. Head. Integument ochreous except ventral and lateral surfaces whitish yellow
and antennal sheaths yellow, densely covered with small round tubercles; frons with two pairs of unbranched short
slender trichomes (Fig. 36A), arising close together; face with pair of unbranched medium-long somewhat stout
trichomes (Fig. 36B). Thorax. Integument light brown except wing sheaths yellow, moderately covered with small
round tubercles; thorax with three medium-long stout anterodorsal trichomes (one posterior trichome somewhat
narrower than two anterior ones) (Fig. 36C), two medium-long stout anterolateral trichomes (anterior trichome
shorter and more slender than posterior one) (Fig. 36D), one medium-long stout mediolateral trichome (Fig. 36E),
and three ventrolateral trichomes (one anterior trichome short and slender, two other trichomes medium-long and
stout) (Fig. 36F), on each side; all unbranched. Gill (Fig. 36G) with eight slender thread-like filaments arranged as
(2+2)+(2+2) from dorsal to ventral, arising from short common basal stalk; all pairs short-stalked except second
pair from above almost sessile; length of filaments variable: upper filaments of first and second pairs from above
shortest (0.84 mm long), two filaments of fourth pair from above longest (1.2–1.3 mm long), and other filaments
intermediate in length (1.0–1.1 mm long); all filaments subequal in thickness to one another, though two filaments

FIGURE 36. Pupa of Simulium (Simulium) sansahoense sp. nov. A, Frontal trichome; B, Facial trichome; C–F, Thoracic
side; outer view); H, Hair-like seta and minute seta on dorsal surface of abdominal segment 1; I, Hair-like seta, minute seta and
stout seta on dorsal surface of abdominal segment 2; J, Terminal hooks (caudal view); K & L, Cocoons (K, dorsal view; L,
lateral view). Scale bars. 1.0 mm for K and L; 0.1 mm for G; 0.02 mm for B–F and J; 0.01 mm for A, I and H.
of fourth pair slightly thicker than others; all filaments dark brown, slightly tapered toward apices, with annular
furrows (annular ridges not well developed), and densely covered with minute tubercles. Abdomen. Dorsally, all
segments unpigmented except segments 1 and 2 entirely light brown, and also segment 3 occasionally with wide
darkened area anteriorly; segment 1 with one unbranched short hair-like seta (Fig. 36H) on each side; segment 2
with one unbranched short hair-like seta and five minute setae, of which three or four are stout (Fig. 36I), on each
side; segments 3 and 4 each with four distinct hooks and one unbranched minute seta on each side; segment 5–7
lacking spine-combs; segment 8 with distinct spine-combs in transverse row on each side; segments 6–9 each with
comb-like groups of minute spines on each side; segment 9 with pair of small conical terminal hooks (Fig. 36J).
Ventrally, segments 3–8 unpigmented, each with comb-like groups of minute spines; segment 4 with few minute
setae on each side; segment 5 with pair of bifid hooks submedially and few minute setae on each side; segments 6
and 7 each with pair of bifid inner and unbranched outer hooks somewhat separated from each other, and few
minute setae on each side. Grapnel-shaped hooklets absent on each side of segment 9. Cocoon (Fig. 36K, L).
Corbicular, moderately woven, ochreous, not extended ventrolaterally, and with several large open spaces
anteriorly; individual threads invisible; 3.5–4.2 long by 1.0–1.3 mm wide.
Mature larva. Body length 5.3–6.0 mm. Body vermilion to ochreous (though intersegmental areas from
thoracic segment 3 to abdominal segment 5 paler) except ventral surface of thorax and abdominal segments 1–4
light grayish-green, and that of abdominal segments 5–9 unpigmented. Head. Cephalic apotome (Fig. 37A)
variable in color patterns: e.g., some cephalic apotome whitish yellow to yellow though medial portion narrowly
darkened along posterior margin, with faint or moderate positive head spots; some others whitish yellow on
anterior three-fifths, with faint anterior spot of mediolongitudinal spots, and light to dark brown on posterior two-
thirds with faint to moderate negative or obscure spots; some others light to medium brown on anterior one-third,
medium to dark brown on posterior two-thirds, with faintly or moderately negative spots, and medial portion
between dark areas yellow, with obscure or faintly positive anterior spot of mediolongitudinal spots; some others
widely yellowish except narrow area along posterior margin darkened and somewhat light brown medially, with
obscure or negative head spots. Lateral surface of head capsule light to medium brown except eye-spot region
whitish and anterior portion near anterior margin and area below eye-spot region yellowish to varying extent, with
spots near posterior margin obscure or faintly or moderately negative, and isolated spot below eye-spot region
often positive; eyebrow distinct. Ventral surface of head capsule (Fig. 37B) light to medium brown except anterior
area near hypostoma and both lateral areas yellowish to varying extent; long spot on each side of postgenal cleft
obscure or faintly positive or negative. Head capsule sparsely covered with minute colorless setae. Antenna
composed of three articles and apical sensillum, much longer than stem of labral fan; length ratio of three articles
(from base to tip) 1.0:1.3–1.4:0.6. Labral fan with 52–56 primary rays. Mandible (Fig. 37C) with mandibular
serration composed of two teeth (one medium-sized, one small); main tooth at obtuse angle against mandible on
apical side; supernumerary serrations absent; comb-teeth decreasing in length from first to third. Hypostoma (Fig.
37D) with nine anterior teeth, of which median tooth is slightly longer than corner teeth, and intermediate teeth on
each side shortest; lateral margins serrate apically; six or seven hypostomal bristles per side lying divergent
posteriorly from lateral margin. Postgenal cleft (Fig. 37B) rounded, 4.0 times length of postgenal bridge; sheath of
subesophageal ganglion weakly pigmented. Cervical sclerites on each side composed of one light-brown rod-like
piece and one light-brown elliptical piece, not fused to occiput. Thorax and Abdomen. Histoblast of pharate pupal
gill with eight short filaments. Thoracic cuticle sparsely covered with minute colorless setae dorsally. Abdominal
segments 1–5 each with pair of small protuberances (Fig. 37E) dorsally; abdominal cuticle sparsely or moderately
covered with minute colorless setae dorsally and dorsolaterally; last abdominal segment moderately covered with
short colorless setae on each side of anal sclerite. Rectal scales present. Rectal organ compound, each lobe with 14–
18 finger-like secondary lobules. Anal sclerite X-shaped, with short broad anterior arms 0.7 times length of
posterior ones; one to five sensilla on base of anal sclerite; four to six sensilla posterior to posterior arms. Last
abdominal segment somewhat swollen laterally but lacking ventral papillae. Posterior circlet with 84–86 rows of
hooklets with up to 16–19 hooklets per row.
in a vial as [UMSRP: Vietnam 034, Holotype, Simulium (S.) sansahoense, Female, coll. Vietnam, 20-XII-2014,
Takaoka et al.], reared from a pupa collected from a river (width 6.0 m, depth 12 cm, water temperature 10.0˚C,
exposed to the sun, elevation 1,194 m, 22˚19’44.349”N/103˚49’49.930”E), moderately flowing from a natural
forest, San Sa Ho, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z.
Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: Four females, six males (all reared from pupae), two pupae and 10

FIGURE 37. Larva of Simulium (Simulium) sansahoense sp. nov. A & B, Head capsules (A, dorsal view; B, ventral view); C,
Mandible; D, Hypostoma; E, Protuberance and minute setae on dorsal surface of abdominal segment 5. Scale bars. 0.1 mm for
A and B; 0.05 mm for D; 0.02 mm for C and E.
mature larvae, in 80% ethano,l same data as those of the holotype, labeled in each vial as [UMSRP: Vietnam 034,
Paratype, Simulium (S.) sansahoense, coll. Vietnam, 20-XII-2014, Takaoka et al.].
Biological notes. The pupae and larvae of this new species were collected from plastic plates in the current.
The associated species were S. (S.) daoense sp. nov. and S. (S.) phuluense sp. nov.
Etymology. The species name sansahoense refers to the name of the locality, San Sa Ho, where this new
species was collected.
Remarks. Simulium (S.) sansahoense sp. nov. is assigned to the S. argentipes species-group of the subgenus
Simulium, defined by Takaoka and Saito (2007), based on the female terminalia (Fig. 34E), pupal gill with eight
filaments (Fig. 36G) and corbicular cocoon (Fig. 36K, L).
This new species is similar to S. (S.) bidentatum (Shiraki) from Japan (Takaoka 1976) and S. (S.) tanae from
Yunnan Province, south China (Xue 1992) by having yellowish legs. However, this new species is distinguished
from S. (S.) bidentatum in the female by the length ratio of the labrum against the clypeus 0.6 (0.7 in S. (S.)
bidentatum), ovipositor valve covered with five to eight short to medium-long hairs (12 or 13 short to medium-long
hairs in S. (S.) bidentatum); in the male by the scutum covered with yellow fine hairs (brassy fine hairs in S. (S.)
bidentatum), and fore and mid femora that are light brown except the apical cap medium-brown (yellow except the
apical cap light to medium-brown in S. (S.) bidentatum). Morphological data of S. (S.) bidentatum used here were
based on specimens collected from Oita, Japan, in May, 2015 by Dr. M. Fukuda.
This new species is distinguished from S. (S.) tanae by the following characters (those of S. (S.) tanae in
parentheses): female body length 2.5–2.6 mm (3.0–3.4 mm), female frons with the ratio of the greatest width
against the height 1.3–1.4 (2.0), spermatheca ellipsoidal (Fig. 34I) (globular), male abdominal segments 2 and 5–7
each with a pair of iridescent dorsolateral spots (segments 2, 6 and 7 each with such spots).
Five other species of the S. argentipes species-group, viz., S. (S.) arboreum Takaoka from the Philippines, S.
(S.) argentipes Edwards from Peninsular Malaysia, S. (S.) beludense Takaoka from Sabah and Sarawak, S. (S.)
canlaonense Delfinado from the Philippines, and S. (S.) minangkabaum Takaoka & Sigit from Sumatra, Indonesia,
differ from this new species by having much darker female legs (Takaoka 1983, 1996, 2006, Takaoka and Davies
1995, Takaoka and Sigit 1997).
2) Simulium christophersi species-group
Simulium (Simulium) atipornae Takaoka, Srisuka & Choochote, 2014
Simulium (Simulium) atipornae Takaoka, Srisuka & Choochote (in Takaoka et al., 2014e): 725–732 (Female, male, pupa and
larva); Takaoka et al., 2015a: 60.
Distribution. Thailand and Vietnam (Thua Thien Hue).

Remarks. Simulium (S.) atipornae was originally described from Thailand and assigned to the Simulium
christophersi species-group, defined by Takaoka et al. (2014e). This species was reported based on one pharate
male collected from Bach Ma National Park, Thua Thien Hue Province, Vietnam (Takaoka et al. 2015a).
3) Simulium griseifrons species-group
Simulium (Simulium) chungi Takaoka & Huang, 2006
Simulium (Simulium) chungi Takaoka & Huang, 2006: 219–227 (Female, male, pupa and larva); Takaoka et al. 2014b: 746.
Distribution. Taiwan and Vietnam (Vinh Phuc).

Remarks. This species was originally described from Taiwan (Takaoka and Huang 2006). The record from
Vietnam was based on 14 females captured in Tam Dao National Park, Vinh Phuc Province, while flying around a
human (Takaoka et al. 2014b). The aquatic stages remain to be collected from Vietnam.
Simulium (Simulium) grossifilum Takaoka & Davies, 1995
Simulium (Simulium) grossifilum Takaoka & Davies, 1995: 105–115 (Female, male, pupa and larva); Takaoka et al., 2014b:
746.
Distribution. Peninsular Malaysia, Thailand and Vietnam (Vinh Phuc).

Remarks. This species was originally described from Peninsular Malaysia (Takaoka and Davies 1995). The
record from Vietnam was based on three females captured in Tam Dao National Park, Vinh Phuc Province, while
flying around a human (Takaoka et al. 2014b). The aquatic stages remain to be collected from Vietnam.
Simulium (Simulium) laocaiense Takaoka & Lau sp. nov.
Female. Body length 3.3 mm. Head. Slightly narrower than thorax. Frons black, shiny, with bluish reflection when
illuminated at certain angles, with several dark stout hairs along lateral margins; frontal ratio 1.3:1.0:0.9;
frons:head ratio 1.0:3.6. Fronto-ocular area well developed, short, directed laterally, and pointed apically. Clypeus
brownish black, white pruinose when illuminated at certain angles, moderately covered with dark-brown medium-
long hairs (though mediolongitudinal portion of upper half widely bare). Labrum 0.7 times length of clypeus.
Antenna composed of scape, pedicel and nine flagellomeres; first flagellomere 1.4 times length of second one;
scape and pedicel whitish yellow, and basal two-thirds of first flagellomere yellow, rest dark brown to brownish
black, when viewed dorsally (first flagellomere entirely yellow or ochreous when viewed ventrally); first to eighth
flagellomeres each with pit-like depression with many sensilla on inner and outer sides. Maxillary palp with five
segments, light to medium brown; proportional lengths of third, fourth, and fifth segments 1.0:1.4:2.8; third
segment (Fig. 38A) with apex somewhat produced medially, with medium-sized ellipsoidal sensory vesicle (0.3
times length of third segment) having medium-sized opening. Maxillary lacinia with14 or 15 inner and 13 or 15
outer teeth. Mandible with 25 inner and 11 or 12 outer teeth. Cibarium (Fig. 38B) with 70 minute processes near
posterodorsal margin. Thorax. Scutum black, slightly shiny and grayish pruinose on shoulders and lateral portions
when illuminated at certain angles, moderately covered with whitish-yellow recumbent short hairs interspersed
with several dark-brown long upright hairs on prescutellar area. Scutellum dark brown, covered with dark-brown
upright long hairs and yellow short hairs. Postnotum brownish black, bare. Pleural membrane bare. Katepisternum

FIGURE 38. Female of Simulium (Simulium) laocaiense sp. nov. A, Third segment of maxillary palp with sensory vesicle (left
side; front view); B, Cibarium (front view); C, Hind basitarsus and second tarsomere (left side; outer view). D, Sternite 8 and
ovipositor valve (right side only; ventral view); E, Genital fork (ventral view); F & G, Paraprocts and cerci (F, ventral view; G,
lateral view). H, Spermatheca. Scale bars. 0.1 mm for C; 0.02 mm for A, B and D–H.
brownish black, longer than deep, bare, and slightly shiny when illuminated at certain angles. Legs. Foreleg: coxa
whitish yellow; trochanter and femur yellow; tibia whitish yellow except apical one-fifth brownish black; tarsus
black, with moderate dorsal hair crest; basitarsus greatly dilated, 5.6 times as long as its greatest width. Midleg:
coxa dark brown; trochanter, femur and tibia yellow except dorsal small portion of apical tip of tibia grayish; tarsus
light to medium brown except basal half or little more of basitarsus yellow (though its border not well defined).
Hind leg: coxa dark brown; trochanter yellow; femur yellow except dorsal portion of apical tip light brown; tibia
yellow with apical cap medium brown; tarsus dark brown except basal two-thirds of basitarsus and basal half of
second tarsomere yellowish white; basitarsus (Fig. 38C) nearly parallel-sided, 6.0 times as long as wide, and 0.9
and 0.7 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 38C) moderately
developed, slightly shorter than its basal width, and 0.4 times as wide as greatest width of basitarsus; pedisulcus
(Fig. 38C) well developed. Claw simple, without basal or subbasal tooth. Wing. Length 3.0 mm. Costa with dark
spinules and hairs; subcosta almost fully haired; basal section of radius bare; R1 with dark-brown spinules and
hairs; R2 with dark-brown hairs; hair tuft on base of radius dark brown; basal cell absent. Halter. White except base
light brown. Abdomen. Basal scale dark brown to brownish black, with fringe of yellowish hairs. Dorsal surface of
abdomen medium-brown to black; tergite 2 shiny and white pruinose when illuminated at certain angles and
tergites 6–9 shiny. Ventral surface medium to dark brown except basal portion of segment 2 white; segment 7 with
large median sternal plate. Terminalia. Sternite 8 (Fig. 38D) with 24–26 dark-brown medium-long to long stout
hairs and six to nine yellow short hairs on each lateral surface. Ovipositor valve (Fig. 38D) wide, somewhat thick
except area along inner margins and wide areas along posterior margin thin and transparent, somewhat rounded
ventrally, densely covered with microsetae (except transparent portions along inner and posterior margins bare)
interspersed with 19 or 20 yellow short hairs; inner margins somewhat concave, moderately separated from each
other. Genital fork (Fig. 38E) of inverted-Y form, with narrow well sclerotized stem; arms of moderate width, each
with heavily sclerotized lateral portion, with short projection directed anterodorsally. Paraproct in ventral view
(Fig. 38F) nearly quadrate, 1.2 times as long as its width, with 33–35 short to medium-long hairs on lateral and
ventral surfaces, and with anterior surface strongly sclerotized and pigmented with 8–14 short sensilla; paraproct in
lateral view (Fig. 38G) 0.72 times as long as wide, and protruding ventrally beyond ventral margin of cercus.
Cercus in lateral view (Fig. 38G) short, rectangular, 0.5 times as long as wide, with numerous short to medium-
long hairs. Spermatheca (Fig. 38H) nearly ovoid, 1.3 times as long as greatest width, well sclerotized except small
portion of junction with duct unsclerotized, with faintly defined reticulate surface patterns; internal setae present;
accessory ducts subequal in thickness to each other, and slightly thicker than major duct.
Male. Unknown.
Pupa. Body length (excluding gill filaments) 3.5–4.0 mm. Head. Integument dark brown except ventral
surface ochreous, and moderately covered with round tubercles except lateral and ventral surfaces and antennal
sheaths bare: frons with two medium-long trichomes (one unbranched and one bifid, or two bifid, or two trifid, or
quadrifid and quinquefid) (Fig. 39A); one female pupa with additional short hair on right side and one minute seta
on left side (Fig. 39B), and one male pupa with additional short trichome on left side (Fig. 39C); face with one
medium-long trichome on each side (one unbranched and one bifid, or two bifid, or trifid and quadrifid) (Fig. 39D)
on each side. Thorax. Integument dark brown except wing sheaths ochreous, moderately covered with round
tubercles; thorax with three anterodorsal trichomes (one unbranched or bifid, one trifid, one quadrifid or
quinquefid) (Fig. 39E), two anterolateral trichomes (one bifid, one trifid or quadrifid) (Fig. 39F), one bifid
mediolateral trichome (Fig. 39G), and three ventrolateral trichomes (all bifid, or one unbranched and two bifid)
(Fig. 39H) on each side; all trichomes medium-long. Gill (Fig. 39I) with six slender thread-like filaments in pairs;
common basal stalk short, with basal fenestra ventrally; dorsal and ventral pairs short stalked and middle pair
almost sessile; outer filament of dorsal pair longest (1.5 mm long), followed by outer filament of middle pair (1.3
mm long), inner filament of dorsal pair and outer filament of ventral pair (1.1 mm long), inner filament of middle
pair (1.0 mm long) and inner filament of ventral pair (0.9 mm long); relative thickness of six filaments from dorsal
to ventral 1.0:0.8:1.0:0.6:0.9:0.5; all filaments dark brown, gradually tapered toward apex; outer filament of dorsal
pair and inner filament of ventral pair divergent basally at angle of 60 degrees; cuticular surface with well-defined
annular ridges and furrows throughout their length except apical portions less distinct, covered with minute
tubercles. Abdomen. Dorsally, entire surface of segment 1 and anterior one-third of segment 2 light brown, other
segments unpigmented; segment 1 smooth (without tubercles), with one unbranched slender medium-long seta
(Fig. 39J) on each side; segment 2 smooth, with one unbranched slender short seta, five unbranched short setae, of

FIGURE 39. Pupa of Simulium (Simulium) laocaiense sp. nov. A–C, Frontal trichomes of different sizes and types; D, Facial
trichome; E–H, Thoracic trichomes (E, anterodorsal; F, anterolateral; G, mediolateral; H, ventrolateral); I, Anterior part of
thorax and gill filaments (right side; outer view); J, Hair-like seta and minute seta on dorsal surface of abdominal segment 1; K,
Hair-like seta, minute seta and stout seta on dorsal surface of abdominal segment 2; L & M, Cocoons (L, dorsal view; M, lateral
view). Scale bars. 1.0 mm for L and M; 0.1 mm for I; 0.02 mm for A–H, J and K.
which four are stout (Fig. 39K), on each side; segments 3 and 4 each with four unbranched hooked spines and one
unbranched short seta on each side; segment 8 with spine-combs in transverse row on each side; segments 5–9 each
with comb-like groups of minute spines; segment 9 without terminal hooks. Ventrally, all segments unpigmented;
segment 4 with one unbranched hook and few short setae on each side; segment 5 with pair of unbranched or bifid
hooks submedially and few unbranched short seta on each side; segments 6 and 7 each with pair of unbranched or
bifid inner and unbranched outer hooks widely spaced and few unbranched short seta on each side; segments 4–8
each with comb-like groups of minute spines. Grapnel-like hooklets absent. Cocoon (Fig. 39L, M). Wall-pocket-
shaped, thickly woven, with short round anterodorsal projection, ochreous, not extended ventrolaterally; individual
threads invisible; 3.1 mm long by 1.6 mm wide.
Mature larva. Body length 8.0 mm. Body ochreous to reddish brown, mottled to varying extent with grayish
black to black pigment except ventral surface of abdominal segments 2–9 unpigmented; intersegmental spaces
from thoracic segment 3 to abdominal segment 4 paler. Head. Cephalic apotome (Fig. 40A) whitish yellow except
large area of triangular shape on posterior half light to medium brown and narrow area along posterior margin dark
brown; head spots moderately positive though posterolateral spots indistinct; lateral surface of head capsule
whitish yellow, with distinct eyebrow, with eye-spot region whitish and two large spots in front of posterior margin
and three isolated spots below eye-spot region positive; ventral surface of head capsule (Fig. 40D) yellow except
areas along posterior margin widely darkened, with dark brown elongate spot on each side of cleft. Antenna
composed of three articles and apical sensillum, slightly longer than stem of labral fan; length ratio of articles
(from base to tip) 1.0:1.2:0.5. Labral fan with 52 primary rays. Mandible (Fig. 40B) with mandibular serration
composed of two teeth (one medium-sized and one small); major tooth at obtuse angle apically against mandible;
FIGURE 40. Larva of Simulium (Simulium) laocaiense sp. nov. A, Head capsule (dorsal view); B, Mandible; C, Hypostoma;
D, Head capsule (ventral view). Scale bars. 0.1 mm for A and D; 0.05 mm for C; 0.02 mm for B.

comb-teeth decreasing in length from first to third; supernumerary serrations absent. Hypostoma (Fig. 40C) with
nine anterior teeth, of which median tooth slightly longer than each corner tooth; lateral margins weakly serrate
apically; nine hypostomal bristles divergent posteriorly from lateral border on each side. Postgenal cleft (Fig. 40D)
medium-long, triangular; sheath of subesophageal ganglion well pigmented, wine-glass-shaped. Cervical sclerites
on each side composed of one light-brown elliptical piece, not fused to occiput. Thorax and Abdomen. Thoracic
and abdominal cuticle bare except last segment of abdomen moderately covered with short colorless setae on each
side of anal sclerite. Rectal scales present. Rectal organ unobservable. Anal sclerite X-shaped, with broadened
anterior arms 0.7 times length of posterior ones; five sensilla on base of anal sclerite and seven sensilla posterior to
posterior arms. Last abdominal segment bulged laterally but lacking ventral papillae. Posterior circlet with 164
rows of hooklets with up to 22 hooklets per row.
in a vial as [UMSRP: Vietnam 035, Holotype, Simulium (S.) laocaiense, Female, coll. Vietnam, 20-XII-2014,
Takaoka et al.], reared from a pupa collected from a stream (width 0.5–2.0 m, depth 3 cm, bottom rocky, water
temperature 11.0˚C, exposed to the sun, elevation 1,853 m, 22˚21’18.609”N/103˚46’27.550”E) moderately flowing
from a forest, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z.
Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: One pupa and three pupal exuviae in 80% ethanol, labeled in each
vial as [UMSRP: Vietnam 035, Paratype, Simulium (S.) laocaiense, coll. Vietnam, 20-XII-2014, Takaoka et al.],
collected from a small stream (width 0.5 m, depth 10 cm, bottom rocky, water temperature 11.0˚C, exposed to the
sun, elevation 1,750 m, 22˚21’43.110”N/103˚47’19.221”E) slowly flowing from a secondary forest, Sapa, Lao Cai
Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau;
one mature larva in 80% ethanol, labeled in a vial as [UMSRP: Vietnam 035, Paratype, Simulium (S.) laocaiense,
Larva, coll. Vietnam, 20-XII-2014, Takaoka et al.], collected from a stream (width 0.5–1.0 m, bottom rocky, water
temperature 9.0˚C, partially shaded, elevation 1,680 m, 22˚22’05.320”N/103˚47’34.403”E) slowly flowing from a
secondary forest, Oguy Ho, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-
Azirun, Z. Ya’cob, C.D. Chen & K. W. Lau.
Biological notes. The pupae of this new species were collected from trailing grasses. Associated species were
S. (N.) laichauense sp. nov., S. (S.) sapaense sp. nov., S. (S.) sp. (S. multistriatum species-group) and S. (S.)
turgidum sp. nov.
Etymology. The species name laocaiense refers to the name of the province, Lao Cai, where this new species
was collected.
Remarks. Simulium (S.) laocaiense sp. nov., though its male is unknown, is placed in the S. griseifrons
species-group, defined by Takaoka and Davies (1996), based on the similarity of the female enlarged ovipositor
valve (Fig. 38D) to that of S. (S.) rudnicki Takaoka & Davies described from Peninsular Malaysia, which is a
member of this species-group (Takaoka and Davies 1995; Takaoka and Choochote 2004c).
This new species is characterized by the simple cocoon with a somewhat produced anterodorsal portion (Fig.
39L, M), by which this new species is distinguished from most species of the S. griseifrons species-group, the
cocoons of which are usually fenestrated or rarely shoe-shaped or simple wall-pocket-shaped.
This new species is similar in the female terminalia to S. (S.) vietnamense Takaoka, Sofian-Azirun and Chen
described from females collected from Vietnam (Takaoka et al. 2014b), but is distinguished from the latter by
lacking hairs on the basal portion of the female radial vein.
Simulium (Simulium) maenoi Takaoka & Choochote, 2002
Simulium (Simulium) maenoi Takaoka & Choochote, 2002: 115–120 (Female, male, pupa and larva); Takaoka et al., 2014b:
746–747.
Distribution. Thailand and Vietnam (Vinh Phuc).

Remarks. This species was originally described from Thailand (Takaoka and Choochote 2002). The record
from Vietnam was based on three females captured in Tam Dao National Park, Vinh Phuc Province, while flying
around a human (Takaoka et al. 2014b). The aquatic stages remain to be collected from Vietnam.
Simulium (Simulium) nigrogilvum Summers, 1911
Simulium nigrogilvum Summers, 1911: 586–588 (Female).

Simulium (Himalayum) nigrogilvum: Takaoka & Suzuki, 1984: 22–27 (Female, male, pupa and larva).
Simulium (Simulium) nigrogilvum: Otsuka et al., 2003: 113–120.
Distribution. Thailand and Vietnam.

