Research Article |
Corresponding author: Atiporn Saeung ( atisaeung.noi@gmail.com ) Academic editor: Art Borkent
© 2020 Hiroyuki Takaoka, Wichai Srisuka, Masako Fukuda, Atiporn Saeung.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Takaoka H, Srisuka W, Fukuda M, Saeung A (2020) Twenty-one new species of the Simulium (Gomphostilbia) asakoae species group (Diptera, Simuliidae) in Thailand, with their genetic relationships. ZooKeys 950: 51-152. https://doi.org/10.3897/zookeys.950.51298
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Females and males reared from pupae, their pupal exuviae and cocoons, and mature larvae of the Simulium (Gomphostilbia) asakoae species group from various localities in Thailand were morphologically examined. A total of 25 species was identified, including two of four known species (Simulium asakoae Takaoka & Davies and S. chiangdaoense Takaoka & Srisuka), one newly confirmed species (S. myanmarense Takaoka, Srisuka & Saeung, originally described from Myanmar), one newly transferred species (S. inthanonense Takaoka & Suzuki formerly of the S. ceylonicum species group), and 21 new species. Descriptions of all 21 new species are given, and the first full description of the male of S. inthanonense, together with the revised descriptions of its female, pupa, and larva, is also provided. Keys to identify all 27 members of this species group from Thailand are given for females, males, pupae, and larvae. The genetic relationships of all but one species were resolved using COI gene sequence-based analysis. All 26 species were divided into nine subgroups, I–IX, each consisting of two, one, four, nine, one, three, two, one and three species, respectively.
Aquatic insects, biodiversity, blackflies, Oriental Region, taxonomy
The Simulium asakoae species group, as defined by
The number of species in this species group in Thailand is expected to be much greater because a high rate of radiation in this species group was suggested by
The S. asakoae species group is separated from the other species groups in the subgenus Gomphostilbia by a combination of the yellowish hair tuft on the base of the radial vein and yellowish forecoxae in the female and male, hind basitarsus enlarged, and ventral plate emarginated on both sides when viewed ventrally in the male (
In general, females of most members of the S. asakoae species group are morphologically similar to one another and are often difficult to identify to species, although certain morphological features, such as the width ratio of the frons against the head, relative length of the sensory vesicle against the length of the third palpal segment, presence or absence of the outer mandibular teeth, relative length of the fore- and hind basitarsi against their greatest width, and relative length of the claw tooth against the claw, are used for species identification (
The possible involvement of undescribed cryptic species, as well as the difficulty of morphological identification of the females, is a serious problem for studies of the vectorial roles in the transmission of parasites and pathogens among species of the S. asakoae species group.
In this study, we first aimed to explore the fauna of the S. asakoae species group in Thailand by morphologically examining numerous females and males reared from pupae, their associated pupal exuviae and cocoons, and mature larvae collected in various provinces in this country, and secondly to molecularly resolve the genetic relationships of Thai members of the S. asakoae species group by a COI gene sequence-based analysis.
The material examined in this study consisted of females and males, their associated pupal exuviae and cocoons, and mature larvae of the S. asakoae species group collected in various localities in Thailand. All specimens were fixed in 80% ethanol.
Methods of morphological observation, terms of features, descriptions, and illustrations followed
Due to close morphological similarities of all new species, the description of the first new species was fully made based on as many morphological characters as possible, whereas those of the other new species were made only for morphological characters differing from those of the first new species.
The holotypes and paratypes of the new species are deposited in the Entomology Section of the Queen Sirikit Botanic Garden, Chiang Mai province, Thailand.
Thoraces of females and males reared from pupae, and whole bodies of mature larvae were used for genetic analysis. The procedures for DNA extraction, COI gene amplification, sequencing, and data analysis followed those of
This paper and the nomenclatural acts have been registered in ZooBank (www.zoobank.org), the official register of the International Commission on Zoological Nomenclature. The Life Science Identifier (LSID) numbers are noted under each of the 21 new species of black flies.
A total of 25 species was identified morphologically, comprising two known species (S. asakoae and S. chiangdaoense Takaoka & Srisuka), one newly confirmed species (S. myanmarense originally described from Myanmar), one newly transferred species (S. inthanonense Takaoka & Suzuki), and 21 new species. Simulium inthanonense, one of three Thai members of the S. ceylonicum species group, is here transferred to the S. asakoae species group based on the male ventral plate, which is emarginated on both sides (Fig.
Simulium asakoae, originally described from Peninsular Malaysia (
Five taxa (S. doisaketense, S. sp. nr. asakoae-2, S. sp. nr. asakoae-3, S. sp. nr. asakoae-4, and S. sp. nr. sheilae-3) reported based on larval salivary gland chromosome studies by
The possibility of the distribution in Thailand of two Myanmar species (S. myanmarense and S. monglaense) was noted by
All 21 new species are here described, and the male of S. inthanonense is fully described for the first time. Keys to identify all 27 members of the S. asakoae species group from Thailand are given for females, males, pupae, and larvae.
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium thungchangense male, QSBG col. no. 8, Thailand, 24-I-2019, by W. Srisuka”, collected from a stream (width 60 cm, depth 10 cm, bed sandy, moderate flow, pH 6.9, 17.9 °C, exposed to the sun, elevation 1,376 m, 18°32'44.4"N, 98°30'53.0"E), at Siribhum waterfall, Chom thong District, Chiang Mai Province, Thailand, 24-I-2019, by W. Srisuka (Coll. No. 8).
Paratypes : 10 females, 10 males (one male for DNA analysis) (with their associated pupal exuviae and cocoons), and 20 mature larvae (two mature larvae for DNA analysis) (in 80% ethanol), same data as for holotype; one male (with its associated pupal exuviae and cocoon) and one pharate male (with its associated pupal exuviae and cocoon) (in 80% ethanol), collected from a stream (18 °C, elevation 1,470 m) from Phu Kha, Samorrophum, Thung Chang District, Nan Province, 2-XII-2004, by W. Choochote.
Female: mandible with several teeth on the outer margin and hind tibia yellowish on little more than the basal half (Fig.
Female of S. thungchangense sp. nov. A third palpal segment with sensory vesicle (right side; anterior view) B cibarium (anterior view) C hind tibia (left side; lateral view) D hind basitarsus and second tarsomere (left side; lateral view) E claw F sternite 8 and ovipositor valves (ventral view) G genital fork (ventral view) H, I paraprocts and cerci (right side H ventral view I lateral view) J spermatheca. Scale bars: 0.1 mm (C, D); 0.02 mm (A, B, F–J); 0.01 mm (E).
Female (N = 10). Body length 2.5–2.6 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 1.8–1.9:1.0:2.5–2.6; frons:head ratio 1.0:4.2–4.7. Fronto-ocular area well developed, narrow, directed dorsolaterally. Clypeus dark brown, densely covered with yellowish white scale-like hairs interspersed with several dark longer hairs on each side. Labrum 0.62–0.68 times length of clypeus. Antenna composed of scape, pedicel and nine flagellomeres, dark brown except scape, pedicel and base of first flagellomere yellow. Maxillary palpus composed of five palpomeres, light to medium brown, proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.6; sensory vesicle (Fig.
Thorax. Scutum dark brown except anterolateral calli ochreous, and three blackish longitudinal vittae (one median, two submedian) faintly visible, thinly pruinose and shiny when illuminated at certain angles, densely covered with 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 when illuminated at certain angles, and bare. Pleural membrane ochreous and bare. Katepisternum longer than deep, medium to dark brown, shiny when illuminated at certain angles, moderately covered with fine yellow and brown short hairs.
Legs.
Foreleg: coxa and trochanter whitish yellow; femur dark yellow to light brown with apical cap medium brown (though extreme tip yellowish); tibia yellowish white except apical one-fourth brownish black and covered with white fine hairs on basal four-fifths; tarsus brownish black, with moderate dorsal hair crest; basitarsus moderately dilated, 6.6–6.9 times as long as its greatest width. Midleg: coxa medium brown except posterolateral surface dark brown; trochanter whitish yellow; femur light to medium brown with basal one-fourth whitish yellow and apical cap medium brown (though extreme tip yellowish); tibia whitish yellow on basal two-fifths and light to dark brown on rest (though whitish yellow on basal half or little more on posterior surface in some females), and covered with yellowish fine hairs on posterior and inner surfaces of basal two-thirds; tarsus dark brown to brownish black though basal one-third of basitarsus dark yellow (its border not well defined). Hind leg: coxa medium brown with apical one-third yellow; trochanter whitish yellow; femur medium brown with base whitish yellow and apical cap dark brown (though extreme tip yellowish white); tibia (Fig.
Wing. Length 2.2–2.4 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 of segment 2 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 segments 2–4 yellow and those of other segments medium to dark brown; sternal plate on segment 7 undeveloped.
Terminalia.
Sternite 8 (Fig.
Male (N = 14). Body length 2.6–3.1 mm.
Head.
Nearly as wide as thorax. Upper eye vermilion, consisting of large facets in 9 (rarely 10) vertical columns and 12 horizontal rows on each side. 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 near lower margin. Antenna composed of scape, pedicel and nine flagellomeres, medium to dark brown except scape, pedicel, and base of first flagellomere yellow; first flagellomere elongate, 1.8 times length of second. Maxillary palpus light brown, with five palpal segments, proportional lengths of third, fourth, and fifth palpomeres 1.0:1.1–1.2:2.7–3.0; third palpomere (Fig.
Male of S. thungchangense sp. nov. A sensory vesicle (right side; anterior view) B hind basitarsus and second tarsomere (left side; lateral 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 half; caudal view) I, J tenth abdominal segments and cerci (right side I lateral view J caudal view). Scale bars: 0.1 mm (B); 0.02 mm (A, C–J).
Thorax. Scutum dark brown to brownish black except anterolateral calli ochreous, shiny and white pruinose when illuminated at certain angles, and densely covered with whitish yellow scale-like recumbent short hairs. Scutellum dark brown, covered with yellow short hairs and dark-brown long upright hairs along posterior margin. Postnotum brownish black, slightly shiny and white pruinose when illuminated at certain angles, and bare. Pleural membrane ochreous and bare. Katepisternum dark brown, longer than deep, shiny and white pruinose when illuminated at certain angles, moderately covered with yellow and brown fine short hairs.
Legs.
Foreleg: coxa whitish yellow; trochanter light brown; femur light brown except apical tip yellowish; tibia whitish yellow except little more than apical one-third dark brown, and covered with white hairs on whitish yellow portion; tarsus brownish black; basitarsus slightly dilated, 8.4–8.7 times as long as its greatest width. Midleg: coxa dark brown except posterolateral surface brownish black; trochanter dark yellow to light brown except base yellow; femur light to medium brown with base yellowish and apical cap dark brown (though apical tip yellow); tibia dark brown except basal one-third (or little more on posterior surface) whitish yellow; tarsus dark brown except basal one-fourth or less of basitarsus dark yellow to light brown (border not well defined). Hind leg: coxa dark brown; trochanter yellowish; femur medium to dark brown with base yellow and apical cap brownish black (though apical tip yellow); tibia dark brown to brownish black except little more than basal two-fifths whitish yellow; tarsus (Fig.
Wing. Length 2.2–2.4 mm. Other characters as in female except subcosta with 9–13 hairs. Halter. Dull white except basal stem darkened.
Abdomen. Basal scale dark brown, with fringe of light-brown hairs. Dorsal surface of abdomen medium brown to brownish black, 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 yellow, those of segments 3 and 4 yellow though sternal plates light brown, and those of other segments medium to dark brown.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 24). Body length 2.5–3.3 mm.
Head. Integument deep 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; face with pair of unbranched (rarely bifid) long trichomes with straight apices; three frontal trichomes on each side arising close together, subequal in length to one another and slightly longer than facial one.
Thorax.
Integument deep yellow, moderately covered with round tubercles, and with three long dorsomedial trichomes with coiled apices, two long anterolateral trichomes (anterior trichome more slender and shorter with straight or coiled apex, posterior one with coiled apex), one medium-long mediolateral trichome with straight apex, and three ventrolateral trichomes (one medium-long, two short) with straight apices, on each side; all trichomes unbranched. Gill (Fig.
