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Acta Tropica 169 (2017) 170–186
Contents lists available at ScienceDirect
Acta Tropica
jo u r n al homep age: www.elsev ier .com/ locate /ac ta t ropica
he black flies (Diptera: Simuliidae) of the Lesser Sunda Archipelago,ndonesia
iroyuki Takaoka a,∗, Mohd Sofian-Azirun a, Zubaidah Ya’cob a, Chee Dhang Chen a,oon Weng Lau a, Van Lun Low b, I. Wayan Suana c
Institute of Biological Sciences, Faculty of Science, University of Malaya, Kuala Lumpur 50603, MalaysiaTropical Infectious Diseases Research and Education Centre (TIDREC), University of Malaya, Kuala Lumpur 50603, MalaysiaFaculty of Mathematics and Natural Science, University of Mataram (UNRAM), Mataram 83125, Indonesia
r t i c l e i n f o
rticle history:eceived 30 December 2016eceived in revised form 15 January 2017ccepted 16 January 2017vailable online 23 January 2017
eywords:lack flyew speciesali
a b s t r a c t
Surveys of pupae and larvae of black flies were carried out in Bali, Lombok, Sumbawa and Flores in theLesser Sunda Archipelago, Indonesia, where 10 species were known. A total of 14 simuliid species includ-ing four new species and five new records of the genus Simulium were collected, bringing the number ofspecies from the Lesser Sunda Archipelago to 19. They are classified into four subgenera: two in Neverman-nia, nine in Gomphostilbia, seven in Simulium and one in Wallacellum. One of four new species, Simulium(Simulium) baliense, is described based on females, males, pupae and larvae from Bali and Lombok. Thisnew species, which is placed in the Simulium striatum species-group of the subgenus Simulium, is closelyrelated to S. (S.) argyrocinctum De Meijere from Java and Sumatra, but it is distinguished from the latterspecies by the smaller number of the male enlarged upper-eye facets and larval abdomen lacking dorsal
ombokumbawandonesia
pairs of conical protuberances. The distribution record of S. (S.) upikae Takaoka & Davies from Flores iscorrected as that of S. (S.) eximium De Meijere. Some aberrant characters of the pupal gill filaments of S.(G.) atratum De Meijere, S. (G.) floresense Takaoka, Hadi & Sigit and S. (G.) sunapii Takaoka, Sofian-Azirun& Suana are illustrated. Characteristics of the fauna of black flies in this archipelago are briefly noted.Keys to all 19 species are provided for females, males, pupae and larvae.
© 2017 Elsevier B.V. All rights reserved.
. Introduction
In the Lesser Sunda Archipelago, Indonesia, the black fly faunaas represented by 10 species, all belonging to the genus Simulium
atreille, of which one species was recorded from Bali, nine fromlores, and four from Timor (Edwards, 1934; Takaoka et al., 2006;dler and Crosskey, 2016; Almet et al., 2016). The biting habits of
hese species remain unknown, although Simulium (Gomphostilbia)tratum De Meijere was reported to feed on the blood of domesticowls in Java (Friederichs, 1925).
We surveyed larvae and pupae of black flies in Bali, Lombok,umbawa and Flores in the Lesser Sunda Archipelago in September014 and February 2016, and collected 14 species of the genusimulium including four new species and five new records. Three
ew species, which all belong to the subgenus Gomphostilbia Ender-ein, were already described (Takaoka et al., 2016b, 2017). Theemaining new species, which is placed in the subgenus Simulium
∗ Corresponding author.E-mail address: [email protected] (H. Takaoka).
ttp://dx.doi.org/10.1016/j.actatropica.2017.01.016001-706X/© 2017 Elsevier B.V. All rights reserved.
Latreille, is here described based on females, males, pupae and lar-vae from Bali and Lombok. Some aberrant pupal gill filaments of S.(G.) atratum De Meijere, S. (G.) floresense Takaoka, Hadi & Sigit andS. (G.) sunapii Takaoka, Sofian-Azirun & Suana are illustrated. Char-acteristics of the fauna of black flies in this archipelago are brieflynoted. Keys to all 19 species are provided for females, males, pupaeand mature larvae.
2. Materials and methods
2.1. New species description and illustration
The methods of collection, description and illustration, andterms for morphological features used here, follow those of Takaoka
(2003) and partially those of Adler et al. (2004).The holotype and paratypes of the new species are depositedat Institute of Biological Sciences, Faculty of Science, University ofMalaya, Kuala Lumpur, Malaysia.
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.2. Systematics
The classification of Adler and Crosskey (2016) is adopted forreatment of 19 species of black flies recorded from the Lesserunda Archipelago. Definitions of subgenera and species-groupsollow those of Takaoka (1983, 2003, 2012), Takaoka and Davies1996) and Takaoka et al. (2014b).
. Results
Genus Simulium LatreilleSubgenus Gomphostilbia EnderleinSimulium asakoae species-groupSimulium (Gomphostilbia) gyorkosae Takaoka and Davies,
996Simulium (Eusimulium) metatarsale var. Edwards, 1934:
19–121 (Female, male and pupa)Simulium (Gomphostilbia) gyorkosae Takaoka and Davies, 1996:
8–32 (Female, male, pupa and larva),Specimens examined. One female, three males, six mature lar-
ae and 19 immature larvae, collected from a small stream (width.5–1.0 m, depth 5–10 cm, bottom sandy, water temperature 22 ◦C,haded, elevation 790 m, 08◦12′16.722′′S/115◦08′18.226′′E), run-ing in a forest, Gitgit, Bali, Indonesia, 27-IX-2014, by H. Takaoka,. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; 12
emales, eight males, 10 mature larvae and 14 immature larvae,ollected from a small stream (width 0.5–1.0 m, depth 10–15 cm,ottom sandy, water temperature 21 ◦C, shaded, elevation 712 m,8◦31′57.775′′S/116◦23′51.041′′E), running in a forest, Otak Koko
oben, Lombok, Indonesia, 25-IX-2014, by H. Takaoka, M. Sofian-zirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I. W. Suana.
Distribution. Bali (New record), Java, Lombok (New record),ulawesi and Sumatra.
Remarks. This species was originally described from Java byakaoka and Davies (1996), and recorded from Sumatra andulawesi (Takaoka et al., 2000; Takaoka, 2003). Simulium (G.)yorkosae is placed in the S. asakoae species-group, defined byakaoka (2012) and is characterized by the yellowish hair tuft of thedult female and male, male hind basitarsus enlarged, and cocoonith a short anterodorsal projection (Takaoka and Davies, 1996).
he males of this species from Bali and Lombok have a slightlymaller number of enlarged upper-eye facets, which is in 10 verti-al columns and 12 horizontal rows (10 or 11 vertical columns and3 or 14 horizontal rows in the original description). This is a newecord of this species from Bali and Lombok.
Simulium (Gomphostilbia) sunapii Takaoka et al., 2016Simulium (Gomphostilbia) sunapii Takaoka, Sofian-Azirun &
uana, in Takaoka et al. (2016b): 1–7 (Female, male, pupa and larva).Distribution. Flores.Remarks. This species was described from females, males,
upae and mature larvae collected from Flores by Takaoka et al.2016b). This record represents the most eastern distribution ofhe S. asakoae species-group in the archipelago. The one pupa waseported to have only seven gill filaments (Fig. 1A) on each sideTakaoka et al., 2016b). In a normal gill of this species, the ventralongest filament should have been divided into two filaments, for
total of eight filaments. This abnormal gill is interpreted to haveailed to form one of two filaments of the ventral pair.
Simulium batoense species-groupSimulium (Gomphostilbia) lemborense Takaoka and Sofian-
zirun, 2017Simulium (Gomphostilbia) lemborense Takaoka & Sofian-Azirun,
n Takaoka et al. (2017): (Female, male, pupa and larva)Distribution. Flores.
a 169 (2017) 170–186 171
Remarks. This species was described from females, males,pupae and mature larvae collected from Flores by Takaoka et al.(2017). This is the only species of the S. batoense species-group inthe archipelago. It is characterized by a narrow female frons andpupal gill with eight filaments, of which two filaments of the ventralpair are three to four times as long as the six other filaments.
Simulium ceylonicum species-groupSimulium (Gomphostilbia) brevilabrum Takaoka et al., 2006Simulium (Gomphostilbia) brevilabrum Takaoka et al., 2006: 8–13
(Female, male, pupa and larva)Distribution. Flores.Remarks. This species was described from females, males,
pupae and mature larvae collected from Flores by Takaoka et al.(2006). It is characterized by the pupal gill with four slender fila-ments.
Simulium (Gomphostilbia) rangatense Takaoka and Sofian-Azirun, 2016
Simulium (Gomphostilbia) rangatense Takaoka & Sofian-Azirun,in Takaoka et al. (2016b): 7–11 (Male and pupa).
Distribution. Flores.Remarks. This species was described from a male and its associ-
ated pupal exuviae collected from Flores by Takaoka et al. (2016b).It is characterized by the pupal gill with six filaments, of which twofilaments of the middle pair have an elongate stalk.