Remarks. This species was assigned to the subgenus Himalayum Lewis (Takaoka and Suzuki 1984) but later
transferred to the subgenus Simulium (Otsuka et al. 2003). The females of this species are known to be
anthropophilic (Choochote et al. 2005) and to transmit an unknown filarial parasite in Thailand (Fukuda et al.
2003). This species was recorded from Vietnam by Crosskey and Howard (1997). However, data including the
material collected and its locality were not given.
Simulium (Simulium) turgidum Takaoka & Pham sp. nov.
Female. Body length 3.3 mm. Head. Nearly as in female of S. (S.) laocaiense sp. nov. except following characters:
Frontal ratio 1.2:1.0:1.0, frons:head ratio 1.0:4.3. Labrum 0.7 times length of clypeus. Antenna: first flagellomere
1.36 times length of second one. Maxillary palp: proportional lengths of third, fourth, and fifth segments
1.0:1.3:3.4; third segment (Fig. 41A) with apex somewhat produced medially, with medium-sized ellipsoidal
sensory vesicle (0.4 times length of third segment) having large or medium-sized opening. Maxillary lacinia
with12–13 inner and 13 or 14 outer teeth. Mandible with 25 inner and 12 or 13 outer teeth. Cibarium (Fig. 41B)
with 38 minute processes near posterodorsal margin. Thorax, Legs, Wing and Halter. As in female of S. (S.)
laocaiense sp. nov. except following characters: Scutum covered with yellow short hairs. Fore tibia whitish yellow
except apical one-fifth brownish black, and little more than apical two-fifths of inner surface light brown to
brownish black. Fore basitarsus greatly dilated, 5.8 times as long as its greatest width. Mid tarsus light to medium
brown except basal one-third to one-half of basitarsus yellow (though its border not well defined). Hind basitarsus
(Fig. 41C) gradually widened toward apical one-third, then slightly narrowed toward apex, 5.6 times as long as
wide, and 0.8 and 0.7 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 41C)
moderately developed, 0.7 times as long as wide, and 0.5 times as wide as greatest width of basitarsus; pedisulcus
(Fig. 41C) well developed. Wing length 3.1 mm. Subcosta haired except near apex bare. Abdomen. Basal scale
medium brown to brownish black, with fringe of yellowish hairs. Dorsal surface of abdomen medium brown to
black; tergite 2 shiny and white pruinose when illuminated at certain angles and tergites 6–9 shiny. Ventral surface
light brown except basal portion of segment 2 white; sternal plate on segment 7 indistinct. Terminalia. Sternite 8
(Fig. 41D) with 11 or 12 dark-brown medium-long to long stout hairs and three to six yellow short hairs on each
lateral surface. Ovipositor valve (Fig. 41D) wide, somewhat thick except narrow areas along inner and posterior
margins thin and transparent, somewhat rounded ventrally, densely covered with microsetae (except narrow
transparent portions along inner and posterior margins bare) interspersed with 22–25 yellow short hairs; inner
margins somewhat separated from each other. Genital fork (Fig. 41E, F) of inverted-Y form, with narrow well
sclerotized stem; arms of moderate width, each with heavily sclerotized lateral portion, with short projection
directed anterodorsally. Paraproct in ventral view (Fig. 41G) nearly quadrate, 1.2 times as long as its width, with 23
or 24 short to medium-long hairs on lateral and ventral surfaces; anterior surface strongly sclerotized and
pigmented, and with four or five short sensilla; paraproct in lateral view (Fig. 41H) 0.7 times as long as wide, and
protruding ventrally beyond ventral margin of cercus. Cercus in lateral view (Fig. 41H) short, rectangular, 0.6 times
as long as wide, with numerous short to medium-long hairs. Spermatheca (Fig. 41I) globular, 1.1 times as long as
greatest width, well sclerotized except small portion of junction with duct unsclerotized, with faintly defined
reticulate surface patterns; internal setae present; accessory ducts subequal in thickness to each other, and slightly
thicker than major duct.
Male. Unknown.
Pupa. Body length (excluding gill filaments) 4.0 mm. Head. Integument dark brown except ventral surface
ochreous, and bare, without tubercles (though surface appearing to have micro-sculptures formed by numerous fine
ridges); frons with two bifid medium-long trichomes (Fig. 42A); face with one bifid or trifid medium-long
trichome on each side (Fig. 42B). Thorax. Integument dark brown except wing sheaths ochreous, and bare except

FIGURE 41. Female of Simulium (Simulium) turgidum sp. nov. A, Third segment of maxillary palp with sensory vesicle (left
ovipositor valve (right side only; ventral view); E, Genital fork (ventral view); F, Arm of genital fork showing anterodorsal
triangular projection; G & H, Paraprocts and cerci (G, ventral view; H, lateral view). I, Spermatheca. Scale bars. 0.1 mm for C;
0.02 mm for A, B and D–I.
dorsal surface of posterior half sparsely to moderately covered with round tubercles and basal portions of gills
densely covered with tubercles having minute secondary projections (though bare surface appearing to have micro-
sculptures formed by numerous fine ridges); thorax with three long anterodorsal trichomes with four to six
branches (Fig. 42C), two long or medium-long anterolateral trichomes with four branches (anterior one little
shorter than posterior one) (Fig. 42D), one unbranched or bifid medium-long mediolateral trichome (Fig. 42E), and
three ventrolateral trichomes (two unbranched medium-long, one bifid in left side (Fig. 42F); one unbranched
medium-long, one quadrifid medium-long, and one unbranched short (Fig. 42G) in right side), on each side. Gill
(Fig. 42H) with six somewhat inflated filaments in pairs; common basal stalk short, with small basal fenestra
ventrally; all pairs almost sessile; outer filament of dorsal pair longest (1.8 mm long), followed by inner filament of
dorsal pair and outer filament of middle pair (1.4 mm long), inner filament of middle pair and outer filament of
ventral pair (1.1 mm long), and inner filament of ventral pair (1.0 mm long); each filament widened from base to
basal one-fifth to one-third, then tapered toward apex; relative thickness of six filaments from dorsal to ventral
when widest portion compared 1.0:0.9:0.9:0.8:0.7:0.6; all filaments dark brown, gradually tapered toward apex;
outer filament of dorsal pair and inner filament of ventral pair divergent basally at angle of 60 degrees; cuticular
surface with well-defined annular ridges and furrows throughout their length forming definite reticulate surface
patterns, covered with minute tubercles. Abdomen. Dorsally, entire surface of segment 1 and anterior half of
segment 2 light brown, other segments unpigmented; segment 1 sparsely covered with minute spines laterally, with
one unbranched slender medium-long seta (Fig. 42I) on each side; segment 2 covered with comb-like groups of
minute spines submedially, with one unbranched slender short seta and five unbranched minute setae, of which
four are stout (Fig. 42J), on each side; segments 3 and 4 each with four unbranched hooked spines and one
unbranched minute seta on each side; segments 7 and 8 each with spine-combs in transverse row on each side;
segments 6–9 each with comb-like groups of minute spines; segment 9 without terminal hooks. Ventrally, all
segments unpigmented; segment 4 with one unbranched hook and few short setae on each side; segment 5 with pair
of unbranched or bifid hooks submedially and few unbranched short seta on each side; segments 6 and 7 each with
pair of bifid inner and unbranched outer hooks widely spaced and few unbranched short seta on each side;
segments 3–8 each with comb-like groups of minute spines. Grapnel-like hooklets absent. Cocoon (Fig. 42K, L).
Wall-pocket-shaped, thickly woven, with anterolateral windows, ochreous, not extended ventrolaterally; individual
threads invisible; 4.0–4.2 mm long by 1.6–1.8 mm wide.
Mature larva. Body length 6.7–7.5 mm. Body ochreous or partially reddish brown, mottled to varying extent
with grayish-black to black pigment except ventral surface of abdominal segments 3–9 unpigmented. Head.
Cephalic apotome (Fig. 43A) whitish yellow on little more than anterior half and light to medium brown on rest
(though small medial area in front of posterior margin dark brown); posterior spot of mediolongitudinal spots and
posterior half of each mediolateral spot dark brown, anterior spot of mediolongitudinal spots light brown, and other
spots indistinct, merged into dark background, or anterior spot of posterolateral spots on each side faintly negative.
Lateral surface of head capsule light to medium brown except eye-spot region, eyebrow and median small area
along anterior margin yellowish white, with faint or distinct negative spots except one small spot below eye-spot
region light brown. Ventral surface of head capsule (Fig. 43B) light to medium brown except postgenal bridge
lighter medially, and elongate spot on each side of cleft indistinct, merged into dark background. Antenna (Fig.
43C) composed of three articles and apical sensillum, slightly longer than stem of labral fan; antenna yellowish
white except third segment dark brown; length ratio of articles (from base to tip) 1.0:1.2–1.3:0.5. Labral fan with
49–51 primary rays. Mandible (Fig. 43D) with mandibular serration composed of two teeth (one medium-sized and
one small); major tooth at obtuse angle apically against mandible; comb-teeth decreasing in length from first to
third; supernumerary serrations absent. Hypostoma (Fig. 43E) with nine anterior teeth, of which median tooth
slightly longer than each corner tooth; lateral margins weakly serrate apically; seven or eight hypostomal bristles
divergent posteriorly from lateral border on each side. Postgenal cleft (Fig. 43B) medium-long, triangular, 1.4
times length of postgenal bridge; sheath of subesophageal ganglion well pigmented, wine-glass-shaped. Cervical
sclerites on each side composed of one light-brown elliptical piece, not fused to occiput. Thorax and Abdomen.
Thoracic and abdominal cuticle bare except last segment of abdomen moderately covered with short colorless setae
on each side of anal sclerite. Rectal scales present. Rectal organ compound, each of three lobes with 23–25 finger-
like secondary lobules. Anal sclerite X-shaped, with broadened anterior arms 0.7–0.9 times length of posterior
ones; no sensilla on base of anal sclerite and 12 sensilla posterior to posterior arms. Last abdominal segment bulged
laterally but lacking ventral papillae. Posterior circlet with 104–110 rows of hooklets with up to 17 or 18 hooklets
per row.

FIGURE 42. Pupa of Simulium (Simulium) turgidum sp. nov. A, Frontal trichome; B, Facial trichome; C–G, Thoracic
trichomes (C, anterodorsal; D, anterolateral; E, mediolateral; F & G, ventrolateral); H, Anterior part of thorax and gill filaments
(right side; outer view); I, Hair-like seta and minute seta on dorsal surface of abdominal segment 1; J, Hair-like seta, minute
seta and stout seta on dorsal surface of abdominal segment 2; K & L, Cocoons (K, dorsal view; L, lateral view). Scale bars. 1.0
mm for K and L; 0.1 mm for H; 0.02 mm for A–G, I and J.
FIGURE 43. Larva of Simulium (Simulium) turgidum sp. nov. A & B, Head capsules (A, dorsal view; B, ventral view); C,
Antenna; D, Mandible; E, Hypostoma. Scale bars. 0.1 mm for A and B; 0.05 mm for C and E; 0.02 mm for D.
in a vial as [UMSRP: Vietnam 036, Holotype, Simulium (S.) turgidum, Female, coll. Vietnam, 20-XII-2014,
Takaoka et al.], reared from a pupa collected from a stream (width 0.5–2.5 m, bottom rocky, water temperature
10.0˚C, exposed to the sun, elevation 1,812 m, 22˚21’27.450”N/103˚46’35.743”E) moderately flowing in a forest,
& K.W. Lau. PARATYPES: One pupa in 80% ethanol, labeled in a vial as [UMSRP: Vietnam 036, Paratype,
Simulium (S.) turgidum, Pupa, coll. Vietnam, 20-XII-2014, Takaoka et al.], collected from a stream (width 0.5 m,
bottom rocky, water temperature 11.0˚C, exposed to the sun, elevation 1,750 m, 22˚21’43.110”N/
103˚47’19.221”E) moderately flowing in a secondary forest, Sapa, Lao Cai Province, northern Vietnam, 20-XII-
2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau; one mature larva, same data as those
of the holotype.
Biological notes. The pupae of S. (S.) turgidum were collected from trailing grasses. Associated species were
S. (M.) nigrofilum sp. nov., S. (N.) laichauense sp. nov., S. (S.) sapaense sp. nov., S. (S.) giayense sp. nov., S. (S.)
sp. (S. multistriatum species-group) and S. (S.) sp. (S. variegatum species-group).
Etymology. The species name turgidum refers to the inflated gill filaments of this new species. The Latin
adjective ‘turgidus’ means ‘inflated’.
Remarks. As for S. (S.) laocaiense sp. nov., S. (S.) turgidum sp. nov. also is placed in the S. griseifrons
species-group based on the similarity of the female enlarged ovipositor valve (Fig. 41D) to that of S. (S.) rudnicki,
which is a member of the S. griseifrons species-group.
In having inflated pupal gill filaments (Fig. 42H), this new species is similar to S. (S.) tumidilfilum Luo, Yang
& Chen described from Hubei, China (Luo et al. 2010), and S. (S.) waterfallum Zhang, Yang & Chen described
from Hainan, China (Zhang et al. 2003), both of which were wrongly placed in the S. multistriatum species-group.
However, this new species is distinguished in the female from the latter two species by the ovipositor valve which

is triangular with 22–25 short hairs (tongue-like with 5–8 short hairs in S. (S.) tumidilfilum and S. (S.) waterfallum),
and in the pupa from S. (S.) tumidilfilum by each inflated filament slightly narrowed at its base (much narrowed in
S. (S.) tumidilfilum), and from S. (S.) waterfallum by abdominal segments 7 and 8 each with spine-combs
(abdominal segments 6–8 each with spine-combs in S. (S.) waterfallum).
Simulium (Simulium) vietnamense Takaoka, Sofian-Azirun & Chen, 2014
Simulium (Simulium) vietnamense Takaoka, Sofian-Azirun & Chen (in Takaoka et al., 2014b): 743–746 (Female).

Remarks. This species was described from two females collected while flying around a human in Tam Dao
National Parks, Vinh Phuc Province (Takaoka et al. 2014b). The male, pupa and larva of this species are unknown.
3) Simulium malyschevi species-group
Simulium (Simulium) obliquum Takaoka & Low sp. nov.
Female. Body length 2.0–2.3 mm. Head. Narrower than thorax. Frons black, shiny, with several dark long stout
hairs along each lateral margin and few similar hairs just above lower margin; frontal ratio 1.3:1.0:1.0; frons:head
ratio 1.0:3.2. Fronto-ocular area well developed, short, directed laterally and slightly upward. Clypeus black, white
pruinose, moderately covered with dark long stout hairs along lateral and ventral margins and middle portion of
upper two-thirds widely bare. Labrum 0.7 times as long as clypeus. Antenna composed of scape, pedicel and nine
flagellomeres, medium to dark brown except scape, pedicel, and flagellomeres 1 and 2 yellow when viewed
anteriorly or ventrally, and medium to dark brown except scape and pedicel and base of flagellomere 1 yellow, and
rest of flagellomere 1 dark yellow to light brown when viewed posteriorly or dorsally. Maxillary palp with five
segments, light brown; proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.0–2.2; third segment (Fig.
44A) of normal size; sensory vesicle (Fig. 44A) small, ellipsoidal (0.2 times length of third segment) having
moderate opening. Maxillary lacinia with 9–12 inner and 12–14 outer teeth. Mandible with 21–26 inner and 11–13
outer teeth. Cibarium (Fig. 44B) with blunt median projection bearing several minute tubercles on posterior margin
and with three or four minute pointed processes near posterior margin on each side. Thorax. Scutum black,
unpatterned, shiny, thinly white pruinose when illuminated at certain angles, moderately covered with dark
recumbent minute hairs intermixed with several dark-brown long upright hairs on prescutellar area. Scutellum
black, white pruinose when illuminated at certain angles, covered with dark-brown upright long hairs. Postnotum
black, white pruinose when illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum black,
longer than deep, bare. Legs. Foreleg: coxa whitish yellow; trochanter yellow; femur deep yellow except apical cap
medium brown; tibia (Fig. 44C) white except inner surface of base light brown, and apical cap dark brown, and
with large white sheen on outer surface when illuminated at certain angles; tarsus brownish black, with moderate
dorsal hair crest; basitarsus greatly dilated, 4.4–4.5 times as long as its greatest width. Midleg: coxa dark brown;
trochanter medium brown except base yellowish white; femur medium brown with extreme base somewhat lighter;
tibia (Fig. 44D) medium to dark brown with extreme base and basal one-third of posterior surface yellowish white
(its border obliquely defined in lateral view), and with white sheen on posterior surface when illuminated at certain
angles; tarsus (Fig. 44E) yellowish white except apical tip of basitarsus, apical one-third of second tarsomere and
most of third tarsomere grayish, fourth tarsomere light brown, and fifth tarsomere medium brown. Hind leg: coxa
dark brown; trochanter yellowish white; femur dark brown except base yellowish white and apical cap brownish
black; tibia (Fig. 44F) medium to dark brown except base and little less than basal half of posterior surface
yellowish white (its border obliquely defined in lateral view) and apical cap brownish black; tarsus (Fig. 44G)
yellowish white except apical one-fourth or little more of basitarsus medium brown, and apical one-third of second
tarsomere and most of third tarsomere grayish, and whole of fourth tarsomere light brown, and fifth tarsomere
medium brown; basitarsus (Fig. 44G) parallel-sided, 6.0–7.2 times as long as wide, and 0.7 and 0.6 times as wide
FIGURE 44. Female of Simulium (Simulium) obliquum sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
side; front view); B, Cibarium (front view); C, Fore tibia (left side; outer view); D, Mid tibia (left side; outer view); E, Mid
tarsus (left side; outer view); F, Hind tibia (left side; outer view); G, Hind tarsus (left side; outer view). H, Claw; I, Abdominal
segment 7 showing cluster of branched hairs; J, Sternite 8 and ovipositor valve (right side only; ventral view); K & L, Genital
forks (K, ventral view; L, lateral view); M & N, Paraprocts and cerci (M, ventral view; N, lateral view). O, Spermatheca. Scale
bars. 0.1 mm for C–G; 0.02 mm for A, B and I–O; 0.01 mm for H.

as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 44G) developed, small, little shorter than its
basal width, and 0.4 times as wide as greatest width of basitarsus; pedisulcus (Fig. 44G) well developed at basal
two-fifths of second tarsomere; claw (Fig. 44H) with small subbasal tooth. Wing. Length 1.9–2.0 mm. Costa with
dark spinules and dull hairs; subcosta bare (except one female with one and two hairs on each subcosta); basal
section of radius bare; R1 with dark spinules and hairs; R2 with hairs; hair tuft on base of radium dark brown; basal
cell absent. Halter. White except base light brown. Abdomen. Basal scale medium brown, with fringe of dull hairs.
Dorsal surface of abdomen medium to dark brown, with short dark hairs; tergite 2 brightly iridescent when
illuminated at certain angles, and tergites 6–9 shiny. Ventral surface of segment 7 (Fig. 44I) with 46–51 branched
stout dark hairs medially. Terminalia. Sternite 8 (Fig. 44J) bare medially, with 8–11 dark medium-long to long
stout hairs and three yellow short hairs on each lateral surface. Ovipositor valve (Fig. 44J) flap-like, bluntly
truncate posteriorly, forming oblique roof-like ridge parallel to posterior margin running obliquely, thin,
membranous, covered with five to eight dark medium-long or long hairs, one to eight yellow short hairs and
numerous microsetae; inner margins not sclerotized, moderately concave medially and widely separated from each
other. Genital fork (Fig. 44K, L) of inverted-Y form, with narrow well sclerotized stem; arms of moderate width,
each with distinct projection with rounded apex directed anterodorsally and triangular projection-like sclerotized
posteromedial corner. Paraproct in ventral view (Fig. 44M) triangular, pointed medially, with unpigmented shallow
depression on ventral surface along anteromedial margin; anteromedial surface moderately sclerotized and with 6–
12 sensilla; paraproct in lateral view (Fig. 44N) 0.6 times as long as wide, with unpigmented shallow depression
narrowing ventrally along anteroventral margin, somewhat protruding beyond ventral margin of cercus, and with
10–12 hairs on ventral and lateral surfaces. Cercus in lateral view (Fig. 44N) short, rounded posteriorly, 0.43 times
as long as wide, and with numerous short to medium-long hairs. Spermatheca (Fig. 44O) large, nearly ovoid, 1.3
times as long as wide, well sclerotized except portion of juncture with duct unsclerotized, with weakly defined
surface patterns near base, and with internal setae; accessory ducts subequal in thickness to each other, and to major
duct.
Male. Body length 2.1–2.3 mm. Head. wider than thorax. Upper eye consisting of large facets in 16–18
vertical columns and in 16–18 horizontal rows. Clypeus brownish black, thickly white pruinose and brightly shiny
when illuminated at certain angles, sparsely covered with dark brown hairs along and near lateral margins (most of
central portion bare). Antenna composed of scape, pedicel and nine flagellomeres, dark brown to brownish black
except scape and pedicel dark yellow, base of first flagellomere yellow, and rest of first flagellomere light brown;
first flagellomere elongate, 1.5–1.7 times length of second one. Maxillary palp composed of five segments, light to
medium brown except first and second segments grayish yellow; proportional lengths of third, fourth, and fifth
segments 1.0:1.2–1.3:2.3; third segment (Fig. 45A, B) of normal size; sensory vesicle (Fig. 45A, B) globular or
ellipsoidal, 0.1–0.2 times length of third segment, and with small opening. Thorax. Scutum black, with whitish
pruinose pattern (brightly iridescent when illuminated at certain angles), i.e., anterior pair of large triangular spots
with pointed posteromedial apex on shoulders, narrow band along each lateral margin and large transverse spot
entirely covering prescutellar area, all these spots connected; scutum uniformly and moderately covered with
brassy recumbent short hairs interspersed with dark-brown long upright hairs on prescutellar area. Scutellum black,
with several dark-brown long upright hairs. Postnotum black, whitish pruinose (brilliantly iridescent when
illuminated at certain angles) and bare. Pleural membrane bare. Katepisternum brownish black, bare. Legs. Color
almost as in female except following characters: Fore tibia (Fig. 45C) medium brown except median outer surface
white and apical cap dark brown. Fore basitarsus greatly dilated, 5.2–5.5 times as long as its greatest width. Mid
femur entirely darkened. Mid and hind tibiae dark brown except base yellowish white. Hind basitarsus (Fig.45D)
yellowish white except apical one-third medium brown, and slightly widened toward apical one-fourth, 5.0–5.2
times as long as wide, and 0.7–0.8 and 0.7–0.8 times as wide as greatest widths of hind tibia and femur,
respectively; calcipala (Fig. 45D) developed, small, little shorter than its basal width, and 0.3 times as wide as
greatest width of basitarsus; pedisulcus (Fig. 45D) well developed. Wing. Length 1.6 mm. Other characters as in
female including bare subcosta. Halter. White except base light brown. Abdomen. Basal scale brownish-black,
with fringe of dark-brown long hairs. Dorsal surface of abdomen dark brown to black, moderately covered with
dark-brown short to medium-long hairs; segments 2, 5, 6 and 7 each with pair of whitish pruinose spots (brightly
iridescent when illuminated at certain angles) dorsolaterally, those on segment 2 broadly connected in middle to
each other. Genitalia. Coxites, styles and ventral plate in ventral view as in Fig. 45E. Style in ventral view (Fig.
45E) elongate, with inner margin sinuous, with slender subapical spine; style in ventrolateral view (Fig. 45F) 1.9
times length of coxite, 2.8 times as long as greatest width near basal one-fourth; style in medial view (Fig. 45G)
flattened dorso-ventrally except basal half gently produced dorsally, which is densely covered with fine short hairs.
Ventral plate in ventral view (Fig. 45E) Y-shaped, with body narrowed posteriorly; arms medium-long and stout,
widely separated from each other; ventral plate in lateral view (Fig. 45H) with body slightly curved ventrally, and
with serrated posterior margin; ventral plate in caudal view (Fig. 45I) with body parallel-sided and rounded
ventrally, bare, with four teeth in each of two vertical rows on posterior surface. Median sclerite (Fig. 45H, J)
weakly sclerotized, plate-like, wide, with blunt apical tip having incision medially. Paramere (Fig. 45K) with wide
base having several hooks apically. Aedeagal membrane (Fig. 45K) moderately covered with minute setae; dorsal
plate (Fig. 45L) horizontal bar-like, weakly sclerotized (except medial portion widely unsclerotized). Abdominal
segment 10 (Fig. 45M, N) without distinct hairs on each side of posterior surface. Cercus (Fig. 45M, N) small,
rounded, with six to eight hairs.
FIGURE 45. Male of Simulium (Simulium) obliquum sp. nov. A & B, Third segments of maxillary palps with different shape
of sensory vesicles (right side; front view); C, Fore tibia (left side; outer view); D, Hind basitarsus and second tarsomere (left
side; outer view); E, Coxites, styles and ventral plate (ventral view); F & G, Styles (right side; F, ventrolateral view; G, medial
view); H, Ventral plate and median sclerite (lateral view); I, Ventral plate (caudal view); J, Median sclerite (caudal view); K,
Paramere and aedeagal membrane (left side; caudal view) ; L, Dorsal plate (caudal view); M & N, Abdominal segment 10 and
cerci (right side; M, lateral view; N, caudal view). Scale bars. 0.1 mm for C and D; 0.02 mm for A, B and E–N.

Pupa. Body length 2.2–2.4 mm. Head. Integument including antennal sheaths grayish yellow, bare except face
sparsely covered with small round tubercles; antennal sheath without any ridges or projections; frons with two
pairs of unbranched slender short trichomes arising close together (Fig. 46A); face with pair of unbranched short
trichomes (Fig. 46B), much longer than frontal trichomes. Thorax. Integument grayish yellow to ochreous, bare on
dorsal and dorsolateral surface of anterior half, and sparsely to moderately covered with small tubercles on dorsal,
dorsolateral and lateral surfaces of posterior half; thorax on each side with two medium-long trichomes
mediodorsally (Fig. 46C), two trichomes (one medium-long, one short) anterolaterally (Fig. 46D), one short
trichome mediolaterally (Fig. 46E), and three trichomes (one medium-long, two short) ventrolaterally (Fig. 46F).
Gill (Fig. 46G) composed of six thread-like short filaments arranged in three pairs (dorsal, middle and ventral)
arising from short common basal stalk; each pair short-stalked; all filaments nearly subequal in length (1.0–1.5
mm) to one another, though two filaments of ventral pair slightly shorter than others, and inner filament of middle
pair slightly longer than outer filament of middle pair; each filament of same thickness (or slightly widened as in
outer filaments of middle and ventral pairs) from base to basal one-third, then gradually tapered toward apex;
relative thickness of six filaments from dorsal to ventral when measured basally 1.0:0.9–1.1:0.7–1.0 :1.1–1.2:0.7–
0.8:0.8–0.9, inner filament of middle pair thickest and outer filament of ventral pair thinnest; all filaments medium
to dark brown, with annular ridges and furrows, densely covered with minute tubercles. Abdomen. Dorsally,
segment 1, basal half of segment 2 and narrow portion along anterior margin of segment 3 light brown, other
segments transparent except basal areas of spine-combs on segments 7–9 yellow; segment 1 with one unbranched
slender short seta (Fig. 46H) on each side; segment 2 with one unbranched short seta and five minute setae (Fig.
46I) on each side; segments 3 and 4 each with four distinct unbranched hooks and one unbranched minute seta on
each side; segments 5 and 6 lacking spine-combs; segments 7–9 each with distinct spine-combs in transverse row
and comb-like groups of minute spines, on each side; segment 9 without terminal hooks. Ventrally, segments 3–9
transparent, each (except segment 9) with comb-like groups of minute spines; segment 4 with few slender minute
setae on each side; segment 5 with pair of bifid stout hooks submedially and few unbranched minute setae on each
side; segments 6 and 7 each with pair of bifid inner and bifid outer stout hooks somewhat separated from each
other, and few unbranched short setae on each side. Grapnel-shaped hooklets absent on each side of segment 9.
Cocoon (Fig. 46J). Simple, wall-pocket-shaped, dull yellowish white, thinly woven, with large anterolateral
window (rarely divided into two open spaces) on each side; posterior two-thirds with floor; individual threads
invisible; 2.6–3.0 mm long by 1.0–1.2 mm wide.
Mature larva. Body length 3.5–4.2 mm. Body color creamy with color markings as follows: thoracic segment
1 with grayish-green transverse band though disconnected medioventrally, grayish green on anterior surface of
proleg (and usually faintly on posterior surface of proleg) (in three larvae, grayish-green portions mottled with
reddish-brown pigment); thoracic segment 2 faintly grayish green on dorsal surface and grayish green mottled with
reddish-brown pigment on ventral surface; thoracic segment 3 with faint to moderate reddish-brown transverse
band on dorsal and dorsolateral surfaces, and grayish green mottled with reddish-brown pigment on ventral
surface; abdominal segments 1–5 each with grayish-green transverse band mottled with reddish-brown pigment,
though dorsomedially disconnected; abdominal segments 6–9 grayish green entirely or partially mottled with
reddish-brown pigment on dorsal and dorsolateral surfaces and on ventral surface of segments 6 and 7, though
dorsomedial portion of segments 7 and 8 often unpigmented; covering of reddish-brown pigment incomplete in
some larvae so that one larva entirely light grayish green, and few larvae showing grayish-green ventral surface of
abdominal segments 1–5. Head. Cephalic apotome (Fig. 47A) yellow except posterior half widely darkened
medially leaving posterior one of mediolongitudinal spots and posterolateral spots pale, hence making these head
spots negative; anterior spot of mediolongitudinal spots and mediolateral spots usually indistinct. Lateral surface of
head capsule yellow except eye-spot region whitish and often somewhat darkened between eye-spot region and
posterior margin; eyebrow not darkened or slightly darkened; spot below eye-spot region rarely recognized as
positive, and large spots in front of posterior margin obscure. Ventral surface of head capsule (Fig. 47B) yellow
except each side of basal portion of postgenal cleft dark brown, and often slightly darkened medially and
posteriorly. Antenna composed of three articles and apical sensillum, slightly longer than stem of labral fan; length
ratio of three articles (from base to tip) 1.0:1.1:0.8–0.9. Labral fan with 40–42 primary rays. Mandible (Fig. 47C)
with mandibular serration composed of two teeth (one medium-sized, one small); main tooth at obtuse angle
against mandible on apical side; supernumerary serrations absent; comb-teeth decreasing in length from first to
third. Hypostoma (Fig. 47D) with nine anterior teeth, of which median and corner teeth subequal in length, and
longer than three intermediate teeth on each side; lateral margins sharply serrate apically; four hypostomal bristles
in row nearly parallel to lateral border on each side. Postgenal cleft (Fig. 47B) long, approaching posterior margin
of hypostoma leaving narrow postgenal bridge. Thorax and Abdomen. Histoblast of pharate pupal gill with six
short filaments. Thoracic and abdominal cuticle with distinct transverse striations and sparsely covered with minute
colorless flattened setae and short unbranched colorless setae (Fig. 47E); abdominal segment 9 with 8–10 short
unbranched colorless setae on each side of anal sclerite; thoracic segment 3 with two small protuberances (one
dorsally and one dorsolaterally) on each side; abdominal segments 1–4 each with six small protuberances (one
dorsally, one dorsolaterally, two laterally, one ventrolaterally, and one ventrally) on each side; abdominal segment 5
with four small protuberances (one dorsally, one dorsolaterally, two laterally), abdominal segment 6 with three
small protuberances (one dorsally, one dorsolaterally and one laterally) on each side; abdominal segment 7 with
two small protuberances (one dorsally and one dorsolaterally) on each side; abdominal segment 8 with one small
protuberance dorsally on each side; all protuberances cone-shaped except ventral ones rounded (Fig. 47F). Rectal
scales present. Rectal organ compound, each lobe with 9 or 10 finger-like secondary lobules. Anal sclerite X-
shaped, widely sclerotized basally, with short broad anterior arms 0.7 times as long as posterior ones, and four to
eight sensilla posterior to posterior arms. Last abdominal segment bulged ventrolaterally but lacking ventral
papillae. Posterior circlet with 70–74 rows of hooklets with up to 13 or 14 hooklets per row.
FIGURE 46. Pupa of Simulium (Simulium) obliquum sp. nov. A, Frontal trichomes; B, Facial trichome; C–F, Thoracic
side; outer view); H, Hair-like seta and minute seta on dorsal surface of abdominal segment 1; I, Hair-like seta, minute seta and
stout seta on dorsal surface of abdominal segment 2; J, Cocoon (lateral view). Scale bars. 0.5 mm for J; 0.1 mm for G; 0.02 mm
for A–F; 0.01 mm for H and I.