Abdomen.
Dorsally, all segments unpigmented or light yellowish except segment 9 and bases of spine-combs of segments 6–8 yellow; segments 1 and 2 sparsely covered with minute tubercles; segment 1 with one unbranched slender short hair-like seta on each side; segment 2 with one unbranched slender short 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 micro-spines on each side; segment 5 with four minute setae on each side; segments 6–8 each with two minute setae on each side; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 20). Body length 5.5–6.5 mm. Body creamy white to light ochreous with following color markings: thoracic segment 1 encircled with distinct ochreous band (though disconnected ventromedially), thoracic segments 2 and 3 ochreous on ventral surface; abdominal segments 1–4 entirely light green or greenish grey, abdominal segment 4 with reddish brown transverse band (though often partially faded, leaving narrow band or small spot(s) dorsally), abdominal segment 5 with distinct reddish brown, W-shaped, transverse band dorsally, abdominal segment 6 often with three distinct, reddish brown spots (one round dorsomedial spot and two lateral spots of various size and shape), dorsal and dorsolateral surface of abdominal segments 5–8 faintly to distinctly covered with pinkish pigment (Fig.
Head.
Head capsule yellow to dark yellow except eye-spot region whitish, sparsely covered with minute setae (though moderately on dorsal surface); head spots faintly positive or indistinct. Antenna composed of three articles and apical sensillum, longer than stem of labral fan; proportional lengths of first, second, and third articles 1.0:0.8:0.7. Labral fan with 29–31 primary rays. Mandible (Fig.
Thorax and abdomen. Cuticle sparsely covered with unpigmented minute setae (though few posterior abdominal segments sparsely covered also with dark minute unbranched setae) dorsally; 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. Rectal scales minute, unpigmented. Rectal organ compound, each of three lobes with seven or eight finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms 1.2 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 80–86 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, thungchangense, refers to the district, Thung Chang, one of the two localities where this species was collected.
Thailand (Chiang Mai and Nan).
This new species is similar to S. chaudinhense Takaoka & Sofian-Azirun described from Vietnam (
Holotype : Male (together with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium puaense male, QSBG col. no. 60, Thailand, 25-VII-2017, by W. Srisuka” reared from a pupa collected from a small waterfall (width 80 cm, depth 3 cm, flast flow, pH 7.2, 20.1 °C, exposed to the sun, elevation 1,157 m, 19°11'10.3"N, 101°04'41.7"E), Nam Dan Village, Pua District, Nan Province, northern Thailand, 25-VII-2017, by W. Srisuka (Coll. No.60).
Paratypes : One female (thorax for DNA analysis) (together with its associated pupal exuviae and cocoon) (in 80% ethanol), reared from a pupa collected from a stream (width 30 cm, depth 5 cm, bed sandy, moderate running, pH 6.3, 22.6 °C, exposed to the sun, elevation 1,097 m, 18°50'03.7"N, 99°22'32.2"E), at Pa Meing Village, Muang Pan District, Lampang Province, northern Thailand, 9-VIII-2016, by W. Srisuka (Coll. No. 86); three males (thorax of one male for DNA analysis) (together with their associated pupal exuviae and cocoons) (in 80% ethanol), same data as for holotype.
Female: small sensory vesicle 0.24 times as long as the third palpal segment (Fig.
Female, male and pupa of S. puaense sp. nov. A, B female C–H male I–L pupa. A sensory vesicle (left side; anterior view) B mandible (left side) C hind basitarsus and second tarsomere (left side; lateral view) D coxites, styles and ventral plate (ventral view) E, F styles (right side; ventrolateral view E gradually tapered toward apex F nearly parallel-sided) G ventral plate and median sclerite (lateral view) H ventral plate (caudal view) I gill filaments (left side; lateral view) J, K terminal hooks (caudal view J normal K right hook abnormal) L cocoon (dorsal view). Scale bars: 1.0 mm (L); 0.1 mm (C, I); 0.02 mm (A, D–H); 0.01 mm (B, J, K).
Female (N = 1). Body length 2.1 mm.
Head.
Frontal ratio 1.9:1.0:2.4 frons:head ratio 1.0:4.4. Labrum 0.56 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.0; sensory vesicle (Fig.
Legs. Foreleg: coxa and trochanter whitish yellow; femur dark yellow to light brown with apical cap medium brown (though extreme tip yellowish); tibia yellowish white except little more than apical one-fourth brownish black; basitarsus moderately dilated, 5.8 times as long as its greatest width. Midleg: tarsus dark brown to brownish black though basal half of basitarsus dark yellow (its border not well defined). Hind leg: coxa light brown; tibia yellowish white on little more than basal half and light brown to brownish black on rest; basitarsus 5.9 times as long as wide, and 0.7 and 0.5 times as wide as greatest widths of tibia and femur, respectively; claw with large basal tooth 0.47 times length of claw.
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except anterior two-thirds of segment 2 ochreous.
Terminalia. Sternite 8 with 20 medium-long to long hairs together with two or three slender short hairs on each side. Paraproct in ventral view with four or five sensilla on anteromedial surface; paraproct in lateral view 0.6 times as long as wide, with 21–23 medium-long to long hairs on ventral and lateral surfaces.
Male (N = 4). Body length 2.1–2.4 mm.
Head. Slightly wider than thorax. Upper eye dark brown, consisting of large facets in nine or ten vertical columns and 12 horizontal rows on each side. Antenna: first flagellomere 1.5–1.6 times length of second. Maxillary palpus light brown, with five palpal segments, proportional lengths of third, fourth, and fifth palpal segments 1.0:1.0:2.0; sensory vesicle 0.13–0.18 times length of third palpal segment.
Legs.
Foreleg: basitarsus moderately dilated, 6.6–7.2 times as long as its greatest width. Hind leg: tibia dark brown to brownish black except basal half whitish yellow; tarsus (Fig.
Wing. Length 2.0–2.1 mm. Subcosta with two to five hairs (though subcosta bare in one male).
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 5). Body length 2.6–3.0 mm.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva. Unknown.
The species name, puaense, refers to the district, Pua, one of the two localities where this species was collected.
Thailand (Lampang and Nan)
This new species is similar to S. vinhphucense Takaoka & Low from Vietnam (
This new species is also similar to S. thungchangense sp. nov. and S. chaudinhense from Vietnam (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium sutheppuiense male, QSBG col. no. 92, Thailand, 7-IX-2017, by W. Srisuka”, collected from a small stream (width 60 cm, depth 15 cm, bed sandy, moderate flow, pH 6.9, 20 °C, partially shaded, elevation 1,395 m, 18°49'09.5"N, 98°53'14.3"E), at Doi Pui Temple, Doi Suthep Pui, Muang District, Chiang Mai Province, Thailand, 7-IX-2017, by W. Srisuka (Coll. No. 92).
Paratypes : One female (thorax for DNA analysis), three males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and five mature larvae (one mature larva for DNA analysis) (in 80% ethanol), same data as for holotype;
Female: mandible with three teeth on the outer margin (Fig.
Female, male, pupa and larva of S. sutheppuiense sp. nov. A female B–F male G–I pupa J larva. A mandible (right side) B hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (left side; lateral view) H terminal hooks (caudal view) I cocoon (dorsal view) J head capsule (ventral view). Scale bars: 1.0 mm (I); 0.1 mm (B, G, J); 0.02 mm (C–F); 0.01 mm (A, H).
Female (N = 1). Body length 2.0 mm.
Head.
Frontal ratio 2.0:1.0:2.5; frons:head ratio 1.0:4.6. Labrum 0.57 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.7; sensory vesicle 0.29–0.32 times length of third palpal segment. Lacinia with ten or eleven inner and 13 or 14 outer teeth. Mandible (Fig.
Legs. Foreleg: basitarsus moderately dilated, 6.3 times as long as its greatest width. Midleg: tarsus dark brown to brownish black though basal half of basitarsus yellow (its border not well defined). Hind leg: coxa light brown; tibia yellowish white on basal three-fifths and light brown to brownish black on rest; basitarsus 5.3 times as long as wide, and 0.8 and 0.6 times as wide as greatest widths of tibia and femur, respectively.
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 ochreous.
Terminalia. Sternite 8 with 25 or 26 medium-long to long hairs together with three or four slender short hairs on each side. Ovipositor valves each moderately covered with microsetae interspersed with two or three short hairs. Paraproct in ventral view with three or four sensilla on anteromedial surface; paraproct in lateral view 0.5 times as long as wide, and with 24 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.5 times as long as wide. Spermatheca 1.5 times as long as its greatest width.
Male (N = 4). Body length 2.0–2.2 mm.
Head. Slightly wider than thorax. Upper eye dark brown, consisting of large facets in eleven vertical columns and 13 horizontal rows. Antenna light to medium brown except scape, pedicel, and base of first flagellomere yellow; first flagellomere elongate, 1.8–1.9 times length of second one. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.6; sensory vesicle 0.15 times length of third palpal segment.
Legs.
Foreleg: tibia whitish yellow except apical three-tenths dark brown; basitarsus slightly dilated, 7.5–7.9 times as long as its greatest width. Hind leg: coxa medium brown; tibia dark brown to brownish black except little less than basal half whitish yellow; tarsus (Fig.
Wing. Length 1.9–2.0 mm. Subcosta with 1–11 hairs (though no hairs in one male).
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 5). Body length 2.5–2.7 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles except dorsolateral surface of posterior half sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, all segments unpigmented except segments 1, 2, and 9 light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 4). Body length 4.7–5.5 mm. Body light ochreous with following color markings: thoracic segment 1 encircled with light to dark brown band (though disconnected ventrally), thoracic segments 2 and 3 ochreous on ventral surface; abdominal segment 4 faintly with reddish brown transverse band (though often entirely faded), abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band dorsally along posterior margin, though often partially faded leaving one round dorsomedial spot and two lateral spots of various size and shape, dorsal and dorsolateral surface of abdominal segments 5–8 faintly to moderately covered with pinkish or reddish brown pigment (Fig.
Head.
Head capsule yellow except eye-spot region whitish; head spots moderately positive or indistinct. Antenna: proportional lengths of first, second, and third articles 1.0:0.7–0.8:0.8. Labral fan with 29 or 30 primary rays. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle sparsely covered with unpigmented minute setae, though few posterior abdominal segments sparsely covered with dark minute unbranched setae dorsally. Rectal organ compound, each of three lobes with nine or ten finger-like secondary lobules. Posterior circlet with 86–89 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, sutheppuiense, refers to the locality name, Doi Suthep Pui, where this species was collected.
Thailand (Chiang Mai).
This new species is similar to S. asakoae in many characters including the presence of teeth on the outer margin of the female mandible, and small number of male upper-eye facets. However, it is distinguished from the latter species in the female by the shorter sensory vesicle relative to the third palpal segment, in the male by the dark brown upper-eye (large) facets, in the pupa by the light yellow dorsum of abdominal segments 1–3, and in the larva by abdominal segments 1–4 light ochreous (Fig.
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium teerachanense male, QSBG col. no. 105, Thailand, 14-VII-2017, by W. Srisuka”, collected from a small stream (width 25 cm, depth 2.5 cm, bed sandy, moderate flow, pH 6.8, 21.9 °C, partially shaded, elevation 974 m, 18°20'35.8"N, 98°01'23.8"E), at Tee Ra Chan Waterfall, Mae La-noi, Mae Hong Son Province, Thailand, 14-VII-2017, by W. Srisuka (Coll. No. 105).
Paratypes : Eight males (thoraces of two males for DNA analysis) (with their associated pupal exuviae and cocoons) and four mature larvae (in 80% ethanol), same data as for holotype. One mature larva (for DNA analysis) collected from a stream (elevation 817m, 19°11'22.0"N, 98°04'12.1"E) , Huai Hee Village, Mae Sarieng, Mae Hong Song Province, northern Thailand, 13-VII-2017, by W. Srisuka (Coll. No. 106).
Male: small number of upper-eye facets in eleven vertical columns and 13 horizontal rows and antenna almost entirely yellow. Larva: postgenal cleft long, 2.8–3.3 times as long as the postgenal bridge (Fig.