Simulium (Gomphostilbia) rutengense Takaoka et al., 2006Simulium (Gomphostilbia) rutengense Takaoka et al., 2006: 13–17
(Male, pupa and larva).Distribution. Flores.Remarks. This species was described from males, pupae and
mature larvae collected from Flores by Takaoka et al. (2006). It ischaracterized by the presence of minute tubercles on pupal abdom-inal segments 1, 2 and 9, and the pupal gill with eight filaments.
Simulium epistum species-groupSimulium (Gomphostilbia) atratum De Meijere, 1913Simulium atratum De Meijere, 1913: 331–332 (Female and
male).Simulium (Gomphostilbia) atratum: Takaoka and Davies, 1996:
18.Simulium (Gomphostilbia) sundaicum Edwards, 1934: Takaoka
and Davies, 1996: 19–22 (Female, male, pupa and larva), syn-onymized by Takaoka (2012).
Specimens examined. One female, four males, six pupae,nine pupal exuviae, two mature larvae and eight immature lar-vae, collected from a stream (width 2–3 m, depth 60 cm, bottomsandy, water temperature 22 ◦C, partially shaded, elevation 538 m,08◦23′56.592′′S/115◦17′52.803′′E), moderately running in a for-est, Sebatu, Lombok, Indonesia, 27-IX-2014, by H. Takaoka, M.Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; 11females, 10 males, nine mature larvae and 152 immature lar-vae, collected from a small stream (width 1.0 m, depth 40–50 cm,bottom sandy, water temperature 20 ◦C, shaded, elevation 830 m,08◦19′53.640′′S/115◦13′42.267′′E), running in a natural forest,Nung nung waterfall, Bali, Indonesia, 27-IX-2014, by H. Takaoka,M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; 12females, 22 males, 20 mature larvae and 100 immature larvae,collected from a small stream (width 0.5–1.0 m, depth 10–15 cm,bottom sandy, water temperature 21 ◦C, shaded, elevation 712 m,08◦31′57.775′′S/116◦23′51.041′′E), running in a forest, Otak KokoJoben, Lombok, Indonesia, 25-IX-2014, by H. Takaoka, M. Sofian-
Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; four femaleand seven males, collected from a small stream (width 1 m, depth20–40 cm, bottom sandy and rocky, water temperature 24 ◦C, par-
172 H. Takaoka et al. / Acta Tropica 169 (2017) 170–186
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Fig. 1. Aberrant pupal gills (lateral view). A, Right gill of S. (G.)
ially shaded, elevation 135 m, 08◦11′45.024′′S/117◦45′50.812′′E),adindi Atas, Perat, Dompu, Sumbawa, Indonesia, 22-II-2016, by. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; 10
emales, 15 males, collected from a waterfall (width 20–30 cm,epth 2–3 cm, rocks, water temperature 20 ◦C, exposed to the sun,levation 1093 m, 08◦41′03.955′′S/120◦19′17.135′′E), Nteer, Mang-arai, Flores, Indonesia, 27-II-2016, by M. Sofian-Azirun, Z. Ya’cob,.D. Chen, K.W. Lau & I.W. Suana.
Distribution. Bali (New record), Flores, Java, Lombok (Newecord), Sumatra, Sumbawa (New record) and Timor.
Remarks. Among 19 species of the Lesser Sunda Archipelago,nly S. (G.) atratum was reported to feed on the blood of domes-ic fowls in Java (Friederichs, 1925). However, its detailed bitingehavior, feeding time, transmission of pathogens and damage to
he poultry farming remain to be investigated.This species was originally described from adult females andales collected from West Java (De Meijere, 1913). Simulium (G.)
ii; B, Left gill of S. (G.) atratum. Scale bars. 0.1 mm for A and B.
sundaicum Edwards was synonymized with this species by Takaoka(2012). The redescription of the female, male, pupa and larvae ofthis species by Takaoka and Davies (1996) was used to identify S.(G.) atratum. In the Archipelago, this species was recorded underthe name of S. (G.) sundaicum from Flores by Takaoka et al. (2006)and from Timor by Almet et al. (2016).
This species is a member of the S. epistum species-group, definedby Takaoka (2012), and is characterized by the hind tibiae of theadult female and male with a subbasal dark spot, male hind basitar-sus narrow and parallel-sided, and pupal gill with eight filaments,of which two filaments of the ventral pair are much longer thanthe six other filaments. This species represents a new record fromBali, Lombok and Sumbawa, where it is one of the most commonsimuliid species, as in Java and Sumatra (Takaoka and Davies, 1996;
Takaoka et al., 2000).The male large upper-eye facets are variable in number, rangingfrom 12 to 15 vertical columns and 14 to 16 horizontal rows (14
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ertical columns and 16 horizontal rows in the redescription byakaoka and Davies (1996)). The dorsal surface of pupal abdominalegments 1–3 are darkened, though pale yellow in the redescriptiony Takaoka and Davies (1996).
One of 20 pupae collected from a stream at Sebatu, Bali, had andditional short branch (0.20 mm and 0.23 mm long) arising fromhe dorsal (or outer) filament of the ventral pair of the gill on eachide (Fig. 1B). A similar additional short branch, which is interpretedo be aberrant, is also known in one pupa of S. (S.) tamorense Takaokand Shrestha from Nepal (2010).
Two of 152 immature larvae collected from Bali were infectedith mermithid nematode larvae.
Simulium (Gomphostilbia) floresense Takaoka et al., 2006Simulium (Gomphostilbia) floresense Takaoka et al., 2006: 2–8
Female, male, pupa and larva).Specimens examined. Three females, three males, collected
rom a fast-flowing river (width 10–15 m, depth 30–50 cm, bottomandy and rocky, water temperature 20 ◦C, exposed to the sun, ele-ation 890 m, 08◦35′27.877′′S/120◦26′05.799′′E), Waegarit, Ruteng,lores, Indonesia, 27-II-2016, by M. Sofian-Azirun, Z. Ya’cob, C.D.hen, K.W. Lau & I.W. Suana.
Distribution. Flores.Remarks. This species was described from adult females, males,
upae and larvae from Flores (Takaoka et al., 2006). It is charac-erized by the outer filament of the ventral pair of the pupal gillhickest and longest of all the eight filaments.
The one pupa of this species collected from Flores shows an aber-ant right gill (Fig. 2B, C), though the left gill is normal, consistingf eight thread-like filaments (Fig. 2A). The right gill is composed ofne short trunk with a round apex, one smaller short branch aris-
ng from the inner surface of the main trunk, and two smaller shortranches with a short stalk arising from the dorsal surface of theain trunk (Fig. 2B, C).
Simulium (Gomphostilbia) merapiense Takaoka and Sofian-zirun, 2016
Simulium (Gomphostilbia) merapiense Takaoka & Sofian-Azirun:n Takaoka et al. (2016a): 76–82 (Female, male, pupa and larva).
This species was described from females, males, pupae andature larvae collected from Java by Takaoka et al. (2016a). It is
haracterized by the pupal gill with eight short filaments, bare pupaead and thoracic integuments and small larval postgenal cleft.
The only mature larva collected from Lombok has some charac-ers which are somewhat different from those of S. (G.) merapienseescribed from Java (Takaoka et al., 2016a). The description of the
arva from Lombok is given below.Mature larva. Body length 3.8 mm. Body blackish except
bdominal segments 2, 3 and 7 grayish, when viewed dorsally.ead capsule sparsely covered with colorless setae. Cephalic apo-
ome dark yellow though anterior two-fifths whitish yellow, withight brown head spots. Lateral surface of head capsule dark yel-ow except eye-spot region white and areas dorsal and posterioro eye-spot region light brown; spots in front of posterior margin
erged into background dark area. Ventral surface of head capsuleight brown except area near anterior margin on each side yellowishnd areas near both lateral margins dark yellow; elongate spot onach side of postgenal cleft obscured. Antenna composed of threerticles and apical sensillum, slightly longer than stem of labral fan;ength ratio of three articles (from base to tip) 1.00:0.72:0.77. Labralan with 32 primary rays. Mandible (Fig. 3A) with mandibular ser-ation composed of two teeth (one medium-sized, one small); main
ooth at acute angle against mandible on apical side; comb-teethradually decreased in length from first to third; supernumeraryerrations absent. Hypostoma (Fig. 3B) with nine anterior teeth, ofhich median and corner teeth subequal in length to each other,a 169 (2017) 170–186 173
followed by three intermediate teeth on each side; lateral marginssmooth; five or six hypostomal bristles slightly divergent posteri-orly from lateral border on each side. Postgenal cleft (Fig. 3C) small,quadrate, 0.38 times length of postgenal bridge; sheath of sube-sophageal ganglion unpigmented. Cervical sclerites on each sidecomposed of light-brown elliptical piece, not fused to occiput. His-toblast of pharate pupal gill with eight short thread-like slenderfilaments, all arising from short common basal stalk, which hasrelatively swollen basal fenestra (Fig. 3D). Thorax and abdomenwithout pair of dorsolateral protuberances. Thoracic cuticle almostbare. Abdominal cuticle sparsely covered with colorless minutesetae except last segment moderately covered with short colorlesssetae on each side of anal sclerite down to base of ventral papillae.Rectal scales not discernible. Rectal organ of three lobes, simple,without secondary lobules (Fig. 3E). Anal sclerite X-shaped, withanterior arms slightly shorter than posterior ones. Last abdominalsegment bulged laterally and having conical ventral papillae. Pos-terior circlet with 82 rows of hooklets with up to 12 hooklets perrow.