FIGURE 47. Larva of Simulium (Simulium) obliquum sp. nov. A & B, Head capsules (A, dorsal view; B, ventral view); C,
Mandible; D, Hypostoma; E, Transverse striations, dorsal protuberance and minute setae on dorsal surface of abdominal
segment 5; F, Entire body showing protuberances (lateral view). Scale bars. 0.5 mm for F; 0.2 mm for A and B; 0.02 mm for C,
D and E.
in a vial as [UMSRP: Vietnam 037, Holotype, Simulium (S.) obliquum, Female, coll. Vietnam, 9-XII-2015,
Takaoka et al.], reared from pupa collected from a river (width 8–10 m, flow moderate, bottom of pebbles, water
temperature 17˚C, exposed to the sun, elevation 162 m, 18˚58’45.918”N/109˚50’10.693”E), fast flowing in a
natural forest, Pu Mat, Nghe An Province, northern Vietnam, 9-XII-2015, H. Takaoka, M. Sofian-Azirun, C.D.
Chen, & K.W. Lau. PARATYPES: 15 females and 15 males, with their associated pupal exuviae and cocoons, and
10 mature larvae, in 80% ethanol, same data as those of the holotype labeled in each vial as [UMSRP: Vietnam
037, Paratype, Simulium (S.) obliquum, Female, coll. Vietnam, 9-XII-2015, Takaoka et al.].
Biological notes. The pupae and larvae of S. (S.) obliquum sp. nov. were collected from grasses trailing in the
current. Associated species were S. (S.) tani (complex) and S. (S.) sp. (S. striatum species-group).
Etymology. The specific name obliquum refers to the ovipositor valve with a slanting posterior margin. The
Latin adjective ‘obliquus’ means slanting.
Remarks. Simulium (S.) obliquum sp. nov. is assigned to the S. malyschevi species-group of the subgenus
Simulium, defined by Rubtsov (1956), in having the female claw with a small subbasal tooth (Fig. 44H), ovipositor
valve with the posterior margin oblique (Fig. 44J), male style without a basal or subbasal projection (Fig. 45G), and
ventral plate Y-shaped (ventral view), with a toothed posterior margin (Fig. 45E).
This new species is similar in the pupa and larva to S. (S.) siripoomense Takaoka & Saito described based on
the pharate female, pupae and larvae collected in Thailand (Takaoka and Saito 1996; Takaoka and Mulla 2000).
However, this new species is distinguished from S. (S.) siripoomense in the female by the length ratio of the
sensory vesicle against the third maxillary palpal segment (Fig. 44A) (0.2 in this new species but 0.4 in S. (S.)
siripoomense), and in the larva by the postgenal cleft not reaching the posterior border of the hypostoma (Fig. 47B)
(widely reaching the posterior border of the hypostoma in S. (S.) siripoomense).
This new species is also similar to S. (S.) hirtipanus Puri, which was described from the female, male and pupa
collected from India (Puri 1932c). This new species differs from the latter species by the following characters
(those of S. (S.) hirtipanus in parentheses): in the female by the frons:head ratio 1.0:3.2 (1.0:2.0), hind trochanter
yellowish white (black), genital fork with a strongly sclerotized projection-like posteromedial corner (Fig. 44K)
(without such sclerotized corner), and the paraproct covered with 10–12 stout hairs (18 stout hairs); in the male by
the hind trochanter yellowish white (black); and in the pupa by the presence of spine-combs on abdominal segment
9 (absent). This species is distinguished from other related species of the S. malyschevi species-group, which have
a cluster of branched stout hairs on female abdominal segment 7 and a pupal gill with six filaments, by the larval
abdomen with multiple pairs of protuberances (Fig. 47F).
4) Simulium multistriatum species-group
Simulium (Simulium) daoense Takaoka & Adler sp. nov.
Female. Body length 3.0–3.2 mm. Head. Narrower than thorax. Frons black, shiny, with several dark, long, stout
hairs along each lateral margin and four or five similar hairs just above lower margin; frontal ratio 1.3–
1.7:1.00:1.5–1.7; frons:head ratio 1.0:4.5–4.6. Fronto-ocular area well developed, short, directed laterally and
slightly upward. Clypeus black, white pruinose, slightly shiny when illuminated at certain angles, moderately
covered with dark-brown long stout hairs except small area near upper margin or mediolongitudinal area of upper
half somewhat widely bare. Labrum 0.8 times as long as clypeus. Antenna composed of scape, pedicel and nine
flagellomeres, medium to dark brown except scape, pedicel, base of first flagellomere yellow when viewed
dorsally, and scape, pedicel, flagellomeres 1 and 2 yellow when viewed ventrally. Maxillary palp with five
segments, proportional lengths of third, fourth, and fifth segments 1.0:1.0:2.1–2.3; third segment (Fig. 48A) of
normal size; sensory vesicle (Fig. 48A) of moderate size (0.4 times length of third segment) having opening of
moderate size near apex. Maxillary lacinia with 13 inner and 14 or 15 outer teeth. Mandible with 24–26 inner and
12–14 outer teeth. Cibarium (Fig. 48B) with blunt medial projection on posterior margin and without minute
processes near base of medial projection. Thorax. Scutum black, shiny when illuminated at certain angles, densely
covered with golden-yellow recumbent short hairs interspersed with several dark-brown long upright hairs on
prescutellar area; when illuminated in front and viewed dorsally, scutum white pruinose with five longitudinal non-
pruinose vittae (one narrow medial vitta, two wider submedial vittae and two wider sublateral vittae though
sublateral vittae somewhat wider than submedial ones), all vittae united with broad transverse band on prescutellar
area; when illuminated from behind, scutum having reversed color pattern. Scutellum dark brown, covered with
dark-brown upright long hairs and golden-yellow short hairs. Postnotum black, white pruinose when illuminated at
certain angles and bare. Pleural membrane bare. Katepisternum longer than deep, black, shiny and white pruinose
when illuminated at certain angles and bare. Legs. Foreleg: coxa whitish yellow; trochanter yellow; femur yellow
except apical cap medium brown on inner surface, and dark yellow except basal one-fifth yellow and apical cap
medium brown on outer surface (in one female, femur light brown except base yellow and apical cap medium
brown); tibia yellowish white except base yellow and apical cap brownish black, with shiny sheen widely on outer

FIGURE 48. Female of Simulium (Simulium) daoense sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
surface when illuminated at certain angles; tarsus black, with moderate dorsal hair crest; basitarsus greatly dilated,
5.5–5.9 times as long as its greatest width. Midleg: coxa brownish black; trochanter dark yellow with base
yellowish white; femur dark yellow with base yellow and apical cap medium brown (in one female, femur light
brown except basal half or little less yellow and apical cap medium brown, and in another female, femur light
brown with apical cap medium brown); tibia white to yellowish white except apical cap medium brown, and with
shiny sheen on posterior surface when illuminated at certain angles; tarsus dark brown except basal half to two-
thirds yellowish white. Hind leg: coxa dark brown; trochanter whitish yellow; femur dark yellow with base yellow
and apical cap medium brown (in one female, femur medium brown except basal half yellow and apical cap dark
brown, and in another female, femur light brown except basal one-sixth yellow and apical cap medium brown);
tibia white to whitish yellow except apical cap dark brown, and with shiny sheen on posterior surface when
illuminated at certain angles; tarsus grayish black except basal two-thirds or little less of basitarsus and basal half
of second tarsomere yellowish white; basitarsus (Fig. 48C) nearly parallel-sided, 5.7–6.5 times as long as wide, and
0.5–0.7 and 0.6 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 48C)
moderately developed, slightly shorter than basal width, and 0.4 times as wide as basitarsus; pedisulcus (Fig. 48C)
well developed; all tarsal claws simple. Wing. Length 2.8–3.0 mm. Costa with dark spinules and hairs; subcosta
haired except near apex bare; basal section of radius bare; R1 with dark spinules and hairs; R2 with hairs; hair tuft
on base of radius dark brown; basal cell absent. Halter. White with basal portion darkened. Abdomen. Basal scale
medium brown, with fringe of dark-brown long hairs. Dorsal surface of abdomen medium to brownish black
except basal half to two-thirds of segment 2 yellowish, with short dark hairs; tergite 2 shiny, white iridescent when
illuminated at certain angles, and tergites 6–8 shiny; ventral surface of segment 2 whitish and those of other
segments light brown; segment 7 with median large sternal plate. Terminalia. Sternite 8 (Fig. 48D) bare medially,
with 25–35 dark medium-long to long stout hairs on each lateral surface. Ovipositor valves (Fig. 48D) rounded
posteomedially, membranous, each moderately covered with microsetae and four or five short to medium-long
hairs, except portion along inner margin narrowly bare; inner margins divergent posteriorly, and so thin and
transparent that they may be overlooked. Genital fork (Fig. 48E) of inverted-Y form, with narrow well-sclerotized
stem; arms of moderate width, each with distinct short and wide projection having rounded apex directed
anterodorsally. Paraproct in ventral view (Fig. 48F) nearly quadrate, densely covered with minute setae and with
32–36 short to medium-long hairs on lateral and ventral surfaces except anterolateral corner widely bare;
anteromedial surface with six short sensilla; paraproct in lateral view (Fig. 48G) much protruded ventrally beyond
ventral margin of cercus, 0.7 times as long as wide. Cercus in lateral view (Fig. 48G) short, rounded posteriorly, 0.5
times as long as wide, and with numerous short to medium-long hairs. Spermatheca (Fig. 48H) large, nearly ovoid,
1.2 times as long as wide, well sclerotized except portion of juncture with duct unsclerotized, with no defined
surface patterns, and with internal setae; accessory ducts subequal in thickness to each other, and to major duct.
Male. Body length 3.6 mm. Head. Slightly wider than thorax. Upper eye medium brown, with large facets in
18 vertical columns and in 18 or 19 horizontal rows. Clypeus black, thickly white pruinose, sparsely or moderately
covered with dark-brown hairs. Antenna composed of scape, pedicel and nine flagellomeres, dark brown except
scape and pedicel dark yellow to light brown, and base of first flagellomere yellow; first flagellomere elongate,
1.8–2.0 times as long as second one. Maxillary palp grayish to dark brown, composed of five segments with
proportional lengths of third, fourth, and fifth segments 1.0:1.3:2.4; third segment (Fig. 49A) of moderate size, with
apex somewhat produced inward; sensory vesicle (Fig. 49A) of moderate size (0.3–0.4 times length of third
segment), ellipsoidal, and with opening of moderate size. Thorax. Scutum black, with white pruinose pattern, i.e.,
anterior pair of rectangular spots on shoulders extended posteriorly along lateral margins and connected to large
transverse spot entirely covering prescutellar area; these pruinose areas shiny and iridescent when illuminated at
certain angles; scutum uniformly and densely covered with golden-yellow recumbent short hairs interspersed with
dark-brown long upright hairs on prescutellar area. Scutellum brownish black, with several dark long upright hairs
and golden-yellow short hairs. Postnotum black, white pruinose when illuminated at certain angles and bare.
Pleural membrane bare. Katepisternum longer than deep, brownish black to black, shiny and white pruinose when
illuminated at certain angles, and bare. Legs. Foreleg: coxa whitish yellow; trochanter medium brown except base
yellowish; femur dark yellow with apical cap dark brown; tibia brownish black except middle one-third light
brown, though outer surface widely white except basal tip and apical cap, and with white sheen when illuminated at
certain angles; tarsus brownish black, with moderate dorsal hair crest; basitarsus moderately dilated, 6.0–7.6 times

FIGURE 49. Male of Simulium (Simulium) daoense sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
view); D, coxite (ventrolateral view); E & F, Styles (right side; E, medial view; F, ventrolateral view); G, Ventral plate and
median sclerite (lateral view); H, Ventral plate (caudal view); I, Median sclerite (caudal view); J, Paramere (right side; caudal
view); K, Aedeagal membrane (caudal view); L, Dorsal plate (caudal view); M & N, Abdominal segment 10 and cerci (right
side; M, lateral view; N, caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–N.
as long as its greatest width. Midleg: coxa brownish black; trochanter dark brown though extreme base yellow;
femur light brown except apical cap medium brown; tibia whitish yellow to yellow except apical cap medium
brown, and with white sheen widely on posterior surface; tarsus dark brown to brownish black except basal one-
third to half of basitarsus yellowish. Hind leg: coxa brownish black; trochanter yellow; femur medium brown
except basal one-third yellow and apical cap brownish black; tibia dark brown to brownish black except basal tip
yellow; tarsus (Fig. 49B) medium to dark brown except basal half or little less of basitarsus whitish yellow (though
base somewhat darkened) and basal one-third of second tarsomere whitish yellow; basitarsus (Fig. 49B) much
enlarged, spindle-shaped, gradually widened from base to apical one-third, then tapered to apex, 3.4–4.1 times as
long as its greatest width, and 0.8–1.0 and 1.1 times as wide as greatest widths of hind tibia and femur,
respectively; calcipala (Fig. 49B) small, 0.8 times as long as width at base, 0.3 times as wide as greatest width of
basitarsus; pedisulcus (Fig. 49B) well developed. Wing. Length 3.0 mm. Other characters as in female except
subcosta haired on basal three-fifths. Halter. White with basal portion darkened. Abdomen. Basal scale dark brown
to brownish black, with fringe of dark long hairs. Dorsal surface of abdomen medium brown to brownish black,
and covered with dark short hairs; segments 2 and 5–7 each with pair of silvery or bluish iridescent spots
dorsolaterally, those on segment 2 broadly connected in middle to each other. Genitalia. Coxites, styles and ventral
plate in ventral view as in Fig. 49C. Coxite in ventrolateral view (Fig. 49D) nearly quadrate, 0.9 times as long as
width. Style in medial view (Fig. 49E) spatulate dorsoventrally, 1.5 times as long as coxite, with long horn-like
basal protuberance having pointed apex in form of stout spine; style in ventrolateral view (Fig. 49F) elongate, 3.2
times as long as its greatest width at base, slightly tapered to middle, then slightly widened to apical one-fourth,
and nearly parallel-sided toward apex; Ventral plate in ventral view (Fig. 49C) with body nearly rectangular
(though somewhat narrowed basally, and rounded on each posterolateral corner), covered with many minute setae
on anterior and anterolateral surfaces of central portion, and several ridges on posterior surface; arms directed
forward and divergent from each other; ventral plate in lateral view (Fig. 49G) having ventromedial process
somewhat curved posteroventrally and with bare round apex, and with serrated posterior margin, and moderately
covered with microsetae on anterolateral surfaces; ventral plate in caudal view (Fig. 49H) nearly triangular, with
several ridges on posterior surface, having ventromedial process nearly parallel-sided. Median sclerite (Fig. 49G, I)
arising near anterior margin of ventral plate and directed dorsally, plate-like, widened from base toward apex, with
round apex, brown basally, but not so well sclerotized apically. Paramere (Fig. 49J) with several distinct hooks.
Aedeagal membrane (Fig. 49K) moderately covered with minute setae, and with weakly sclerotized dorsal plate in
form of horizontal bar (Fig. 49L). Abdominal segment 10 (Fig. 49M, N) without distinct hair on ventral and lateral
surfaces. Cercus (Fig. 49M, N) small, rounded, with 11 or 12 distinct hairs.
Pupa. Body length 3.4–3.6 mm. Head. Integument ochreous, densely and elaborately covered with small
round tubercles; antennal sheath bare except basal portion covered with small tubercles; frons with two unbranched
slender short trichomes (Fig. 50A) on each side; face with unbranched slender short trichome (Fig. 50B) on each
side. Thorax. Integument dark yellow to ochreous, densely and elaborately covered with small round tubercles;
thorax with three long anterodorsal trichomes (all unbranched, or two unbranched and one bifid or quadrifid, or one
unbranched and two bifid or trifid, or two bifid and one trifid) (Fig. 50C), two anterolateral trichomes (anterior
trichome unbranched, medium-long, and posterior trichome unbranched or rarely bifid, long) (Fig. 50D), one
unbranched medium-long mediolateral trichome (Fig. 50E), and three unbranched ventrolateral trichomes (one
short, slender, and two others long, stout) (Fig. 50F), on each side. Gill (Fig. 50G) with eight slender thread-like
short filaments in four pairs (though two middle pairs sharing short stalk) arising from short common basal stalk;
all pairs short-stalked; gill filaments divergent basally, upper filament of dorsal pair forming an angle of 90 degrees
or little more against lower filament of ventral pair when viewed laterally; filaments slightly shortened from dorsal
to ventral, with upper filament of dorsal pair longest (about 2.0 mm long), and lower filament of ventral pair
shortest (about 1.1 mm long); relative thickness of eight filaments from dorsal to ventral when basal portions were
compared 1.0:0.8–0.9:0.8:0.6–0.7:0.7–0.9:0.6–0.7:0.6:0.6; all filaments light to medium brown, tapered toward
apex (though nearly parallel-sided or slightly widened from base for short distance except upper filament of middle
inner pair), with distinct annular ridges and furrows forming definite reticulate surface patterns, and densely
covered with minute tubercles. Abdomen. Dorsally, segment 1 light brown and sparsely covered with minute
tubercles, and often segments 2–4 each light brown narrowly along anterior margin, other segments unpigmented
except basal portions of spine-combs on segment 8 yellow; segment 1 with one unbranched slender short seta (Fig.
50H) on each side; segment 2 with one unbranched slender short seta and five unbranched short setae, of which
four are stout (one is often bifid) (Fig. 50I), on each side; segments 3 and 4 each with four distinct hooked spines

FIGURE 50. Pupa of Simulium (Simulium) daoense sp. nov. A, Frontal trichomes; B, Facial trichome; C–F, Thoracic
side; outer view); H, Hair-like seta on dorsal surface of abdominal segment 1; I, Hair-like seta, minute seta and stout setae on
dorsal surface of abdominal segment 2; J & K, Cocoons (J, dorsal view; K, lateral view). Scale bars. 1.0 mm for J and K ; 0.1
mm for G; 0.02 mm for A–F, H and I.
and one unbranched short seta on each side; segment 8 with distinct spine-combs in transverse row on each side;
segments 5, 6, 7 and 9 lacking spine-combs; segments 6–9 each with comb-like groups of minute spines on each
side; segment 9 without terminal hooks. Ventrally, segments 3–9 unpigmented, each (except segment 9) with
comb-like groups of minute spines; segment 4 with few unbranched slender minute setae on each side; segment 5
with pair of bifid stout hooks submedially and few unbranched short setae on each side; segments 6 and 7 each with
pair of bifid inner and unbranched or bifid outer stout hooks somewhat separated from each other, and few
unbranched short setae on each side. Grapnel-shaped hooklets absent on each side of segment 9. Cocoon (Fig. 50J,
K). Wall-pocket-shaped, tightly and thickly woven, ochreous, with large anterolateral window on each side, and
not extended ventrolaterally; individual threads invisible; 4.3–4.9 mm long by 1.8–2.2 mm wide.
Mature larva. Body length 6.0–6.2 mm. Body light brown except thorax dark gray to grayish black. Head. In
one larva, cephalic apotome (Fig. 51A) yellowish white on anterior two-fifths and yellow on posterior three-fifths,
with darkened narrow area along posterior margin; head spots faintly positive though posterior spot of
posterolateral spots on each side obscured; lateral surface of head capsule yellow except eye-spot region yellowish
white and eyebrow darkened and area between eye-spot and posterior margin somewhat darkened; spots in front of
posterior margin obscured or faintly negative except one small spot faintly positive; one small spot below eye-spot
region faintly positive; ventral surface of head capsule (Fig. 51B) yellow except submedian areas somewhat
darkened longitudinally; spots on each side of postgenal cleft faintly positive; in another larva, cephalic apotome
yellowish white on anterior half or little more, and light brown on rest with darkened narrow area along posterior
margin; head spots obscured though anterior spot of mediolongitudinal spots and both mediolateral spots faintly
positive; lateral surface of head capsule light brown except eye-spot region yellowish white and area between eye-
spot region and posterior margin somewhat darkened; spots in front of posterior margin obscured, and one small
spot below eye-spot region moderately positive; ventral surface of head capsule light brown except postgenal
bridge yellow; spots on each side of postgenal cleft obscured. Head capsule sparsely covered with minute setae on
dorsal, lateral and ventral surfaces. Antenna composed of three articles and apical sensillum, slightly longer than
stem of labral fan; length ratio of three articles (from base to tip) 1.0:1.3–1.4:0.7. Labral fan with 42 primary rays.
Mandible (Fig. 51C) with serration composed of two teeth (one medium-sized, one small); main tooth at obtuse
angle against mandible on apical side; supernumerary serrations absent; comb-teeth decreasing in length from first
to third. Hypostoma (Fig. 51D) with nine anterior teeth, of which median and corner teeth subequal in length to
each other; three intermediate teeth on each side shortest; lateral margins moderately serrate apically; five or six
hypostomal bristles per side divergent posteriorly from lateral margin. Postgenal cleft (Fig. 51B) large, rounded,
3.2 times as long as postgenal bridge; sheath of subesophageal ganglion weakly pigmented, forming indefinite
patterns. Cervical sclerites on each side composed of one medium-brown elliptical piece, not fused to occiput.
Thorax and Abdomen. Histoblast of pharate pupal gill with eight short filaments. Thoracic and abdominal cuticle
moderately covered with minute colorless setae interspersed with short similar setae, which are much longer than
minute setae (Fig. 51E), dorsally and dosolaterally; last segment of abdomen moderately covered with short
colorless setae on each side of anal sclerite. Rectal scales present. Rectal organ compound, each lobe with 17–22
finger-like secondary lobules. Anal sclerite X-shaped, with short broad anterior arms with forked apices, 0.7 times
as long as posterior ones and junction with deep unsclerotized incision posteriorly; 10–15 sensilla on base of anal
sclerite; nine sensilla posterior to posterior arms. Last abdominal segment lacking ventral papillae. Posterior circlet
with 88–93 rows of hooklets with up to 16–18 hooklets per row.
in a vial as [UMSRP: Vietnam 038, Holotype, Simulium (S.) daoense, Female, coll. Vietnam, 22-XII-2014,
Takaoka et al.], reared from a pupa collected from a small shallow stream (width 0.5 m, bottom of pebbles, water
temperature 7.0˚C, partially shaded, elevation 1,315 m, 22˚23’03.208”N/103˚50’58.990”E) moderately flowing
from a forest, Sapa, northern Vietnam, 22-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen &
K.W. Lau. PARATYPES: Eight females, two males (all reared from pupae), two pupae, one pupal exuviae and two
mature larvae, in 80% ethanol, same data as those of the holotype, labeled in each vial as [UMSRP: Vietnam 038,
Holotype, Simulium (S.) daoense, coll. Vietnam, 22-XII-2014, Takaoka et al.].
Biological notes. The pupae and larvae of S. (S.) daoense sp. nov. were collected from grass leaves and roots
trailing in the current. Associated species were S. (G.) hongthaii, S. (S.) sapaense sp. nov. and S. (S.) phuluense sp.
nov.

FIGURE 51. Larva of Simulium (Simulium) daoense sp. nov. A & B, Head capsules (A, dorsal view; B, ventral view); C,
Mandible; D, Hypostoma; E, Minute setae on dorsal surface of abdominal segment 7. Scale bars. 0.1 mm for A and B; 0.05 mm
for D; 0.02 mm for C and E.

Etymology. The species name daoense refers to the land in which the Dao tribe lives.
Remarks. Simulium (S.) daoense sp. nov. is assigned to the S. multistriatum species-group of the subgenus
Simulium, redefined by Takaoka and Davies (1996), based on the female scutum with five longitudinal vittae,
female ovipositor valves thin and unpigmented along inner margins (Fig. 48D), male style with elongate basal
protuberance (Fig. 49E), pupal gill with eight filaments (Fig. 50G), and cocoon with an anterolateral window on
each side (Fig. 50J, K).
This new species is characterized by the bare basal portion of the radial vein of the female wing, ellipsoidal
male sensory vesicle (Fig. 49A), unbranched mediolateral and ventrolateral trichomes of the pupal thorax (Fig.
50E, F), pupal gill filaments divergent at an angle of about 90 degrees when viewed laterally (Fig. 50G), and
cocoon with an enlarged window on each side (Fig. 50J, K).
This new species is similar in the arrangement of the pupal gill filaments to the following species but is
distinguished from them by the following characters (those of other species in parentheses); from S. (S.)
xialongtanense Chen, Luo & Yang described from Hubei, China (Chen et al. 2006) by the male upper-eye facets in
18 vertical columns and in 18 or 19 horizontal rows (in 12 vertical columns and 12 horizontal rows), male style
with a basal protuberance without minute spines (Fig. 49E) (with spines), and spine-combs on pupal abdominal
segment 8 (spine-combs on pupal abdominal segments 7 and 8); from S. (S.) bifengxiaense Huang, Zhang & Chen
described from Sichuan, China (Huang et al. 2013) by the female cibarium without tubercles (Fig. 48B) (with
about 15 tubercles) and male ventral plate with its posterior margin nearly straight when viewed ventrally (Fig.
49C) (posterior margin round); from S. (S.) chongquingense Zhu & Wang from Chong Quing, China (Zhu and
Wang 1995) by yellow hairs on the male scutum (dark hairs); from S. (S.) uncum Zhang & Chen from Guizhou,
China (Zhang and Chen 2001) by the male upper-eye facets in 18 vertical columns and in 18 or 19 horizontal rows
(in 16 vertical columns and 13 horizontal rows); from S. (S.) chainarongi Kuvangkadilok & Takaoka from
Thailand (Takaoka and Kuvangkadilok 1999) by the male ventral plate with its posterior margin nearly straight
when viewed ventrally (Fig. 49C) (posterior margin round) and wall-pocket-shaped cocoon (Fig. 50J, K) (shoe-
shaped cocoon); from S. (S.) demolaense Takaoka & Somboon from Bhutan (Takaoka and Somboon 2008) by the
ellipsoidal male sensory vesicle (Fig. 49A) (globular), male hind basitarsus with its greatest width at apical one-
third (Fig. 49B) (greatest width at the middle) and ventral plate covered with many minute setae (Fig. 49C) (a few
setae); from S. (S.) hirtinervis Edwards from Peninsular Malaysia (Takaoka and Davies 1995) by the bare basal
portion of the female radial vein (haired); from S. (S.) malayense Takaoka and Davies from Peninsular Malaysia
(Takaoka and Davies 1995) by the pupal head densely covered with tubercles (bare) and cocoon with a large
anterolateral window on each side (Fig. 50J, K) (without such a window); and from S. (S.) sakishimaense Takaoka
from the Ryukyu Islands, Japan (Takaoka 1977) by the male upper-eye facets in 18 or 19 horizontal rows (in 15
horizontal rows) and spine-combs on pupal abdominal segment 8 (spine-combs on pupal abdominal segments 7
and 8).
Four related Indian species, S. (S.) barraudi Puri, S. (S.) dentatum Puri, S. (S.) digitatum Puri, and S. (S.)
novolineatum Puri (Puri 1932b, 1933a), differ from this new species by the shape and hairlessness of the male
ventral plates.
The remaining species of the S. multistriatum species-group have the pupal gill filaments much diverged at an
angle of about 120 degrees when viewed laterally, or shoe-shaped cocoons, or enlarged basal fenestra at the base of
the pupal gill, all differing from those of this new species.
Simulium (Simulium) hirtinervis Edwards, 1928
Simulium (Simulium) hirtinervis Edwards, 1928: 63–64 (Female): Takaoka and Davies, 1995: 115–120 (Female, male, pupa
and larva).
Distribution. Peninsular Malaysia and Vietnam (Ha Tay and Hoa Binh).
Remarks. This species was originally described from Peninsular Malaysia based on females by Edwards
(1928) and later its female was redescribed and its male, pupa and larva were described by Takaoka and Davies
(1995). This species was recorded from Vietnam by Pham (1998).
Simulium (Simulium) lacduongense Takaoka & Ya’cob, 2015
Simulium (Simulium) lacduongense Takaoka & Ya’cob (in Takaoka et al., 2015a): 60–67 (Female, male, pupa and larva).