Male, pupa and larva of S. teerachanense sp. nov. A–E male F–H pupa I larva. A hind basitarsus and second tarsomere (left side; lateral view) B coxites, styles and ventral plate (ventral view) C style (right side; ventrolateral view) D ventral plate and median sclerite (lateral view) E ventral plate (caudal view) F gill filaments (right side; lateral view) G terminal hooks (caudal view) H cocoon (dorsal view) I head capsule (ventral view). Scale bars: 1.0 mm (H); 0.1 mm (A, F, I); 0.02 mm (B–E); 0.01 mm (G).
Male (N = 9). Body length 2.0–2.2 mm.
Head. Somewhat wider than thorax. Upper eye dark brown, consisting of large facets in eleven vertical columns and 13 horizontal rows. Antenna entirely yellow, though few apical flagellomeres slightly darkened in some males; first flagellomere elongate, 1.8 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.5; sensory vesicle small, globular or ellipsoidal (0.16–0.17 times length of third palpal segment).
Legs.
Foreleg: tibia whitish yellow except apical three-tenths dark brown; basitarsus slightly dilated, 7.3–7.5 times as long as its greatest width. Hind leg: coxa light brown; tibia dark brown to brownish black except little more than basal two-fifths whitish yellow; tarsus (Fig.
Wing. Length 1.9–2.0 mm. Subcosta with one to four hairs, though no hairs in three males.
Abdomen. Dorsal surface of abdomen medium brown to brownish black, except most of segment 2 yellow to light ochreous.
Genitalia.
Ventral plate in ventral view (Fig.
Pupa (N = 9). Body length 2.4–2.6 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles, except dorsolateral surface of posterior half sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, all segments unpigmented except segments 1, 2, and 9 and bases of spine-combs of segments 6–8 light yellow; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 4). Body length 4.5–5.5 mm. Body dull ochreous with following color markings: thoracic segment 1 encircled with dark brown band (though disconnected ventrally), abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band dorsally along posterior margin, though that on segment 6 often partially faded leaving one round dorsomedial spot and two lateral spots of various size and shape), dorsal and dorsolateral surface of abdominal segments 5–8 faintly covered with pinkish or reddish brown pigments (Fig.
Head.
Head capsule yellow except eye-spot region whitish; head spots indistinct except those on lateral and ventral surfaces faintly positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.7–0.8:0.8–1.0. Labral fan with 28–30 primary rays. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle sparsely covered with unpigmented minute setae, though few posterior abdominal segments sparsely covered with dark minute unbranched setae dorsally. Rectal organ compound, each of three lobes with 7–11 finger-like secondary lobules. Posterior circlet with 90–95 rows of hooklets with up to 14 hooklets per row.
Female. Unknown.
The species name, teerachanense, refers to the name of the waterfall, Tee Ra Chan, where this species was collected.
Thailand (Mae Hong Son).
This new species is similar to S. roslihashimi described from Peninsular Malaysia (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium maewongense male, QSBG col. no. 80, Thailand, 12-VII-2018, by W. Srisuka”, collected from a small stream (width 20 cm, depth 2 cm bed sandy, slow flow, 19.5 °C, partially shaded, elevation 1,322 m, 16°06'02.9"N, 99°06'23.8"E, 98°30'53.0"E), at Chon Yen, Mae Wong National Park, Klong Lan District, Kham Phaeng Phet Province, Thailand, 12-VII-2018, by W. Srisuka (Coll. No. 80).
Paratypes : Three females, four males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and 14 mature larvae (two mature larvae for DNA analysis) (in 80% ethanol), same data as for holotype; one female (with its associated pupal exuviae and cocoon). (in 80% ethanol), collected from a stream of Klong Nam Lai (width 1.4 m, depth 13 cm, bed sandy, moderate flow, pH 6.23, 25.8 °C, exposed to the sun, elevation 196 m, 16°12'28.3"N, 99°15'47.8"E), at Klong Lan District, Kham Phaeng Phet Province, Thailand, 27-VI-2013, by W. Srisuka (Coll. No. 144).
Female: mandible lacking teeth on the outer margin. Male: upper-eye (large) facets in eleven vertical columns and 13 or 14 horizontal rows. Pupa: thorax bare except the anterior two-fifths or half and small area of the dorsal surface near the posterior margin moderately covered with tubercles, and cocoon with a short anterodorsal projection or bulge (Fig.
Male and pupa of S. maewongense sp. nov. A–D male E–G pupa. A hind basitarsus and second tarsomere (left side; lateral view) B coxites, styles and ventral plate (ventral view) C ventral plate and median sclerite (lateral view) D ventral plate (caudal view) E gill filaments (right side; lateral view) F terminal hooks (caudal view) G cocoon (dorsal view). Scale bars: 1.0 mm (G); 0.1 mm (A, E); 0.02 mm (B–D); 0.01 mm (F).
Female (N = 4). Body length 2.1–2.2 mm.
Head. Frontal ratio 1.8–2.0:1.0:2.3–3.3. Frons: head ratio 1.0:4.3–5.4. Labrum 0.68 times length of clypeus. Maxillary palpus: proportional length of third, fourth and fifth palpal segments 1.0:1.0–1.1:2.2–2.3; sensory vesicle medium-long, 0.36 times as long as third palpal segment. Lacinia with 8–11 inner and 12 or 13 outer teeth. Mandible with 21–23 teeth on inner margin and lacking teeth on outer margin (though outer margin undulated).
Legs. Fore basitarsus 6.3–6.5 times as long as its greatest width. Hind basitarsus 5.9–6.1 times as long as its greatest width and 0.7 and 0.6 times as wide as greatest width of hind tibia and femur, respectively. Calcipala nearly as long as wide and 0.56 times as wide as greatest width of hind basitarsus.
Wing. Length 2.2–2.3 mm.
Terminalia. Sternite 8 with 17–22 medium to long stout hairs and three to five short slender hairs. Ovipositor valve with one to four short hairs. Paraproct 0.5 times as long as wide, and with 22–25 short to medium-long hairs on outer surface. Cercus 0.4 times as long as wide. Spermatheca ellipsoidal, 1.4 times as long as its greatest width.
Male (N = 5). Body length 2.6 mm.
Head. Upper eye dark brown, consisting of large facets in eleven vertical columns and 13 or 14 horizontal rows on each side. Antenna: first flagellomere 1.6 times as long as second. Maxillary palpus: proportional length of third, fourth and fifth palpal segments 1.0:1.2:2.8; sensory vesicle small, globular or ellipsoidal, 0.19–0.21 times as long as palpal segment 3.
Legs.
Foreleg: basitarsus 7.7 times as long as its greatest width. Hind leg: basitarsus (Fig.
Wing. Length 2.2 mm. Subcosta haired except near apex bare.
Genitalia.
Ventral plate in ventral view (Fig.
Pupa (N = 9). Body length 3.0 mm.
Thorax.
Integument almost bare except anterior two-fifths to half moderately covered with round tubercles and small dorsal area near posterior margin sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, segments 1 and 2 bare. Terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 12). Body length 5.5–6.5 mm. Body creamy white to light ochreous with following color markings: thoracic segment 1 encircled with ochreous or reddish brown band (though disconnected ventromedially), thoracic segments 2 and 3 ochreous on ventral surface; abdominal segments 1 and 2 entirely grey, dorsal and dorsolateral surface of abdominal segment 4 with faint reddish brown band or spot (though completely faded in some larvae), abdominal segments 5 and 6 each with reddish brown, W-shaped, transverse band near posterior margin of dorsal and dorsolateral surface, which is always distinct on abdominal segment 5 but is faded to varying extent on abdominal segment 6 leaving one small round dorsomedial spot and two distinct lateral areas, abdominal segments 7 and 8 distinctly covered with reddish brown pigment on each dorsolateral surface (Fig.
Head. Head spots faintly (or rarely moderately) positive or indistinct. Antenna: proportional lengths of first, second, and third articles 1.0:0.8:0.7–0.8. Labral fan with 27–29 primary rays. Hypostoma: median tooth little longer than each corner tooth. Postgenal cleft small, rounded, 0.9–1.2 times length of postgenal bridge.
Abdomen. Rectal organ compound, each of three lobes with 6–8 finger-like secondary lobules. Anal sclerite with anterior arms 1.1–1.2 times as long as posterior ones. Posterior circlet with 91–96 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, maewongense, refers to the name of the national park, Mae Wong, where this species was collected.
Thailand (Kham Phaeng Phet).
Among 36 species of the S. asakoae species group, S. gyorkosae Takaoka & Davies from Indonesia (
The larval body color pattern (Fig.
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium loeiense male, QSBG col. no. 43, Thailand, 6-IX-2018, by W. Srisuka”, collected from a small stream (width 40 cm, depth 2 cm, bed sandy, slow flow, pH 5.8, 19.5 °C, partially shaded, elevation 1,525 m, 17°16'51.7"N, 101°31'02.5"E), at Khok Nok Kra Ba, Phu Luang, Phu Ruea District, Loei Province, Thailand, 6-IX-2018, by W. Srisuka (Coll. No. 43).
Paratypes : Five females, five males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and 15 mature larvae (two mature larvae for DNA analysis) (in 80% ethanol), same data as for holotype.
Female: mandible with several teeth on the outer margin. Male: small number of upper-eye facets in eleven vertical columns and 13 horizontal rows, presence of many hairs on subcosta and much widened hind basitarsus (Fig.
Female, male, pupa and larva of S. loeiense sp. nov. A female B–E male F–H pupa I larva. A genital fork (ventral view) B hind basitarsus and second tarsomere (left side; lateral view) C coxites, styles and ventral plate (ventral view) D ventral plate and median sclerite (lateral view) E ventral plate (caudal view) F gill filaments (right side; lateral view) G terminal hooks (caudal view) H cocoon (dorsal view) I head capsule (ventral view). Scale bars: 1.0 mm (H); 0.1 mm (B, F, I); 0.02 mm (A, C–E); 0.01 mm (G).
Female (N = 5). Body length 1.9–2.2 mm.
Head. Frontal ratio 1.9:1.0:2.7–2.9; frons:head ratio 1.0:4.7–4.8. Labrum 0.7 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1–1.2:2.6; sensory vesicle 0.26–0.32 times length of third palpal segment. Lacinia with 10–12 inner and 12–16 outer teeth. Mandible with 19–21 inner teeth and three to four outer teeth at some distance from tip.
Legs. Foreleg: trochanter yellow to dark yellow; femur light brown with apical cap medium brown (though extreme tip yellowish); tibia white except apical one-fourth brownish black; basitarsus moderately dilated, 6.1–6.7 times as long as its greatest width. Midleg: trochanter dark yellow; femur light brown except base and extreme apical tip yellowish); tibia whitish on basal one-third, greyish on middle one-third and dark brown on apical one-third (though whitish on basal two-thirds on posterior surface); tarsus dark brown to brownish black except basal one-fourth to half yellow (its border not well defined). Hind leg: femur light to medium brown with base whitish yellow and apical cap dark brown (though extreme tip yellowish); tibia white to yellowish white on basal two-thirds or little less and brownish black on rest; tarsus brownish black except little more than basal two-thirds (though base light brown) and basal half of second tarsomere white; basitarsus 5.6–6.3 times as long as wide, and 0.7–0.8 and 0.6 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as width at base, and 0.53–0.57 times as wide as greatest width of basitarsus.
Wing. Length 2.0–2.2 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except anterior four-fifths of segment 2 ochreous.
Terminalia.
Sternite 8 with 21–26 medium-long to long hairs together with three or four slender short hairs on each side. Ovipositor valve moderately covered with microsetae interspersed with two to four short hairs. Genital fork (Fig.
Male (N = 6). Body length 2.3–2.6 mm.
Head. Upper eye bright medium brown, consisting of (large) facets in eleven vertical columns and 13 horizontal rows. Antenna light to medium brown except scape, pedicel, and base of first flagellomere yellow; first flagellomere elongate, 1.5 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.4–2.5; sensory vesicle globular or ellipsoidal (0.14–0.19 times length of third palpal segment).