Specimen examined. One larva, collected from a stream (width2–3 m, depth 50 cm, water temperature 23 ◦C, partially shaded,elevation 599 m, 08◦32′01.474′′S/116◦20′29.394′′E), moderatelyflowing in a forest, Benang Stokel waterfall, Lombok, Indonesia, 25-IX-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W.Lau & I.W. Suana.
Distribution. Lombok (New record) and Java.Remarks. The unique larva collected from Lombok is identified
as S. (G.) merapiense based on the same arrangement of the pharatepupal gill filaments (Fig. 3D) and similar body color pattern. How-ever, there are some differences in certain features (characters oflarvae from Java in parentheses): length ratio of the postgenal cleftagainst the postgenal bridge (Fig. 3C) 0.38 (0.27), rectal organ sim-ple (Fig. 3E) (compound, each lobe with one or two short nipple-likesecondary lobules) and number of rows of the posterior circlet 82(96–102).
A combination of the small postgenal cleft and simple rectalorgan in the larva from Lombok, though occurring rarely in lar-vae of species of the subgenus Gomphostilbia, is also observed inS. (G.) guniki Takaoka, which was originally described from Sabah,Malaysia (Takaoka, 2001), and was placed in the S. darjeelingensespecies-group, defined by Takaoka (2012). However, this larva isdistinguished from that of S. (G.) guniki by the relatively short com-mon basal stalk of the pharate pupal gill with somewhat inflatedbasal fenestra (Fig. 3D), and second antennal article shorter thanthe first one (its relative length is 0.72) (second and first antennalarticles of S. (G.) guniki are equal in length).
Subgenus Nevermannia EnderleinSimulium feuerborni species-groupSimulium (Nevermannia) feuerborni Edwards, 1934Simulium feuerborni Edwards, 1934: 129–132 (Male, pupa and
larva).Simulium (Nevermannia) feuerborni: Takaoka and Davies, 1996:
10–13 (Female, male, pupa and larva).Distribution. Bali, Java and Sumatra.Remarks. This species was originally described from males,
pupae and larvae collected from Java and Bali (Edwards, 1934). InBali, pupae and larvae of this species were collected from a water-fall at Ljemampeh, Padangombo river at Baturiti, and Baturiti riverbetween Baturiti and Batunge, all at high elevations (1000–1100 mabove sea level) (Edwards, 1934). In our survey, this species wasnot collected.
Simulium ruficorne species-groupSimulium (Nevermannia) aureohirtum Brunetti, 1911Simulium aureohirtum Brunetti, 1911: 283–288 (Male).
174 H. Takaoka et al. / Acta Tropica 169 (2017) 170–186
Fig. 2. Pupal gills of S. (G.) floresense. A, Normal gill (left side; lateral view); B and C, Aberrant gills (right side; B, lateral view; C, dorsal view). Scale bars. 0.1 mm for A–C.
Fig. 3. Larvae of S. (G.) merapiense. A, Mandible; B, Hypostoma; C, Head capsule showing postgenal cleft (ventral view); D, Basal portion of pharate pupal gill (left side; dorsalview); E, Rectal organ with three simple lobes (dorsal view). Scale bars. 0.05 mm for C; 0.02 mm for A, B, D and E.
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Simulium (Nevermannia) aureohirtum: Ogata (1956): 61–62;gata (1966): 129; Takaoka and Roberts (1988): 194–195; Takaoka
2003): 37–45 (Female, male, pupa and larva).Simulium (Eusimulium) aureohirtum: Puri (1933): 1–7 (Female,
ale, pupa and larva); Takaoka (1979): 382–384 (Female, male,upa and larva).
Specimens examined. One female, one pupal exuviae and oneature larva, collected from a river (width 5–8 m, depth 5 cm,
ottom sandy, water temperature 21 ◦C, exposed to the sun, eleva-ion 137 m, 08◦35′46.850′′S/116◦12′18.327′′E), Narmada, Lombok,ndonesia, 23-IX-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob,.D. Chen, K.W. Lau & I.W. Suana.
Distribution. India, Bhutan, China, Guam, Indonesia (Java, Flo-es, Halmahera, Lombok (New record), Sumatra, Sulawesi andimor), Japan, Malaysia, Nepal, Pakistan, Philippines, Sri Lanka,aiwan, Thailand and Vietnam.
Remarks. This species is characterized by the unique female andale genitalia, pupal gill with six filaments and larval head capsuleith bold head spots (Takaoka, 2003). As noted previously (Takaoka
t al., 2014a), the female autogeny and larval adaptability to rela-ively high water temperature and slow flow, together with otheractors, might explain why this species is widely distributed in theriental Region and extends into the Palearctic and Australasianegions. Given its wide geographical range, it might be a complexf species.
Subgenus Simulium LatreilleSimulium christophersi species-groupSimulium (Simulium) nebulicola Edwards, 1934Simulium (Simulium) nebulicola Edwards, 1934: 114–115 (Male);
akaoka and Davies (1996): 71–75 (Female, male, pupa and larva).Specimens examined. Three males, five pupal exuviae, col-
ected from a cascading stream (width 20–30 cm, depth 2–3 cm,ottom rocks, water temperature 20 ◦C, exposed to the sun, eleva-ion 1093 m, 08◦41′03.955′′S/120◦19′17.135′′E), Nteer, Manggarai,lores, Indonesia, 27-II-2016, by M. Sofian-Azirun, Z. Ya’cob, C.D.hen, K.W. Lau & I.W. Suana.
Distribution. Java and Flores.Remarks. This species was described from a single male from
ast Java by Edwards (1934), and its female, pupa and larva wereescribed by Takaoka and Davies (1996).
This species was placed in the S. christophersi species-group,ecently defined by Takaoka et al. (2014b). It was recorded onlyrom Flores in the Lesser Sunda Archipelago (Takaoka et al., 2006).
Three males reared from pupae collected from Flores have thenlarged upper-eye facets in 16 vertical columns and 17 horizontalows, slightly greater in number than those of males from Java (in5 vertical columns and 16 horizontal rows). The pupae have theorsal surface of abdominal segment 1 medium brown and that ofegment 2 with a light brown area covered with numerous comb-ike groups of minute spines anterosubmedially (similar to that of. (S.) javaense–Fig. 4B) on each side; abdominal segments 5–9 haveomb-like groups of minute spines in transverse rows on each side.one of these characters are mentioned in the original description.
Simulium eximium species-groupSimulium (Simulium) eximium De Meijere, 1913Simulium eximium De Meijere, 1913: 130 (Female and male)Simulium (Simulium) eximium: Edwards (1934): 104–106
Female, male, pupa and larva); Takaoka and Davies (1996): 48–54Female, male, pupa and larva).
Specimens examined. One pharate male (with associated
upal exuviae), collected from a river (width 8–15 m, depth0–60 cm, water temperature 20 ◦C, partially shaded, elevation27 m, 08◦19′42.039′′S/115◦13′44.116′′E), moderately running innatural forest, Nung Nung waterfall, Bali, Indonesia, 27-IX-
a 169 (2017) 170–186 175
2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen,K.W. Lau & I.W. Suana; 10 immature larvae, collected from astream (width 0.5–2.5 m, depth 5–15 cm, bed of pebbles andsands, water temperature 20 ◦C, partially shaded, elevation 1010 m,08◦15′26.684′′S/115◦04′12.625′′E), moderately running in a for-est, Munduk, Bali, Indonesia, 27-IX-2014, by H. Takaoka, M.Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana;One pupa, collected from a stream (width 0.5–3.0 m, depth0.5–1.0 m, water temperature 23 ◦C, partially shaded, elevation630 m, 08◦32′01.474′′S/116◦20′29.394′′E), moderately running ina forest, Benang Stokel waterfall, Lombok, Indonesia, 25-IX-2014,by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau &I.W. Suana; One female, three pupae and one pupal exuviae col-lected from a fast-flowing river (width 10–15 m, depth 30–50 cm,bottom of sands and rocks, water temperature 20 ◦C, exposed tothe sun, elevation 890 m, 08◦35′27.877′′S/120◦26′05.799′′E), Wae-garit, Ruteng, Flores, Indonesia, 27-II-2016, by M. Sofian-Azirun, Z.Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana.
Distribution. Bali (New record), Flores, Java and Lombok (Newrecord).
Remarks. This species was originally described from adultfemales and males collected from East Java (De Meijere, 1913) andits pupa and larva were described by Edwards (1934). This specieswas placed in the S. eximium species-group and its female, male,pupa and larva were redescribed by Takaoka and Davies (1996),who pointed out the difference in certain morphological characters,such as the number of the male upper-eye facets and the presenceor absence of the apical spine of the style, between East and WestJavan populations, and further noted the variation in the numberand length of the gill filaments observed in the pupae collected fromWest Java.
The one pharate male collected from Bali is characterized byenlarged upper-eye facets in 15 vertical and 16 horizontal rows, andits associated pupal exuviae has a different number of gill filamentson each side (10 filaments on the left side and nine filaments on theright side), both characters agreeing with those of the West Javanpopulations (Takaoka and Davies, 1996).