Remark. This species was described from Lam Dong Province, Vietnam (Takaoka et al. 2015a).
Simulium (Simulium) laui Takaoka & Sofian-Azirun, 2015
Simulium (Simulium) laui Takaoka & Sofian-Azirun, 2015 (in Takaoka et al., 2015a): 67–74 (Female, male, pupa and larva).

Remarks. Simulium (S.) laui was described from Lam Dong Province, Vietnam (Takaoka et al. 2015a). This
species is similar to S. (S.) lacduongense although it is distinguished from the latter by the bare basal portion of the
female radial vein (Takaoka et al. 2015a).

Simulium (Simulium) malayense Takaoka & Davies, 1995
Simulium (Simulium) malayense Takaoka & Davies, 1995: 120–123 (Female, pupa and larva).
Distribution. Peninsular Malaysia, Thailand and Vietnam (Bac Thai, Hoa Binh and Ha Tay).
Remark. This species was originally described from Peninsular Malaysia by Takaoka and Davies (1995) and
later recorded from three provinces in northernVietnam by Pham (1998, 1999).
5) Simulium nobile species-group
Simulium (Simulium) nodosum Puri, 1933

Simulium (Simulium) nodosum Puri. 1933b: 813–817 (Female, pupa and larva); Lewis, 1974: 34–37 (Female, male, pupa and
larva).
Specimens examined. Two males (reared from pupae) and five mature larvae, collected from a slow-flowing
stream (width 1.5–3.0 m, depth 10–15 cm, bottom sandy, water temperature 18.0˚C, exposed to the sun, elevation
73 m, 19˚30’45.497”N/105˚09’09.261”E), Chau Dinh, Quy Chau, Nghe An Province, northern Vietnam, 11-XII-
2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W. Lau.
Distribution. India, Thailand, Taiwan, South China, Vietnam (Ha Tay, Hoa Binh, Vinh Phuc, Lam Dong and
Nghe An).
Remarks. Simulium (S.) nodosum was originally described by Puri (1933b) and redescribed by Lewis (1974).
It was recorded from Vietnam by Crosskey and Howard (1997) and Pham (1998). The females of this species were
reported to be a human-biter and to transmit an Onchocerca species in Thailand (Takaoka et al. 2003).
6) Simulium striatum species-group
Simulium (Simulium) chiangmaiense Takaoka & Suzuki, 1984
Simulium (Simulium) chiangmaiense Takaoka & Suzuki, 1984: 38–39 (Female, male and pupa).
Distribution. Thailand, China (Hainan) and Vietnam (Lang Son).

Remarks. Simulium (S.) chiangmaiense was originally described from Thailand (Takaoka and Suzuki 1984). It
was recorded from Lang Son Province in northern Vietnam by Pham (1999).
Simulium (Simulium) jeffreyi Takaoka & Davies, 1995
Simulium (Simulium) jeffreyi Takaoka & Davies, 1995: 124–130 (Female, male, pupa and larva).
Distribution. Peninsular Malaysia and Vietnam (Lang Son).

Remarks. Simulium (S.) jeffreyi was originally described from Peninsular Malaysia (Takaoka and Davies
1995). It was recorded from Lang Son Province in northern Vietnam by Pham (1999).
Simulium (Simulium) nakhonense Takaoka & Suzuki, 1984
Simulium (Simulium) nakhonense Takaoka & Suzuki, 1984: 33–37 (Female, male, pupa and larva).
Distribution. Thailand, China (Hainan) and Vietnam (Bac Giang, Bac Thai, Ha Tay, Lang Son and Yen Bai).
Remarks. Simulium (S.) nakhonense was originally described from Thailand (Takaoka and Suzuki 1984). This
species was recorded from five provinces in northern Vietnam by Pham (1998, 1999).
Simulium (Simulium) quinquestriatum (Shiraki, 1935)
Stilboplax 5-striatum Shiraki, 1935: 27–33 (Female).

Simulium (Simulium) quinquestriatum (Shiraki), Anonymous, 1974: 192; Takaoka 1977: 205–209 (Female, male, pupa and
larva).
Distribution. China, Korea, Japan, Taiwan, Thailand and Vietnam (Bac Giang, Bac Thai, Ha Tay, Hoa Binh, Lang
Son and Yen Bai).
Remarks. Simulium (S.) quinquestriatum was originally described from Taiwan (Shiraki 1935). It was
recorded from six provinces in northern Vietnam by Pham (1998, 1999).
Simulium (Simulium) thailandicum Takaoka & Suzuki, 1984
Simulium (Simulium) thailandicum Takaoka & Suzuki, 1984: 37–38 (Male and pupa).
Distribution. China (Yunnan), Thailand, and Vietnam (Bac Giang, Ha Tay, Lang Son and Yen Bai).
Remarks. Simulium (S.) thailandicum was originally described from Thailand (Takaoka and Suzuki 1984). It
was recorded from four provinces in Vietnam by Pham (1998, 1999).
Simulium (Simulium) tavanense Takaoka & Sofian-Azirun sp. nov.
Female. Body length 2.2–2.4 mm. Head. Slightly narrower than thorax. Frons black, shiny with bluish reflection
when illuminated at certain angles, with several dark stout hairs along lateral margins and several hairs just above
lower margin; frontal ratio 1.4:1.0:1.2; frons:head ratio 1.0:3.6. Fronto-ocular area well developed, short, directed
laterally, and rounded apically. Clypeus brownish-black, slightly shiny and gray pruinose when illuminated at
certain angles, moderately covered with dark-brown medium-long hairs (though mediolongitudinal portion of
upper half widely bare). Labrum 0.7 times length of clypeus. Antenna composed of scape, pedicel and nine
flagellomeres, medium to dark brown except scape, pedicel, and first and second flagellomeres yellow when
viewed anteriorly (medium brown to dark brown except scape, pedicel and base of first flagellomere yellow when
viewed posteriorly). Maxillary palp with five segments, medium brown except first and second segments ochreous
or light brown, and third segment dark brown; proportional lengths of third, fourth, and fifth segments 1.0:1.0:2.5;
third segment (Fig. 52A) of normal size, with medium-sized ellipsoidal sensory vesicle (0.42 times length of third
segment) having opening of moderate size. Maxillary lacinia with 12 inner and 13 or 14 outer teeth. Mandible with
23 inner and 13 outer teeth. Cibarium (Fig. 52B) with 31 minute processes near posterodorsal margin. Thorax.
Scutum black, shiny, densely covered with whitish-yellow to yellow recumbent short hairs and sparsely with
several dark-brown long upright hairs on prescutellar area; scutum gray pruinose with five non-pruinose
longitudinal vittae (one medial, two submedial, and two lateral), medial and submedial vittae well defined from
anterior margin to posterior portion and submedial and lateral vittae united widely near anterior margin), all vittae
united with transverse non-pruinose band on prescutellar area, when illuminated in front and viewed dorsally;
scutum gray pruinose except four non-pruinose longitudinal vittae, when illuminated posteriorly and viewed
dorsally. Scutellum medium brown, covered with dark-brown upright long hairs and yellow short hairs. Postnotum
brownish black, shiny, white pruinose when illuminated at certain angles, and bare. Pleural membrane bare.
Katepisternum brownish black, longer than deep, shiny, white pruinose when illuminated at certain angles, and
bare. Legs. Foreleg: coxa and trochanter yellowish white; femur light brown except inner surface of basal portion
widely whitish yellow, and apical tip medium brown; tibia dark brown to brownish black; tarsus black, with
brownish black; trochanter light brown except base whitish; femur medium brown except apical cap dark brown;
tibia light to medium brown with apical cap dark brown and base whitish, and with whitish sheen widely on
posterior surface when illuminated at certain angles; tarsus light to medium brown except basal five-sixths of
basitarsus and base of second tarsomere whitish. Hind leg: coxa brownish black; trochanter yellowish white; femur
dark brown to brownish black except base whitish yellow; tibia dark brown except base yellowish white and with

FIGURE 52. Female of Simulium (Simulium) tavanense sp. nov. A, Third segment of maxillary palp with sensory vesicle (left
whitish sheen on basal half or more of posterior surface when illuminated at certain angles; tarsus medium brown
except basal two-thirds of basitarsus and basal half of second tarsomere yellowish white; basitarsus (Fig. 52C)
nearly parallel-sided, 6.3 times as long as wide, and 0.7 and 0.6–0.7 times as wide as greatest widths of hind tibia
and femur, respectively; calcipala (Fig. 52C) moderately developed, slightly shorter than wide, and 0.4 times as
wide as greatest width of basitarsus; pedisulcus (Fig. 52C) well developed. Claw simple, without tooth. Wing.
Length 2.2 mm. Costa with dark spinules and hairs; subcosta haired except near apex bare; basal section of radius
with four or five hairs on apical half or little more; R1 with dark brown spinules and hairs; R2 with dark-brown
hairs; hair tuft on base of radius dark brown; basal cell absent. Halter. White except base darkened. Abdomen.
Basal scale dark brown, with fringe of pale hairs. Dorsal surface of abdomen medium brown to brownish black,
with light to dark-brown short hairs; tergite 2 shiny and silvery iridescent when illuminated at certain angles and
tergites 6–9 shiny. Ventral surface of seventh segment with pair of weakly sclerotized submedian sternal plates.
Terminalia. Sternite 8 (Fig. 52D) with posterior margin concave medially in form of reversed-U shape, bare
medially, with 15–20 dark-brown medium-long to long stout hairs and 9–11 yellow short to medium-long hairs on
each lateral surface. Ovipositor valve (Fig. 52D) triangular, with ventrally produced lobe near inner margin,
membranous except narrow area along inner margin slightly sclerotized, covered with 23–26 short yellow hairs and
numerous microsetae; inner margins slightly sinuous, somewhat separated from each other. Genital fork (Fig. 52E)
of inverted-Y form, with narrow well-sclerotized stem; arms of moderate width, each with moderately sclerotized
lateral portion. Paraproct in ventral view (Fig. 52F) rounded, subequal in length to greatest width, strongly
pigmented on anterior surface, with 41–44 yellow and dark short to medium-long hairs on lateral and ventral
surfaces, and with six to eight short sensilla on anteromedial surface; paraproct in lateral view (Fig. 52G) nearly 0.5
times as long as wide, and much protruding ventrally beyond ventral margin of cercus. Cercus in lateral view (Fig.
52G) short, 0.5 times as long as wide, with numerous medium-long hairs, and rounded posteriorly. Spermatheca
(Fig. 52H) nearly ovoid, 1.2 times as long as greatest width, well sclerotized except portion of junction with duct
widely unsclerotized, without definite reticulate patterns on its surface; internal setae present; accessory ducts
subequal in thickness to each other, and slightly thicker than major duct.
Male. Body length 2.9 mm. Head. Nearly as wide as thorax. Upper-eye large facets in 18 vertical columns and
19 horizontal rows. Clypeus black, thickly white pruinose and iridescent when illuminated at certain angles, with
dark-brown hairs along and near lateral margins (most of central portion bare). Antenna composed of scape,
pedicel and nine flagellomeres, dark brown except scape and pedicel light brown and base of first flagellomere
yellow; first flagellomere elongate, 1.6 times length of second one. Maxillary palp with five segments, medium-
brown except first and second segments ochreous and fifth segment grayish light-brown; proportional lengths of
third, fourth, and fifth segments 1.0:1.3:2.9; third segment (Fig. 53A) of normal size; sensory vesicle (Fig. 53A)
ellipsoidal, 0.2–0.3 times length of third segment, and with small opening. Thorax. Scutum black, with whitish-
pruinose pattern, i.e., anterior pair of large spots on shoulders extended posteriorly along lateral margins and
connected to large transverse spot entirely covering prescutellar area, anterior pair of large spots divided into
anterior half and posterior half, either of which disappears depending on direction of lights; all these spots
brilliantly iridescent when illuminated at certain angles; scutum uniformly and moderately covered with yellow
recumbent short hairs and with several dark-brown long upright hairs on prescutellar area. Scutellum medium
brown, with dark-brown long upright hairs and yellow short hairs. Postnotum brownish black, shiny and whitish
pruinose when illuminated at certain angles and bare. Pleural membrane bare. Katepisternum longer than deep,
brownish black, and bare. Legs. Foreleg: coxa yellowish white; trochanter light brown; femur light brown with
apical tip medium brown; tibia medium brown to brownish black; tarsus black, with moderate dorsal hair crest;
basitarsus greatly dilated, 5.8–6.3 times as long as its greatest width. Midleg: coxa brownish black; trochanter
medium brown except base whitish; femur medium brown with apical cap dark brown; tibia medium to dark brown
except extreme base yellowish white; tarsus medium brown except basal two-thirds of basitarsus yellowish white.
Hind leg: coxa brownish black; trochanter yellowish white; femur medium brown except base whitish yellow and
apical cap dark brown; tibia dark brown to brownish black except base yellowish white; tarsus medium brown
except basal half of basitarsus and basal one-third of second tarsomere whitish yellow; basitarsus (Fig. 53B)
enlarged, slightly widened toward apical one-third, then slightly narrowed, 3.9–4.0 times as long as wide, and 0.8–
0.9 and 0.9–1.0 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 53B)
developed, small, slightly shorter than its basal width, and 0.3 times as wide as greatest width of basitarsus;
pedisulcus (Fig. 53B) well developed. Wing. Length 2.3 mm. Other characters as in female except subcosta with
hairs on basal half or little more, and basal portion of radius with four or five hairs on apical half or little less.
Halter. White except base darkened.Abdomen. Basal scale brownish black, with fringe of dark-brown long hairs.
Dorsal surface of abdomen dark brown to brownish black, moderately covered with dark-brown short to medium-
long hairs; segments 2, 5, 6 and 7 each with pair of whitish pruinose spots (brilliantly iridescent when illuminated
at certain angles) dorsolaterally, those on segment 2 broadly connected in middle to each other. Genitalia. Coxites,
styles and ventral plate in ventral view as in Fig. 53C; coxite in ventrolateral view (Fig. 53D) rectangular, 0.76
times as long as wide; style in ventrolateral view (Fig. 53E) 1.6 times length of coxite, 3.4 times as long as greatest
width near base, somewhat narrowed to little more than basal half, then slightly widened and nearly parallel-sided,
with subapical spine; style in medial view (Fig. 53F) somewhat flattened dorsoventally, with short basal
protuberance directed dorsomedially, with several cone-like spines along its anterior margin. Ventral plate in
ventral view (Fig. 53C) with body broad, with lateral margins gently convex, anterior margin deeply concave, and
posterior margin nearly straight or slightly concave; body bearing prominent median process sharply narrowed to
round tip; body covered with minute setae medially; arms short, stout, divergent from base; ventral plate in lateral

FIGURE 53. Male of Simulium (Simulium) tavanense sp. nov. A, Third segment of maxillary palp with sensory vesicle (left
view); D, coxite (right side; ventrolateral view); E & F, Styles (right side; E, ventrolateral view; F, medial view); G, Ventral
plate and median sclerite (lateral view); H, Ventral plate (caudal view); I, Median sclerite (caudal view); J, Paramere (right side;
caudal view); K, Aedeagal membrane (caudal view); L & M, Abdominal segment 10 and cerci (right side; L, lateral view; M,
caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–M.
view (Fig. 53G) with median process abruptly bent ventrally at nearly right angle; ventral plate in caudal view (Fig.
53H) in form of equilateral triangle, and bare. Median sclerite in lateral view (Fig. 53G) arising anterior to
anteromedian portion of ventral plate, and in middle curved dorsally; median sclerite in caudal view (Fig. 53I)
nearly parallel-sided. Paramere in caudal view (Fig. 53J) enlarged basally, with several hooks apically. Aedeagal
membrane in caudal view (Fig. 53K) sparsely covered with minute setae; dorsal plate not sclerotized. Abdominal
segment 10 (Fig. 53L, M) with two to five hairs and 10–12 shorter hairs on ventral surface and two to seven hairs
on lateral surface; cercus (Fig. 53L, M) small, with 16 or 17 distinct hairs.
Pupa. Body length 2.5–3.0 mm. Head. Integument light ochreous, moderately covered with relatively large
tubercles having few to several secondary projections on frons (Fig. 54A), and moderately with round or cone-like
small ones without secondary projections on face; frons with two pairs of unbranched or bifid short to medium-
long trichomes with straight apices (Fig. 54B); face with pair of unbranched medium-long trichomes with straight
apices (Fig. 54C), as long as or longer or shorter than frontal trichomes. Thorax. Integument light ochreous,
moderately covered with relatively large tubercles (similar to those on frons) except lateral surfaces and dorsal
surface of posterior half of thorax moderately covered with relatively smaller round or cone-like tubercles; thorax
on each side with three long trichomes with two to five branches (Fig. 54D) anterodorsally, two bifid or trifid long
trichomes (Fig. 54E) anterolaterally, one unbranched or bifid medium-long trichome (Fig. 54F) mediolaterally, and
three trichomes with coiled or straight apices (two unbranched or bifid long, one bifid short) (Fig. 54G)
ventrolaterally. Gill (Fig. 54H) with 10 thread-like filaments arranged as 2+[(1+2)+(2+1)]+2 from dorsal to ventral;
dorsal and ventral pairs with short stalk, two middle triplets with short stalk; all filaments subequal in length (0.7–
1.0 mm) and thickness, though ventral filament of ventral pair slightly thicker than other filaments (relative
thickness of filaments from dorsal to ventral when compared basally 1.0:1.0:0.8–1.0:0.8–0.9:0.9–1.0:0.8 –1.0:0.8–
1.0:1.0–1.1:1.0–1.3:1.1–1.3); all filaments light brown, covered with sharply-defined annular ridges and furrows
and densely covered with minute tubercles, relatively larger ones on ridges and smaller one on interridges.
Abdomen. Dorsally, all segments nearly transparent except segments 1 and 9 light yellow; segment 1 without
tubercles, with one unbranched short seta (Fig. 54I) on each side; segment 2 with one unbranched short seta and
five minute setae, of which four are stout and one slender (Fig. 54J) on each side; segments 3 and 4 each with four
distinct hooks and one short spinous seta on each side; all setae and hooks unbranched; segments 5, 6, 7 and 9
lacking spine-combs; segment 8 with distinct spine-combs in transverse row; segments 6–9 each with comb-like
groups of minute spines on each side; segment 9 with pair of cone-like terminal hooks (Fig. 54K). Ventrally, all
segment unpigmented except segment 9 yellowish; segments 4–8 each with comb-like groups of minute spines;
segment 5 with pair of bifid stout hooklets submedially and few unbranched minute setae on each side; segments 6
and 7 each with pair of bifid inner and unbranched outer stout hooklets somewhat separated from each other and
few unbranched minute setae on each side. Grapnel-shaped hooklets absent on each side of segment 9. Cocoon
(Fig. 54L). Light ochreous, shoe-shaped, with several small to large open spaces anterolaterally on each side;
posterior half with floor; posterior half thickly woven and individual threads almost invisible; 3.0–3.6 mm long by
1.2–1.5 mm wide; height 0.8–1.0 mm.
Mature larva. Body length 5.3–6.0 mm. Body dark gray. Abdomen in lateral view gradually widened from
segment 1 to segment 7, then narrowed to segment 9. Head. Cephalic apotome (Fig. 55A) whitish yellow on
anterior two-fifths, dark yellow to ochreous on posterior three-fifths, with area along posterior margin darkened;
head spots obscured except mediolateral spots often faintly negative; lateral surface of head capsule yellow to
ochreous (in one larva, widely medium brown), except eye-spot region whitish and areas above and posterior to
eye-spot region darkened; eyebrow distinct; spots near posterior margin and below eye-spot region obscured or
faintly negative; ventral surface of head capsule (Fig. 55B) yellow to ochreous; elongate spots on each side of
postgenal cleft faintly negative. Antenna composed of three articles and apical sensillum, slightly longer than stem
of labral fan; length ratio of three articles (from base to tip) 1.0:1.3:0.5–0.7. Labral fan with 53–56 primary rays.
Mandible (Fig. 55C) with mandibular serration composed of two teeth (one medium-sized, one small); main tooth
at right angle against mandible on apical side; comb-teeth gradually decreased in length from first to third;
supernumerary serrations absent (though one minute tooth present between two mandibular teeth in left mandible
of one larva). Hypostoma (Fig. 55D) with nine anterior teeth, of which median and corner teeth subequal in length
to each other, followed by three intermediate teeth on each side; lateral margins serrate apically; six or seven
hypostomal bristles divergent posteriorly from lateral border on each side. Postgenal cleft (Fig. 55B) large,
rounded, 3.7–4.0 times length of postgenal bridge; sheath of subesophageal ganglion weakly or moderately
pigmented. Cervical sclerites on each side composed of light-brown elliptical piece, not fused to occiput. Thorax

FIGURE 54. Pupa of Simulium (Simulium) tavanense sp. nov. A, Tubercles with minute secondary projections on frons; B,
Frontal trichomes; C, Facial trichome; D–G, Thoracic trichomes (D, anterodorsal; E, anterolateral; F, mediolateral; G
ventrolateral); H, Anterior part of thorax and gill filaments (right side; outer view); I, Hair-like seta on dorsal surface of
abdominal segment 1; J, Hair-like seta, minute seta and stout seta on dorsal surface of abdominal segment 2; K, Terminal hooks
(caudal view); L, Cocoon (lateral view). Scale bars. 1.0 mm for L ; 0.1 mm for H; 0.02 mm for A–G and I–K.
FIGURE 55. Larva of Simulium (Simulium) tavanense sp. nov. A & B, Head capsules (A, dorsal view; B, ventral view); C,
Mandible; D, Hypostoma; E, Protuberance and minute setae on dorsal surface of abdominal segment 7. Scale bars. 0.1 mm for
A and B; 0.05 mm for D; 0.02 mm for C and E.
and Abdomen. Histoblast of pharate pupal gill with 10 thread-like slender filaments. Thoracic cuticle sparsely
covered with minute colorless setae. Abdominal segments 1–7 each with pair of cone-like dorsolateral
protuberances (Fig. 55E). Abdominal cuticle sparsely covered with minute colorless setae (Fig. 55E), and
moderately covered with short colorless setae on each side of anal sclerite. Rectal scales present. Rectal organ of
three lobes, each with 12–15 finger-like secondary lobules. Anal sclerite X-shaped, with short broad anterior arms
0.65 times length of posterior ones and with forked apices; three or four sensilla on base of anal sclerite; four
sensilla posterior to posterior arms. Last abdominal segment not bulged laterally and lacking ventral papillae.
Posterior circlet with 134–138 rows of hooklets with up to 21–23 hooklets per row.
a vial as [UMSRP: Vietnam 039, Holotype, Simulium (S.) tavanense, Male, coll. Vietnam, 21-XII-2014, Takaoka et
al.], collected from a stream (width 1.0 m, depth 8 cm, bottom of pebbles, water temperature 13˚C, exposed to the
sun, elevation 1,048 m, 22˚18’33.255’’N/103˚53’12.129’’E), flowing near forest, Tavan, Sapa, Lao Cai Province,
northern Vietnam, 21-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
PARATYPES. One male (with its associated pupal exuviae and cocoon) and three mature larvae, in 80% ethanol,
same data as those of the holotype labeled in a vial as [UMSRP: Vietnam 039, Paratype, Simulium (S.) tavanense,
coll. Vietnam, 21-XII-2014, Takaoka et al.]; two females (all with their associated pupal exuviae and cocoons) in
80% ethanol, labeled in each vial as [UMSRP: Vietnam 039, Paratype, Simulium (S.) tavanense, Female, coll.
Vietnam, 21-XII-2014, Takaoka et al.], collected from a stream (width 1.0 m, depth 7 cm, bottom sandy, water
temperature 17˚C, exposed to the sun, elevation 999 m, 22˚18’23.788’’N/103˚53’42.780’’E), slowly flowing in a

grassland, Tavan, Sapa, Lao Cai Province, northern Vietnam, 21-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z.
Ya’cob, C.D. Chen & K.W. Lau.
Biological notes. The pupae and larvae of this new species were collected from grass leaves trailing in the
current. Associated species were S. (G.) hongthaii, S. (S.) daoense sp. nov., S. (S.) doipuiense (complex) and S. (S.)
phuluence sp. nov.
Etymology. The species name tavanense refers to the name of the locality, Ta Van, where this new species was
collected.
Remarks. Simulium (S.) tavanense sp. nov. is assigned to the S. striatum species-group, defined by Takaoka
and Davies (1996), by the unique shape of the female terminalia and male genitalia (Figs. 52D–H, 53C). This new
species is characterized by having a haired basal portion of the radius of the female and male, male scutum covered
with yellow short hairs, and pupal gill with 10 slender filaments of almost the same thickness, arranged as
2+[(1+2)+(2+1)]+2 filaments from dorsal to ventral (Fig. 54H), and larval body with paired protuberances (Fig.
55E).
None of known species of the S. striatum species-group have hairs on the basal portion of the radial vein and
on the subcosta in the male, as in this new species.
On the other hand, five species—S. (S.) grisescens Brunetti from India (Brunetti 1911; Puri 1932d), S. (S.)
quinquestriatum (Shiraki) from Taiwan (Shiraki 1935; Takaoka 1979), S. (S.) pingtungense Huang & Takaoka from
Taiwan (Huang and Takaoka 2008), S. (S.) taythienense Takaoka, Sofian-Azirun & Ya’cob from Vietnam (Takaoka
et al. 2014a) and S. (S.) wuzhishanense Chen from China (Chen 2003) —have a haired basal portion of the radial
vein in the female and a similar arrangement of the pupal gill filaments. Apart from the haired subcosta and haired
basal portion of the radial vein in the male, this new species is distinguished from S. (S.) grisescens by the male
hind basitarsus, which is 0.87 times as wide as the hind tibia (almost the same width as the hind tibia in S. (S.)
grisescens), and from the latter three species by the number of upper-eye facets, which are in 18 vertical columns
and 19 horizontal rows in this new species but in 16 horizontal rows in S. (S.) quinquestriatum, in 19 vertical
columns and 20 horizontal rows in S. (S.) pingtungense and in 12 vertical columns and 14 horizontal rows in S. (S.)
wuzhishanense. This new species differs from S. (S.) grisescens, S. (S.) pingtungense and S. (S.) taythienense by the
yellow short hairs on the male scutum (copper-colored short hairs in S. (S.) grisescens and brassy short hairs in S.
(S.) pingtungense and S. (S.) taythienense) and from S. (S.) pingtungense and S. (S.) wuzhishanense by the presence
of paired dorsal protuberances on the larval abdomen (protuberances absent in the latter two species).
This new species is distinguished from S. (S.) xuandai Takaoka, Sofian-Azirun & Ya’cob described from a
single male reared from a pupa from Vietnam (Takaoka et al. 2014a) by the male upper-eye facets in 18 vertical
columns and 19 horizontal rows (20 vertical columns and 20 horizontal rows in S. (S.) xuandai).
Simulium (Simulium) taythienense Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Simulium) taythienense Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014a): 353–361 (Female, male, pupa
and larva).
Distribution. Vietnam (Lam Dong and Vinh Phuc).