Legs.
Foreleg: trochanter dark yellow; femur light brown except apical tip yellowish; tibia whitish except apical one-third dark brown and inner margin dark yellow to light brown, and covered with white hairs on whitish yellow portion; tarsus brownish black; basitarsus somewhat dilated, 7.1–7.6 times as long as its greatest width. Midleg: trochanter yellow; femur light brown with base yellowish and apical cap medium brown (though apical tip yellow); tibia light to medium brown except basal one-third (or little more on posterior surface) whitish; tarsus dark brown except basal one-fourth to one-third of basitarsus dark yellow (border not well defined). Hind leg: coxa light brown; trochanter yellowish; femur light to medium brown with base yellow and apical cap dark brown (though apical tip yellow); tibia medium to dark brown except little less than basal half yellowish white; tarsus (Fig.
Wing. Length 2.0–2.1 mm. Subcosta with three to six hairs.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 11). Body length 2.5–2.8 mm.
Thorax.
Gill (Fig.
Abdomen.
Segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 13). Body length 5.0–5.5 mm. Body creamy white to light ochreous with following color markings: thoracic segment 1 encircled with distinct ochreous band (though disconnected ventromedially and dorsomedially), thoracic segments 2 and 3 ochreous or dark grey on ventral surface; abdominal segments 1–4 entirely greenish grey, abdominal segments 5, 7, and 8 dark grey on dorsal and dorsolateral surface; abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band along posterior margin dorsally, though that on abdominal segment 6 often partially faded, leaving three distinct, reddish brown spots (one round dorsomedial spot and two lateral spots of various size and shape), dorsal and dorsolateral surface of abdominal segments 7 and 8 faintly to distinctly covered with reddish brown pigment (Fig.
Head.
Head capsule yellow except narrow areas along posterior margin and surrounding areas of postgenal cleft somewhat darkened, and eye-spot region whitish; head spots distinctively positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.8:0.9. Labral fan with 35 or 36 primary rays. Hypostoma: four hypostomal bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig.
Abdomen. Rectal organ compound, each of three lobes with 10–15 finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms 0.9 times as long as posterior ones. Posterior circlet with 76–86 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, loeiense, refers to the name of the province, Loei, where this species was collected.
Thailand (Loei).
This new species is similar to S. asakoae in many characters including the presence of teeth on the outer margin of the female mandible, and small number of male upper-eye facets. However, it is distinguished from the latter species by the dorsum of pupal abdominal segments 1 and 2 yellowish (the dorsum of pupal abdominal segments 1 and 2 darkened in S. asakoae).
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium maelanoiense male, QSBG col. no. 105, Thailand, 14-VII-2017, by W. Srisuka”, collected from a small stream (width 25 cm, depth 2.5 cm, bed sandy, moderate flow, pH 6.8, 21.9 °C, partially shaded, elevation 974 m, 18°20'35.8"N, 98°01'23.8"E), at Tee La Chan Waterfall, Mae La Noi District, Mae Hong Son Province, Thailand, 14-VII-2017, by W. Srisuka (Coll. No. 105).
Paratypes : Two females (thoraces for DNA analysis) (with its associated pupal exuviae and cocoon) (in 80% ethanol), collected from a stream (width 110 cm, depth 12 cm, bed sandy, moderate flow, pH 6.9, 18.3 °C, partially shaded, elevation 1,446 m, 18°51'38.8"N, 99°22'15.2"E), at Kiew Fin, Muang Pan District, Lampang Province, Thailand, 6-IV-2018, by W. Srisuka (Coll. No. 36); two males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons) (in 80% ethanol), same data as for holotype.
Female: mandible with one tooth on the outer margin (Fig.
Female, male and pupa of S. maelanoiense sp. nov. A female B–F male G–I pupa. A mandible (left side) B hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (right side; lateral view) H terminal hooks (caudal view) I cocoon (dorsal view). Scale bars: 1.0 mm (I); 0.1 mm (B, G), 0.02 mm (C–F); 0.01 mm (A, H).
Female (N = 2). Body length 2.0 mm.
Head.
Frontal ratio 1.9:1.0:2.7; frons:head ratio 1.0:4.9. Labrum 0.70 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.1; sensory vesicle 0.26–0.32 times length of third palpal segment. Lacinia with ten or eleven inner and 12 or 13 outer teeth. Mandible (Fig.
Legs. Foreleg: basitarsus moderately dilated, 6.6 times as long as its greatest width. Midleg: tarsus dark brown to brownish black though basal half of basitarsus dark yellow (its border not well defined). Hind leg: coxa light brown; tibia yellowish white on basal three-fifths and light brown to brownish black on rest; tarsus brownish black except basal three-fifths (though base light brown) and basal half of second tarsomere yellowish white; basitarsus 6.9 times as long as wide, and 0.7 and 0.5 times as wide as greatest widths of tibia and femur, respectively; calcipala 0.6 times as wide as greatest width of basitarsus; claw with large basal tooth 0.44 times length of claw.
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except anterior four-fifths of segment 2 ochreous.
Terminalia. Sternite 8 with 27 medium-long to long hairs together with two to four slender short hairs on each side. Ovipositor valves each moderately covered with microsetae interspersed with one to three short hairs. Paraproct in ventral view with four sensilla on anteromedial surface; paraproct in lateral view 0.6 times as long as wide, with 19 or 20 medium-long to long hairs on ventral and lateral surfaces.
Male (N = 3). Body length 2.3–2.4 mm.
Head. Somewhat wider than thorax. Upper eye medium brown, consisting of large facets in eleven (rarely ten) vertical columns and 13 horizontal rows on each side. Antenna light to medium brown except scape, pedicel, and base of first flagellomere whitish yellow; first flagellomere elongate, 1.9 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.6; sensory vesicle 0.16–0.21 times length of third palpal segment.
Legs.
Foreleg: tibia yellowish white except apical three-tenths dark brown; basitarsus moderately dilated, 6.6–7.8 times as long as its greatest width. Hind leg: coxa light brown; tibia dark brown to brownish black except little less than basal half yellowish white; tarsus (Fig.
Wing. Length 2.0–2.1 mm. Subcosta with 2–13 hairs, though rarely without hair.
Genitalia.
Style in ventrolateral view (Fig.
Pupa (N = 5). Body length 2.5–3.0 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles except dorsal surface of posterior half sparsely covered with tubercles, and dorsolateral surface of posterior half almost bare. Gill (Fig.
Abdomen.
Dorsally, segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva. Unknown.
The species name, maelanoiense, refers to the district, Mae La Noi, where this species was collected.
Thailand (Lampang and Mae Hong Son).
Simulium maelanoiense sp. nov. is most similar in the male and pupa to S. sutheppuiense sp. nov. in many characters including the number of male upper-eye facets, relative width of the hind basitarsus compared to the hind tibia and femur, shape of the ventral plate when viewed caudally, and arrangement of the pupal gill. However, this new species is barely distinguished in the female from S. sutheppuiense sp. nov. by the number of the outer teeth of the mandible (one tooth in this new species versus three teeth in S. sutheppuiense sp. nov.) and the length ratio of the labrum against the clypeus (0.7 in this new species versus 0.6 in S. sutheppuiense sp. nov.).
Holotype : Male (thorax for DNA analysis) (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium phapeungense male, QSBG col. no. 107, Thailand, 12-VII-2017, by W. Srisuka”, collected from a stream (width 120 cm, depth 2 cm, bed sandy, fast flow, pH 7.6, 21.2 °C, partially shade, elevation 1,034 m, 19°36'58.5"N, 97°59'48.2"E), at Pha Peung, Muang District, Mae Hong Son Province, Thailand, 12-VII-2017, by W. Srisuka (Coll. No.107).
Paratype : One male (thorax for DNA analysis) (with its associated pupal exuviae and cocoon) (in 80% ethanol), same data as for holotype; five mature larvae (one mature larva for DNA analysis) (in 80% ethanol), collected from a stream (width 80 cm, depth 2.5 cm, moderate flow, pH 7.2, 20.1 °C, partially shaded, elevation 1,157 m, 19°11'10.3"N, 101°04'41.7"E), Nam Dan Village, Pua District, Nan Province, Thailand, 25-VII-2017, by W. Srisuka (Coll. No. 60).
Male: small number of brown upper-eye (large) facets in eleven vertical columns and 13 horizontal rows, and widened hind basitarsus (Fig.
Male, pupa and larva of S. phapeungense sp. nov. A–E male F–H pupa I, J larva A hind basitarsus and second tarsomere (left side; lateral view) B coxites, styles and ventral plate (ventral view) C style (right side; ventrolateral view) D ventral plate and median sclerite (lateral view) E ventral plate (caudal view) F gill filaments (right side; lateral view) G terminal hooks (caudal view) H cocoon (dorsal view) I head capsule showing postgenal cleft (ventral view) J postgenal cleft (ventral view). Scale bars: 1.0 mm (H); 0.1 mm (F, I, J); 0.02 mm (A–E); 0.01 mm (G).
Male (N = 2). Body length 2.2 mm.
Head. Somewhat wider than thorax. Upper eye dark brown, consisting of large facets in eleven vertical columns and 13 horizontal rows on each side. Antenna: first flagellomere elongate, 1.5 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpomeres 1.0:1.3:2.7; sensory vesicle ellipsoidal, 0.18–0.21 times length of third palpomere.
Legs.
Foreleg: tibia whitish except apical three-tenths dark brown; basitarsus somewhat dilated, 6.9 times as long as its greatest width. Midleg: tarsus dark brown except basal one-third of basitarsus dark yellow (border not well defined). Hind leg: coxa light brown; tibia light to dark brown except little less than basal half whitish; tarsus (Fig.
Wing. Length 2.0 mm. Subcosta with five hairs.
Genitalia.
Style in ventral view (Fig.
Pupa (N = 2). Body length 2.5 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles except dorsal surface of posterior two-thirds sparsely covered with tubercles and dorsolateral surfaces of posterior two-thirds almost bare. Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish except bases of spine-combs of segments 6–8 yellow; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 4). Body length 4.8–5.2 mm. Body creamy white to light ochreous with following color markings: thoracic segment 1 encircled with distinct grey band (though disconnected ventromedially), thoracic segments 2 and 3 grey on ventral surface; abdominal segments 1, 3, 4, and 5 light grey, abdominal segment 4 with reddish brown small spots dorsolaterally in one larva, abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band dorsally (though that on abdominal segment 6 partially faded, leaving one round dorsomedial spot and two lateral spots of various size and shape), dorsal and dorsolateral surface of abdominal segments 7 and 8 faintly greyish partially, overlaid by reddish brown pigment (Fig.
Head.
Head capsule yellow except narrow portion along posterior margin moderately darkened connected to dark posterolateral spots; head spots faintly to moderately positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.8:0.8. Labral fan with 30–33 primary rays. Hypostoma: lateral margin with four to six hypostomal bristles per side. Postgenal cleft (Fig.
Abdomen. Rectal organ compound, each of three lobes with 11–14 finger-like secondary lobules. Posterior circlet with 85–87 rows of hooklets with up to 14 or 15 hooklets per row.
Female. Unknown.
The species name, phapeungense, refers to the name of the village, Pha Peung, where this species was collected.
Thailand (Mae Hong Son and Nan).
Simulium phapeungense sp. nov. is similar to S. loeiense sp. nov. described above by having the small number of brown upper-eye (large) facets in eleven vertical columns and 13 horizontal rows and a widened hind basitarsus (Fig.
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium nanthaburiense male, QSBG col. no. 60, Thailand, 25-VII-2017, by W. Srisuka”, collected from a stream (width 80 cm, depth 2.5 cm, moderate flow, pH 7.2, 20.1 °C, partially shaded, elevation 1,157 m, 19°11'10.3"N, 101°04'41.7"E), Nam Dan Village, Pua District, Nan Province, Thailand, 25-VII-2017, by W. Srisuka (Coll. No. 60).