A female with its associated pupal exuviae, three pupae and onepupal exuviae of the S. eximium species-group collected from Floresin February 2016 were identified as S. (S.) eximium by the shape ofthe paraproct and relative lengths of 10 pupal gill filaments (i.e.,ventral paired filaments are not shortened, as in S. (S.) upikae). Oneof five pupae from Flores has nine filaments on the left side and 10filaments on the right side.
Reexamination of the female and pupae of the species identifiedas S. (S.) upikae Takaoka & Davies from Flores (Takaoka et al., 2006)shows that most morphological characters including the shape ofthe paraproct, number of tubercles on the cibarium, and relativelengths of the pupal gill filaments agree with those of S. (S.) eximium.For this reason, the previous distribution record of S. (S.) upikaefrom Flores is here corrected as that of S. (S.) eximium, although thecocoon is rather similar in shape to that of S. (S.) upikae.
Simulium melanopus species-groupSimulium (Simulium) iridescens De Meijere, 1913Simulium iridescens De Meijere, 1913: 333 (Female and male)Simulium (Simulium) iridescens: Edwards (1934): 112–114
(Female, male, pupa and larva); Takaoka and Davies (1996): 67–70(Female, male, pupa and larva).
Specimens examined. One female, five males, five maturelarvae and 44 immature larvae, collected from a stream(width 0.5–2.5 m, depth 5–15 cm, bottom of pebbles and sands,
water temperature 20 ◦C, partially shaded, elevation 1010 m,08◦15′26.684′′S/115◦04′12.625′′E), moderately running in a forest,Munduk, Bali, Indonesia, 27-IX-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; One female,
176 H. Takaoka et al. / Acta Tropica 169 (2017) 170–186
Fig. 4. Pupa of S. (S.) javaense. A, Gill filaments (left side; lateral view); B, Dorsal surface of abdominal segment 2 showing dark area with minute spines, together with ones A; 0.0
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our pupae, collected from a stream (width 0.5–3.0 m, depth.5–1.0 m, water temperature 23 ◦C, partially shaded, elevation30 m, 08◦32′01.474′′S/116◦20′29.394′′E), moderately running in aorest, Benang Stokel waterfall, Lombok, Indonesia, 25-IX-2014, by. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W.uana.
Distribution. Bali (New record), Java, Lombok (New record) andumatra.
Remarks. This species was originally described from adultemales and males collected from East Java (De Meijere, 1913) andts pupa and larva were described by Edwards (1934). It was placedn the S. melanopus species-group, defined by Takaoka (1983) andts female, male, pupa and larva were redescribed by Takaoka andavies (1996). The difference in the number of the male upper-ye facets was noted between the East and West Javan populationsnumbers of vertical columns: 13 or 14 for East Java versus 18 for
est Java) (Takaoka and Davies, 1996). The males collected fromali have the enlarged upper-eye facets in 16–18 vertical columnsnd 18 horizontal rows, similar to West Javan males. The pupaerom Bali and Lombok have the dorsal surface of abdominal seg-
ent 1 medium brown and that of segment 2 with a light brownrea covered with numerous comb-like groups of minute spines
nterosubmedially (similar to that of S. (S.) javaense–Fig. 4B) onach side; segment 6 has comb-like groups of minute spines onach side.5 mm for B.
Simulium (Simulium) javaense Takaoka and Hadi, 1991Simulium iridescens var. Edwards, 1934: 114 (Male, pupa and
larva).Simulium (Simulium) javaense Takaoka and Hadi (1991):
363–368 (Female, male, pupa and larva).Specimens examined. 16 females, 11 males, 12 mature larvae
and 29 immature larvae, collected from a stream (width 0.2 m,depth 8–10 cm, bottom rocky, water temperature 23 ◦C, exposedto the sun, elevation 599 m, 08◦23′56.429′′S/115◦17′50.434′′E),fast running near a forest, Sebatu, Bali, Indonesia, 27-IX-2014,by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W.Lau & I.W. Suana; three females, two males, two pupae,five mature larvae and 34 immature larvae, collected froma small stream (width 0.2–0.5 m, depth 5–7 cm, bed sandy,water temperature 22 ◦C, exposed to the sun, elevation 884 m,08◦19′46.331′′S/115◦13′23.372′′E), moderately running in paddyfields, Nung Nung, Bali, Indonesia, 27-IX-2014, by H. Takaoka, M.Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana.
Distribution. Bali (New record) and Java.Remarks. The pupae of S. (S.) javaense from Bali have gills with
much more inflated filaments (Fig. 4A) than those from Java (rela-tive greatest thickness of gill filaments against the interspiracular
trunk 2.4–3.6 in pupae from Bali versus 1.3–2.0 in pupae from Java).The difference in thickness of the pupal gill filaments is interpretedto be intraspecific variation. Morphological pupal characters, whichwere not mentioned in the original description, include the dor-
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al surface of abdominal segment 1 entirely darkened and withoutubercles, that of segment 2 partially darkened anterosubmedially,.e., a darkened area densely covered with numerous comb-likeroups of minute spines (Fig. 4B) on each side, that of segment
with comb-like groups of minute spines on each side, basal por-ions of spine-combs of segments 7 and 8 and those of comb-likeroups of minute spines of segment 9 somewhat darkened.
Simulium nobile species-groupSimulium (Simulium) nobile De Meijere, 1907Simulium nobile De Meijere, 1907: 206 (Male)Simulium (Simulium) nobile: Edwards (1934): 115 (Female, male,
upa and larva); Takaoka and Davies (1996): 61–66 (Female, male,upa and larva).
Specimens examined. One female, one male, one pupal exu-iae and three mature larvae, collected from a stream (width 0.2 m,epth 8–10 cm, bottom rocky, water temperature 23 ◦C, exposedo the sun, elevation 599 m, 08◦23′56.429′′S/115◦17′50.434′′E), fastunning near a forest, Sebatu, Bali, Indonesia, 27-IX-2014, by H.akaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I. W.uana.
Distribution. Bali (New record), Java and Sumatra.Remarks. This species was originally described from adult males
ollected from Central Java (De Meijere, 1907) and its female, pupand larva were briefly described by Edwards (1934). It was placed inhe S. nobile species-group and its female, male, pupa and larva wereedescribed by Takaoka and Davies (1996). The previous distribu-ion record of this species from Sabah and Peninsular Malaysia wasorrected using DNA sequence-based analyses (Low et al., 2016).
Simulium (Simulium) timorense Takaoka et al., 2006Simulium (Simulium) timorense Takaoka et al., 2006: 18–24
Female, male, pupa and larva).Specimens examined. 12 females, 11 males, five mature larvae,
ollected from a stream (width 5–8 m, depth 5 cm, bottom sandy,ater temperature 21 ◦C, exposed to the sun, elevation 137 m,
8◦35′46.850′′S/116◦12′18.327′′E), Narmada, Lombok, Indonesia,3-IX-2014, by H. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen,.W. Lau & I.W. Suana; eight females, nine males, 27 pupae and fiveature larvae, collected from a river (width 6–8 m, depth 30 cm,
ottom rocky, water temperature 24.5 ◦C, exposed to the sun, eleva-ion 72 m, 08◦28′32.854′′S/118◦48′23.844′′E), Sungai Toloweki, Kelunger, Sumbawa, Indonesia, 22-II-2016, by M. Sofian-Azirun, Z.a’cob, C.D. Chen, K.W. Lau & I. W. Suana.
Distribution. Flores, Lombok (New record), Sumbawa (Newecord) and Timor.
Remarks. This species was originally described from Timor andlores, Indonesia, and placed in the S. nobile species-group (Takaokat al., 2006). This species is distinguished from S. (S.) nobile byacking dorsal pairs of conical protuberances on larval abdominalegments 1–5.