Remark. This species was described from Vinh Phuc Province in northern Vietnam (Takaoka et al. 2014a),
and also recorded from Lam Dong Province in southern Vietnam (Takaoka et al. 2015a).
Simulium (Simulium) xuandai Takaoka, Sofian-Azirun & Ya’cob, 2014
Simulium (Simulium) xuandai Takaoka, Sofian-Azirun & Ya’cob (in Takaoka et al., 2014a): 361–364 (Male and pupa).

Remarks. This species was described from a male reared from a pupa collected from Tam Dao, Vinh Phuc
Province, northern Vietnam (Takaoka et al. 2014a). The female and larva remain to be collected.
7) Simulium tuberosum species-group
Simulium (Simulium) congi Takaoka & Sofian-Azirun, 2015
Simulium (Simulium) congi Takaoka & Sofian-Azirun (in Takaoka et al., 2015a): 80–84 (Male, pupa and larva).

Remarks. This species was described from one male reared from a pupa, one pharate male and one mature
larva collected from Lam Dong Province, southern Vietnam (Takaoka et al. 2015a). The female of this species
remains to be collected.
Simulium (Simulium) doipuiense Takaoka & Choochote, 2005 (complex)
Simulium (Simulium) doipuiense Takaoka & Choochote, 2005c: 106–110 (Female, male, pupa and larva).
Simulium (Simulium) rufibasis (not Brunetti): Takaoka et al., 2014b: 747.
Specimens examined. One female, three males (all reared from pupae), collected from a stream (width 1.0 m,
depth 7 cm, streambed sandy, water temperature 17˚C, exposed to the sun, elevation 999 m, 22˚18’23.788’’N/
103˚53’42.780’’E), slowly flowing in a grassland, Tavan, Sapa, Lao Cai Province, northern Vietnam, 21-XII-2014,
by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau; one female, collected while flying around a
human, along a road traversing a mountain slope (elevation 997 m, 21˚28’104”N/105˚51’38.687”E), Tam Dao
National Park, Vinh Phuc Province, Vietnam, 9-IX-2013, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen
& K.W. Lau.
Distribution. Vietnam (Lao Cai and Vinh Phuc).
Remarks. Simulium (S.) doipuiense was described from Thailand (Takaoka and Choochote 2005c) and was
shown to be a species complex consisting of two cytoforms (A and B) by chromosome analysis (Tangkawanit et al.
2009). According to the original description (Takaoka and Choochote 2005c), the female of this species is
characterized by having a pair of clustered medium-long hairs on the ventral surface of abdominal segment 7,
antenna dark brown except the scape, pedicel and first flagellomere yellow, scutum covered with yellow short
hairs, hind tibia yellowish white except the apical two-fifths medium to dark brown, and stem of the genital fork
without a round apex. The females collected from Vietnam are almost identical to those of S. (S.) doipuiense from
Thailand except the following numerical differences (characters of S. (S.) doipuiense from Thailand in
parentheses): the relative length of the height of the frons against its narrowest width 1.1 (1.3), relative length of
the fore basitarsus against its greatest width 5.1 (4.9), relative width of the hind basitarsus against the hind tibia and
femur 0.8 (0.7) and 0.7(0.6), respectively. The males from Vietnam are almost identical to those of S. (S.)
doipuiense from Thailand including the number of upper-eye facets in 17 or 18 vertical columns and 18 horizontal
rows and the fewer number of spines on the basal protuberance of the style (only several spines), but there is a
difference in the relative length of the fore basitarsus against its greatest width (6.7–7.1 in the males from Vietnam
versus 5.9–6.1 in those from Thailand). The pupae from both countries are almost identical in many characters
including the frons moderately covered with round tubercles.
The female collected from Tam Dao National Park previously reported as S. (S.) rufibasis Brunetti by Takaoka
et al. (2014b) was reexamined. Our morphological examination shows that the female differs from that of S. (S.)
rufibasis by the antenna dark brown except the scape, pedicel and the base of the first flagellomere yellow, scutum
covered with yellow short hairs and hind basitarsus whitish yellow on the basal two-thirds and darkened on the
rest. According to the redescription of Puri (1932a), the female of S. (S.) rufibasis has the antennae almost entirely
darkened, scutum with brassy short hairs and hind basitarsus whitish yellow on the basal one-third to half and
darkened on the rest. The characters of the female are most similar to those of S. (S.) doipuiense, though its hind
tibia is yellowish white on the basal half and darkened on the rest, slightly differing from the color of the hind tibia
of S. (S.) doipuiense from Thailand which is yellowish white on the basal three-fifths and darkened on the rest. The
female collected from Tam Dao National Park is here treated as S. (S.) doipuiense (complex).

Simulium (Simulium) fuscicoxae Takaoka & Ya’cob sp. nov.
Simulium (Simulium) rufibasis (not Brunetti): Takaoka et al., 2015a: 85.
The female and its associated pupal exuviae and cocoon previously reported from Lam Dong Province, southern
Vietnam as S. (S.) rufibasis were reexamined. The result shows that this species is distinguished from S. (S.)
rufibasis and other related species. It is here described as a new species.
Female. Body length 2.0 mm. Head. Slightly narrower than thorax. Frons black, shiny, with several dark stout
hairs along lateral margins; frontal ratio 1.3:1.0:1.2; frons:head ratio 1.0:3.8. Fronto-ocular area well developed,
short, directed laterally, and rounded apically. Clypeus black, grayish pruinose, shiny, moderately covered with
dark-brown medium-long hairs though mediolongitudinal portion of upper half widely bare. Labrum 0.68 times
length of clypeus. Antenna composed of scape, pedicel and nine flagellomeres, brownish black except scape and
pedicel light to medium brown and base of first flagellomere dark yellow. Maxillary palp with five segments,
medium brown except third segment dark brown; proportional lengths of third, fourth, and fifth segments
1.0:1.2:2.2; third segment (Fig. 56A, B) of moderate size, with medium-sized ellipsoidal sensory vesicle (0.3 times
length of third segment) having large opening. Maxillary lacinia with 13 or 14 inner and 15 or 16 outer teeth.
Mandible with 26 inner and 12 outer teeth. Cibarium (Fig. 56C) with 82 minute processes, of which several are
vestigial, near posterodorsal margin. Thorax. Scutum black, shiny, white pruinose when illuminated at certain
angles, moderately covered with dark short fine hairs interspersed with several dark-brown long upright hairs on
prescutellar area. Scutellum dark brown, covered with dark-brown upright long and short hairs. Postnotum black,
shiny and white pruinose when illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum
longer than deep, brownish black, shiny and white pruinose when illuminated at certain angles, and bare. Legs.
Foreleg: coxa light brown; trochanter medium brown with base whitish; femur medium brown with apical cap
dark; tibia whitish except apical cap dark brown and base light brown, with median outer surface widely white and
with bright white sheen when illuminated at certain angles; tarsus brownish black, with moderate dorsal hair crest;
basitarsus greatly dilated, 5.1 times as long as its greatest width. Midleg: coxa brownish black; trochanter medium
except basal extreme whitish; femur medium brown except apical cap dark brown; tibia light brown except basal
two-fifths whitish and apical cap dark brown, and with whitish sheen widely on posterior surface when illuminated
at certain angles; tarsus light to medium brown except basal half of basitarsus whitish. Hind leg: coxa dark brown;
trochanter light brown; femur medium to dark brown except base light brown; tibia (Fig. 56D) medium to dark
brown except basal two-fifths white and with whitish sheen widely on posterior surface when illuminated at certain
angles; tarsus medium brown except little more than basal half of basitarsus (though base light brown) and basal
half of second tarsomere white; basitarsus (Fig. 56E) nearly parallel-sided, 5.4 times as long as wide, and 0.8 and
0.7 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 56E) moderately
developed, little shorter than wide, and 0.4 times as wide as greatest width of basitarsus; pedisulcus (Fig. 56E) well
developed; claw simple, without tooth. Wing. Length 1.9 mm. Costa with dark spinules and hairs; subcosta haired
except apical half bare; basal section of radius bare; R1 with dark spinules and hairs; R2 with dark hairs; hair tuft on
base of radius dark brown; basal cell absent. Halter. White except base darkened. Abdomen. Basal scale medium
brown, with fringe of dull hairs. Dorsal surface of abdomen medium brown to brownish black, with light to dark
brown short hairs; tergite 2 and 6–9 shiny when illuminated at certain angles. Ventral surface medium brown to
brownish black; segment 7 with pair of submedian sternal plates each bearing 18 or 19 stout medium-long to long
hairs, of which seven or eight hairs in cluster (Fig. 56F). Terminalia. Sternite 8 (Fig. 56G) well sclerotized and bare
medially, covered with eight to ten long and medium-long stout hairs and few short fine hairs on each side.
Ovipositor valve (Fig. 56G) nearly tongue-like, rounded posteromedially, membranous except inner margin weakly
sclerotized, densely covered with microsetae together with six or seven short fine hairs; inner margin slightly
sinuous. Genital fork (Fig. 56H) of inverted-Y form; stem slender and well sclerotized, with widened apex three
times as wide as middle of stem; arms of moderate width, each with moderately sclerotized projection directed
forwardly from lateral ridge. Paraproct in ventral view (Fig. 56I) pointed posteromedially, moderately concave
anterolaterally, and with strongly sclerotized anteromedial surface bearing few to several sensilla; paraproct in
lateral view (Fig. 56J) somewhat produced ventrally, 0.5 times as long as wide, covered with 21–24 short to
medium-long hairs and numerous microsetae on lateral and ventral surfaces. Cercus in lateral view (Fig. 56J) short,
rounded posteriorly, 0.5 times as long as its greatest width, and covered with medium-long and short hairs.
Spermatheca (Fig. 56K) ovoidal, 1.2 times as long as wide, moderately sclerotized except duct and narrow area of
juncture with duct unsclerotized, without discernible reticulate surface patterns; minute internal setae present;
accessory ducts subequal in thickness to each other and slightly thicker than main duct.
FIGURE 56. Female of Simulium (Simulium) fuscicoxae sp. nov. A & B, Third segments of maxillary palps with sensory
vesicles (right side; A, front view; B, lateral view); C, Cibarium (front view); D, Hind tibia (left side; outer view); E, Hind
basitarsus and second tarsomere (left side; outer view). F, Cluster of hairs on sternite 7 (left half); G, Sternite 8 and ovipositor
valve (right side only; ventral view); H, Genital fork (ventral view); I & J, Paraprocts and cerci (I, ventral view; J, lateral view).
K, Spermatheca. Scale bars. 0.1 mm for D and E; 0.02 mm for A–C and F–K.
Pupa. Body length 2.3 mm. Head. Integument including antennal sheaths yellow and moderately covered with
round tubercles (Fig. 57A); frons with two pairs of unbranched trichomes (anterior trichome medium-long,
posterior one long) (Fig. 57B); face with pair of unbranched medium-long trichomes (Fig. 57C), which are
subequal in length to anterior ones of frontal trichomes. Thorax. Integument yellow, moderately covered with
round tubercles except dorsal surface of posterior half covered with cone-shaped tubercles; thorax with two long
anterodorsal trichomes (posterior trichome broken, not measureable but probably as long as or slightly shorter than
anterior one) (Fig. 57D), two anterolateral trichomes (anterior trichome medium-long, posterior one long) (Fig.
57E), one medium-long mediolateral trichome (Fig. 57F), and three ventrolateral trichomes (one short and two
medium-long) (Fig. 57G), on each side; all trichomes unbranched. Gill (Fig. 57H) with six slender thread-like
filaments in three pairs (one dorsal, one middle and one ventral) arising from short common basal stalk; basal
fenestra small; all pairs short-stalked; stalks of dorsal and ventral pairs lying at angle of 90 degrees when viewed

FIGURE 57. Pupa of Simulium (Simulium) fuscicoxae sp. nov. A, Frons moderately covered with tubercles (left half; front
view); B, Frontal trichomes; C, Facial trichome; D–G, Thoracic trichomes (D, anterodorsal; E, anterolateral; F, mediolateral; G
ventrolateral); H, Anterior part of thorax and gill filaments (right side; outer view); I, Hair-like seta on dorsal surface of
abdominal segment 1; J, Hair-like seta and minute seta on dorsal surface of abdominal segment 2; K, Cocoon (lateral view).
Scale bars. 0.5 mm for K ; 0.1 mm for H; 0.02 mm for A–G, I and J.
laterally; filaments decreasing in length and thickness from dorsal to ventral, with longest filament 1.3 mm long
and shortest filament 0.8 mm long; dorsal filament of dorsal pair thickest, and two filaments of ventral pair
thinnest; relative thickness of each filament from dorsal to ventral when measured basally 1.0:1.0:0.8:0.8:0.7:0.7;
all filaments light brown, tapered toward apex, with moderately developed annular ridges and furrows forming
definite reticulate surface patterns except apical half of dorsal pair of filaments and middle and ventral pairs of
filaments with weakly or indistinct ridges and furrows, and densely covered with minute tubercles. Abdomen.
Dorsally, all segments unpigmented; segment 1 with one unbranched medium-long seta (Fig. 57I) on each side;
segment 2 with one unbranched short seta and five minute setae (Fig. 57J) on each side; segments 3 and 4 each
with four distinct hooked spines and one unbranched spinous short seta on each side; segments 5–7 lacking spine-
combs; segments 6 with comb-like groups of minute spines on each side; segments 8 and 9 each with distinct
spine-combs in transverse row and comb-like groups of minute spines on each side; segment 9 without terminal
hooks. Ventrally, segments 4–8 unpigmented and segment 9 somewhat yellowish; segments 3–8 each with comb-
like groups of minute spines; segment 4 with one unbranched or bifid stout hook and few slender minute setae on
each side; segment 5 with pair of bifid stout hooks submedially and few unbranched short setae on each side;
segments 6 and 7 each with pair of bifid inner and unbranched outer stout hooks somewhat separated from each
other, and few unbranched short setae on each side. Grapnel-shaped hooklets absent on each side of segment 9.
Cocoon (Fig. 57K). Wall-pocket-shaped, roughly woven anteriorly with many small open spaces in webs,
ochreous, and not extended ventrolaterally; individual threads visible; 2.0 mm long by 1.0 mm wide.
Male and mature larva. Unknown.
in a vial as [UMSRP: Vietnam 040, Holotype, Simulium (S.) fuscicoxae, Female, coll. Vietnam, 24-IV-2014,
Takaoka et al.], reared from a pupa collected from a small stream (width 1–2 m, water temperature 16.0˚C, exposed
to the sun, elevation 1,492 m, 12˚07’59.430”N/108˚35’42.001”E), slowly flowing in open land, 45 km from Dalat,
Da Chais, Lac Duong, Lam Dong Province, southern Vietnam, 24-IV-2014, by H. Takaoka, M. Sofian-Azirun, Z.
Ya’cob, C.D. Chen & K.W. Lau.
Biological notes. The pupa of this new species was collected from a slender tree root trailing in the current.
Associated species were S. (G.) confertum, S. (S.) chamlongi and S. (S.) nodosum.
Etymology. The species name fuscicoxae refers to the dark fore coxae of the female of this new species.
Remarks. Simulium (S.) fuscicoxae sp. nov. is assigned to the S. tuberosum species-group of the subgenus
Simulium, defined by Rubtsov (1956), based on the unpatterned female scutum, simple claw, paired clustered hairs
on the seventh sternite (Fig. 56F), and ordinary ovipositor valves (Fig. 56G), pupal gill with six slender filaments
(Fig. 57H) and simple wall-pocket-shaped cocoon (Fig. 57K).
This new species is characterized in the female by a sensory vesicle having a large opening (Fig. 56A), dark
short hairs on the scutum, darkened fore coxae, a pair of clustered stout hairs on the ventral surface of abdominal
segment 7 (Fig. 56F), and in the pupa by lacking spine-combs on the dorsal surface of abdominal segment 7. By
having the darkened female fore coxae and pupal abdominal segment 7 lacking spine-combs, this new species is
distinguished from seven of eight related species with a pair of clustered stout hairs on female sternite 7: S. (S.)
rufibasis Puri and S. (S.) ramosum Puri from India, S. (S.) doipuiense (complex), S. (S.) manooni Takaoka &
Choochote, S. (S.) weji Takaoka, all from Thailand, S. (S.) neorufibais Sun, and S. (S.) xinbinen Sun, both from
China (Chen and An 2003; Puri 1932a; Takaoka 2001a; Takaoka and Choochote 2005c). The remaining species, S.
(S.) setsukoae Takaoka & Choochote, from Thailand, which has the darkened female fore coxae, differs from this
new species by having a longer female sensory vesicle (0.4–0.5 times the length of the third maxillary palpal
segment) with a moderate-sized opening, and triangular ovipositor valves, each with a pointed posteromedian tip
(Takaoka and Choochote 2004a).
Simulium (Simulium) rosliramlii Takaoka & Chen sp. nov.
Female. Body length 2.5–2.9 mm. Head. Slightly narrower than thorax. Frons black, shiny, with several dark stout
hairs along lateral margins; frontal ratio 1.3:1.0:0.9–1.1; frons:head ratio 1.0:3.4–3.8. Fronto-ocular area well
developed, short, directed laterally, and rounded apically. Clypeus black, grayish pruinose, shiny, moderately
covered with dark-brown medium-long hairs though mediolongitudinal portion of upper half widely bare. Labrum

FIGURE 58. Female of Simulium (Simulium) rosliramlii sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Cibarium (front view); C, Hind tibia (left side; outer view); D, Hind basitarsus and second tarsomere
(left side; outer view). E, cluster of hairs on sternite 7 (right half); F, Sternite 8 and ovipositor valve (right side only; ventral
view); G, Genital fork (ventral view); H & I, Paraprocts and cerci (H, ventral view; I, lateral view). J, Spermatheca. Scale bars.
0.1 mm for C and D; 0.02 mm for A, B and E–J.
0.7–0.8 times length of clypeus. Antenna composed of scape, pedicel and nine flagellomeres, dark brown to
brownish black (though base of first flagellomere light brown). Maxillary palp with five segments, medium brown
except third segment dark brown; proportional lengths of third, fourth, and fifth segments 1.0:1.0–1.1:2.3–2.4;
third segment (Fig. 58A) of moderate size, with medium-sized ellipsoidal sensory vesicle (0.3–0.4 times length of
third segment) having medium-sized or large opening. Maxillary lacinia with11–14 inner and 11–17 outer teeth.
Mandible with 33–35 inner and 12 or 13 outer teeth. Cibarium (Fig. 58B) with 98–110 minute processes, of which
several are vestigial, near posterodorsal margin. Thorax. Scutum black, shiny, white pruinose when illuminated at
certain angles, moderately covered with yellow short fine hairs interspersed with several dark-brown long upright
hairs on prescutellar area. Scutellum dark brown, covered with dark-brown upright long and short hairs. Postnotum
black, shiny and white pruinose when illuminated at certain angles, and bare. Pleural membrane bare.
Katepisternum longer than deep, brownish black, shiny and white pruinose when illuminated at certain angles, and
bare. Legs. Foreleg: coxa yellow; trochanter medium brown with base yellow; femur medium brown with apical
cap dark brown; tibia whitish except apical cap dark brown and base light brown, with median outer surface widely
white and with bright white sheen when illuminated at certain angles; tarsus black, with moderate dorsal hair crest;
basitarsus moderately dilated, 6.4–7.2 times as long as its greatest width. Midleg: coxa brownish black; trochanter
medium to dark brown; femur dark brown except apical cap brownish black; tibia dark brown except basal two-
fifths whitish and apical cap brownish black, and with whitish sheen widely on posterior surface when illuminated
at certain angles; tarsus light to medium brown except basal half of basitarsus whitish yellow. Hind leg: coxa dark
brown; trochanter dark yellow to light brown; femur dark brown except base dark yellow to light brown; tibia (Fig.
58C) medium to dark brown except basal half or little more or less yellowish white, and apical cap brownish black,
and with whitish sheen widely on posterior surface when illuminated at certain angles; tarsus medium brown
except little less than basal two-thirds of basitarsus (though base light brown) and basal half of second tarsomere
yellowish white; basitarsus (Fig. 58D) nearly parallel-sided, 6.3–6.9 times as long as wide, and 0.7–0.8 and 0.7
times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 58D) moderately developed,
slightly shorter than wide, and 0.4 times as wide as greatest width of basitarsus; pedisulcus (Fig. 58D) well
developed; claw simple, without tooth. Wing. Length 2.5–2.7 mm. Costa with dark spinules and hairs; subcosta
haired except apical one-fourth to two-fifths bare; basal section of radius bare; R1 with dark spinules and hairs; R2
with dark hairs; hair tuft on base of radius dark brown; basal cell absent. Halter. White except base darkened.
Abdomen. Basal scale dark brown, with fringe of dull hairs. Dorsal surface of abdomen dark brown to brownish
black, with light to dark brown short hairs; tergite 2 and 6–9 shiny and iridescent when illuminated at certain
angles. Ventral surface medium brown to brownish black; segment 7 with pair of submedian sternal plates each
bearing 15–20 stout long hairs (Fig. 58E). Terminalia. Sternite 8 (Fig. 58F) well sclerotized and bare medially,
covered with four to seven long and medium-long stout hairs and few short fine hairs on each side. Ovipositor
valve (Fig. 58F) nearly tongue-like, rounded posteromedially, membranous except inner margin weakly
sclerotized, densely covered with microsetae together with seven or eight short fine hairs; inner margin slightly
sinuous. Genital fork (Fig. 58G) of inverted-Y form; stem slender and well sclerotized, with widened apex three to
four times as wide as middle of stem; arms of moderate width, each with moderately sclerotized projection directed
forwardly from lateral ridge. Paraproct in ventral view (Fig. 58H) pointed posteromedially, moderately concave
anterolaterally, and with strongly sclerotized anteromedial surface bearing six to eight sensilla; paraproct in lateral
view (Fig. 58I) somewhat produced ventrally, 0.7 times as long as wide, covered with 26–30 short to medium-long
hairs and numerous microsetae on lateral and ventral surfaces. Cercus in lateral view (Fig. 58I) short, rounded
posteriorly, 0.6 times as long as its greatest width, and covered with medium-long and short hairs. Spermatheca
(Fig. 58J) ovoidal, 1.2 times as long as wide, moderately sclerotized except duct and narrow area of juncture with
duct unsclerotized, without discernible reticulate surface patterns; minute internal setae present; accessory ducts
subequal in thickness to each other and slightly thicker than main duct.
in 20–22 vertical columns and in 21 or 22 horizontal rows. Clypeus black, thickly white pruinose, shiny when
illuminated at certain angles, covered with dark-brown hairs along lateral margins and near ventral margin (medial
portion of upper half bare). Antenna composed of scape, pedicel and nine flagellomeres, brownish black; first
flagellomere elongate, 1.6–1.9 times as long as second one. Maxillary palp grayish to dark brown, composed of
five segments with proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.3–2.6; third segment (Fig.
59A) of moderate size; sensory vesicle (Fig. 59A) small, ellipsoidal (0.2–0.3 times as long as third segment), and

FIGURE 59. Male of Simulium (Simulium) rosliramlii sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
view); D & E, Styles (right side; D, medial view; E, ventrolateral view); F, Ventral plate and median sclerite (lateral view); G,
Ventral plate (caudal view); H, Median sclerite (caudal view); I, Paramere and aedeagal membrane (left side; caudal view); J,
Dorsal plate (caudal view); K & L, Abdominal segment 10 and cerci (right side; K, lateral view; L, caudal view). Scale bars. 0.1
mm for B; 0.02 mm for A and C–L.
with small opening. Thorax. Scutum black, with white pruinose pattern, i.e., anterior pair of crescent spots (though
not sharply pointed posteriorly) on shoulders extended posteriorly along lateral margins and narrowly connected
near base of wings to large transverse posterior spot entirely covering prescutellar area; these pruinose areas silvery
when illuminated at certain angles; scutum uniformly and moderately covered with brassy recumbent short hairs
interspersed with dark-brown long upright hairs on prescutellar area. Scutellum brownish black, with several dark-
brown long upright hairs. Postnotum black, white pruinose when illuminated at certain angles and bare. Pleural
membrane bare. Katepisternum longer than deep, black, white pruinose when illuminated at certain angles, and
bare. Legs. Foreleg: coxa medium brown; trochanter medium to dark brown; femur medium to dark brown with
apical cap brownish black; tibia dark brown except median large portion widely white on outer surface, and with
white sheen widely on outer surface when illuminated at certain angles; tarsus brownish black to black, with
brownish black; trochanter dark brown; femur dark brown with apical cap brownish black; tibia dark brown except
apical cap brownish black and extreme base light brown; tarsus medium brown except basal one-third of basitarsus
light brown. Hind leg: coxa dark brown; trochanter medium brown; femur dark brown except apical cap brownish
black and extreme base light brown; tibia dark brown except apical cap brownish black and basal tip dark yellow to
light brown; tarsus (Fig. 59B) dark brown except basal one-third or little less of basitarsus whitish yellow (though
extreme base somewhat darkened) and basal one-third of second tarsomere whitish yellow; basitarsus (Fig. 59B)
much enlarged, gradually widened from base to apical one-third, then slightly narrowed to apex, 3.5–3.7 times as
long as its greatest width, and 1.1 and 1.3–1.4 times as wide as greatest widths of hind tibia and femur,
respectively; calcipala (Fig. 59B) small, slightly shorter than wide, 0.3 times as wide as greatest width of
basitarsus; pedisulcus (Fig. 59B) well developed. Halter. White except basal portion darkened. Wing. Length 2.5–
2.6 mm. Other characters as in female except subcosta bare. Abdomen. Basal scale brownish black, with fringe of
dark long hairs. Dorsal surface of abdomen brownish black to black, with dark short hairs; segments 2, 6 and 7
each with pair of silvery spots dorsolaterally, those on segment 2 broadly connected in middle to each other.
Genitalia. Coxites, styles and ventral plate in ventral view as in Fig. 59C. Coxite in ventrolateral view subquadrate,
0.7 times as long as wide. Style in medial view (Fig. 59D) 2.1 times as long as coxite, somewhat produced dorsally
near basal one-third forming low protuberance having several small cone-like spines on its dorsal surface. Style in
ventrolateral view (Fig. 59E) elongate, 2.7 times as long as its greatest width at base (or at basal one-third), slightly
constricted subbasally, tapered from basal one-third to apex. Ventral plate in ventral view (Fig. 59C) well
sclerotized, with body subquadrate (though slightly narrowed basally, and rounded on each posterolateral corner),
posterior margin somewhat concave, and with round projection directed ventrally, which is covered with many
minute setae; arms divergent from each other; ventral plate in lateral view (Fig. 59F) with posterior margin serrated
along basal half or little more; ventral plate in end view (Fig. 59G) rounded ventrally, with lateral margin serrated
along basal two-thirds, with numerous setae on posterior surface except each side of basal portion bare. Median
sclerite in lateral view (Fig. 59F) arising just anterior to ventral plate, directed dorsally; median sclerite in caudal
view (Fig. 59H) plate-like, gradually widened from base to apical one-third, then narrowed to apex, with round
apex. Paramere in caudal view (Fig. 59I) with several distinct hooks and several smaller ones. Aedeagal membrane
(Fig. 59I) moderately covered with minute setae, and with moderately sclerotized dorsal plate in form of horizontal
bar (Fig. 59J). Abdominal segment 10 (Fig. 59K, L) without hair or with one or two hairs on ventral surface and
three or four hairs on lateral surface on each side. Cercus (Fig. 59K, L) small, rounded, with six to nine distinct
hairs.
Pupa. Body length 2.6–3.2 mm. Head. Integument including antennal sheaths yellow and bare except face
moderately covered with round tubercles (Fig. 60A); frons with two pairs of unbranched long trichomes (Fig.
60A); face with pair of unbranched long trichomes (Fig. 60A), which are slightly shorter than frontal trichomes.
Thorax. Integument yellow and widely bare on dorsal surface of anterior half except narrow area along
mediolongitudinal line moderately covered with round tubercles; integument moderately covered with round
tubercles on ventrolateral surfaces near gills, and sparsely or moderately covered with cone-like tubercles on dorsal
surface of posterior half; thorax with two long anterodorsal trichomes (anterior trichome somewhat longer than
posterior one) (Fig. 60B), two long anterolateral trichomes (anterior trichome slightly thinner and shorter than
posterior one) (Fig. 60C), one short or medium-long mediolateral trichome (Fig. 60D), and three ventrolateral
trichomes (one short and two medium-long) (Fig. 60E), on each side; all trichomes unbranched except posterior
trichome of anterolateral trichomes in one pupa bifid. Gill (Fig. 60F) with six slender thread-like filaments in three