Paratypes : One female (thorax for DNA analysis) and 10 males (thorax of one male for DNA analysis) (with their associated pupal exuviae) (in 80% ethanol), same date and data as the holotype; one female (thorax for DNA analysis) (with their associated pupal exuviae) (in 80% ethanol) collected from a stream (width 120 cm, depth 10 cm, bed sandy, moderate flow, pH 6.63, 17.1 °C, partially shaded, elevation 1,154 m, 19°03'36.8"N, 99°19'15.7"E), Huai Mor Nuea Village, Doi Saket, Chiang Mai Province, northern Thailand, 2-II-2019, by W. Srisuka and A. Saeung (Coll. No. 49).
Female: mandible with two distinct teeth on the outer margin (11A). Male: small number of upper-eye facets in 12 vertical columns and 13 or 14 horizontal rows.
Female (N = 2). Body length 2.0 mm.
Head.
Frontal ratio 1.7:1.0:2.1; frons:head ratio 1.0:4.3. Labrum 0.64 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.4; sensory vesicle 0.26–0.29 times length of third palpal segment. Maxillary lacinia with ten or eleven inner and 13 or 14 outer teeth. Mandible (Fig.
Legs. Foreleg: coxa and trochanter whitish yellow; femur dark yellow to light brown with apical cap medium brown (though extreme tip yellowish); tibia yellowish white except little less than apical three-tenths brownish black; basitarsus moderately dilated, 6.0 times as long as its greatest width. Midleg: tarsus dark brown to brownish black though basal half or little less of basitarsus dark yellow (its border not well defined). Hind leg: coxa light brown with apical one-third yellow; tibia yellowish white on basal two-thirds and brownish black on rest; basitarsus 6.2 times as long as wide, and 0.65 and 0.53 times as wide as greatest widths of tibia and femur, respectively; calcipala slightly longer than width at base, and 0.56 times as wide as greatest width of basitarsus; claw with large basal tooth 0.45 times length of claw.
Female, male and pupa of S. nanthaburiense sp. nov. A female B–F male G–I pupa. A mandible (right side) B hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (right side; lateral view) H terminal hooks (caudal view) I cocoon (dorsal view). Scale bars: 1.0 mm (I); 0.1 mm (B, G); 0.02 mm (C–F); 0.01 mm (A, H).
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 ochreous.
Terminalia. Sternite 8 with 22 or 23 medium-long to long hairs together with three to six slender short hairs on each side. Paraproct in ventral view with three or four sensilla on anteromedial surface; paraproct in lateral view 0.6 times as long as wide, with 27–29 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.5 times as long as wide. Spermatheca 1.6 times as long as its greatest width.
Male (N = 11). Body length 2.0–2.3 mm.
Head. Somewhat wider than thorax. Upper eye dark brown, consisting of large facets in 12 vertical columns and 13 or 14 horizontal rows. Antenna light to medium brown except scape, pedicel and base of first flagellomere yellowish white; first flagellomere elongate, 1.8 times length of second. Maxillary palp light brown, with five palpal segments, proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1.2:2.3; sensory vesicle 0.19 times length of third palpal segment.
Legs.
Foreleg: trochanter yellow to dark yellow; tibia whitish except apical three-tenths dark brown and subbasal portion dark yellow to light brown (though outer surface whitish); basitarsus moderately dilated, 6.8–7.4 times as long as its greatest width. Midleg: tarsus dark brown except basal one-fourth to one-third of basitarsus dark yellow to light brown (border not well defined). Hind leg: coxa light brown; tarsus (Fig.
Wing. Length 1.9–2.0 mm. Subcosta with 1–10 hairs, though no hair in two males.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 13). Body length 2.5–2.7 mm.
Head.
Integument yellow. Thorax. Integument yellow, moderately covered with round tubercles except dorsolateral surface of posterior half sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva. Unknown.
The species name, nanthaburiense, refers to the historical name of Nan Province, Nanthaburi, where this species was collected.
Thailand (Nan).
This new species is similar to S. brinchangense described from Peninsular Malaysia (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium nanoiense male, QSBG col. no. 66, Thailand, 5-VIII-2017, by W. Srisuka”, collected from a stream (width 1 m, depth 3 cm, bed sandy, moderate flow, pH 6.2, 19.6 °C, partially shaded, elevation 1,349 m, 18°16'40.4"N, 100°30'19.0"E), Khun Sathan Village, Na Noi District, Nan Province, Thailand, 5-VIII-2017, by W. Srisuka (Coll. No. 66).
Paratypes : One female, nine males (thoraces of two males for DNA analysis) (with their associated pupal exuviae and cocoons), and 10 mature larvae (one mature larva for DNA analysis) (in 80% ethanol), same data as for holotype.
Diagnosis. Female: small sensory vesicle 0.22–0.24 times as long as the third palpal segment (Fig.
Female, male, pupa and larva of S. nanoiense sp. nov. A, B female C–G male H–J pupa K larva. A sensory vesicle (right side; anterior view) B mandible (right side) C hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (right side; lateral view) I terminal hooks (caudal view) J cocoon (dorsal view) K head capsule (ventral view). Scale bars: 1.0 mm (J); 0.1 mm (C, H, K); 0.02 mm (A, D–G); 0.01 mm(B, I).
Female (N = 1). Body length 2.0 mm.
Head.
Frontal ratio 1.8:1.0:2.7; frons:head ratio 1.0:5.0. Labrum 0.66 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.4; sensory vesicle (Fig.
Legs. Foreleg: tibia yellowish white except apical three-tenths brownish black; basitarsus moderately dilated, 7.1 times as long as its greatest width. Hind leg: coxa light brown; tibia yellowish white on little more than basal half and light brown to brownish black on rest; tarsus brownish black except basal three-tenths (though base light brown) and basal half of second tarsomere yellowish white; basitarsus 6.2 times as long as wide, and 0.7 and 0.6 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as width at base, and 0.6 times as wide as greatest width of basitarsus.
Wing. Length 2.2 mm.
Abdomen. Basal scale ochreous, with fringe of whitish yellow hairs. Dorsal surface of abdomen medium to dark brown except anterior four-fifths of segment 2 ochreous.
Terminalia. Sternite 8 bare medially, with 27 or 28 medium-long to long hairs together with two to four slender short hairs on each side. Ovipositor valves each moderately covered with microsetae interspersed with two short hairs. Paraproct in ventral view with four or five sensilla on anteromedial surface; paraproct in lateral view 0.6 times as long as wide, with 19–21 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.5 times as long as wide. Spermatheca 1.5 times as long as its greatest width.
Male (N = 10). Body length 2.1–2.3 mm.
Head. Slightly wider than thorax. Upper eye medium brown, consisting of large facets in 12 or 13 vertical columns and 14 (rarely 13) horizontal rows. Antenna: first flagellomere elongate, 1.7 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.3:2.6; sensory vesicle 0.14–0.17 times length of third palpal segment.
Legs.
Foreleg: tibia whitish yellow except little more than apical one-third dark brown, and small subbasal area on inner and lateral surfaces light brown; basitarsus somewhat dilated, 7.0–7.1 times as long as its greatest width. Midleg: tarsus dark brown except basal one-fourth to one-third of basitarsus dark yellow (border not well defined). Hind leg: coxa light brown; femur light to medium brown with base yellow and apical cap brownish black (though apical tip yellow); tibia dark brown to brownish black except basal half or little less whitish yellow; tarsus (Fig.
Wing. Length 1.9–2.0 mm. Subcosta bare in three males but with one to seven hairs in six males.
Genitalia.
Style in ventrolateral view (Fig.
Pupa (N = 11). Body length 2.4–2.6 mm.
Head. Integument yellow. Thorax. Integument yellow, moderately covered with round tubercles except dorsal and dorsolateral surfaces of posterior half almost bare or sparsely covered with tubercles, though small dorsal area near posterior margin sparsely to moderately covered with tubercles.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 9). Body length 5.0–5.6 mm. Body creamy white with following color markings: thoracic segment 1 encircled with dark grey band (though disconnected ventromedially), dorsal surface of thoracic segments 1–3 often light grey except posterior half of segment 3 light ochreous; ventral surface of thoracic segment 2 dark grey; ventral surface of thoracic segment 3 light grey on anterior one-third and light ochreous on posterior two-thirds; abdominal segments 1–3 entirely grey, abdominal segments 7 and 8 light grey dorsally, abdominal segment 4 with purplish transverse band (though often partially faded, leaving narrow band or small spot(s) dorsomedially), abdominal segments 5 and 6 each with distinct, dark purplish, W-shaped, transverse band dorsally (though that on segment 6 often faded partially, leaving one distinct round dorsomedial spot and two dorsolateral spots), dorsal and dorsolateral surfaces of abdominal segments 7 and 8 each covered with dark purplish pigment, though dorsomedial portions usually faded (Fig.
Head.
Head spots faintly or moderately positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.7:0.8. Labral fan with 33–38 primary rays. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle sparsely covered with unpigmented minute setae dorsally except 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. Rectal organ compound, each of three lobes with ten or eleven finger-like secondary lobules. Anal sclerite with anterior arms slightly longer than posterior ones. Posterior circlet with 85–88 rows of hooklets with up to 14 hooklets per row.
The species name, nanoiense, refers to the district, Na Noi, where this species was collected.
Thailand (Nan).
This new species is similar to S. brinchangense described from Peninsular Malaysia (
This new species is similar in larval body color to S. junkumae sp. nov. but is distinguished from the latter species by the dorsal and dorsolateral surfaces of larval abdominal segments 7 and 8 light grey (Fig.
Holotype: Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium muangpanense male, QSBG col. no. 86, Thailand, 9-VIII-2016, by W. Srisuka”, collected from a stream (width 30 cm, depth 5 cm, bed sandy, moderate running, pH 6.3, 22.6 °C, exposed to the sun, elevation 1,097 m, 18°50'03.7"N, 99°22'32.2"E), at Pa Meing Village, Muang Pan District, Lampang Province, northern Thailand, 9-VIII-2016, by W. Srisuka (Coll. No. 86).
Paratypes : Three females (thorax of one female for DNA analysis), four males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and five mature larvae (one mature larva for DNA analysis) (in 80% ethanol), same data as for holotype.
Female: mandible with two or three outer teeth (Fig.
Female, male, pupa and larva of S. muangpanense sp. nov. A female B–F male G–I pupa J, K larva. A mandible (right side) B hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (left side; lateral view) H terminal hooks (caudal view) I cocoon (dorsal view) J head capsule (ventral view) K postgenal cleft. Scale bars: 1.0 mm (I); 0.1 mm (B, G, J, K); 0.02 mm (C–F); 0.01 mm (A, H).
Female (N = 3). Body length 2.0–2.1 mm.
Head.
Frontal ratio 1.8:1.0:2.1; frons:head ratio 1.0:4.0. Labrum 0.61 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.0:2.5; sensory vesicle medium sized, ellipsoidal (0.29–0.31 times length of third palpal segment), with small opening. Lacinia with nine or ten inner and 14 outer teeth. Mandible (Fig.
Legs. Foreleg: basitarsus moderately dilated, 6.4 times as long as its greatest width. Midleg: femur light brown with basal one-fourth whitish yellow and apical cap medium brown (though extreme tip yellowish); tarsus dark brown to brownish black though little less than basal half of basitarsus yellow (its border not well defined). Hind leg: coxa light brown with apical one-third yellowish white; tibia white to yellowish white on basal two-thirds and light brown to brownish black on rest; basitarsus 6.04 times as long as wide, and 0.76 and 0.58 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as or lightly longer than width at base, and 0.5–0.6 times as wide as greatest width of basitarsus.
Wing. Length 2.0 mm.
Terminalia. Sternite 8 with 18–21 medium-long to long hairs together with four slender short hairs on each side. Ovipositor valve moderately covered with microsetae interspersed with two or three short hairs. Paraproct in ventral view with four or five or six sensilla on anteromedial surface; paraproct in lateral view 0.6 times as long as wide, with 24–28 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.46 times as long as wide.
Male (N = 5). Body length 2.1–2.5 mm.
Head. Slightly wider than thorax. Upper eye dark brown, consisting of large facets in 13 vertical columns and 14 horizontal rows on each side. Antenna: first flagellomere 1.5–1.7 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments1.0:1.2:2.9–3.1; sensory vesicle globular or ellipsoidal, 0.15–0.18 times length of third palpal segment.