Simulium striatum species-groupSimulium (Simulium) baliense Takaoka & Sofian-Azirun sp.
ov.Female. Body length 2.1–2.4 mm. Head. Slightly narrower than
horax. Frons black, shiny with bluish reflection when illumi-ated at certain angles, with several dark stout hairs along lateralargins and few hairs just above lower margin; frontal ratio
.31:1.00:1.43; frons:head ratio 1.00:4.31. Fronto-ocular area welleveloped, short, directed laterally, and rounded apically. Clypeusrownish black, slightly shiny and gray pruinose when illuminatedt certain angles, moderately covered with dark-brown medium-
ong hairs. Labrum 0.69 times length of clypeus. Antenna composedf scape, pedicel and nine flagellomeres, medium to dark brownxcept scape, pedicel, and first and second flagellomeres yellowa 169 (2017) 170–186 177
when viewed anteriorly (medium brown to dark brown exceptscape, pedicel and base of first flagellomere yellow when viewedposteriorly). Maxillary palp with five segments, medium brownexcept first and second segments ochreous or light brown, andthird segment dark brown; proportional lengths of third, fourth,and fifth segments 1.00:1.11:2.24; third segment (Fig. 5A) of nor-mal size, with medium-sized ellipsoidal sensory vesicle (0.33–0.35times length of third segment) having large or medium-sized open-ing. Maxillary lacinia with 10 inner and 12 or 13 outer teeth.Mandible with 24 inner and 13 outer teeth. Cibarium (Fig. 5B) with18 min processes near posterodorsal margin. Thorax. Scutum black,shiny, densely covered with whitish recumbent short hairs andsparsely with several dark-brown long upright hairs on prescutel-lar area; scutum gray pruinose with five non-pruinose longitudinalvittae (one medial, two submedial, and two lateral), medial andsubmedial vittae well defined from anterior margin to posteriorportion and submedial and lateral vittae united widely near ante-rior margin, all vittae united with transverse non-pruinose bandon prescutellar area, when illuminated in front and viewed dor-sally; scutum gray pruinose except four non-pruinose longitudinalvittae, when illuminated posteriorly and viewed dorsally. Scutel-lum medium brown, covered with dark-brown upright long hairsand yellow short hairs. Postnotum brownish black, shiny, whitepruinose when illuminated at certain angles, and bare. Pleuralmembrane bare. Katepisternum dark brown, longer than deep,shiny when illuminated at certain angles, and bare. Legs. Foreleg:coxa yellowish white; trochanter yellowish white except apical halfsomewhat darkened and ventral surface light brown; femur lightbrown except apical cap medium brown; tibia medium to darkbrown; tarsus brownish black, with moderate dorsal hair crest;basitarsus moderately dilated, 6.5 times as long as its greatestwidth. Midleg: coxa brownish black; trochanter medium brownexcept base whitish yellow; femur medium brown except apicalcap dark brown; tibia medium brown with apical cap dark brownand extreme base whitish yellow; tarsus medium brown exceptbasal two-thirds of basitarsus whitish yellow. Hind leg: coxa darkbrown; trochanter whitish yellow; femur medium brown exceptbase whitish yellow; tibia medium to dark brown except basewhitish yellow; tarsus medium brown except little more than basalhalf of basitarsus and basal half of second tarsomere whitish yel-low; basitarsus (Fig. 5C) nearly parallel-sided, 5.70 times as long aswide, and 0.68 and 0.60 times as wide as greatest widths of hindtibia and femur, respectively; calcipala (Fig. 5C) moderately devel-oped, slightly shorter than wide, and 0.47 times as wide as greatestwidth of basitarsus; pedisulcus (Fig. 5C) well developed. Claw sim-ple, without tooth. Wing. Length 2.0 mm. Costa with dark spinulesand hairs; subcosta haired except near apex bare; basal sectionof radius fully haired; R1 with dark brown spinules and hairs; R2with 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 light-brown hairs. Dor-sal surface of abdomen medium brown to brownish black, withlight to dark brown short hairs; tergite 2 shiny and silvery irides-cent when illuminated at certain angles and tergites 6–9 shiny.Ventral surface of seventh segment with pair of weakly sclero-tized submedian sternal plates. Terminalia. Sternite 8 (Fig. 5D) withposterior margin concave medially in form of reversed-U shape,bare medially, with 17–22 dark-brown medium-long to long stouthairs and 16–19 yellow short hairs on each lateral surface. Ovipos-itor valves (Fig. 5D, E) rectangular, with ventrally produced lobenear inner margin, membranous except narrow area along innermargin slightly sclerotized, covered with 23–25 short yellow hairs
and numerous microsetae; inner margins slightly sinuous, some-what separated from each other. Genital fork (Fig. 5F) of inverted-Yform, with long, narrow well-sclerotized stem; arms of narrowwidth, each with short, moderately sclerotized, triangular projec-
178 H. Takaoka et al. / Acta Tropica 169 (2017) 170–186
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ig. 5. Female of S. (S.) baliense sp. nov. A, Third segment of maxillary palp showinecond tarsomere of hind leg (left side; outer view); D and E, Sternite 8 and oviparaprocts and Cerci (right side; G, ventral view; H, lateral view); I, Spermatheca. S
ion directed dorsally. Paraproct in ventral view (Fig. 5G) rounded,ubequal in length to greatest width, strongly pigmented on ante-ior surface, with 33–35 yellow and dark short to medium-longairs on lateral and ventral surfaces, and with four or five shortensilla on anteromedial surface; paraproct in lateral view (Fig. 5H)early 0.48 times as long as wide, and much protruding ventrallyeyond ventral margin of cercus. Cercus in lateral view (Fig. 5H)hort, 0.56 times as long as wide, with numerous medium-longairs, and rounded posteriorly. Spermatheca (Fig. 5I) nearly ovoid,.31 times as long as greatest width, well sclerotized except portionf junction with duct moderately unsclerotized, without definiteeticulate patterns on its surface; internal setae present; accessoryucts subequal in thickness to each other, and slightly thicker thanajor duct.
Male. Body length 2.5 mm. Head. Slightly wider than thorax.pper-eye large facets in 14 or 15 vertical columns and 15 horizon-
al rows. Clypeus black, thickly white pruinose and iridescent whenlluminated at certain angles, with dark-brown hairs along and near
ory vesicle (right side; frontal view); B, Cibarium (frontal view); C, Basitarsus and valves (D, ventral view; E, lateral view); F, Genital fork (ventral view); G and H,ars. 0.1 mm for C; 0.02 mm for A, B, D–I.
lateral margins. Antenna composed of scape, pedicel and nine flag-ellomeres, medium brown except scape and pedicel light brownand base of first flagellomere yellow; first flagellomere elongate,1.75 times length of second one. Maxillary palp with five segments,medium brown except first and second segments ochreous andfifth segment grayish light-brown; proportional lengths of third,fourth, and fifth segments 1.00:1.28:2.88; third segment (Fig. 6A)of normal size; sensory vesicle (Fig. 6A) ellipsoidal, 0.23–0.26 timeslength of third segment, and with small opening. Thorax. Scutumbrownish black, with whitish pruinose pattern, i.e., anterior pair oflarge spots on shoulders extended posteriorly along lateral marginsand connected to large transverse spot entirely covering prescutel-lar area, anterior pair of large spots divided into anterior half andposterior half, either of which disappears depending on direction
of lights; all these spots brilliantly iridescent when illuminated atcertain angles; scutum uniformly and moderately covered withyellow recumbent short hairs and with several dark-brown longupright hairs on prescutellar area. Scutellum medium brown, with
H. Takaoka et al. / Acta Tropica 169 (2017) 170–186 179
Fig. 6. Male of S. (S.) baliense sp. nov. A, Third segment of maxillary palp showing sensory vesicle (right side; frontal view); B, Basitarsus and second tarsomere of hind leg (leftside; outer view); C, Coxites, styles, ventral plate and median sclerite (ventral view); D, Coxite and style (right side; ventrolateral view); E, Style (right side; medial view); F,S view)I erci (C
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tyle showing short basal projection with serrated margin (right side; posteodorsal, Paramere (right side; posteroventral view); J and K, Abdominal segment 10 and c–K.
ark-brown long upright hairs and yellow short hairs. Postnotumrownish black and bare. Pleural membrane bare. Katepisternum
onger than deep, brownish black, and bare. Legs. Coloration nearlys in female. Fore basitarsus moderately dilated, 5.64 times as longs its greatest width. Hind basitarsus (Fig. 6B) enlarged, 0.9 timess wide as greatest width of hind tibia, which is as wide as that ofind femur; calcipala (Fig. 6B) developed, small, as long as its basalidth; pedisulcus (Fig. 6B) well developed. Wing. Length not mea-
urable. Other characters as in female except subcosta and basalortion of radius bare. Abdomen. Basal scale brownish black, with
ringe of light-brown long hairs. Dorsal surface of abdomen darkrown to brownish black, moderately covered with dark-brownhort to medium-long hairs; segments 2, 5, 6 and 7 each withair of whitish pruinose spots (brilliantly iridescent when illumi-
; G, Ventral plate and median sclerite (lateral view); H, Ventral plate (caudal view);right side; J, lateral view; K, caudal view). Scale bars. 0.1 mm for B; 0.02 mm for A,
nated at certain angles) dorsolaterally, those on segment 2 broadlyconnected in middle to each other. Genitalia. Coxites, styles, ven-tral plate and median sclerite in ventral view as in Fig. 6C; coxitein ventrolateral view (Fig. 6D) rectangular, 0.75 times as long aswide; style in ventrolateral view (Fig. 6D) 1.74 times length ofcoxite, 2.90 times as long as greatest width at base, nearly parallel-sided from base to basal one-fourth, then narrowed to middle,then nearly parallel-sided, with rounded apex having subapicalspine; style in medial view (Fig. 6E) somewhat flattened dorsoven-tally, with short basal protuberance directed dorsomedially; style
in posterodorsal view (Fig. 6F) with short basal protuberance hav-ing several spines along its anterior margin. Ventral plate in ventralview (Fig. 6C) with body broad, with lateral margins slightly nar-rowed posteriorly, anterior margin deeply concave, and posterior
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argin rounded; body bearing prominent median process sharplyarrowed to round tip; body covered with minute setae medially;rms short, stout, divergent from base; ventral plate in lateral viewFig. 6G) with median process abruptly bent ventrally at nearly rightngle; ventral plate in caudal view (Fig. 6H) in form of equilateralriangle, and bare. Median sclerite in ventral view (Fig. 6C) arisingnterior to anteromedian portion of ventral plate; median scleriten lateral view (Fig. 6G) slightly curved dorsally. Paramere in ventraliew (Fig. 6I) enlarged basally, with several hooks apically. Aedea-al membrane sparsely covered with minute setae; dorsal plate notclerotized. Abdominal segment 10 (Fig. 6J, K) with several shorterairs together with numerous minute setae on ventral surface andhree to five short hairs on lateral surface; cercus (Fig. 6J, K) small,ound with 14 or 15 short to medium-long hairs.