FIGURE 60. Pupa of Simulium (Simulium) rosliramlii sp. nov. A, Frons bare except near face moderately covered with
tubercles (left half; front view); B–E, Thoracic trichomes (B, anterodorsal; C, anterolateral; D, mediolateral; E ventrolateral); F,
Anterior part of thorax and gill filaments (right side; outer view); G, Hair-like seta on dorsal surface of abdominal segment 1; H,
Hair-like seta and minute seta on dorsal surface of abdominal segment 2; I, Terminal hooks (caudal view); J, Bifid hooklet on
ventral surface of abdominal segment 4. Scale bars. 0.1 mm for F; 0.05 mm for A; 0.02 mm for B–E and G–J.
pairs (one dorsal, one middle and one ventral) arising from short common basal stalk; basal fenestra small; dorsal
pair with medium-long stalk, and middle and ventral pairs short-stalked; stalks of dorsal and ventral pairs lying at
angle of 67–90 degrees when viewed laterally; filaments decreasing in length and thickness from dorsal to ventral,
with longest filament 1.6–2.2 mm long and shortest filament 1.2–1.5 mm long; dorsal filament of dorsal pair
thickest, and two filaments of ventral pair thinnest; relative thickness of each filament from dorsal to ventral when
measured basally 1.0:0.9:0.8:0.8:0.7:0.7; all filaments medium brown, tapered toward apices, with distinct annular
ridges and furrows forming definite reticulate surface patterns (except stalks and short basal portions of four
filaments of middle and ventral pair smooth), and densely covered with minute tubercles. Abdomen. Dorsally, all
segments unpigmented except segment 1 light yellowish; segment 1 with one unbranched medium-long seta (Fig.
60G) on each side; segment 2 with one unbranched short seta and five spinous minute setae (Fig. 60H) on each
side; segments 3 and 4 each with four distinct hooked spines and one unbranched spinous minute seta on each side;
segments 5 and 6 lacking spine-combs; segments 6 with comb-like groups of minute spines on each side; segments
7–9 each with distinct spine-combs in transverse row (though those on segment 7 small in number) and comb-like
groups of minute spines on each side; segment 9 having pair of small weakly developed terminal hooks (Fig. 60I)
or not. Ventrally, segments 4–8 unpigmented and segment 9 somewhat yellowish; segments 3–8 each with comb-
like groups of minute spines; segment 4 with one unbranched or bifid stout hook (Fig. 60J) and few slender minute
setae on each side; segment 5 with pair of bifid stout hooks submedially and few unbranched short setae on each
side; segments 6 and 7 each with pair of bifid inner and unbranched outer stout hooks somewhat separated from
each other, and few unbranched short setae on each side. Grapnel-shaped hooklets absent on each side of segment
9. Cocoon. Wall-pocket-shaped, moderately woven, thickly woven, ochreous, and not extended ventrolaterally;
individual threads visible; 3.0–3.5 mm long by 1.5–1.6 mm wide.
Mature larva. Unknown.
in a vial as [UMSRP: Vietnam 041, Holotype, Simulium (S.) rosliramlii, Female, coll. Vietnam, 20-XII-2014,
11.0˚C, exposed to the sun, elevation 1,728 m, 22˚22’22.719”N/103˚45’24.852”E) fast-flowing in forest, Lai Chau,
& K.W. Lau. PARATYPES: Five females and five males, in 80% ethanol, same data as those of the holotype,
labeled in each vial as [UMSRP: Vietnam 041, Paratype, Simulium (S.) rosliramlii, coll. Vietnam, 20-XII-2014,
Takaoka et al.].
Biological notes. The pupae of this new species were collected from a plastic sheet in the current. Associated
species were S. (G.) eshimai sp. nov., S. (N.) laichauense sp. nov. and S. (N.) tayense sp. nov.
Etymology. The species name rosliramlii is in honor of Dr. Rosli Ramli, Head of Institute of Biological
Sciences, University of Malaya, who supported our current research project on black flies in Vietnam.
Remarks. Simulium (S.) rosliramlii sp. nov. is assigned to the S. tuberosum species-group of the subgenus
Simulium, based on the male genitalia, of which the ventral plate is quadrate or subquadrate, having a round
projection directed ventrally with serrated posterolateral margins when viewed ventrally (Fig. 59C, F, G), and the
style bears several cone-like spines subbasally on the dorsal surface (Fig. 59D).
This new species is characterized in the female by having a pair of clustered stout hairs on the ventral surface
of abdominal segment 7 (Fig. 58E), and in the pupa by lacking tubercles on the frons (Fig. 60A) and anterodorsal
surface of the thorax. Among species of the S. tuberosum species-group, only S. (S.) pradyai Takaoka described
from Bhutan has a similar combination of these characters (Takaoka and Somboon 2008). However, this new
species is distinguished in the female from S. (S.) pradyai by the following characters (those of S. (S.) pradyai in
parentheses): length ratio of the height of the frons against its narrowest width 0.9–1.1 (1.2), antenna entirely
darkened (scape, pedicel and base of the first flagellomere yellow), ratio of the length of the fore basitarsus against
its greatest width 6.4–7.2 (5.4), and ratio of the length of the hind basitarsus against its greatest width 6.3–6.9 (5.6).
The pupa of this new species is distinguished from that of S. (S.) pradyai by the short stalk of the ventral pair of
filaments (about half as long as that of the dorsal pair) (Fig. 60F) (the stalk of the ventral pair of filaments is about
twice as long as that of the dorsal pair in S. (S.) pradyai).

Simulium (Simulium) giayense Takaoka & Lau sp. nov.
Female. Nearly as in female of S. (S.) rosliramlii sp. nov. except following characters. Body length 2.7 mm. Head.
Frontal ratio 1.3–1.4:1.0:1.2. Frons:head ratio 1.0:3.8–4.3. Labrum 0.6–0.7 times length of clypeus. Antenna dark
brown to brownish black except base of first flagellomere dark yellow. Maxillary palp: length ratio of third, fourth
and fifth segments 1.0:1.1:2.4; sensory vesicle oblong, 0.4 times as long as third segment, with opening of
moderate size. Maxilla with 12–14 inner and 15 or 16 outer teeth. Mandible with 33–35 inner and 13–15 outer
teeth. Cibarium with 82–107 minute processes. Legs. Fore basitarsus 5.9–6.0 times as long as its greatest width.
Hind basitarsus 6.3–6.5 times as long as wide, and 0.7–0.8 and 0.6 times as wide as greatest widths of hind tibia
and femur, respectively. Terminalia. Sternite 8 with nine stout long hairs together with four or five short hairs on
each side; ovipositor valves each with five or six short hairs; paraproct with 35 or 36 short to long hairs on ventral
and lateral surfaces.
Male. Nearly as in male of S. (S.) rosliramlii sp. nov. except following characters. Body length 3.0 mm. Head.
Nearly as wide as thorax. Upper eye with large facets in 20 vertical columns and in 20 horizontal rows. Antenna:
first flagellomere 1.7 times as long as second one. Maxillary palp: length ratio of third, fourth and fifth segments
1.0:1.2:2.8; sensory vesicle ellipsoidal, 0.2 times as long as third segment. Thorax. Scutum covered with yellow
short hairs. Legs. Fore basitarsus 8.0 times as long as its greatest width. Hind basitarsus (Fig. 61A) enlraged,
gradually widened from base to apical one-third, then narrowed, 3.4 times as long as its greatest width, 1.0 and 1.2
times as wide as greatest widths of hind tibia and femur, respectively; second tarsomere dark brown except basal
half yellowish white. Genitalia. Abdominal segment 10 with one or three short hairs on each lateral surface. Cerci
round, each nine short to medium-long hairs.
Pupa. Nearly as in pupa of S. (S.) rosliramlii sp. nov. except following characters. Body length 2.8–3.2 mm.
Head. Integument ochreous, densely covered with round tubercles (Fig. 61B). Thorax. Integument ochreous,
densely covered with round tubercles except dorsal surface of posterior portion moderately covered with small
tubercles. Mediodorsal trichomes (Fig. 61C) long, subequal in length to each other, and nearly as long as posterior
trichome of anterolateral trichomes (Fig. 61D); mediolateral trichome as in Fig. 61E, and ventrolateral trichomes as
in Fig. 61F. Gill (Fig. 61G) with six slender filaments in three pairs, each short-stalked, arising from short common
basal stalk; all filaments grayish brown, decreasing in length and thickness from dorsal to ventral; dorsal filament
of dorsal pair longest (2.4 mm long), and ventral filament of ventral pair shortest (1.3 mm long); relative thickness
of filaments from dorsal to ventral 1.0:0.9:0.8:0.8:0.7:0.7; stalk of dorsal pair of filaments lying at 60 degree
against stalk of ventral pair when viewed laterally. Abdomen. Terminal hooks absent.
in a vial as [UMSRP: Vietnam 042, Holotype, Simulium (S.) giayense, Female, coll. Vietnam, 20-XII-2014,
Takaoka et al.], reared from a pupa collected from a small stream (width 0.5 m, bottom rocky, water temperature
11.0˚C, exposed to the sun, elevation 1,750 m, 22˚21’43.110”N/103˚47’19.221” E) moderately flowing in a
secondary forest, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z.
Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: One female and one male (both reared from pupae), in 80%
ethanol, same data as those of the holotype labeled in a vial as [UMSRP: Vietnam 042, Paratype, Simulium (S.)
giayense, coll. Vietnam, 20-XII-2014, Takaoka et al.].
species were S. (M.) nigrofilum sp. nov., S. (N.) tayense sp. nov., S. (S.) laocaiense sp. nov., S. (S.) sapaense sp.
Etymology. The species name giayense refers to the land in which the Giay tribe lives.
Remarks. This new species is similar in the female to S. (S.) rosliramlii sp. nov. but is distinguished in the
male by the scutum covered with yellow short hairs, and in the pupa by the frons and dorsal surface of the anterior
half of the thorax densely covered with tubercles (Fig. 61B). This new species is distinguished from S. (S.)
rufibasis from India (Puri 1932a) by the female and male scuta covered with yellow short hairs (brassy or copper-
colored short hairs in S. (S.) rufibasis), female hind basitarsus yellowish white on the basal three-fifths (yellowish
white on the basal half or basal one-third in S. (S.) rufibasis), and male abdomen with segments 2, 6 and 7 each
having a pair of shiny dorsolateral patches (segments 2, 5, 6 and 7 each with such shiny patches in S. (S.) rufibasis).
The arrangement of the pupal gill filaments is similar to that of S. (S.) manooni Takaoka & Choochote from
Thailand (Takaoka and Choochote 2005c), from which this new species is distinguished by the pupal frons densely
covered with tubercles (Fig. 61B) (pupal frons moderately covered with tubercles in S. (S.) manooni).
FIGURE 61. Male and pupa of Simulium (Simulium) giayense sp. nov. A, Male; B–G, Pupa. A, Hind basitarsus and second
tarsomere (left side; outer view); B, Frons densely covered with tubercles (left half; front view); C–E, Thoracic trichomes (C,
mediodorsal; D, anterolateral; E, mediolateral; F ventrolateral); G, Anterior part of thorax and gill filaments (right side; outer
view). Scale bars.0.1 mm for A and G; 0.05 mm for B; 0.02 mm for C–F.

Simulium (Simulium) huense Takaoka & Ya’cob (Substitute name)
Simulium (Simulium) cavum Takaoka & Ya’cob (in Takaoka et al., 2015a) (preoccupied name): 74–79 (Male, pupa and larva).
The substitute name huense is here given for the species which was described as Simulium (Simulium) cavum
Takaoka & Ya’cob, 2015 from central Vietnam (Takaoka et al. 2015a). The name cavum is preoccupied by an
African species Simulium (Metomphalus) cavum Gibbins, 1938 (Adler and Crosskey 2016). Thus, cavum Takaoka
& Ya’cob, 2015 is a junior primary homonym of cavum Gibbins, 1938.
Remark. This species was described from Lam Dong Province, southern Vietnam (Takaoka et al. 2015a).
Simulium (Simulium) sapaense Takaoka & Low sp. nov.
Female. Body length 2.5 mm. Head. Slightly narrower than thorax. Frons black, shiny, with bluish reflection when
illuminated at certain angles, with several dark stout hairs along lateral margins; frontal ratio 1.5:1.0:1.4;
frons:head ratio 1.0:3.6. Fronto-ocular area well developed, short, directed laterally, and rounded apically. Clypeus
black, shiny, with bluish reflection when illuminated at certain angles, moderately covered with dark-brown
medium-long hairs though mediolongitudinal portion widely bare. Labrum 0.6 times length of clypeus. Antenna
composed of scape, pedicel and nine flagellomeres, dark brown, except scape, pedicel and first flagellomere yellow
when viewed ventrally, but dark brown except scape, basal half of pedicel and basal half of first flagellomere
yellow when viewed dorsally. Maxillary palp with five segments, medium brown except third segment dark brown;
proportional lengths of third, fourth, and fifth segments 1.0:1.1:2.4; third segment (Fig. 62A) of moderate size, with
medium-sized ellipsoidal sensory vesicle (0.3 times length of third segment) having medium-sized opening.
Maxillary lacinia with 15 or 16 inner and 18 outer teeth. Mandible with 25–27 inner and 12 outer teeth. Cibarium
(Fig. 62B) with 70 minute processes, of which several are vestigial, near posterodorsal margin. Thorax. Scutum
black, shiny with bluish reflection and white pruinose when illuminated at certain angles, moderately covered with
dark short fine hairs interspersed with several dark-brown long upright hairs on prescutellar area. Scutellum dark
brown, covered with dark-brown upright long and short hairs. Postnotum black, shiny and white pruinose when
illuminated at certain angles, and bare. Pleural membrane bare. Katepisternum longer than deep, brownish black,
shiny and white pruinose when illuminated at certain angles, and bare. Legs. Foreleg: coxa yellow; trochanter
yellow except dorsal surface partially medium brown; femur medium brown with apical cap dark brown, though
basal portion light brown on outer surface and widely yellowish on basal half on inner surface; tibia medium brown
except apical cap dark brown, with median outer surface widely white and with bright white sheen when
illuminated at certain angles; tarsus black, with moderate dorsal hair crest; basitarsus greatly dilated, 5.0 times as
long as its greatest width. Midleg: coxa brownish black; trochanter medium brown except base yellow; femur dark
brown except apical cap brownish black; tibia dark brown except base whitish yellow and apical cap brownish
black, and with whitish sheen widely on posterior surface when illuminated at certain angles; tarsus light to
medium brown except basal five-sixths of basitarsus and basal half of second tarsomere whitish yellow and fifth
tarsomere brownish black. Hind leg: coxa medium brown; trochanter yellow; femur medium to dark brown except
extreme base yellowish; tibia medium to dark brown except base whitish yellow, apical cap brownish black, and
with whitish sheen widely on posterior surface when illuminated at certain angles; tarsus medium brown except
basal two-thirds of basitarsus (though base medium brown) and basal half of second tarsomere whitish yellow;
basitarsus (Fig. 62C) nearly parallel-sided, 6.2 times as long as wide, and 0.8 and 0.6 times as wide as greatest
widths of hind tibia and femur, respectively; calcipala (Fig. 62C) moderately developed, nearly as long as wide,
and 0.4 times as wide as greatest width of basitarsus; pedisulcus (Fig. 62C) well developed; claw simple, without
tooth. Wing. Length 2.5 mm. Costa with dark spinules and hairs; subcosta haired except near apex bare; basal
section of radius bare; R1 with dark-brown spinules and hairs; R2 with dark-brown hairs; hair tuft on base of radius
dark brown; basal cell absent. Halter. White except base darkened. Abdomen. Basal scale dark brown, with fringe
of yellow hairs. Dorsal surface of abdomen medium brown to brownish black, with dark short hairs; tergite 2 and
6–9 shiny and iridescent when illuminated at certain angles. Ventral surface medium brown to brownish black;
segment 7 with large median sternal plate. Terminalia. Sternite 8 (Fig. 62D) well sclerotized and bare medially,
covered with 12 or 13 long and medium-long stout hairs and few short fine hairs on each side. Ovipositor valve
(Fig. 62D) nearly triangular, rounded posteromedially, membranous except inner margin weakly sclerotized,
densely covered with microsetae together with 12 or 13 short fine hairs; inner margin nearly straight or slightly
concave medially. Genital fork (Fig. 62E) of inverted-Y form; stem slender and well sclerotized; arms of moderate
width, each with strongly-sclerotized projection directed forwardly from lateral ridge. Paraproct in ventral view
(Fig. 62F) pointed posteromedially, moderately concave anterolaterally, and with strongly sclerotized anteromedial
surface; seven and four sensilla on anterodorsal concave portion and anteromedial surface, respectively; paraproct
in lateral view (Fig. 62G) somewhat produced ventrally, 0.7 times as long as wide, covered with 16 or 17 short to
medium-long hairs and numerous microsetae on lateral and ventral surfaces. Cercus in lateral view (Fig. 62G)
short, rounded posteriorly, 0.3 times as long as its greatest width, and covered with medium-long and short hairs.
Spermatheca (Fig. 62H) ovoidal, 1.3 times as long as wide, moderately sclerotized except duct and narrow area of
juncture with duct unsclerotized, without discernible reticulate surface patterns; minute internal setae present;
accessory ducts subequal in thickness to each other and slightly thicker than main duct.
FIGURE 62. Female of Simulium (Simulium) sapaense sp. nov. A, Third segment of maxillary palp with sensory vesicle (right

in 17 or 18 vertical columns and in 19 or 20 horizontal rows. Clypeus black, thickly white pruinose, silvery or
bluish, shiny when illuminated at certain angles, covered with dark-brown hairs along lateral margins and near
ventral margin (medial portion widely bare). Antenna composed of scape, pedicel and nine flagellomeres, medium
brown to brownish black except base of first flagellomere yellowish; first flagellomere elongate, 1.7 times as long
as second one. Maxillary palp grayish to dark brown, composed of five segments with proportional lengths of third,
fourth, and fifth segments 1.0:1.3:2.9; third segment (Fig. 63A) of moderate size; sensory vesicle (Fig. 63A) small,
ellipsoidal (0.3 times as long as third segment), and with small opening. Thorax. Scutum black, with white
pruinose pattern, i.e., anterior pair of crescent spots (though not sharply pointed posteriorly) on shoulders extended
posteriorly along lateral margins but disconnected near base of wings to large transverse posterior spot entirely
covering prescutellar area; these pruinose areas silvery or bluish iridescent when illuminated at certain angles;
scutum uniformly and moderately covered with brassy recumbent short hairs interspersed with dark-brown long
upright hairs on prescutellar area. Scutellum brownish black, with several dark-brown long upright hairs.
Postnotum black, white pruinose when illuminated at certain angles and bare. Pleural membrane bare.
Katepisternum longer than deep, black, white pruinose when illuminated at certain angles, and bare. Legs. Foreleg:
coxa yellow; trochanter light brown except base yellow; femur light to medium brown with apical cap dark brown;
tibia medium brown except median large portion widely white on outer surface, and with bright white sheen widely
on outer surface when illuminated at certain angles; tarsus brownish black to black, with moderate dorsal hair crest;
basitarsus greatly dilated, 5.2 times as long as its greatest width. Midleg: coxa brownish black; trochanter medium
brown except base yellow; femur medium brown with apical cap dark brown; tibia medium brown except apical
cap dark brown and extreme base whitish yellow; tibia with white sheen on posterior surface when illuminated at
certain angles; tarsus medium brown except basal three-fourths of basitarsus and base of second tarsomere whitish
yellow. Hind leg: coxa dark brown; trochanter dark yellow; femur dark brown except apical cap brownish black
and extreme base light brown; tibia dark brown except apical cap brownish black and basal tip yellow; tarsus (Fig.
63B) medium brown except little less than basal half of basitarsus (though extreme base darkened) and little less
than basal half of second tarsomere whitish yellow; basitarsus (Fig. 63B) much enlarged, gradually widened from
base to apical one-fourth, then slightly narrowed to apex, 3.9 times as long as its greatest width, and 0.9–1.0 and
1.0 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 63B) small, slightly
shorter than wide, 0.3 times as wide as greatest width of basitarsus; pedisulcus (Fig. 63B) well developed. Halter.
White except basal portion darkened. Wing. Length 2.1–2.5 mm. Other characters as in female except subcosta
bare. Abdomen. Basal scale brownish black, with fringe of dark long hairs. Dorsal surface of abdomen brownish
black to black, with dark short hairs; segments 2, 6 and 7 each with pair of silvery or bluish iridescent spots
dorsolaterally, those on segment 2 broadly connected in middle to each other. Genitalia. Coxites, styles and ventral
plate in ventral view as in Fig. 63C. Coxite in ventrolateral view (Fig. 63D) 0.8 times as long as wide. Style in
ventrolateral view (Fig. 63E) elongate, 3.0 times as long as its greatest width at basal one-third, nearly parallel-
sided from base to basal one-third, then tapered toward middle, and slightly tapered to apex; style in medial view
(Fig. 63F) 1.7 times as long as coxite, much produced dorsally near base forming round protuberance having
several small cone-like spines on its apex. Ventral plate in ventral view (Fig. 63C) well sclerotized, with body
subquadrate (though slightly narrowed basally, and rounded on each posterolateral corner), 0.6 times as long as
wide, posterior margin somewhat concave, and with round projection directed ventrally which is covered with
many minute setae; arms divergent from each other; ventral plate in lateral view (Fig. 63G) with posterior margin
weakly serrated along basal two-thirds; ventral plate in caudal view (Fig. 63H) rounded ventrally, with lateral
margin serrated along basal two-thirds, with numerous setae on posterior surface except most of basal portion bare.
Median sclerite in lateral view (Fig. 63G) arising just anterior to ventral plate, directed dorsally; median sclerite in
caudal view (Fig. 63I) plate-like, gradually widened from base to middle, then nearly parallel-sided to apex, with
round apex, brown basally, but not so well sclerotized apically. Paramere in caudal view (Fig. 63J) with several
distinct hooks and several smaller ones. Aedeagal membrane moderately covered with minute setae, and with
moderately sclerotized dorsal plate in form of horizontal bar (Fig. 63K). Abdominal segment 10 (Fig. 63L, M) with
three to five hairs on ventral surface and three to five hairs on lateral surface on each side. Cercus (Fig. 63L, M)
small, rounded, with five to seven distinct hairs.
Pupa. Body length 2.5–2.8 mm. Head. Integument dark brown except ventral surface light ochreous,
moderately covered with relatively larger tubercles having minute secondary projections on frons (Fig. 64A), and
relatively smaller tubercles on antennal sheaths and face; frons with two pairs of unbranched short slender
trichomes, of which one is shorter than other (Fig. 64B), arising close together; face with pair of unbranched short
FIGURE 63. Male of Simulium (Simulium) sapaense sp. nov. A, Third segment of maxillary palp with sensory vesicle (left
view); D, Coxite (right side; ventrolateral view); E & F, Styles (right side; E, ventrolateral view; F, medial view); G, Ventral
plate and median sclerite (lateral view); H, Ventral plate (caudal view); I, Median sclerite (caudal view); J, Paramere (left side;
caudal view) ; K, Dorsal plate (caudal view); L & M, Abdominal segment 10 and cerci (right side; L, lateral view; M, caudal
view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–M.

FIGURE 64. Pupa of Simulium (Simulium) sapaense sp. nov. A, Tubercles with minute secondary projections on frons; B,
Frontal trichomes; C, Facial trichome; D, Anterior portion of thorax near base of gill showing pit-like organ surrounded by
tubercles (dorsal view); E–H, Thoracic trichomes (E, anterodorsal; F, anterolateral; G, mediolateral; H ventrolateral); I,
Anterior part of thorax and gill filaments (right side; outer view); J, Hair-like seta on dorsal surface of abdominal segment 1; K,
Hair-like seta, minute seta and stout seta on dorsal surface of abdominal segment 2. Scale bars. 0.1 mm for I; 0.05 mm for D;
0.02 mm for B–H; 0.01 mm for A, J and K.
trichomes (Fig. 64C). Thorax. Integument dark brown except wing sheaths ochreous, moderately covered with
relatively larger tubercles (similar to those on frons though somewhat smaller) on anterior half, and relatively
smaller tubercles on posterior half, and with pit-like organ near base of each gill (Fig. 64D); thorax with two
anterodorsal trichomes of similar or different length (Fig. 64E), two anterolateral trichomes (Fig. 64F), one
mediolateral trichome (Fig. 64G), and three ventrolateral trichomes (Fig. 64H), on each side; all trichomes short
and unbranched. Gill (Fig. 64I) with six slender thread-like filaments in three pairs (one dorsal, one middle and one
ventral) arising directly from thorax and lacking common basal stalk; dorsal and middle pairs short-stalked, and
ventral pair sessile; lower filament of ventral pair directed downward and somewhat backward, then curved
forward, other filaments directed forward and downward, then curved forward and directed forward or somewhat
upward to various extent; two filaments of dorsal pair and inner filament of middle pair subequal in length to one
another (2.0 mm long), outer filament of middle pair and outer or upper filament of ventral pair subequal in length
to each other (1.9 mm long) and lower filament of ventral pair shortest (1.5 mm long); relative thickness of six
filaments from dorsal to ventral when compared basally 1.0:0.9:0.7:0.9:0.7:0.7; upper filament of dorsal pair
gradually tapered from base toward apex, other filaments nearly subequal in thickness from base to one-half or
more, then slightly tapered toward apex; all filaments dark brown, with distinct annular ridges and furrows forming
definite reticulate surface patterns, and densely covered with minute tubercles. Abdomen. Dorsally, all segments
unpigmented except segment 1 entirely medium brown; segment 1 with one unbranched short seta (Fig. 64J) on
each side; segment 2 with one unbranched short seta and five minute setae, of which three are stout (Fig. 64K), on
each side; segments 3 and 4 each with four distinct hooks and one unbranched spinous short seta on each side;
segment 5 and 6 lacking spine-combs; segments 7–9 each with distinct spine-combs in transverse row; segments 6–
9 each with comb-like groups of minute spines on each side; segment 9 without terminal hooks. Ventrally,
segments 3–8 unpigmented, each with comb-like groups of minute spines; segment 4 with one unbranched hook
and few slender minute setae on each side; segment 5 with pair of bifid hooks submedially and few unbranched
short setae on each side; segments 6 and 7 each with pair of bifid inner and unbranched outer hooks somewhat
separated from each other, and few unbranched short setae on each side. Grapnel-shaped hooklets absent on each
side of segment 9. Cocoon. Wall-pocket-shaped, thickly woven (in particular anterior margin), medium brown, not
extended ventrolaterally, and with thick anterodorsal margin; individual threads invisible; 2.5–2.7 mm long by 1.2–
1.3 mm wide.
a vial as [UMSRP: Vietnam 043, Holotype, Simulium (S.) sapaense, Male, coll. Vietnam, 20-XII-2014, Takaoka et
al.], reared from a pupa collected from a small stream (width 0.5–1.0 m, depth 7 cm, bottom rocky, water
temperature 9.0˚C, partially shaded, elevation 1,680 m, 22˚22’05.320”N/103˚47’34.403”E), slowly flowing from a
secondary forest, Sapa, Lao Cai Province, northern Vietnam, 20-XII-2014, by H. Takaoka, M. Sofian-Azirun, Z.
Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: One female, one pupal exuviae, same data as those of the holotype,
labeled in each vial as [UMSRP: Vietnam 043, Paratype, Simulium (S.) sapaense, coll. Vietnam, 20-XII-2014,
Takaoka et al.]; one male labeled in a vial as [UMSRT: Vietnam 043, Paratype, Simulium (S.) sapaense, Male, coll.
Vietnam, 20-XII-2014, Takaoka et al.] reared from a pupa collected from a stream (width 0.5 m, depth 10 cm,
streambed rocky, water temperature 11.0˚C, exposed to the sun, elevation 1,750 m, 22˚21’43.110”N/
103˚47’19.221”E) moderately flowing in a secondary forest, Sapa, Lao Cai Province, northern Vietnam, 20-XII-
2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau.
Biological notes. The pupae of this new species were collected from dead tree leaves in the current. Associated
species were S. (M.) nigrofilum sp. nov., S. (N.) tayense sp. nov., S. (S.) laocaiense sp. nov., S. (S.) giayense sp.
Etymology. The species name sapaense refers to the name of the district, Sapa, where this new species was
collected.
Remarks. Simulium (S.) sapaense sp. nov. is assigned to the S. tuberosum species-group of the subgenus
Simulium, based on the male genitalia (Fig. 63C). This new species is characterized by having the pit-like organ on
the thoracic integument near the base of the gill (Fig. 64D). Seven species of the S. tuberosum species-group, i.e., S.
(S.) brevipar Takaoka & Davies from Peninsular Malaysia and Thailand, S. (S.) tiomanense Takaoka, Sofian-
Azirun & Belabut from Peninsular Malaysia, S. (S.) tianchi Chen, Zhang & Yang from China, S. (S.) sigiti Takaoka
& Hadi from Java, S. (S.) yuphae Takaoka & Choochote from Thailand, and S. (S.) huense Takaoka & Ya’cob