Legs.
Foreleg: femur light brown except apical cap medium brown (though apical tip yellowish); tibia light brown on basal one-third (though basal tip yellow and outer surface narrowly yellowish white), yellowish white on middle one-third, and brownish black on apical one-third; basitarsus moderately dilated, 6.7–7.3 times as long as its greatest width. Midleg: femur light brown with base yellowish on inner surface and apical cap medium brown (though apical tip yellowish); tibia medium brown except basal one-third (or little more on posterior surface) yellowish white; tarsus dark brown except base of basitarsus dark yellow to light brown (border not well defined). Hind leg: coxa light brown with apical one-third yellowish; femur light to medium brown with base whitish yellow and apical cap brownish black (though apical tip yellow); tibia dark brown to brownish black except basal half or little less whitish yellow; tarsus dark brown except basal two-fifths of basitarsus and little less than basal half of second tarsomere whitish yellow; basitarsus (Fig.
Wing. Length 1.6–1.9 mm. Other characters as in female except subcosta with 0–6 hairs.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 8). Body length 2.3–2.6 mm.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, all segments pale yellowish; segments 1 and 2 almost bare or sparsely covered with minute tubercles; segment 9 with pair of flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 4). Body length 4.8–5.0 mm. Body ochreous with following color markings: thoracic segment 1 encircled with grey band (though disconnected ventromedially), thoracic segments 2 and 3 grey on ventral surface (though often faded on segment 3); abdominal segments 1 and 4 entirely grey, abdominal segments 5, 7, and 8 grey on dorsal and dorsolateral surface; abdominal segment 4 with faint narrow reddish brown transverse band on dorsal and dorsolateral surface in one larva; abdominal segment 5 with distinct reddish brown, W-shaped, transverse band dorsally, abdominal segment 6 often with three reddish brown spots (one round dorsomedial spot and two lateral spots of various size and shape), and abdominal segments 7 and 8 distinctly covered with reddish brown pigment usually on dorsolateral surface (Fig.
Head.
Head capsule yellow with narrow area along posterior margin darkened; head spots faintly or moderately positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.7–0.8:0.8–0.9. Labral fan with 31 or 32 primary rays. Hypostoma with five hypostomal bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle almost bare except abdominal segments 5–8 moderately covered with unbranched dark minute setae (though setae rarely bifid) on dorsal and dorsolateral surface. Rectal organ compound, each of three lobes with seven to nine finger-like secondary lobules. Anal sclerite with anterior arms 1.1–1.2 times as long as posterior ones. Posterior circlet with 76–80 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, muangpanense, refers to the district, Muang Pan, where this species was collected.
Thailand (Lampang).
The male of this new species is similar to S. phulocense from Vietnam (
Simulium tanahrataense from Peninsular Malaysia (
The larva of this new species is almost the same in body color pattern as S. monglaense described from Myanmar (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium maehongsonense male, QSBG col. no. 107, Thailand, 12-VII-2017, by W. Srisuka”, collected from a stream (width 120 cm, depth 2 cm, bed sandy, fast flow, pH 7.6, 21.2 °C, partially shade, elevation 1,034 m, 19°36'58.5"N, 97°59'48.2"E), at Pha Peung, Muang District, Mae Hong Son Province, Thailand, 12-VII-2017, by W. Srisuka (Coll. No. 107).
Paratypes : Two females (thorax of one female for DNA analysis), seven males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and six mature larvae (one mature larva for DNA analysis) (in 80% ethanol), same data as for holotype.
Female: short sensory vesicle and mandible lacking distinct outer teeth (Fig.
Female, male, pupa and larva of S. maehongsonense sp. nov. A female B–F male G–I pupa J larva. A mandible (right side) B hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (left side; lateral view) H terminal hooks (caudal view) I cocoon (dorsal view) J head capsule (ventral view). Scale bars: 1.0 mm (I); 0.1 mm (B, G, J); 0.02 mm (C–F); 0.01 mm (A, H).
Female (N = 2). Body length 1.9 mm.
Head.
Frontal ratio 1.9:1.0:2.1; frons:head ratio 1.0:4.2. Labrum 0.52 times length of clypeus. Maxillary palpus: proportional length of third, fourth and fifth palpal segments 1:1.1:2.2; sensory vesicle medium-long (0.26 times length of third palpal segment. Lacinia with 9–11 inner and 11–13 outer teeth. Mandible (Fig.
Legs. Foreleg: tibia yellowish white except apical three-tenths brownish black; basitarsus moderately dilated, 5.5 times as long as its greatest width. Midleg: tarsus dark brown though basal half of basitarsus yellow (its border not well defined). Hind leg: coxa light brown; tibia yellowish white on basal two-thirds and light brown to brownish black on rest; basitarsus 6.7 times as long as wide, and 0.6 and 0.5 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 0.46 times length of claw.
Wing. Length 1.9 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 ochreous (though narrow portion along posterior margin somewhat darkened).
Terminalia. Sternite 8 bare medially, with 20–22 medium-long to long hairs together with two or three slender short hairs on each side. Ovipositor valves each moderately covered with microsetae interspersed with two or three short hairs. Paraproct in lateral view 0.6 times as long as wide, with 18 medium-long to long hairs on ventral and lateral surfaces. Spermatheca ellipsoid, 1.3 times as long as its greatest width.
Male (N = 8). Body length 2.0–2.4 mm.
Head. Slightly wider than thorax. Upper eye dark brown, consisting of large facets in 13 vertical columns and 14 horizontal rows on each side. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.8; sensory vesicle 0.17 times length of third palpal segment.
Legs.
Foreleg: tibia yellowish white except l apical one-third dark brown, and inner and lateral surfaces of subbasal portion light brown; basitarsus moderately dilated, 6.5–7.3 times as long as its greatest width. Hind leg: coxa light brown; tibia dark brown to brownish black except little less than basal half yellowish white; tarsus dark brown except little less than basal half of basitarsus and basal one-third of second tarsomere yellowish; basitarsus (Fig.
Wing. Length 1.9–2.0 mm. Subcosta with four to nine hairs in four males but bare in four males.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 10). Body length 2.5–2.7 mm.
Head. Integument yellow. Thorax. Integument yellow.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 5). Body length 4.5–5.5 mm. Body light ochreous with following color markings: thoracic segment 1 encircled with distinct greyish band (though disconnected ventromedially), thoracic segment 2 greyish and thoracic segment 3 ochreous on ventral surface; abdominal segment 1 light grey, abdominal segments 5 and 6 each with distinct reddish brown spot of various size and shape dorsomedially, and dorsal and dorsolateral surface of abdominal segments 4–8 faintly to moderately covered with pinkish to reddish brown pigments (Fig.
Head.
Head spots moderately positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.8:0.7–0.9. Labral fan with 38–40 primary rays. Hypostoma with four to six hypostomal bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig.
Abdomen. Posterior circlet with 88–96 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, maehongsonense, refers to the province, Mae Hong Son, where this species was collected.
Thailand (Mae Hong Son).
The male of this new species is similar to S. phulocense from Vietnam (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium chaowaense male, QSBG col. no. 164, Thailand, 16-III-2017, by W. Srisuka”, collected from a small stream (width 40 cm, depth 3.5 cm, bed sandy, moderate flow, pH 7.3, 21.9 °C, partially, elevation 582 m, 18°45'30.2"N, 100°20'11.4"E), Chao Wa Waterfall, Song District, Phrae Province, Thailand, 16-III-2017, by W. Srisuka (Coll. No. 164).
Paratypes : Three males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and six mature larvae (one mature larva for DNA analysis) (in 80% ethanol), same data as in the holotype
Male: number of upper-eye facets in 14 or 15 vertical columns and 14 or 15 horizontal rows, and moderately widened hind basitarsus (Fig.
Male, pupa and larva of S. chaowaense sp. nov. A–D male E–G pupa H larva. A hind basitarsus and second tarsomere (left side; lateral view) B coxites, styles and ventral plate (ventral view) C ventral plate and median sclerite (lateral view) D ventral plate (caudal view) E gill filaments (right side; lateral view) F terminal hooks (caudal view) G cocoon (dorsal view) H head capsule (ventral view). Scale bars: 1.0 mm (G); 0.1 mm (A, E, H); 0.02 mm (B–D); 0.01 mm (F).
Male (N = 4). Body length 2.0–2.3 mm.
Head. Distinctly wider than thorax. Upper eye dark brown, consisting of large facets in 14 or 15 vertical columns and 14 or 15 horizontal rows on each side. Antenna light to medium brown except scape, pedicel, and base of first flagellomere yellow; first flagellomere elongate, 1.8 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.0–1.2:2.5–2.8; sensory vesicle globular or ellipsoidal, 0.16–0.17 times length of third palpal segment.
Thorax. Scutum with faint longitudinal vittae (one median and two submedian).
Legs.
Foreleg: tibia whitish yellow except basal one-fourth light brown and apical one-third dark brown; basitarsus moderately dilated, 7.3–7.4 times as long as its greatest width. Hind leg: coxa light brown; femur light to medium brown with base yellow and apical cap dark brown (though apical tip yellow); tibia dark brown except little less than basal half yellow; tarsus (Fig.
Wing. Length 1.6–1.7 mm. Subcosta with 2–8 hairs.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 4). Body length 2.3–2.5 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles except posterior half sparsely covered with small tubercles on dorsolateral and lateral surfaces. Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 5). Body length 4.1–4.5 mm. Body with following color markings: thoracic segment 1 encircled with greyish band (though disconnected ventromedially), ventral surface of thoracic segment 2 grey and that of thoracic segment 3 ochreous; abdominal segments 1–4 encircled with grey transverse band, and abdominal segments 5–8 greyish dorsally and dorsolaterally; abdominal segments 5 and 6 each overlaid with light reddish brown transverse band, appearing W-shaped, on dorsal and dorsolateral surfaces (though often faded out to varying extent, leaving small round medial spot and dorsolateral spots, or only small medial spot), and also overlaid with pair of round spots on ventral surface; abdominal segments 7 and 8 each overlaid with light reddish brown pigments to varying extent on dorsal and dorsolateral surfaces (completely faded out in one larva) (Fig.
Head.
Head capsule yellow to dark yellow except eye-spot region whitish, sparsely covered with minute setae (though moderately on dorsal surface); head spots faintly to moderately positive. Antenna: proportional lengths of first, second, and third articles 1.00:0.65–0.75:0.81–0.95. Labral fan with 34–38 primary rays. Hypostoma with row of nine apical teeth, of which median tooth little longer than each corner tooth; lateral margin smooth; four hypostomal bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle very sparsely covered with unpigmented minute setae (though few posterior abdominal segments sparsely to moderately covered with dark minute unbranched or bifid setae (rarely trifid setae) 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. Rectal scales minute, unpigmented. Rectal organ compound, each of three lobes with seven to nine finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms 1.1 times as long as posterior ones. Posterior circlet with 73–78 rows of hooklets with up to 13 or 14 hooklets per row.
Female. Unknown.
The species name, chaowaense, refers to the name of the waterfall, Chao Wa, where this species was collected.
Thailand (Phrae).
This new species is similar to S. tanahrataense described from males and their associated pupal exuviae collected from Peninsular Malaysia (
This new species is similar to S. confertum from Vietnam (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium pitasawatae male, QSBG col. no. 49, Thailand, 2-II-2019, by W. Srisuka”, collected from a stream (width 120 cm, depth 10 cm, bed sandy, moderate flow, pH 6.6, 17.1 °C, partially shaded, elevation 1,154 m, 19°03'36.8"N, 99°19'15.7"E), Huai Mor Nuea Village, Doi Saket, Chiang Mai, Thailand, 2-II-2019, by W. Srisuka and A. Saeung (Coll. No. 49).
Paratypes : Three females, eight males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and 10 mature larvae (one mature larva for DNA analysis) (in 80% ethanol), same data as for holotype.
Female: small sensory vesicle and mandible with three to five distinct teeth on outer margin (Fig.