Pupa. Body length 2.5 mm. Head. Integument yellow, denselyovered with round tubercles except most of antennal sheathsare; frons with two pairs of unbranched medium-long slen-er trichomes with straight apices (Fig. 7A); face with pair ofnbranched medium-long slender trichomes with straight apicesFig. 7B), as long as or slightly longer than frontal trichomes.horax. Integument light ochreous, densely covered with roundubercles except dorsal surface of posterior area of thorax mod-rately covered with cone-like tubercles; thorax on each sideith three long stout trichomes (combination of unbranched,
ifid and trifid trichomes) (Fig. 7C) anterodorsally, two long tri-homes (anterior one unbranched and slender, posterior one stoutnd unbranched, or bifid or trifid) (Fig. 7D) anterolaterally, oneifid medium-long trichome (Fig. 7E) mediolaterally, and threenbranched trichomes (two long and stout, one short and slender)Fig. 7F) ventrolaterally. Gill (Fig. 7G) with 10 thread-like fila-
ents arranged as 2 + (2 + 2) + 2 + 2 filaments from dorsal to ventral;ll short-stalked; all filaments subequal in length (0.7–1.0 mm),hough ventral filament of fourth pair from dorsal longest; rela-ive thickness of filaments from dorsal to ventral when comparedasally 1.00–1.15:1.15–1.25:1.00–1.25:1.00–1.25:1.25–1.37:1.25–.37:1.00–1.25:1.50–1.55:1.00:1.00; all filaments light brown, cov-red with sharply-defined annular ridges and furrows and denselyovered with minute tubercles. Abdomen. Dorsally, all segmentsearly transparent except segment 1 light yellow; segment 1 with-ut tubercles, with one unbranched short seta (Fig. 7H) on eachide; segment 2 with one unbranched short seta and five minuteetae, of which three or four are stout and one or two are slen-er (Fig. 7I) on each side; segments 3 and 4 each with four distinctooks and one short spinous seta on each side; all setae and hooksnbranched; segments 5, 6, 7 and 9 lacking spine-combs; seg-ent 8 with distinct spine-combs in transverse row; segments 7–9
ach with comb-like groups of minute spines on each side; seg-ent 9 with pair of cone-like terminal hooks (Fig. 7J). Ventrally,
ll segment unpigmented except segment 9 yellowish; segments–8 each with comb-like groups of minute spines; segment 5ith pair of bifid stout hooklets submedially and few unbranchedinute setae on each side; segments 6 and 7 each with pair of
ifid inner and unbranched outer stout hooklets somewhat sepa-ated from each other and few unbranched minute setae on eachide. Grapnel-shaped hooklets absent on each side of segment 9.ocoon (Fig. 7K). Light ochreous, shoe-shaped, with several small to
arge open spaces anterolaterally on each side; posterior half withoor; posterior half thickly woven and individual threads almost
nvisible; 3.0–3.5 mm long by 1.1–1.2 mm wide; height 0.5–0.7 mm.Mature larva. Body length 4.5–5.5 mm. Body dark gray except
horacic segment 3 creamy with ventral surface and parts of lateralurface ochreous. Abdomen in lateral view gradually widened from
egment 1 to segment 7, then narrowed to segment 9. Head capsuleparsely covered with colorless setae. Cephalic apotome variablen color pattern: in seven of 11 larvae examined, cephalic apotomehitish yellow with darkened areas along lateral margins and pos-
a 169 (2017) 170–186
terior margin and faintly to moderately positive head spots exceptposterolateral spots always indistinct (though all spots indistinct inone larva); in four other larvae, cephalic apotome entirely darkenedwith or without whitish-yellow portion medially on anterior three-fifths, and head spots obscured except mediolateral spots faintlynegative in two larvae. Lateral surface of head capsule whitish yel-low except little less than dorsal half darkened, with or withoutpositive spots below eye-spot region and near posterior marginin four larvae, or almost entirely darkened, with faintly or clearlynegative spots in seven larvae; eyebrow always distinct. Ventralsurface of head capsule yellow to dark yellow with faintly positiveor indistinct elongate spot on each side of postgenal cleft in threelarvae, or almost entirely darkened, with elongate spot on each sideof postgenal cleft faintly or clearly negative or obscured. Antennacomposed of three articles and apical sensillum, slightly longerthan stem of labral fan; length ratio of three articles (from baseto tip) 1.00:1.28–1.31:0.64–0.67. Labral fan with 42–46 primaryrays. Mandible (Fig. 8A) with mandibular serrations composed oftwo teeth (one medium-sized, one small); main tooth at right angleagainst mandible on apical side; comb-teeth gradually decreased inlength from first to third; supernumerary serrations absent. Hypos-toma (Fig. 8B) with nine anterior teeth, of which median and cornerteeth subequal in length to each other, followed by three interme-diate teeth on each side; lateral margins serrate apically; five or sixhypostomal bristles divergent posteriorly from lateral border oneach side. Postgenal cleft (Fig. 8C) large, rounded, 3.2 times lengthof postgenal bridge; sheath of subesophageal ganglion weakly ormoderately pigmented. Cervical sclerites on each side composedof light-brown elliptical piece, not fused to occiput. Histoblast ofpharate pupal gill with 10 thread-like slender filaments. Thoraxand abdomen without pair of dorsolateral protuberances. Thoraciccuticle almost bare. Abdominal cuticle almost bare except last seg-ment moderately covered with short spinous colorless setae oneach side of anal sclerite. Rectal scales not discernible. Rectal organof three lobes, each with six or seven finger-like secondary lobules.Anal sclerite X-shaped, with short broad anterior arms 0.55 timeslength of posterior ones; base of anal sclerite with deep unscle-rotized cleft posteromedially. Last abdominal segment not bulgedlaterally and lacking ventral papillae. Posterior circlet with 102–104rows of hooklets with up to 18–22 hooklets per row.
Type material. HOLOTYPE: Female (with associated pupalexuviae and cocoon, in 80% ethanol), collected from a river(width 8–15 m, depth 20–60 cm, water temperature 20 ◦C, par-tially shaded, elevation 727 m, 08◦19′42.039′′S/115◦13′44.116′′E),moderately running in a natural forest, Nung Nung waterfall,Bali, Indonesia, 27-IX-2014, by H. Takaoka, M. Sofian-Azirun,Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana. PARATYPES: Twofemales, one pharate male, eight mature larvae and 51 imma-ture larvae, same data as those of the holotype; two femalesand one pupal exuviae and four immature larvae, collectedfrom a small stream (width 0.2 m, depth 8–10 cm, bottomrocky, water temperature 23 ◦C, exposed to the sun, eleva-tion 599 m, 08◦23′56.429′′S/115◦17′50.434′′E), fast running neara forest, Sebatu, Bali, Indonesia, 27-IX-2014, by H. Takaoka, M.Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W. Suana; fiveimmature larvae, collected from a stream (width 0.5–3.0 m, depth0.5–1.0 m, water temperature 23 ◦C, partially shaded, elevation630 m, 08◦32′01.474′′S/116◦20′29.394′′E), moderately running in aforest, Benang Stokel waterfall, Lombok, Indonesia, 25-IX-2014, byH. Takaoka, M. Sofian-Azirun, Z. Ya’cob, C.D. Chen, K.W. Lau & I.W.Suana.
Biological notes. The pupae and larvae of this new species
were collected from grass leaves trailing in the current. Associatedspecies were S. (G.) atratum, S. (S.) eximium, S. (S.) iridescens, S. (S.)javaense and S. (S.) nobile.Distribution. Bali and Lombok.
H. Takaoka et al. / Acta Tropica 169 (2017) 170–186 181
Fig. 7. Pupa of S. (S.) baliense sp. nov. A, Frontal trichomes; B, Facial trichome; C–F, Thoracic trichomes (C, mediodorsal; D, anterolateral; E, mediolateral; F, ventrolateral); G,Gill filaments (right side; outer view); H, Short seta on dorsal surface of abdominal segment 1; I, Short seta, minute slender seta and minute spinous seta; J, Terminal hooks( mm f
i
tutbwafd
fgft
caudal view); K, Cocoon (lateral view). Scale bars. 1.0 mm for K; 0.1 mm for G; 0.02
Etymology. The species name baliense refers to the name of thesland, Bali, where this new species was collected.
Remarks. Simulium (S.) baliense sp. nov. is assigned to the S. stria-um species-group, defined by Takaoka and Davies (1996), by thenique shape of the female terminalia (Fig. 5D–H) and male geni-alia (Fig. 6C). This new species is characterized by having a hairedasal portion of the radius of the female, male scutum coveredith yellow short hairs, and pupal gill with 10 slender filaments of
lmost the same thickness, arranged as 2 + (2 + 2) + 2 + 2 filamentsrom dorsal to ventral (Fig. 7G), and larval body without pairedorsal protuberances.