(substitute name for cavum) and S. (S.) lowi sp. nov. from Vietnam, have a similar pit-like organ (Takaoka and
Davies 1995; Takaoka et al. 2012b, 2015a; Takaoka and Choochote 2005d; Takaoka and Hadi 1991; Chen et al.
2003b). This new species is distinguished in the male from S. (S.) tianchi and S. (S.) huense by having the upper-
eye facets in 17 vertical columns (in 15 vertical columns in S. (S.) tianchi and 21 vertical columns in S. (S.)
huense), and in the pupa from S. (S.) brevipar, S. (S.) huense, S. (S.) lowi sp. nov., S. (S.) sigiti, S. (S.) tiomanense
and S. (S.) yuphae by the vicinity of the pit-like organ covered with tubercles (Fig. 64D) (vicinity of the pit-like
organ is bare in the latter five species), and also from S. (S.) tianchi and S. (S.) sigiti by having six gill filaments of
different length and thickness (Fig. 64I) (six gill filaments are subequal in length and thickness to one another in
the latter two known species). Although whether its pupa has a similar pit-like organ is not known, S. (S.)
nigrifacies Datta from India (Datta 1974) differs from this new species by lacking spine-combs on abdominal
segment 7.
Simulium (Simulium) lowi Takaoka & Adler sp. nov.
Simulium (Simulium) brevipar (not Takaoka & Davies): Takaoka et al., 2014a: 365.
A chromosome analysis reveals that the species previously reported from Tam Dao National Park, northern
Vietnam as S. (S.) brevipar from Malaysia consists of two distinct species: S. (S.) yuphae from Thailand (Takaoka
and Choochote 2005d) and S. (S.) sp., which is clearly distinguished chromosomally from S. (S.) brevipar from
Malaysia (Adler et al. 2016). Our DNA analysis using the COII gene shows that the specimens from Vietnam are
distinct from S. (S.) brevipar from Malaysia (Low et al. unpublished data). A subsequent morphological
reexamination of two females and one male, together with their associated pupal exuviae, on which the record of S.
(S.) brevipar from Vietnam was based, shows that they are almost identical to those of S. (S.) brevipar but are
distinguished from the latter by a few features such as the shape of the female frons, relative size of the female
labrum against the clypeus, and shape of the male ventral plate and style. Based on the results from morphological,
molecular and chromosomal analyses, this species is here described as new.
Female. Body length 2.0 mm. Nearly similar to that of S. (S.) sapaense sp. nov. except following characters.
Head. Frontal ratio 1.4–1.5:1.0:1.3–1.4; frons:head ratio 1.0:3.4–3.5. Labrum 0.7–0.8 times length of clypeus.
Proportional lengths of third, fourth, and fifth segments 1.0:1.0:2.2; sensory vesicle medium-long (0.2–0.3 times
length of third segment) having small or medium-sized opening. Maxillary lacinia with 14–17 inner and 16–18
outer teeth. Mandible with 26 inner and eight or nine outer teeth. Cibarium with 67 minute processes, of which
several are vestigial, near posterodorsal margin. Legs. Foreleg: trochanter yellow; femur dark yellow basally,
becoming darker toward apex with apical cap dark brown, though widely yellowish on basal half on inner surface;
basitarsus greatly dilated, 4.8 times as long as its greatest width. Midleg: tarsus light brown except apical tips of
basitarsus and second tarsomere yellowish white and fifth tarsomere dark brown. Hind leg: tarsus medium brown
except basal seven-tenths of basitarsus (though base slightly darkened) and basal half of second tarsomere whitish
yellow, and fourth and fifth tarsomeres dark brown; basitarsus (Fig. 65A) nearly parallel-sided, 6.1 times as long as
wide, and 0.7 and 0.6 times as wide as greatest widths of hind tibia and femur, respectively. Wing. Length 1.9 mm.
Subcosta haired except apical one-fourth bare. Terminalia. Sternite 8 well sclerotized and bare medially, covered
with six to nine long and medium-long stout hairs and few short fine hairs on each side. Ovipositor valve densely
covered with microsetae together with seven to 15 short fine hairs. Paraproct in ventral view with six sensilla on
anteromedial surface; paraproct in lateral view 0.8 times as long as wide, covered with 19 short to medium-long
hairs and numerous microsetae on lateral and ventral surfaces. Cercus in lateral view short, rounded posteriorly, 0.5
times as long as its greatest width, and covered with medium-long and short hairs. Spermatheca ovoidal, 1.19–1.22
times as long as wide.
Male. Body length 2.3 mm. Nearly as in male of S. (S.) sapaense sp. nov. except following characters. Head.
Slightly wider than thorax. Upper eye medium brown, with large facets in 18 vertical columns and in 18 horizontal
rows. Antenna medium brown to brownish black except scape, pedicel dark yellow, base of first flagellomere
whitish yellow; first flagellomere elongate, 1.9 times as long as second one. Maxillary palp grayish to dark brown,
composed of five segments with proportional lengths of third, fourth, and fifth segments 1.0:1.2:2.4; third segment
of moderate size; sensory vesicle small, ellipsoidal (0.2 times as long as third segment), and with small opening.
FIGURE 65. Female, male and pupa of Simulium (Simulium) lowi sp. nov. (except D, F and J are characters of S. brevipar
from Peninsular Malaysia). A, Female; B–F, Male; G–J, Pupa. A, Hind basitarsus and second tarsomere (left side; outer view);
B, Hind basitarsus and second tarsomere (left side; outer view); C & D, Styles (right side; ventrolateral view); E & F, Ventral
plates (ventral view); G, Anterior portion of thorax near base of gill showing bare area surrounding pit-like organ (dorsal view);
H, Anterior part of thorax and gill filaments (right side; outer view); I & J, Basal portions of gill filaments (right side; outer
view). Scale bars. 0.1 mm for A, B and H; 0.05 mm for G, I and J; 0.02 mm for C–F.
Legs. Foreleg: trochanter yellow partially darkened on apical half; femur medium brown except basal portion dark
yellow to light brown and apical cap dark brown; basitarsus greatly dilated, 4.5 times as long as its greatest width.
Midleg: coxa brownish black; trochanter medium brown except base yellow; femur medium brown with apical cap
dark brown though basal extreme somewhat paler; tarsus light brown except basal tips of basitarsus, second and
third tarsomeres whitish yellow, and fifth tarsomere dark brown. Hind leg: trochanter yellow; femur dark brown

except apical cap brownish black and extreme base whitish yellow; tibia dark brown except apical cap brownish
black and basal tip whitish yellow; tarsus (Fig. 65B) medium brown except little more than basal half of basitarsus
whitish yellow (though extreme base darkened) and basal half of second tarsomere whitish yellow; basitarsus (Fig.
65B) much enlarged, gradually widened from base to apical one-third, then slightly narrowed to apex, 3.7 times as
long as its greatest width, and 0.9 and 1.0 times as wide as greatest widths of hind tibia and femur, respectively.
Wing. Length 1.8 mm. Genitalia. Style in ventrolateral view (Fig. 65C) elongate, 3.1 times as long as its greatest
width at base, gradually tapered toward apex. Ventral plate in ventral view (Fig. 65E) with arms directed forward.
Abdominal segment 10 with three or four hairs on ventral surface and three or four hairs on lateral surface on each
side. Cercus with seven distinct hairs.
Pupa. Body length 2.3–2.5 mm. Nearly as in S. (S.) sapaense sp. nov. except following characters. Thorax.
Integument: surrounding area of pit-like organ near base of each gill bare (Fig. 65G). Gill (Fig. 65H) with six
slender thread-like filaments directed forward and downward, decreasing in length from dorsal to ventral, with
dorsal filament of dorsal pair longest (1.4 mm) and ventral filament of ventral pair shortest (0.8 mm); relative
thickness of six filaments from dorsal to ventral when compared basally 1.0:0.7:0.6:0.6:0.6:0.6.
a vial as [UMSRP: Vietnam 044, Holotype, Simulium (S.) lowi, Male, coll. Vietnam, 8-XI-2013, Takaoka et al.],
reared from a pupa collected from a small stream (width 0.5 m, bottom rocky, water temperature 17˚C, shaded,
elevation 975 m, 21˚27’501’’N/105˚38’941’’E), slow-flowing in bushes along the road near the Tam Dao National
Park, Vinh Phuc Province, northern Vietnam, 8-XI-2013, by H. Takaoka, M. Sofian-Azirun & Z. Ya’cob.
PARATYPES: Two males (with their associated pupal exuviae and cocoons) in 80% ethanol, same data as those of
the holotype, labeled in each vial as [UMSRP: Vietnam 044, Paratype, Simulium (S.) lowi, Male, coll. Vietnam, 8-
XI-2013, Takaoka et al.].
Biological notes. The pupae of this new species were collected from dead tree leaves in the current. Associated
species were S. (G.) hongthaii, S. (G.) sanchayense sp. nov., S. (G.) vinhphucense sp. nov. and S. (N.) sp. (S.
feuerborni species-group).
Etymology. The species name lowi is in honor of Dr. V.L. Low, senior lecturer, Tropical Infectious Diseases
Research and Education Centre, University of Malaya, who greatly contributed to phylogenetic studies of black
flies.
Remarks. This new species is morphologically distinguished from S. (S.) brevipar in the male by the arms of
the ventral plate not divergent when viewed ventrally (Fig. 65E) (moderately divergent in S. (S.) brevipar, as shown
in Fig. 65D), and style with its greatest width basally when viewed ventrolaterally (Fig. 65C) (at the basal one-third
in S. (S.) brevipar, as shown in Fig. 65F), and in the pupa by the dorsal filament of the dorsal pair gradually tapered
toward the apex (Fig. 65I) (abruptly tapered basally, then gradually tapered in S. (S.) brevipar, as shown in Fig.
65J).
Simulium (Simulium) tani Takaoka & Davies, 1995 (complex)
Simulium (Simulium) tani (complex): Morphoform ‘a’: Takaoka et al., 2015a: 85–86.
Specimens examined. Seven females and 15 males (all reared from pupae), collected from a river (width 8–10 m,
depth 10–15 cm, bottom of pebbles, water temperature 17˚C, exposed to the sun, elevation 162 m,
18˚58’45.918”N/109˚50’10.693”E), fast flowing in a natural forest, Pu Mat, Nghe An Province, northern Vietnam,
9-XII-2015, H. Takaoka, M. Sofian-Azirun, C.D. Chen, & K.W. Lau; four females and four males (all reared from
pupae), collected from a stream (width 1.0–1.5 m, depth 10–15 cm, bottom of pebbles, water temperature 17˚C,
partially shaded, elevation 162 m, 18˚58’45.918”N/109˚50’10.693”E), moderate to fast flowing near a natural
forest, Pu Mat, Nghe An Province, northern Vietnam, 9-XII-2015, H. Takaoka, M. Sofian-Azirun, C.D. Chen, &
W. K. Lau; 13 females and 14 males (all reared from pupae), collected from a stream (width 1.5–3.0 m, depth 10–
15 cm, bottom of sands and pebbles, water temperature 18˚C, exposed to the sun, elevation 73 m, 19˚30’45.487”N/
105˚09’29.261”E), moderately flowing near a natural forest, Chau Dinh, Quy Chau, Nghe An Province, northern
Vietnam, 9-XII-2015, H. Takaoka, M. Sofian-Azirun, C.D. Chen & K. W. Lau.
Distribution. Vietnam (Lam Dong and Nghe An).
Remarks. This species is a species complex comprising 11 cytoforms, of which nine are reported from
Thailand (Adler and Crosskey 2016). In Vietnam, two morphoforms (‘a’ and ‘b’) were recognized, both of which
are morphologically distinguished from S. (S.) tani (= cytoform K) originally described from Peninsular Malaysia
(Takaoka et al. 2015a). Morphoform ‘a’ is chromosomally divided into two cytoforms (Adler et al. 2016) and is
also shown to include an additional possible cryptic taxon by a DNA sequence-based analysis (Low et al. 2016).
Simulium (Simulium) suoivangense Takaoka & Pham sp. nov.
Simulium (Simulium) tani (complex): Morphoform ‘b’: Takaoka et al., 2015a: 85–86.
Morphoform ‘b’ of S. (S.) tani (complex) reported from southern Vietnam (Takaoka et al. 2015a) is here described
as a distinct species and a new name S. suoivangense is given.
Female. Body length 2.4–2.6 mm. Head. Slightly narrower than thorax. Frons black, shiny with bright
reflection when illuminated at certain angles, with several dark stout hairs along lateral margins; frontal ratio 1.3–
1.4:1.0:1.2–1.3; frons:head ratio 1.0:3.5–4.0. Fronto-ocular area well developed, short, directed laterally, and
rounded apically. Clypeus black, slightly shiny, moderately covered with dark-brown medium-long hairs though
mediolongitudinal portion widely bare. Labrum 0.5–0.6 times length of clypeus. Antenna composed of scape,
pedicel and nine flagellomeres; antenna dark brown, except scape, pedicel and base of first flagellomere yellow.
Maxillary palp with five segments, medium brown except segment 3 dark brown; proportional lengths of third,
fourth, and fifth segments 1.0:1.2–1.3:2.6–2.8; third segment (Fig. 66A) of moderate size, with long ellipsoidal
sensory vesicle (0.5–0.6 times length of third segment) having medium-sized opening. Maxillary lacinia with 14–
16 inner and 19 or 20 outer teeth. Mandible with 31 or 32 inner and 11 outer teeth. Cibarium (Fig. 66B) with 82–90
minute processes near posterodorsal margin. Thorax. Scutum black, shiny with bright reflection when illuminated
at certain angles, moderately covered with dark short fine hairs interspersed with several dark-brown long upright
hairs on prescutellar area. Scutellum dark brown, shiny when illuminated at certain angles, covered with dark-
brown upright long hairs. Postnotum brownish black, shiny when illuminated at certain angles, and bare. Pleural
membrane bare. Katepisternum longer than deep, brownish black, shiny when illuminated at certain angles, and
bare. Legs. Foreleg: coxa whitish yellow; trochanter light brown with base whitish yellow; femur light to medium
brown with apical cap dark brown; tibia medium brown except apical cap dark brown, with median outer surface
widely white and with bright white sheen when illuminated at certain angles; tarsus black, with moderate dorsal
hair crest; basitarsus greatly dilated, 4.3–4.6 times as long as its greatest width. Midleg: coxa dark brown with
posterolateral surface brownish black; trochanter medium brown with base whitish yellow; femur medium to dark
brown except apical cap brownish black; tibia medium brown except base yellowish white and apical cap brownish
black, and with whitish sheen widely on posterior surface when illuminated at certain angles; tarsus medium brown
except basal three-fourths or two-thirds of basitarsus and base of second tarsomere yellowish white. Hind leg: coxa
medium to dark brown; trochanter whitish yellow; femur dark brown except base whitish yellow; tibia medium to
dark brown except base yellowish white and with whitish sheen widely on posterior surface when illuminated at
certain angles; tarsus medium brown except little more than basal two-thirds of basitarsus and basal half of second
tarsomere yellowish white; basitarsus (Fig. 66C) nearly parallel-sided, 5.4–5.6 times as long as wide, and 0.8 and
0.7 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig. 66C) moderately
developed, nearly as long as wide, and 0.4 times as wide as greatest width of basitarsus; pedisulcus (Fig. 66C) well
developed; claw simple, without tooth. Wing. Length 1.7–2.0 mm. Costa with dark spinules and hairs; subcosta
haired except apical half to one-fifth bare; basal section of radius bare; R1 with dark brown spinules and hairs; R2
with dark-brown hairs; hair tuft on base of radius dark brown; basal cell absent. Halter. White except basal portion
darkened. Abdomen. Basal scale dark brown, with fringe of yellow hairs. Dorsal surface of abdomen medium
brown to brownish black, with dark short hairs; tergite 2 shiny and silvery iridescent when illuminated at certain
angles and tergites 6–9 shiny and iridescent when illuminated at certain angles. Ventral surface of segment 7 with
pair of weakly sclerotized submedian sternal plates. Terminalia. Sternite 8 (Fig. 66D) well sclerotized and bare
medially, covered with 9–13 long and medium-long stout hairs and three to six short fine hairs on each side.
Ovipositor valve (Fig. 66D) nearly triangular, rounded posteromedially, membranous except inner margin weakly

sclerotized, densely covered with microsetae together with seven to nine short fine hairs; inner margin nearly
straight or sinuous or slightly concave medially. Genital fork (Fig. 66E) of inverted-Y form; stem slender and well
sclerotized; arms of moderate width, each with distinct projection directed forward from lateral ridge. Paraproct in
ventral view (Fig. 66F) pointed posteromedially, moderately concave anterolaterally, and with strongly sclerotized
anteromedial surface bearing five sensilla; paraproct in lateral view (Fig. 66G) somewhat produced ventrally and
slightly posteriorly, 0.6 times as long as wide, covered with 18–22 medium-long stout hairs and numerous
microsetae on lateral and ventral surfaces. Cercus in lateral view (Fig. 66G) short, rounded posteriorly, 0.6 times as
long as its greatest width, and covered with medium-long and short stout hairs. Spermatheca (Fig. 66H) ovoidal,
1.2–1.3 times as long as wide, well sclerotized except duct and narrow area of juncture with duct unsclerotized,
without discernible reticulate surface patterns; minute internal setae present; accessory ducts subequal in thickness
to each other and slightly thicker than main duct.
FIGURE 66. Female of Simulium (Simulium) suoivangense sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Cibarium (front view); C, Hind basitarsus and second tarsomere (left side; outer view). D, Sternite 8
and ovipositor valve (right side only; ventral view); E, Genital fork (ventral view); F & G, Paraprocts and cerci (F, ventral view;
G, lateral view). H, Spermatheca. Scale bars. 0.1 mm for C; 0.02 mm for A, B and D–H.
Male. Body length 2.5–2.7 mm. Head. Slightly wider than thorax. Upper eye medium brown, consisting of
large facets in 17 or 18 vertical columns and in 18 horizontal rows. Clypeus black, thickly white pruinose, silvery
shiny when illuminated at certain angles, covered with dark-brown hairs along and near lateral margins (most of
central portion bare). Antenna composed of scape, pedicel and nine flagellomeres, brownish black except scape,
pedicel and base of first flagellomere dark yellow to light brown; first flagellomere elongate, 1.5 times as long as
second one. Maxillary palp grayish to dark brown, composed of five segments with proportional lengths of third,
fourth, and fifth segments 1.0:13:2.7–2.8; third segment (Fig. 67A) widened apically; sensory vesicle (Fig. 67A)
small (0.2 times as long as third segment), ellipsoidal, and with small opening. Thorax. Scutum black, with white
pruinose pattern, i.e., anterior pair of triangular spots (not sharply pointed posteriorly) on shoulders extended
posteriorly along lateral margins but not connected to large transverse spot entirely covering prescutellar area;
these pruinose areas silvery or bluish iridescent when illuminated at certain angles; scutum uniformly and densely
covered with brassy recumbent short hairs interspersed with dark-brown long upright hairs on prescutellar area.
Scutellum brownish black, with several dark-brown long upright hairs. Postnotum brownish black, white pruinose
when illuminated at certain angles and bare. Pleural membrane bare. Katepisternum longer than deep, brownish
black, thinly white pruinose and shiny when illuminated at certain angles, and bare. Legs. Color as in female except
hind tibia with basal small portion yellow, and hind basitarsus yellowish white on basal half or little less. Fore
basitarsus greatly dilated, 5.4–6.1 times as long as its greatest width. Hind basitarsus (Fig. 67B) much enlarged,
gradually widened from base to apical two-fifths, then slightly narrowed toward apex, 3.6–3.8 times as long as its
greatest width, and 0.8–0.9 and 0.9 times as wide as greatest widths of hind tibia and femur, respectively; calcipala
(Fig. 67B) small, nearly as long as wide, 0.2 times as wide as greatest width of basitarsus; pedisulcus (Fig. 67B)
well developed. Halter. White except basal portion darkened. Wing. Length 1.6–1.7 mm. Other characters as in
female except subcosta bare. Abdomen. Basal scale brownish black, with fringe of pale (though basally dark) long
hairs. Dorsal surface of abdomen brownish black to black, with dark short hairs; segments 2 and 5–7 each with pair
of silvery or bluish iridescent spots dorsolaterally, those on segment 2 broadly connected in middle to each other.
Genitalia. Coxite in ventral view (Fig. 67C) nearly quadrate, covered with stout hairs near posterior margin. Style
in ventral view (Fig. 67C) elongate, gradually tapered from base to middle, then nearly parallel-sided, with apical
spine; style in ventrolateral view (Fig. 67D) elongate, 2.6 times as long as its greatest width at basal one-third,
slightly widened from base to basal one-third, much tapered toward apical one-third, then slightly widened toward
tip; style in medial view (Fig. 67E) 1.6 times as long as coxite, much produced dorsally near base and with 8–12
small cone-like spines on dorsal surface. Ventral plate in ventral view (Fig. 67C) well sclerotized, with body
subquadrate (though rounded on each posterolateral corner), 0.7 times as long as wide, posterior margin somewhat
convex, and with round projection directed ventrally which is covered with many minute setae; arms directed
forward and somewhat divergent from each other; ventral plate in lateral view (Fig. 67F) with posterior margin
serrated along basal two-thirds; ventral plate in caudal view (Fig. 67G) rounded ventrally, with lateral margin
serrated along basal two-thirds, with numerous setae on posterior surface except both areas of dorsolateral corners
bare. Median sclerite (Fig. 67F, H) plate-like, gradually widened from base toward near apex, with round apex,
brown basally, but not sclerotized apically. Paramere (Fig. 67I) with several distinct hooks and several smaller
ones. Aedeagal membrane (Fig. 67J) moderately covered with minute setae, and with moderately sclerotized dorsal
plate in form of horizontal bar (Fig. 67K). Abdominal segment 10 (Fig. 67L, M) with four to eight hairs on lateral
surface and one to three hairs on ventral surface on each side. Cercus (Fig. 67L, M) small, rounded, with seven to
ten distinct hairs.
Pupa. Body length 2.0–2.5 mm. Head. Integument including antennal sheaths yellow; frons moderately
covered with tubercles, of which some or most having minute secondary projections, and with two pairs of
unbranched short trichomes; face with pair of unbranched short trichomes slightly longer and stouter than frontal
trichomes. Thorax. Integument yellow, moderately covered with tubercles, of which some on anterodorsal surface
have minute secondary projections; thorax on each side with two short anterodorsal trichomes, two short
anterolateral trichomes (one sometimes slightly thinner and shorter than other), one short mediolateral trichome,
and three short ventrolateral trichomes (one slightly shorter and thinner than two others); all trichomes unbranched.
Gill (Fig. 68A) with six slender thread-like filaments in three pairs (one dorsal, one middle and one ventral); no
common basal stalk and basal fenestra small; all pairs short-stalked, and stalks of dorsal and ventral pairs lying at
angle of 70 degrees when viewed laterally; filaments decreasing in length from dorsal to ventral, with longest
dorsal filament 1.6–1.8 mm long and shortest ventral filament 1.1–1.5 mm long; relative thickness of filaments
from dorsal to ventral when compared basally 1.0:0.9:0.9:0.8:0.9:0.7; all filaments medium brown, tapered toward

FIGURE 67. Male of Simulium (Simulium) suoivangense sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Hind basitarsus and second tarsomere (left side; outer view); C, Coxites, styles and ventral plate
(ventral view); D & E, Styles (right side; D, ventrolateral view; E, medial view); F, Ventral plate and median sclerite (lateral
view); G, Ventral plate (caudal view); H, Median sclerite (caudal view); I, Paramere (left side; caudal view); J, Aedeagal
membrane (caudal view); K, Dorsal plate (caudal view); L & M, Abdominal segment 10 and cerci (right side; L, lateral view;
M, caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–M.
basal half, then nearly of same thickness up to tip, with distinct annular ridges and furrows, and densely covered
with minute tubercles. Abdomen. Dorsally, all segments unpigmented except segment 1, basal half of segment 2,
and segment 9 light yellowish; segment 1 with one unbranched short seta on each side; segment 2 with one
unbranched short seta and five spinous short setae on each side; segments 3 and 4, each with four distinct hooked
spines and one unbranched spinous short seta on each side; segments 5 and 6 lacking spine-combs; segments 7–9
each with distinct spine-combs in transverse row and comb-like groups of minute spines on each side; segment 9
without terminal hooks. Ventrally, segments 4–8 unpigmented and segment 9 somewhat yellowish; segments 4–8
each with comb-like groups of minute spines; segment 4 with one unbranched stout hooklet and few slender minute
setae on each side; segment 5 with pair of bifid stout hooks submedially and few unbranched short setae on each
side; segments 6 and 7 each with pair of bifid inner and unbranched outer stout hooks somewhat separated from
each other, and few unbranched short setae on each side. Grapnel-shaped hooklets absent on each side of segment
9. Cocoon. Wall-pocket-shaped, moderately woven, thin, ochreous, and not extended ventrolaterally; individual
threads visible; 2.7–3.1 mm long by 1.0–1.2 mm wide.
FIGURE 68. Pupa and larva of Simulium (Simulium) suoivangense sp. nov. A, Pupa; B–D, Larva. A, Anterior portion of
thorax and gill filaments (left side; outer view); B, Mandible; C, Hypostoma; D, Head capsule (ventral view). Scale bars. 0.1
mm for A and D; 0.02 mm for B and C.
Mature larva. Body length 4.6–5.0 mm. Thoracic segment 1 and abdominal segments 1–4 each encircled by
broad dark-gray transverse band and abdominal segments 5–9 dark gray on dorsal and dorsolateral surfaces;
thoracic segments 2 and 3 often light gray on dorsal surface and dark gray on ventral surface, and abdominal
segments 4–6 each with dark mediolongitudial band on ventral surface; abdominal segments 6 and 7 each with thin
grayish transverse band ventrally. Head. Cephalic apotome yellow (though rarely dark yellow) with small medial
area near posterior margin sometimes darkened; head spots almost indistinct or faintly positive. Lateral surface of
head capsule yellow except eye-spot region whitish and areas anterior and posterior to eye-spot region somewhat
darkened. Ventral surface of head capsule (Fig. 68D) yellow, though often widely darkened medially; long spot on
each side of postgenal cleft clearly negative. Antenna composed of three articles and apical sensillum, slightly
longer than stem of labral fan; length ratio of three articles (from base to tip) 1.0:0.9–1.0:0.7–0.8. Labral fan with
37–41 main rays. Mandible (Fig. 68B) with mandibular serrations composed of two teeth (one medium-sized, one
small); main tooth at right angle against mandible on apical side; supernumerary serrations absent; comb-teeth

decreasing in length from first to third. Hypostoma (Fig. 68C) with nine anterior teeth, of which corner teeth most
prominent, slightly longer than median tooth, and intermediate teeth shortest; lateral margins serrate apically; four
or five hypostomal bristles nearly parallel to lateral border on each side. Postgenal cleft (Fig. 68D) arrow-head
shaped, long, 4.2–5.3 times length of postgenal bridge; subesophageal ganglion not pigmented. Cervical sclerite on
each side composed of one medium-brown elongate piece and one light-brown elliptical piece, not fused to
occiput. Thorax and Abdomen. Histoblast of pharate pupal gill with six short filaments. Thoracic and abdominal
cuticle almost bare except few posterior segments moderately covered with colorless minute setae dorsally, and last
segment of abdomen moderately covered with colorless short setae on each side of anal sclerite. Rectal scales
present. Rectal organ compound, each lobe with 10–12 finger-like secondary lobules. Anal sclerite X-shaped, with
short broad anterior arms 0.6–0.8 times as long as posterior ones; one or two sensilla on juncture area; five sensilla
posterior to posterior arms. Last abdominal segment somewhat swollen laterally but lacking ventral papillae.
Posterior circlet with 75–80 rows of hooklets with up to 15 or 16 hooklets per row.
in a vial as [UMSRP: Vietnam 045, Holotype, Simulium (S.) suoivangense, Female, coll. Vietnam, 24-IV-2014,
Takaoka et al.], reared from a pupa collected from a river (width 4–6 m, water temperature 18.0˚C, exposed to the
sun, elevation 1,452 m, 12˚10’56.408”N/108˚40’48.152”E), moderately flowing in open land, Suoi Vang, Lac
Duong District, Lam Dong Province, southern Vietnam, 24-IV-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob,
C.D. Chen & K.W. Lau. PARATYPES: 11 females and five males and two mature larvae, same data as those of the
holotype, labeled in each vial as [UMSRP: Vietnam 045, Paratype, Simulium (S.) suoivangense, coll. Vietnam, 24-
IV-2014, Takaoka et al.]; 10 females and seven males (all reared from pupae) in 80% ethanol, labeled in each vial
as [UMSRP: Vietnam 045, Paratype, Simulium (S.) suoivangense, Male, coll. Vietnam, 9-XII-2015, Takaoka et al.],
collected from a river (width 4–5 m, depth 20–30 cm, bottom of pebbles, water temperature 15.0˚C, partially
shaded, elevation 223 m, 18˚58’03.270”N/104˚48’07.811”E), rapidly flowing in a natural forest, Pu Mat National
Park, Nghe An Province, northern Vietnam, 9-XII-2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W.
Lau.
Biological notes. The pupae and larvae of this new species were collected from leaves of trailing grasses and
fallen tree leaves in the current. Associated species were S. (G.) asakoae and S. (S.) laui.
Distribution. Vietnam (Lam Dong and Nghe An).
Etymology. The species name suoivangense refers to the locality name, Suoi Vang, where this new species
was collected.
Remarks. This new species is characterized by lacking pupal terminal hooks, differing from all 11 cytoforms
and morphoform ‘a’ among S. (S.) tani (complex), all of which have pupal terminal hooks (Takaoka et al. 2015a).
It is also distinguished from cytoform ‘K’ (typical S. tani from Peninsular Malaysia) in the female by the relatively
smaller size of the sensory vesicle to the third maxillary palpal segment (0.5 versus 0.6). The recognition of
morphoform ‘b’ as a distinct species is supported by DNA sequence-based and chromosomal analyses (Adler et al.
2016; Low et al. 2016).
Simulium (Simulium) xuandei Takaoka & Pham, 2015
Simulium (Simulium) xuandei Takaoka & Pham (in Takaoka et al., 2015a): 86–93 (Female, male, pupa and larva).
Specimens examined. Three females and two males (all reared from pupae), collected from a stream (width 1.5–
4.0 m, depth 10–20 cm, bottom of pebbles and rocks, water temperature 18.0˚C, exposed to the sun, elevation 107
m, 19˚30’10.540”N/105˚09’17.144”E), moderately flowing near a natural forest, Chau Dinh, Quy Chau, Nghe An
Province, northern Vietnam, 11-XII-2015, by H. Takaoka, M. Sofian-Azirun, C.D. Chen & K.W. Lau.
Distribution. Vietnam (Lam Dong and Nghe An)
Remarks. This species was described from Lam Dong Province, southern Vietnam (Takaoka et al. 2015a), and
also recorded from Nghe An Province, northern Vietnam by our surveys. The molecular data show that this species
includes two additional cryptic taxa (Low et al. 2016).
8) Simulium variegatum species-group
Simulium (Simulium) chamlongi Takaoka & Suzuki, 1984
Simulium (Simulium) chamlongi Takaoka & Suzuki, 1984: 27–30 (Female, male, pupa and larva); Crosskey and Howard, 1997;
Pham, 1999: 335; Takaoka et al., 2015a: 93.
Distribution. Thailand and Vietnam (Bac Thai and Lam Dong).