Female, male, pupa and larva of S. pitasawatae sp. nov. A female B–F male G–J pupa K larva. A mandible (left side) B hind basitarsus and second tarsomere (left side; lateral 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, H gill filaments (left side; lateral view) I terminal hooks (caudal view) J cocoon (dorsal view) K head capsule (ventral view). Scale bars: 1.0 mm (J); 0.1 mm (B, G, H, K); 0.02 mm (C–F); 0.01 mm (A, I).
Female (N = 3). Body length 2.0 mm.
Head.
Frontal ratio 1.7–1.8:1.0:2.3–2.5; frons:head ratio 1.0:4.2–4.9. Labrum 0.61–0.64 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1–1.2:2.0–2.3; sensory vesicle medium sized, ellipsoidal (0.32–0.35 times length of third palpal segment). Lacinia with nine or ten inner and 12–14 outer teeth. Mandible (Fig.
Legs. Foreleg: basitarsus moderately dilated, 6.3–6.7 times as long as its greatest width. Hind leg: coxa medium brown; tibia yellowish white on basal three-fifths and light brown to brownish black on rest; basitarsus 5.7–6.0 times as long as wide, and 0.7–0.8 and 0.6 times as wide as greatest widths of tibia and femur, respectively; claw with large basal tooth 0.43 times length of claw.
Wing. Length 2.0–2.1 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except anterior one-fifths of segment 2 ochreous.
Terminalia. Sternite 8 bare medially, with 19–23 medium-long to long hairs together with two to four slender short hairs on each side. Paraproct in ventral view with four sensilla on anteromedial surface; paraproct in lateral view 0.6–0.7 times as long as wide, with 18–20 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.5 times as long as wide.
Male (N = 9). Body length 2.1–2.3 mm.
Head. Somewhat wide than thorax. Upper eye dark brown, consisting of large facets in 14 (rarely 13) vertical columns and 15 horizontal rows on each side. Antenna: first flagellomere elongate, 1.8 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.4; sensory vesicle small, ellipsoidal (0.22–0.25 times length of third palpal segment).
Legs.
Foreleg: tibia light grey to light brown except basal tip and outer surface of basal two-thirds whitish, and apical one-third dark brown; basitarsus 7.9–8.3 times as long as its greatest width. Hind leg: coxa medium brown; tarsus (Fig.
Wing. Length 2.0–2.1 mm. Subcosta bare in three males but with one to five hairs in six males.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 12). Body length 2.4–2.7 mm.
Head. Integument yellow, Thorax. Integument yellow, moderately covered with round tubercles except dorsal and dorsolateral surfaces of posterior half sparsely covered with tubercles.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without tubercles; segment 5 with one spine (same size as those on segments 6–9) on one side in two pupae; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 9). Body length 4.3–5.4 mm. Body light ochreous with following color markings: thoracic segment 1 encircled with light to dark brown (rarely reddish brown) band (though disconnected ventrally), thoracic segments 2 and 3 ochreous on ventral surface; abdominal segment 4 with reddish brown transverse band (though often entirely faded out), abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band dorsally along posterior margin, though often partially faded out leaving one round dorsomedial spot and two lateral spots of various size and shape), dorsal and dorsolateral surface of abdominal segments 5–8 faintly to moderately covered with pinkish or reddish brown pigments (Fig.
Head.
Head capsule yellow except eye-spot region whitish; head spots moderately positive. Antenna: proportional lengths of first, second, and third articles 1.0:0.7–0.8:0.8–1.0. Labral fan with 23 or 24 primary rays. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle sparsely covered with unpigmented minute setae, though few posterior abdominal segments sparsely covered also with dark minute unbranched setae dorsally. Rectal organ compound, each of three lobes with 7–11 finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms 1.1 times as long as posterior ones. Posterior circlet with 81–85 rows of hooklets with up to 14 or 15 hooklets per row.
The species name, pitasawatae, is in honor of Associate Prof. Dr. Benjawan Pitasawat, Head of Department of Parasitology, Faculty of Medicine, Chiang Mai University, Thailand, who kindly supported A. Saeung in collections of black flies.
Thailand (Chiang Mai).
This new species is similar to S. tamdaoense Takaoka, Sofian-Azirun & Ya’cob described from Vietnam (
The larva of this new species is similar in the body color pattern to S. (G.) sutheppuiense sp. nov. but is barely distinguished from the latter species by the labral fan with 23 or 24 primary rays (29 or 30 primary rays in S. (G.) sutheppuiense sp. nov.).
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium banluangense male, Thailand, 12-VI-2001, by W. Choochote & H. Takaoka”, reared from a pupa collected from a stream (width 1.0 m, 25 °C, exposed to the sun, elevation 804 m, 18°32'33.3"N, 98°35'32.5"E), at Ban Luang, Chiang Mai Province, Thailand, 12-VI-2001, by W. Choochote and H. Takaoka.
Male: upper-eye (large) facets in 14 vertical columns and 14 or 15 horizontal rows on each side, and hind basitarsus (Fig.
Male and pupa of S. banluangense sp. nov. A–D male E–H pupa. A hind basitarsus and second tarsomere (left side; lateral view) B coxites, styles and ventral plate (ventral view) C ventral plate and median sclerite (lateral view) D ventral plate (caudal view) E, F gill filaments (lateral view E right side F left side) G terminal hooks (caudal view) H cocoon (dorsal view). Scale bars: 1.0 mm (H); 0.1 mm (A, E, F); 0.02 mm (B–D); 0.01 mm (G).
Male (N = 1). Body length 2.4 mm.
Head. Slightly wider than thorax. Upper eye medium brown, consisting of 14 vertical columns and 14 or 15 horizontal rows of large facets on each side. Antenna: first flagellomere elongate, 1.9 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:3.0; sensory vesicle 0.21–0.24 times length of third palpal segment.
Legs.
Foreleg: tibia light brown except median large area of outer surface of basal three-fourths whitish and apical one-fourth dark brown; tarsus brownish black; basitarsus moderately dilated, 6.4 times as long as its greatest width. Midleg: tarsus dark brown except basal one-fourth to two-fifths of basitarsus dark yellow to light brown (border not well defined). Hind leg: tibia dark brown to brownish black except little less than basal half whitish yellow; tarsus (Fig.
Wing. Length 2.1 mm. Subcosta with seven hairs.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 1). Body length 3.0 mm.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, segments 1 and 2 without minute tubercles; terminal hooks (Fig.
Cocoon
(Fig.
Female and Larva. Unknown.
The species name, banluangense, refers to the name of the locality, Ban Luang, where this species was collected.
Thailand (Chiang Mai).
Simulium doisaketense described based on pupal and larval specimens from Thailand (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium junkumae male, QSBG col. no. 92, Thailand, 7-IV-2017, by W. Srisuka”, collected from a small stream (width 40 cm, depth 3 cm, bed sandy, moderate flow, pH 6.2, 19 °C, partially shaded, elevation 1,395 m, 18°49'09.5"N, 98°53'14.3"E), at Doi Pui Temple, Doi Suthep Pui, Muang District, Chiang Mai Province, Thailand, 7-IV-2017, by W. Srisuka (Coll. No. 92).
Paratypes : Five females, five males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and six mature larvae (two mature larvae for DNA analysis) (in 80% ethanol), same data as for holotype.
Female: relatively narrower frons against the head width (1.0:4.7–5.5), and mandible without teeth on the outer margin (Fig.
Female, male, pupa and larva of S. junkumae sp. nov. A, B female C–I male J–L pupa M larva. A mandible (right side) B, C hind basitarsus and second tarsomere (left side; lateral view B female C male) D coxites, styles and ventral plate (ventral view) E, F styles (right side; ventrolateral view) G ventral plate (ventral view) H ventral plate and median sclerite (lateral view) I ventral plate (caudal view) J gill filaments (right side; lateral view) K terminal hooks (caudal view) L cocoon (dorsal view) M head capsule (ventral view). Scale bars: 1.0 mm (L); 0.1 mm (B, C, J, M); 0.02 mm (D–I); 0.01 mm (A, K).
Female (N = 5). Body length 2.0–2.3 mm.
Head.
Frontal ratio 1.8–1.9:1.0:2.5–3.2; frons:head ratio 1.0:4.7–5.5. Labrum 0.64–0.67 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.0–1.1:2.0–2.1; sensory vesicle medium sized, ellipsoidal (0.27–0.32 times length of third palpal segment), with medium opening. Lacinia with ten or eleven inner and 14 or 15 outer teeth. Mandible (Fig.
Legs.
Foreleg: trochanter dark yellow to light brown; femur light brown with apical cap medium brown (though extreme tip yellowish); basitarsus moderately dilated, 6.7–7.0 times as long as its greatest width. Midleg: trochanter dark yellow except basal half yellow; tarsus dark brown to brownish black except basal one-fourth of basitarsus dark yellow (its border not well defined). Hind leg: tibia yellowish white on little more than basal half and light brown to brownish black on rest; basitarsus (Fig.
Wing. Length 2.1–2.3 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except anterior four-fifths of segment 2 whitish. Ventral surface of segment 2 whitish and those of other segments light to dark brown; sternal plate on segment 7 undeveloped.
Terminalia. Sternite 8 bare medially, with 25–28 medium-long to long hairs together with two or six slender short hairs on each side. Ovipositor valve triangular, thin, membranous, moderately covered with microsetae interspersed with two to four short hairs; inner margins nearly straight or slightly sinuous, somewhat sclerotized, and moderately separated from each other. Paraproct in ventral view with five or six sensilla on anteromedial surface; paraproct in lateral view somewhat produced ventrally beyond ventral tip of cercus, 0.6 times as long as wide, with 25–28 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.4–0.5 times as long as wide. Spermatheca ellipsoidal, 1.3 times as long as its greatest width.
Male (N = 6). Body length 2.2–2.3 mm.
Head. Slightly wider than thorax. Upper eye dark brown, consisting of large facets in 15–17 vertical columns and 15–17 horizontal rows on each side. Antenna: first flagellomere elongate, 1.6–1.8 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.0:2.4–2.6; sensory vesicle ellipsoidal, 0.17–0.23 times length of third palpal segment.
Legs.
Foreleg: tibia light grey to light brown except basal tip and outer surface of basal two-thirds whitish, and apical one-third dark brown; basitarsus moderately dilated, 7.8–8.1 times as long as its greatest width. Midleg: basitarsus dark brown except basal tip yellow. Hind leg: coxa medium brown; tibia dark brown to brownish black except little more than basal one-third whitish yellow; tarsus (Fig.
Wing. Length 2.2–2.4 mm. Subcosta with 7–12 hairs.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 11). Body length 2.4–2.8. mm.
Thorax.
Integument moderately covered with round tubercles except dorsal and dorsolateral surfaces of posterior half sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, all segments light yellow; segments 1 and 2 without tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 4). Body length 4.9–5.4 mm. Body creamy white with following color markings: thoracic segment 1 encircled with distinct reddish brown band (though disconnected ventromedially), thoracic segments 2 and 3 ochreous on ventral surface; abdominal segments 1–3 entirely grey, abdominal segment 4 with reddish brown transverse band dorsally (though often partially to completely faded, leaving narrow band or small spot(s) dorsally), abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band (of which central and dorsolateral parts marked) along posterior margin on dorsal and dorsolateral surfaces (though band on abdominal segment 6 often partially faded, leaving one round dorsomedial spot and two larger dorsolateral lateral spots); abdominal segments 7 and 8 covered with reddish brown pigments on dorsal and dorsolateral surfaces (though central portion often faded to varying extent) (Fig.
Head.
Head capsule yellow except eye-spot region whitish, sparsely covered with minute setae (though moderately on dorsal surface); head spots faintly to moderately positive or indistinct. Antenna: proportional lengths of first, second, and third articles 1.0:0.7–0.8:0.7–0.8. Labral fan with 24 or 25 primary rays. Postgenal cleft (Fig.
Abdomen. Rectal organ compound, each of three lobes with 8–11 finger-like secondary lobules. Anal sclerite: anterior arms 1.0–1.2 times as long as posterior ones. Posterior circlet with 81–84 rows of hooklets with up to 14 hooklets per row.