This new species is similar to S. (S.) argyrocinctum De Meijere,
rom Java and Sumatra in having a similar arrangement of the pupalill filaments, but differs from the latter species in the female by therontal ratio 1.31:1.00:1.43 (1.4–1.5:1.0:1.6–1.7 in S. (S.) argyrocinc-um) and fore basitarsus 6.5 times as long as its greatest width (4.8or A–F and J; 0.01 mm for H and I.
times in S. (S.) argyrocinctum), in the male by the enlarged upper-eye facets in 14 or 15 vertical columns and 15 horizontal rows (in17 or 18 vertical columns and 17 or 18 horizontal rows in S. (S.)argyrocinctum) and fore basitarsus 5.64 times as long as its greatestwidth (6.5 times in S. (S.) argyrocinctum), in the pupa by the facialtrichomes as long as or slightly longer than the frontal trichomes(twice as long as the frontal trichomes in S. (S.) argyrocinctum)and in the larva by the body without paired dorsal protuberances(with paired dorsal protuberances on thoracic segments 1–3 andabdominal segments 1–8 in S. (S.) argyrocinctum). Simulium (S.)thailandicum Takaoka & Suzuki from Thailand also has a similararrangement of the pupal gill, but the two ventral pairs of filaments
are distinctively thinner than the other dorsal pairs of filaments,and the cephalic and thoracic integument have variously shapedtubercles (Takaoka and Suzuki, 1984).
182 H. Takaoka et al. / Acta Tropica 169 (2017) 170–186
F showing postgenal bridge (ventral view). Scale bars. 0.1 mm for C; 0.02 mm for A and B.
olApsd
4
itwTcaso41i1BotSsw(
aatNtot
Table 1The geographical distributions of 19 species of black flies in the Lessar SundaArchipelago.
Species Islands*
Java Bali Lombok Sumbawa Sumba Flores Timor
S. gyorkosae + +** +**
S. sunapii +S. lemborense +S. brevilabrum +S. rutengense +S. rangatense +S. atratum + +** +** +** + +S. floresense +S. merapiense + +**
S. feuerborni + +S. aureohirtum + +** + +S. nebulicola + +S. iridescens + +** +**
S. javaense + +**
S. eximium + +** +** +**
S. baliense sp. nov +** +**
S. nobile + +**
S. timorense +** +** + +S. sp. + +
the geographical distributions of the two species of the S. striatum
ig. 8. Larva of S. (S.) baliense sp. nov. A, Mandible; B, Hypostoma; C, Head capsule
Subgenus Wallacellum TakaokaSimulium (Wallacellum) sp.Simulium (Wallacellum) sp. Takaoka et al., 2006: 25 (Larva).Distribution. Flores and Timor.Remarks. This unnamed species was recorded from Flores based
n a single mature larva, and was assigned to the subgenus Wal-acellum by Takaoka et al. (2006) and also recorded from Timor bylmet et al. (2016). The larva is characterized by the four pharateupal gill filaments, postgenal cleft with a narrow anterior exten-ion, and simple rectal organ. Adult specimens are needed forefinitive identification.
. Discussion
The Lesser Sunda Archipelago except Bali biogeographically liesn Wallacea, the intermediate area between the Oriental and Aus-ralasian Regions, delimited in the east by Weber’s Line and in theest by Wallace’s Line (Audley-Charles, 1981; Dickerson, 1928).
he fauna of black flies in this archipelago is essentially Oriental,ontaining two Oriental subgenera, Gomphostilbia and Wallacellum,nd lacking the Australasian subgenus Morops Enderlein. It is noto rich in the number of species (19 species), compared with thosef neighboring areas such as Java, Borneo and Sulawesi, where 23,0 and 46 species are recorded, respectively (Takaoka and Davies,996; Takaoka, 2003; Takaoka et al., 2016a). At the species level,
t has a high affinity with the fauna of Java, sharing 10 (52.6%) of9 species (Table 1), reflecting the close geographical distance ofali and Lombok to Java, whereas it has a low affinity with thosef Borneo and Sulawesi, sharing only one (S. (N.) aureohirtum) andwo (S. (N.) aureohirtum and S. (G.) gyorkosae) species, respectively.imulium aureohirtum is known as the most widely distributedpecies in the Oriental Region, extending its distribution north-ard to the Palaearctic Region and eastward to Australasian Region
Adler and Crosskey, 2016).The fauna of black flies in the Lesser Sunda Archipelago is char-
cterized by a moderate rate of endemism, 47.4% or 9/19 species,nd the diversity of lineages consisting of four species-groups ofhe subgenus Gomphostilbia, two species-groups of the subgenus
evermannia, five species-groups of the subgenus Simulium, andhe subgenus Wallacellum. Flores is a hotspot for insular speciationf the subgenus Gomphostilbia in this archipelago, and also markshe eastern or southern boundary of geographical distributions of
* For comparison, Java is included, though not belonging to Lesser SundaArchipelago; Sumba Island remains unsurveyed.
** New records.
certain lineages: the eastern boundary for the S. epistum, S. christo-phersi and S. eximium species-groups, and the southern boundaryfor the subgenus Wallacellum.
In this archipelago, the fauna of black flies is likely to be dividedbetween Lombok and Flores, although four species (S. (G.) atratum,S. (N.) aureohirtum, S. (S.) eximiun and S. (S.) timorense) are widelydistributed across the archipelago from west to east (Table 1).Wallace’s line running between Bali and Lombok is unlikely tobe associated with the separation of the geographical distribu-tions of two species of any simuliid species-groups except the S.nobile species-group, of which S. (S.) nobile is distributed from Baliwestward and S. (S.) timorense from Lombok eastward. Similarly,
species-group are separated from each other between Java and Bali:Sumatra and Java for S. (S.) argyrocinctum and Bali and Lombok forS. (S.) baliense sp. nov.
Tropica 169 (2017) 170–186 183
…
S. lembo rense
… ...3
..……………………… 4
………………………… 5
…… S. floresense
………………
……
S. atratum
………… S. brevilab rum
… ……………………. 6
al segment…S. merap iense
ill ary palpal segment.......7
S. sunap ii
S. gyorkosae
……………………… 9
……. 10
S. au reoh irtum
mere yellow.. S. fe uerbo rni
…….. 11
………. 13
..... S. nobil e& S. ti morense
mere yellow...................12
. iridescens & S. javaense
S. nebuli cola
ther, nea rly
S. bali ense s p. nov.
rated from
S . ex imium
……….. 2
………. 9
……….. 3
……….. 6
S. lembo rense
………... 4
……..… 5
K
…………………………2.
…………………………8
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...
…
.
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.
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each other
H. Takaoka et al. / Acta
Females*
1. Katepisternum haired………………………………………………
Katepisternum bare…………………………………………………
2. Hind tibia darkened exce pt base yellow………………….. Hind tibia yellowish on basal two-fifths or more……………….…
3. Hind tibia with subbasal dark spot……………………….…………
Hind tibia without sub basal dark spot…… ……………………… …
4. Frons: head ratio 1:6.7………………………….……………………
Frons: head ratio 1:4.4…………………………………..…… …… .
5. Hind tibia yellowish on basal half…………………..…… ………… .
Hind tibia yellowish on basal three-fifths ……………………………
6. Sensory vesicle enlarged, 0.65 tim es length of third maxil lary palp
Sensory vesicle medium-size d, 0.25 –0.34 tim es length of third max
7. Frons: head ratio 1:5.8…………………..……………..……… Frons: head ratio 1:4.9 ………………………………….…..
8. Claw with large basal too th……………………………..……………
Claw without too th or with s mall subbasal too th……………………
9. Antenn a yellowish except first flagellomere darkened………
Antenna dark brown exce pt sca pe, pedicel and base of first flagello
10. Claw with s mall basal too th…………………………………………
Claw without too th…………………………………………………
11. Antenna yellow with apica l two flagellomeres darkened.................
Antenna dark brown exce pt sca pe, pedicel and base of first flagello
12. Tergites 5–8 shiny………………………………… S
Tergites 6–8 shiny……………………………………………….. 13. Inner margins of ovipositor valves narr owly separated from ea ch o
parall el-sided…………………………………… ..……… Inner margins of ovipositor valves dee ply conca ve and widely sepa
medially................. …………………………………………...…..
Males**
1. Katepisternum haired………………………………………………
Katepisternum bare…………………………………………………
2. Hind basitarsus slender, not enlarged…… …………………………
Hind basitarsus enlarged……………………………………………
3. Hind tibia darkened exce pt base yellow………………………... Hind tibia yellowish on basal two-fifths or more……………….…
4. Hind tibia with subbasal dark spot……………………….…………
eys to 19 spec ies of black flies in the Less er Sund a Archipelago
…………
……
………
………
………………
…
…………
…
Hind tibia without sub basal dark spot…… ……………………... 5. Fore basitarsus 8.9 times as long as its greatest width.............. …..
Fore basitarsus 9.1 times as long as its greatest width................ …….. …
………
……
S. merap iense
S. floresense....
S. atratum…………
………
…… .