Remarks. This species was originally described from Thailand by Takaoka and Suzuki (1984) and later
recorded from Vietnam by Crosskey and Howard (1997), Pham (1999) and Takaoka et al. (2015a). No detailed
information for the locality was given by Crosskey and Howard (1997).
Simulium (Simulium) phuluense Takaoka & Sofian-Azirun sp. nov.
Female. Body length 2.6–3.1 mm. Head. Narrower than thorax. Frons black, shiny, with several dark long stout
hairs along each lateral margin and three to five similar hairs on each side just above lower margin; frontal ratio
1.3:1.0:1.2–1.4; frons:head ratio 1.0:3.6–4.3. Fronto-ocular area well developed, short, directed laterally and
slightly upward. Clypeus black, white pruinose, slightly shiny when illuminated at certain angles, moderately
covered with dark-brown long stout hairs except mediolongitudinal area of upper half somewhat widely bare.
Labrum 0.7 times as long as clypeus. Antenna composed of scape, pedicel and nine flagellomeres, medium to dark
brown except scape, pedicel and base of first flagellomere yellow when viewed dorsally, and scape, pedicel and
basal four-fifths of flagellomere yellow when viewed ventrally. Maxillary palp with five segments, grayish brown
except segments 1 and 2 yellow; proportional lengths of third, fourth, and fifth segments 1.0:1.3:2.8–3.1; third
segment (Fig. 69A) of normal size; sensory vesicle (Fig. 69A) of moderate size, (0.3–0.4 times length of third
segment) having opening of moderate size. Maxillary lacinia with 13 or 14 inner and 16 or 17 outer teeth. Mandible
with 30–32 inner and 16 outer teeth. Cibarium (Fig. 69B) with blunt medial projection on posterior margin and
with about 30 small pointed processes and about 28 minute ones near base of medial projection. Thorax. Scutum
black, white pruinose along lateral margins, shiny when illuminated at certain angles, densely covered with golden-
yellow recumbent short hairs interspersed with several dark-brown long upright hairs on prescutellar area.
Scutellum dark brown, covered with dark-brown upright long hairs and golden-yellow short hairs. Postnotum
brownish black, white pruinose when illuminated at certain angles and bare. Pleural membrane bare.
Katepisternum longer than deep, brownish black, shiny and white pruinose when illuminated at certain angles and
bare. Legs. Foreleg: coxa whitish yellow; trochanter yellow; femur yellow except apical cap light brown; tibia
yellowish white except basal one-fourth light brown and apical cap brownish black, with shiny sheen widely on
outer surface when illuminated at certain angles; tarsus brownish black, with moderate dorsal hair crest; basitarsus
greatly dilated, 5.6–5.8 times as long as its greatest width. Midleg: coxa brownish black; trochanter yellow; femur
yellow with apical cap light brown; tibia (Fig. 69C) whitish yellow to dark yellow except apical cap medium brown
and narrow portion of outer surface along posterior margin widely light to medium brown, and with shiny sheen on
posterior surface when illuminated at certain angles; tarsus medium brown except basal half to two-thirds of
basitarsus yellow. Hind leg: coxa dark brown; trochanter yellow; femur yellow with apical cap medium brown;
tibia (Fig. 69D) yellow on little less than basal half, and gradually darkened toward apex and apical cap dark brown
on outer and posterior surface, and widely yellow except apical cap dark brown on inner surface, with shiny sheen
on posterior surface when illuminated at certain angles; tarsus medium brown except basal two-thirds of basitarsus
and basal half of second tarsomere yellow; basitarsus (Fig. 69E) nearly parallel-sided, 6.2–6.3 times as long as
wide, and 0.7–0.8 and 0.6–0.7 times as wide as greatest widths of hind tibia and femur, respectively; calcipala (Fig.
69E) moderately developed, slightly shorter than basal width, and 0.5 times as wide as basitarsus; pedisulcus (Fig.
69E) well developed; claw (Fig. 69F) with small subbasal tooth. Wing. Length 2.7–3.1 mm. Costa with dark
spinules and hairs; subcosta haired except near apex bare; basal section of radius bare; R1 with dark spinules and
hairs; R2 with hairs; hair tuft on base of radial vein dark brown; basal cell absent. Halter. White with basal portion

FIGURE 69. Female of Simulium (Simulium) phuluense sp. nov. A, Third segment of maxillary palp with sensory vesicle
(right side; front view); B, Cibarium (front view); C, Fore tibia (left side; outer view); D, Hind tibia (left side; outer view); E,
Hind basitarsus and second tarsomere (left side; outer view). F, Claw; G, Sternite 8 and ovipositor valve (right side only; ventral
view); H, Genital fork (ventral view); I & J, Paraprocts and cerci (I, ventral view; J, lateral view). K, Spermatheca. Scale bars.
0.1 mm for C– E; 0.02 mm for A, B and F–K.
darkened. Abdomen. Basal scale light brown, with fringe of pale long hairs. Dorsal surface of abdomen medium
brown except segment 2 light brown, with short dark hairs; tergite 2 shiny, white iridescent when illuminated at
certain angles, and tergites 6–8 shiny; ventral surface of segment 2 whitish and those of others light brown;
segment 7 with median large sternal plate having numerous hairs. Terminalia. Sternite 8 (Fig. 69G) bare medially,
with 22–25 dark medium-long to long stout hairs on each lateral surface. Ovipositor valves (Fig. 69G) wide, with
inner margins widely concave, with posteromedial apices pointed anteriorly, each moderately covered with
microsetae and 35 or 36 short to medium-long hairs except small area of posteromedial apex bare; inner margins
widely concave and darkened except near apex unpigmented. Genital fork (Fig. 69H) of inverted-Y form, with
narrow, well sclerotized stem; arms of moderate width, each with distinct short projection directed anterodorsally.
Paraproct in ventral view (Fig. 69I) oblong, widely depressed on ventral surface, unpigmented except medial
surface somewhat darkened, sparsely covered with minute setae and sensilla; paraproct in lateral view (Fig. 69J)
much protruded ventrally beyond ventral margin of cercus, 0.7 times as long as wide, with numerous short to
medium-long hairs on lateral and ventral surfaces; anteromedial surface with 10 or 11 short sensilla. Cercus in
lateral view (Fig. 69J) short, trapezoid, 0.48 times as long as wide, and with numerous short to medium-long hairs.
Spermatheca (Fig. 69K) large, nearly ovoid, 1.3 times as long as wide, well sclerotized except portion of juncture
with duct unsclerotized, with no defined surface patterns, and with internal setae; accessory ducts subequal in
thickness to each other, and to major duct.
Male. Body length 3.0–3.3 mm. Head. Nearly as wide as thorax. Upper eye medium brown, consisting of large
facets in 23 or 24 vertical columns and in 23 or 24 horizontal rows. Clypeus black, thickly white pruinose,
moderately covered with dark-brown hairs. Antenna composed of scape, pedicel and nine flagellomeres, dark
brown except base of first flagellomere light brown; first flagellomere elongate, 1.6 times as long as second one.
Maxillary palp medium brown, composed of five segments with proportional lengths of third, fourth, and fifth
segments 1.0:1.2:2.5; third segment (Fig. 70A) of moderate size, with apex somewhat produced inward; sensory
vesicle (Fig. 70A) small (0.2 times length of third segment), ellipsoidal, and with small opening. Thorax. Nearly as
in male of S. (S.) daoense sp. nov. Legs. Foreleg: coxa yellow; trochanter light brown; femur light to medium
brown with apical cap dark brown; tibia brownish black except middle one-third medium brown, though outer
surface widely white except basal tip and apical cap, and with white sheen when illuminated at certain angles;
tarsus brownish black, with moderate dorsal hair crest; basitarsus moderately dilated, 6.7–7.7 times as long as its
greatest width. Midleg: dark brown to brownish black except extreme base of tibia whitish yellow and basal half of
basitarsus yellow. Hind leg: coxa brownish black; trochanter light brown; femur medium brown except base light
brown or dark yellow and apical cap brownish black; tibia dark brown to brownish black except basal tip whitish
yellow; tarsus (Fig. 70B) medium brown except basal half to two-fifths of basitarsus yellow (though base
somewhat darkened) and little less than basal half of second tarsomere yellow; basitarsus (Fig. 70B) much
enlarged, spindle-shaped, gradually widened from base to apical one-third, then tapered to apex, 4.3 times as long
as its greatest width, and 0.8 and 0.9–1.0 times as wide as greatest widths of hind tibia and femur, respectively;
calcipala (Fig. 70B) small, slightly shorter than width at base, 0.3 times as wide as greatest width of basitarsus;
pedisulcus (Fig. 70B) well developed. Wing. Length 2.7–2.9 mm. Other characters as in female including subcosta
with 12–24 hairs though apical one-fifth and one-third bare. Halter. White with basal portion darkened. Abdomen.
Basal scale dark brown to brownish black, with fringe of dark long hairs. Dorsal surface of abdomen dark brown to
brownish black, and covered with dark short hairs; segments 2 and 5–7 each with pair of silvery or bluish iridescent
spots dorsolaterally, those on segment 2 broadly connected in middle to each other. Genitalia. Coxites, styles and
ventral plate in ventral view as Fig. 70C. Coxite in ventral view nearly quadrate, 1.2 times as long as width. Style in
ventrolateral view (Fig. 70D) elongate, 3.48 times as long as its greatest width near base, slightly tapered to apex;
style in medial view (Fig. 70E) 1.6 times as long as coxite, spatulate dorsoventrally except basal half much
produced dorsally in form of isosceles triangular. Ventral plate in ventral view (Fig. 70C) Y-shaped, body with
several teeth on each side of posterior surface, and with ventrally produced process covered with several minute
setae on its ventroposterior surface; arms widely divergent basally and convergent apically; ventral plate in lateral
view (Fig. 70F): body with serrated posterior margin, and sparsely covered with minute setae on each anterolateral
surface, and having ventrally produced process with anterolateral surface bare except several minute setae near
base; arm wide with apex curved ventrally; ventral plate in caudal view (Fig. 70G): body and ventrally produced
process gradually tapered ventrally, with round apex, having seven to nine teeth on each side of posterior surface,
and several minute setae on posterior surface of ventrally produced process. Median sclerite (Fig. 70F, H) arising

FIGURE 70. Male of Simulium (Simulium) phuluense sp. nov. A, Third segment of maxillary palp with sensory vesicle (right
view); D & E, Styles (D, right side and ventrolateral view; E, left side and medial view); F, Ventral plate and median sclerite
(lateral view); G, Ventral plate (caudal view); H, Median sclerite (caudal view); I, Paramere (right side; caudal view) ; J,
Aedeagal membrane (right half; caudal view); K, Dorsal plate (caudal view); L & M, Abdominal segment 10 and cerci (right
side; L, lateral view; M, caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A and C–M.
near anterior margin of ventral plate and directed dorsally, plate-like, widened from base toward apical one-third,
then nearly parallel-sided or slightly narrowed toward apex, brown basally, but not so well sclerotized apically.
Paramere (Fig. 70I) broad basally, with several long hooks and shorter ones. Aedeagal membrane (Fig. 70J)
densely covered with minute setae, and with partially sclerotized dorsal plate in form of horizontal bar (Fig. 70K).
Abdominal segment 10 (Fig. 70L, M) without distinct hair on ventral and lateral surfaces. Cercus (Fig. 70L, M)
small, rounded, with 14–16 distinct hairs.
Pupa. Body length 2.7–3.2 mm. Head. Integument yellow to ochreous, moderately covered with small round
tubercles except antennal sheath bare; frons with pair of unbranched slender short trichomes (Fig. 71A) on each
side; face with unbranched slender short trichome (Fig. 71B) on each side. Thorax. Integument yellow to ochreous,
moderately covered with small round tubercles; thorax with two long anterodorsal trichomes (Fig. 71C), two
anterolateral trichomes (anterior trichome medium-long, and posterior trichome long) (Fig. 71D), one medium-
long mediolateral trichome (Fig. 71E), and three ventrolateral trichomes (one short, two others medium-long) (Fig.
71F), on each side; all unbranched and with straight apices. Gill (Fig. 71G) with six slender thread-like short
filaments in three pairs; all pairs short-stalked, arising from short common basal stalk; gill filaments divergent
basally, upper filament of dorsal pair lying at angle of 120 degrees against lower filament of ventral pair when
viewed laterally; filaments subequal in length to one another (2.2–2.4 mm long) (though lower filament of middle
pair and upper filament of ventral pair somewhat longer (2.7 mm long); relative thickness of six filaments from
dorsal to ventral when basal portions were compared 1.0:0.9:1.0:1.0:0.8:0.8; all filaments light to medium brown,
tapered toward apex, with distinct annular ridges and furrows forming definite reticulate surface patterns, and
densely covered with minute tubercles (relatively larger ones on ridges). Abdomen. Dorsally, segment 1 and basal
half of segment 2 light brown and without minute tubercles, and other segments unpigmented except segment 9
and basal portions of spine-combs on segments 7 and 8 yellow; segment 1 with one unbranched slender short seta
(Fig. 71H) on each side; segment 2 with one unbranched slender short seta and five unbranched short setae, of
which four are stout (Fig. 71I), on each side; segments 3 and 4 each with four distinct hooked spines and one
unbranched short seta on each side; segments 7 and 8 each with distinct spine-combs in transverse row on each
side; segments 5, 6 and 9 lacking spine-combs; segments 6–9 each with comb-like groups of minute spines on each
side; segment 9 with cone-shaped terminal hooks (Fig. 71J). Ventrally, segments 3–9 unpigmented, each (except
segment 9) with comb-like groups of minute spines; segment 4 with one unbranched slender minute seta, two stout
hooklets (Fig. 71K) on each side; segment 5 with pair of bifid stout hooks submedially and few unbranched short
setae on each side; segments 6 and 7 each with pair of bifid inner and unbranched outer stout hooks somewhat
separated from each other, and few unbranched short setae on each side. Grapnel-shaped hooklets absent on each
side of segment 9. Cocoon. Wall-pocket-shaped, tightly and thickly woven, ochreous, not so extended
ventrolaterally; individual threads invisible; 3.5–4.5 mm long by 1.6–2.2 mm wide.
Mature larva. Body length 7.4–8.4 mm. Body whitish to light grayish, and thoracic segment 1 encircled with
pinkish to reddish-brown transverse band though disconnected ventromedially, and other thoracic segments and
abdominal segments pinkish to light ochreous dorsally to varying extent. Head. Cephalic apotome yellowish white
to yellow with somewhat darkened narrow area along posterior margin; head spots faintly positive though anterior
spot of mediolongitudinal spots often less distinct; lateral surface of head capsule yellow except eye-spot region
yellowish white and eyebrow darkened and area between eye-spot region and posterior margin always darkened
though spots in front of posterior margin obscured or faintly negative; one small spot below eye-spot region
indistinct; ventral surface of head capsule (Fig. 72A) yellow to light brown, with spots on each side of postgenal
cleft obscured or faintly negative. Head capsule sparsely covered with minute setae on dorsal, lateral and ventral
surfaces. Antenna composed of three articles and apical sensillum, slightly longer than stem of labral fan; length
ratio of three articles (from base to tip) 1.0:1.1–1.2:0.6–0.7. Labral fan with 49–51 primary rays. Mandible (Fig.
72B) with mandibular serration composed of two teeth (one medium-sized, one small); main tooth at obtuse angle
against mandible on apical side; supernumerary serrations absent; comb-teeth decreasing in length from first to
third. Hypostoma (Fig. 72C) with nine anterior teeth, of which corner teeth slightly longer than median tooth; three
intermediate teeth on each side shortest; lateral margins moderately serrate apically; six hypostomal bristles per
side divergent posteriorly from lateral margin. Postgenal cleft (Fig. 72A) medium-sized, 2.1 times as long as
postgenal bridge, pointed apically; sheath of subesophageal ganglion usually weakly pigmented, wine-glass-
shaped, or not pigmented. Cervical sclerites on each side composed of anterior dark slender rod-like piece and
posterior dark elliptical piece close together, and not fused to occiput.Thorax and Abdomen. Histoblast of pharate

FIGURE 71. Pupa of Simulium (Simulium) phuluense sp. nov. A, Frontal trichomes; B, Facial trichome; C–F, Thoracic
trichomes (C, anterodorsal; D anterolateral; E, mediolateral; F ventrolateral); G, Anterior part of thorax and gill filaments (right
side; outer view); H, Hair-like seta on dorsal surface of abdominal segment 1; I, Hair-like seta, minute seta and stout seta on
dorsal surface of abdominal segment 2; J, Terminal hooks (caudal view); K, Hooklets and short seta on ventral surface of
abdominal segment 4. Scale bars. 0.1 mm for G; 0.02 mm for A–F and H–J.
FIGURE 72. Larva of Simulium (Simulium) phuluense sp. nov. A, Head capsule (ventral view); B, Mandible; C, Hypostoma.
Scale bars. 0.1 mm for A; 0.04 mm for C; 0.02 mm for B.
pupal gill with six thread-like filaments. Thoracic and abdominal cuticle bare except last segment of abdomen
moderately covered with short colorless setae on each side of anal sclerite. Rectal scales present. Rectal organ
compound, each lobe with 18–21 finger-like secondary lobules. Anal sclerite X-shaped, with short broad anterior
arms with forked apices, 0.8 times as long as posterior ones; no or one sensillum on base of anal sclerite; four to
nine sensilla posterior to posterior arms. Last abdominal segment lacking ventral papillae. Posterior circlet with
88–95 rows of hooklets with up to 13 or 14 hooklets per row.
in a vial as [UMSRP: Vietnam 046, Holotype, Simulium (S.) phuluense, Female, coll. Vietnam, 23-XII-2014,
Takaoka et al.], reared from a pupa collected from a small shallow stream (width 2.0 m, depth 15 cm, bottom of
pebbles, water temperature 9.0˚C, partially shaded, elevation 1,192 m, 22˚23’03.208”N/103˚50’58.990”E)
moderately flowing in a natural forest, Sapa, Lao Cai Province, northern Vietnam, 23-XII-2014, by H. Takaoka, M.
Sofian-Azirun, Z. Ya’cob, C.D. Chen & K.W. Lau. PARATYPES: Three females, four males (all reared from
pupae) and five mature larvae in 80% ethanol, same data as those of the holotype, labeled in each vial as [UMSRP:
Vietnam 046, Paratype, Simulium (S.) phuluense, coll. Vietnam, 23-XII-2014, Takaoka et al.].
Biological notes. The pupae and larvae of S. (S.) phuluense sp. nov. were collected from grass leaves and roots
trailing in the current. This species is one of the dominant species in Sapa, being found in 12 of 23 streams
surveyed. Associated species were S. (G.) hongthaii, S. (S.) daoense sp. nov., S. (S.) doipuiense (complex) and S.
(S.) sapaense sp. nov.
Etymology. The species name phuluense refers to the land in which the tribe Phulu lives.
Remarks. Simulium (S.) phuluense sp. nov. is assigned to the S. variegatum species-group of the subgenus
Simulium, redefined by Takaoka (2003), based on the pleural membrane bare, female claw with a small subbasal
tooth (Fig. 69F), ovipositor valves with inner margins widely concave (Fig. 69G), male style without basal
protuberance (Fig. 70E), pupal gill with six filaments (Fig. 71G).
This new species is characterized by a greater number of the upper-eye facets in 23 or 24 vertical columns and
23 or 24 horizontal rows, and haired subcosta in the male. No other related species of the S. variegatum species-
group have such a combination of characters except S. (S.) oitanum (Shiraki) (reported as S. (Odagmia) aokii) from
Koshiki Island, Nansei Islands, Japan, which has the male subcosta with 10–12 hairs, although it has the male
upper-eye facets in about 16 horizontal rows (Takaoka 1976). Simulium (S.) zunyiense Chen, Xiu & Zhang, from
Guizhou Province, China, has a similar greater number of male upper-eye facets (in 22 vertical columns and 26

horizontal rows), but differs by having a bare male subcosta, style with a short subbasal protuberance, and
triangular larval postgenal cleft (Chen et al. 2012). The females of S. (S.) jingfui Cai, An & Li described from a
female and a male from Sichuan Province, China, and S. (S.) liubaense Liu & An described from a female from
Shaanxi Province, China, differ by having dark fore coxa, and an ovipositor valve rounded apically (Cai et al.
2008; Liu and An 2009).
Acknowledgements
We are grateful to Prof. Emeritus H.S. Yong, Academician, University of Malaya, Prof. R. Hashim, Deputy Dean
of Faculty of Science, University of Malaya, and Assoc. Prof. R. Ramli, Head of Institute of Biological Sciences,
University of Malaya, for their support and encouragement.
Thanks are due to staffs of local offices of Lao Cai, Lam Dong and Thua Thien Hue Provinces, the National
Institute of Food Control, Ministry of Health, Vietnam, for their kind arrangement and cooperation during our
surveys. We are grateful to Dr. M. Fukuda, Oita University, Japan, Dr. Y. Otsuka, Kagoshima University, Japan, Dr.
W. Srisuka, Queen Sirikit Botanic Garden, and Dr. A. Saeung, Chiang Mai University, Thailand, for their help in
providing Japanese and Thai samples of black flies needed for this study.
All field surveys conducted in Vietnam were financially supported by research grants of University of Malaya
(UMRG Project No. RP003A/13SUS and RP021A/16SUS).
Finally, assistance provided by Mrs. Asako Takaoka during this study is highly appreciated.
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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.
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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.
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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.
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Simuliidae). Japanese Journal of Tropical Medicine and Hygiene, 20 (2), 135–142.
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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:
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Zootaxa 4261 (1): 001–165 ISSN 1175-5326 (print edition)

The black flies (Diptera: Simuliidae) of Vietnam

HIROYUKI TAKAOKA1,5, MOHD SOFIAN-AZIRUN1, ZUBAIDAH YA’COB1, CHEE DHANG

Accepted by L. Hernandez-Triana: 15 Feb. 2017; published: 5 May 2017

FIRST PUBLISHED IN 2017 BY

© 2017 Magnolia Press

ISSN 1175-5326 (Print edition)

2 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.

Key words: black fly, Simulium, new species, fauna, Vietnam

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6 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.

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Genus Simulium Latreille

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Keys to 10 subgenera of the genus Simulium in the Oriental Region

I. Subgenus Gomphostilbia Enderlein

12 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.

Keys to 25 species of the subgenus Gomphostilbia

14 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.

1) Simulium asakoae species-group

Simulium (Gomphostilbia) asakoae Takaoka & Davies, 1995

16 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.

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Simulium (Gomphostilbia) chaudinhense Takaoka & Sofian-Azirun sp. nov.

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Simulium (Gomphostilbia) confertum Takaoka & Sofian-Azirun, 2015

Simulium (Gomphostilbia) fuscidorsum Takaoka & Ya’cob, 2015

Distribution. Vietnam (Lam Dong).

Simulium (Gomphostilbia) hmongense Takaoka & Ya’cob sp. nov.

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Simulium (Gomphostilbia) hongthaii Takaoka, Sofian-Azirun & Ya’cob, 2014

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Simulium (Gomphostilbia) phulocense Takaoka & Chen, 2015

Simulium (Gomphostilbia) quychauense Takaoka & Chen sp. nov.

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Simulium (Gomphostilbia) sanchayense Takaoka & Lau sp. nov.

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Simulium (Gomphostilbia) tamdaoense Takaoka, Sofian-Azirun & Ya’cob, 2014

Distribution. Vietnam (Vinh Phuc).

Simulium (Gomphostilbia) unii Takaoka & Pham sp. nov.

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Simulium (Gomphostilbia) thituyenae Takaoka & Pham, 2015

Distribution. Vietnam (Lam Dong).

Simulium (Gomphostilbia) vinhphucense Takaoka & Low sp. nov.

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2) Simulium batoense species-group

Simulium (Gomphostilbia) dachaisense Takaoka & Lau, 2015

Distribution. Vietnam (Lam Dong).

Simulium (Gomphostilbia) duolongum Takaoka & Davies, 1995

Distribution. Peninsular Malaysia, Sumatra and Vietnam (Bac Thai).

Simulium (Gomphostilbia) lamdongense Takaoka & Sofian-Azirun, 2015

Distribution. Vietnam (Lam Dong).

Simulium (Gomphostilbia) longlanhense Takaoka & Ya’cob, 2015

Distribution. Vietnam (Lam Dong).

52 · Zootaxa 4261 (1) © 2017 Magnolia Press TAKAOKA ET AL.