1 Tropica 169 (2017) 170–186
…….. 7
…….. 8
ws......................................... S. sunap ii
rows….............................. S. gyorkosae
S. brevilab rum & S. ruteng ense
S. ranga tense
…… 10
……... 11
S. au reoh irtum
re yellow.......................... S. feuerborni
S. nobil e & S. ti morense
………. 12
……… 13
……… 14
S. bali ense s p. nov.
.. S . ex imium
th, and
S. nebuli cola
all y and
................... S. iridescens & S . javaense
P
S. brevilab rum
………. 2
………. 3
…………. 9
S. nobil e & S . ti morense
………….. 4
…………. 5
…………. 6
S. au reoh irtum
S. feuerbo rni
S. ranga tense
................ 7
S. javaense
……….... 8
fil aments........................... S. nebuli cola
ger than other filaments... S. iridescens………… 10
………. 16
S. gyorkosae
………………………………
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.
.
. .
. .
. .
. .
.
.
.
.
.
. .
.
.
.
.
.
brownish blac k on apical
brownish blac k on apical
84 H. Takaoka et al. / Acta
6. Hair tuft on base of radial vein yellow…… …………………………… Hair tuft on base of radial vein darkened………………………………
7. Upp er-eye facets in eight or nine vertical columns and 12 horizontal ro
Upp er-eye facets in 10 or 11 vertical columns and 12 or 13 horizontal
8. Subcosta haired……………………………..….
Subcosta bare…………………………………………………… ..
9. Coxite longer than style…………………………………………………
Coxite shorter than style………………………………………… ..…… 10. Antenn a yellowish except first flagellomere darkened……… .……
Antenna dark brown except scape, pedicel and base of first flagellome
11. Hind basitarsus s lender, much narr ower than tibia………
Hind basitarsus enlarge, nea rly as wide as tibia…………………………
12. Scutum covered with yellow short hairs…………………………………
Scutum covered with dark short hairs…… ……………………………… 13. Ventral plate saddle-shaped………………………… ..…………
Ventral plate pea r-shaped……………………………………………… 14. Fore femur yellow on basal half, gradu all y darkened to apical one-four
one-fourth...........................................……… ..…………………… .… Fore femur dark yellow to li ght brown at base, gradu ally darkened apic
cap...........................................................................................................
upae***
1. Gill with four filaments……………………… ..…………………… Gill with six, eight, nine or 10 filaments…………………………………
2. Gill with s ix filaments……………………………………………………
Gill with eight, nine or 10 fil aments……… ……………………………
3. Cocoon shoe-shaped……………….…… ………………..
Cocoon wall-pocket-shaped……………………………………………
4. Cocoon with short or medium-long anterodorsal projec tion……………
Cocoon without anterodorsal projec tion…………………..……………
5. All gill filaments widely divergent…………………… .………..… All gil l filaments arising close together and direc ted forward……….
6. Stalk of mi ddle pair of gill filaments elongate…………………….… Stalk of mi ddle pair of gill filaments short.........................................
7. Gill with s ix inflated filaments…………………………………….…… Gill with six slender threa d-li ke filaments………………………………
8. Dorsal filament of dorsal pair of gill filaments much longer than otherDorsal filament of dorsal pair of gill filaments as long as or sli ghtly lon
9. Gill with eight filaments……………………….………………………
Gill with nine or 10 fil aments……………………………………………
10. Cocoon with s hort anterodorsal projec tion…………………………..
………………………
……
……
………………
………
…
……
……………
……
…
…
…
Cocoon without anterodorsal projec tion………………………………………
11. All gill filaments s hort and of same thickness……………....………
All gil l filaments otherwise……………………………………………………
…………
…. 11
S. mer ap iense
… 12
……………………………
…………………
……………………………
.
Tropica 169 (2017) 170–186 185
S . sunap ii
S. lembo rense
. S. atratum
S. bali ense s p. nov. S. ex imium
L……. . 2
…….. . 11
…….. 3……….. ... 5
err ations
S. aureoh irtum
err ations consisting of one large……….. 4
S. feuerbo rni
S. merap iense
…… .. S . sp.
………. 6
………… 7…………. 8 S. ruteng ense
S. gyorkosae
S. fl oresense
………. 9
S. lembo rense
……. 10
S. atratum
. S . sunap ii
……… 12
……….. 13
… S. nobil e
S. ti morense
………. 14
……… 15
nt...... .. S. floresense………
……… .. 14
r filaments.................................15 S. ruteng ense
S. bali ense s p. nov.
S. ex imium
……........................ S. nebuli cola……. 16
S. iridescens
S. j avaense
*
……………..…………....................13
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…
consisting of two large
H. Takaoka et al. / Acta
Frons moderately covered with tubercles……………………………… .
15. Termi nal hook cone-like ………………………………………… ..… Termi nal hook widened with s err ate ourter margin………………… .….
16. Abdomi nal segment 8 with spine -combs………………………
Abdomi nal segment 8 without spine-combs…………………… ..…… .
arvae ****1. Ventral papil la present……………………………………………………
Ventral papil la absent…………………………………………………… 2. Postgenal cleft short, shorter than postgenal bridge………………………
Postgenal cleft medium-long to long, longer than postgenal bridg e…… 3. Postgenal cleft about 0.8 tim es length of postgenal bridg e; mandibular s
tee th.......................................………………………………………
Postgenal cleft about 0.4 tim es length of postgenal bridg e; mandibular stoo th and one small too th.......................................………………………
4. Body creamy, with reddish-brown markings…………………………
Body blac kish, without such reddish-brown markings………………
5. Rec tal organ sim ple, without sec ondary lobules……………………… ..
Rec tal organ compound, with sec ondary lobules………………………… 6. Postgenal cleft medium-long, about twice as long as postgenal bridge…
Postgenal cleft long, more than 2.5 tim es as long as postgenal bridge… 7. Rec tal organ with one to three short, nipple-li ke sec ondary lobules…
Rec tal organ with four or five fing er-li ke sec ondary lobules…………
8. First and sec ond antenn al articles equ al in leng th to eac h other…..… First antennal article longer than sec ond article...………………………
9. Postgenal cleft 2.6–3.0 tim es length of postgenal bridge…………… ..
Postgenal cleft 4.7–5.0 tim es length of postgenal bridge…………………
10. Posterior circlet with about 76 rows of hoo klets……………………… . Posterior circlet with about 94 rows of hoo klets………… .…………… .
11. Postgenal cleft rea ching posterior margin of hypostoma………………… Postgenal cleft not reac hing posterior margin of hypostoma……………
12. Abdomi nal segments 1–5 eac h with pair of do rsal protuberances……… Abdomi nal segments 1–5 without dorsal protuberances………… ..…
13. Postgenal cleft rounded apicall y…………………………………………
Postgenal cleft pointed apicall y…………………………… ..……………
12. Outer filament of ventral pair of gill filaments thicker than inn er filameTwo fil aments of ventral pair of gil l equ al in thickness to eac h other……
13. Two fil aments of ventral pair of gill sli ghtly longer than other filaments
Two fil aments of ventral pair of gil l more than 1.5 tim es as long as othe14. Frons densely covered with tubercles…… ………………………… ..
14. Posterior circlet with 102–104 rows of hooklets……… ..…..….. Posterior circlet with about 210 rows of hooklets……………………… ..
15. Rec tal organ with seven to nine fing er-li ke secondary lobules per lobe…… Rec tal organ with one to three fing er-li ke s econdary lobules per lobe……
16. Pharate pupal gil l with six slender threa d-li ke fil aments………… ..….. Pharate pupal gil l with s ix inflated filaments…………………… ..…… ..
…
……
……………
……
…
……
…
……
…
…
…
…………
…
…
…
*Females of S. ruteng ense, S. rangatense and S. sp. are unknown.**The male of S. sp. is unknown.***The pupa of S. sp. is unknown.***The larvae of S. brevilab rum , S. rangatense and S. ruteng ense are unknown.
1 Tropic
A
Cig
R
A
A
A
A
D
D
D
E
F
L
T
Takaoka, H., 2003. The Black Flies (Diptera: Simuliidae) of Sulawesi, Maluku andIrian Jaya. Kyushu University Press, Fukuoka, Japan, xxii+581 pp.
86 H. Takaoka et al. / Acta
cknowledgements
We are grateful to Prof. Peter H. Adler, Clemson University,lemson, SC, USA, for reading the current manuscript and provid-
ng valuable comments. This work was supported by the researchrant from University of Malaya (RP021A/16SUS).
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udley-Charles, M.G., 1981. Geological history of the region of Wallace’s line. In:Whitmore, T.C. (Ed.), Wallace’s Line and Plate Tectonics. Clarendon Press,Oxford, UK, pp. 24–35 (91 pp.).
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ickerson, R.E., 1928. Distribution of Life in the Philippines. Philippine Bureau ofSciences Monograph 21, Manila, Philippines, 322 pp.
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ow, V.L., Takaoka, H., Pramual, P., Adler, P.H., Ya’cob, Z., Chen, C.D., Yotopranoto,S., Zaid, A., Hadi, U.K., Sofian-Azirun, M., 2016. Three taxa in one: cryptic
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