The species name junkumae is in honor of Assistant Prof. Dr. Anuluck Junkum, Department of Parasitology, Faculty of Medicine, Chiang Mai University, Thailand, for her kind help in the field and laboratory works of black flies.
Thailand (Chiang Mai).
This new species is similar to S. hongthaii Takaoka, Sofian-Azirun & Ya’cob described from Vietnam (
The larva of S. sp. nr. asakoae-2 reported from Thailand by
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium kiewfinense male, QSBG col. no. 36, Thailand, 6-IV-2018, by W. Srisuka”, collected from a stream (width 1.1 m, depth 12 cm, bed sandy, moderate flow, pH 6.9, 18.3 °C, partially shaded, elevation 1,446 m, 18°51'38.8"N, 99°22'15.2"E), at Kiew Fin, Muang Pan District, Lampang Province, Thailand, 6-IV-2018, by W. Srisuka (Coll. No. 36).
Paratypes : Two males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons) (in 80% ethanol), same data as for holotype.
Male: greater number of upper-eye facets in 16 or 17 vertical columns and 16 or 17 horizontal rows one each side.
Male (N = 3). Body length 2.0–2.1 mm.
Head. Somewhat wider than thorax. Upper eye medium brown, consisting of large facets in 16 or 17 vertical columns and 16 or 17 horizontal rows. Antenna composed of scape, pedicel and nine flagellomeres, medium to dark brown except scape, pedicel, and base of first flagellomere yellow; first flagellomere elongate, 1.8 times length of second. Maxillary palpus light brown, with five palpal segments, proportional lengths of third, fourth, and fifth palpomeres 1.0:1.0:2.6; sensory vesicle 0.19 times length of third palpal segment.
Legs.
Foreleg: tibia whitish except apical three-tenths dark brown and subbasal portion light brown; basitarsus moderately dilated, 6.6–7.1 times as long as its greatest width. Midleg: tarsus dark brown except basal one-third of basitarsus dark yellow to light brown (border not well defined). Hind leg: coxa light brown; tarsus (Fig.
Wing. Length 1.9–2.0 mm. Subcosta bare.
Genitalia.
Coxite in ventral view (Fig.
Male and pupa of S. kiewfinense sp. nov. A–E male F–H pupa. A hind basitarsus and second tarsomere (left side; lateral view) B coxites, styles and ventral plate (ventral view) C style (right side; ventrolateral view) D ventral plate and median sclerite (lateral view) E ventral plate (caudal view) F gill filaments (left side; lateral view) G terminal hooks (caudal view) H cocoon (dorsal view). Scale bars: 1.0 mm (H); 0.1 mm (A, F); 0.02 mm (B–E); 0.01 mm (G).
Pupa (N = 3). Body length 2.5–2.7 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles except dorsolateral surface of posterior half almost bare. Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Female and mature larva. Unknown.
The species name, kiewfinense, refers to the local name, Kiew Fin, where this species was collected.
Thailand (Lampang).
This new species is similar to S. junkumae sp. nov. described above and S. hongthaii from Vietnam (
Holotype. Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium huaimorense male, QSBG col. no. 49, Thailand, 2-II-2019, by W. Srisuka”, collected from a stream (width 1.2 m, depth 10 cm, bed sandy, moderate flow, pH 6.6, 17.1 °C, exposed to the sun, elevation 1,154 m, 19°03'36.8"N, 99°19'15.7"E), at a coffee plantation, Huai Mor, Doi Saket, Chiang Mai Province, Thailand, 2-II-2019, by W. Srisuka (Coll. No. 49).
Paratypes. One female, two males (thorax of one male for DNA analysis) (with its associated pupal exuviae and cocoon) (in 80% ethanol), and three mature larvae (in 80% ethanol), collected from a small stream (width 30 cm, depth 2 cm, bed sandy, moderate flow, pH 7.3, 19.8 °C, partially shaded, elevation 1,440 m, 19°54'04.1"N, 99°34'26.6"E), at coffee plantation, Pha Lung Village, Muang District, Chiang Rai Province, northern Thailand, 30-VIII-2018, by W. Srisuka (Coll. No. 122).
Male: upper-eye large facets in 16 vertical columns and 17 horizontal rows. Pupa: extremely short primary stalks of the dorsal and middle triplets of filaments (Fig.
Female, male, pupa and larva of S. huaimorense sp. nov. A female B–F male G–I pupa J larva. A mandible (right side) B hind basitarsus and second tarsomere (left side; lateral 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 gill filaments (right side; lateral view) H terminal hooks (caudal view) I cocoon (dorsal view) J head capsule (ventral view). Scale bars: 1.0 mm (I); 0.1 mm (B, G, J); 0.02 mm (C–F); 0.01 mm (A, H).
Female (N = 1). Body length 2.0 mm.
Head.
Frontal ratio 1.9:1.0:2.6; frons:head ratio 1.0:4.8. Labrum 0.67 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.3; sensory vesicle ellipsoidal, medium long (0.27–0.28 times length of third palpal segment). Lacinia with nine or ten inner and 14 or 15 outer teeth. Mandible (Fig.
Legs. Foreleg: Basitarsus moderately dilated, 6.5 times as long as its greatest width. Hind leg: coxa light brown; tibia yellowish white on basal two-thirds and light brown to brownish black on rest; basitarsus 6.2 times as long as wide, and 0.7 and 0.6 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as width at base, and 0.6 times as wide as greatest width of basitarsus; claw with large basal tooth 0.46 times length of claw.
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except anterior five-sixths of segment 2 ochreous
Terminalia. Sternite 8 bare medially, with 23 or 24 medium-long to long hairs together with three or four slender short hairs on each side. Ovipositor valve moderately covered with microsetae interspersed with two or three short hairs; inner margins nearly straight or slightly sinuous, Paraproct with four sensilla on anteromedial surface; paraproct in lateral view 0.6 times as long as wide, with 24 or 25 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.5 times as long as wide. Spermatheca ellipsoidal, 1.3 times as long as its greatest width.
Male (N = 3). Body length 2.0 mm.
Head. Slightly wider than thorax. Upper eye dark brown, consisting of large facets in 16 vertical columns and 17 horizontal rows on each side. Antenna: first flagellomere elongate, 1.9 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.2; sensory vesicle small, ellipsoidal, 0.22 times length of third palpal segment.
Legs.
Foreleg: coxa whitish yellow; trochanter light brown; femur light brown except apical tip yellowish; tibia whitish yellow except inner and lateral surface of basal one-fifth light brown and little more than apical one-third dark brown; basitarsus 7.0–8.0 times as long as its greatest width. Midleg: trochanter light brown except base yellow; femur light to medium brown with base yellowish and apical cap dark brown (though apical tip yellow). Hind leg: coxa light brown except apical one-third whitish yellow; femur medium to dark brown with basal tip yellow and apical cap brownish black (though apical tip yellow); tibia dark brown to brownish black except little less than basal half whitish yellow and subbasal small portion somewhat dark yellow; tarsus (Fig.
Wing. Length 2.0 mm. Subcosta bare or with two hairs.
Abdomen. Ventral surface of segment 2 white, those of segments 3 and 4 light brown although sternal plates medial brown, and those of other segments medium to dark brown.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 4). Body length 2.5 mm.
Thorax.
Integument yellow, moderately covered with round tubercles except dorsal and dorsolateral surface of posterior half sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without minute tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 3). Body length 4.5–5.3 mm. Body creamy white with following color markings: thoracic segment 1 encircled with distinct reddish brown band (though disconnected ventromedially), thoracic segments 2 and 3 ochreous on ventral surface; abdominal segments 1–4 entirely grey, abdominal segments 7 and 8 light grey on dorsal and dorsolateral surfaces; abdominal segments 5 and 6 each with distinct reddish brown, W-shaped, transverse band (of which central and dorsolateral parts marked) along posterior margin of dorsal and dorsolateral surfaces (though band on abdominal segment 6 often partially faded, leaving one round dorsomedial spot and two larger dorsolateral lateral spots); abdominal segments 7 and 8 covered with reddish brown pigments on dorsal and dorsolateral surfaces (though central portion often faded out to varying extent) (Fig.
Head.
Head capsule yellow except eye-spot region whitish, sparsely covered with minute setae (though moderately on dorsal surface); head spots faintly to moderately positive. Antenna: proportional lengths of first, second, and third articles 1.00:0.72–0.75:0.75–0.82. Labral fan with 24–26 primary rays. Postgenal cleft (Fig.
Abdomen. Rectal organ compound, each of three lobes with 11–13 finger-like secondary lobules. Anal sclerite: anterior arms nearly as long as or slightly longer than posterior ones. Posterior circlet with 86–92 rows of hooklets with up to 14 hooklets per row.
The species name, huaimorense, refers to the locality name, Huai Mor, where this species was collected.
Thailand (Chiang Mai and Chiang Rai).
Simulium huaimorense sp. nov. is similar to S. myanmarense described from Myanmar (
This new species is distinguished from S. maewongense sp. nov. and four other related species (noted under S. maewongense sp. nov.), which have a cocoon with a short anterodorsal projection (Fig.
This new species is similar in the higher number of male upper-eye facets and the arrangement of the pupal gill filaments to S. kiewfinense sp. nov., and S. junkumae sp. nov. from Thailand and S. hongthaii from Vietnam (
This new species is distinguished from S. doisaketense from Doi Saket, Chiang Mai Province (
Holotype : Female (thorax for DNA analysis) (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium songense male, QSBG col. no. 60, Thailand, 25-VII-2017, by W. Srisuka”, collected from a small stream (width 80 cm, depth 2.5, fast flow, pH 7.2, 20.1 °C, partially shaded, elevation 1,157 m, 19°11'10.3"N, 101°04'41.7"E), at Nam Dan Village, Pua District, Nan Province, northern Thailand, 25-VII-2017, by W. Srisuka (Coll. No. 60).
Paratype : One female (thorax for DNA analysis) (with its associated pupal exuviae and cocoons) (in 80% ethanol), collected from a small stream (width 40 cm, depth 3.5 cm, bed sandy, moderate flow, pH 7.36, 21.9 °C, partially shaded, elevation 582 m, 18°45'30.2"N, 100°20'11.4"E), at Chao Wa Waterfall, Song District, Phrae Province, Thailand, 16-III-2017, by W. Srisuka (Coll. No.164).
Female: mandible without distinct teeth on the outer margin (Fig.
Female (N = 2). Body length 1.9 mm.
Head.
Frontal ratio 1.9:1.0:2.6; frons:head ratio 1.0:4.9. Labrum 0.65 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1:2.6; sensory vesicle 0.26–0.29 times length of third palpal segment. Lacinia with nine or ten inner and 13 outer teeth. Mandible (Fig.
Legs. Foreleg: basitarsus moderately dilated, 5.5 times as long as its greatest width. Midleg: tarsus dark brown to brownish black though basal half of basitarsus dark yellow or light brown (its border not well defined). Hind leg: coxa light brown); tibia yellowish white on basal two-thirds and light brown to brownish black on rest; tarsus brownish black except basal seven-tenths (though base light brown) and basal half of second tarsomere yellowish white; basitarsus 6.2 times as long as wide, and 0.7 and 0.5 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as width at base, and 0.6 times as wide as greatest width of basitarsus; claw with large basal tooth 0.46 times length of claw.
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 ochreous (though narrowly darkened along posterior margin).
Terminalia. Sternite 8 bare medially, with 28– 31 medium-long to long hairs together with three or four slender short hairs on each side. Ovipositor valves each moderately covered with microsetae interspersed with three to four short hairs. Paraproct in ventral view with four or five sensilla on anteromedial surface; paraproct in lateral view 0.5 times as long as wide, with 29–32 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view 0.5 times as long as wide.
Pupa (N = 2). Body length 3.0 mm.
Head. Integument yellow.
Thorax.
Integument yellow, moderately covered with round tubercles except dorsolateral surface of posterior half sparsely covered with tubercles. Gill (Fig.
Abdomen.
Dorsally, all segments light yellowish; segments 1 and 2 without tubercles; segment 9 with pair of wide flat terminal hooks (Fig.
Cocoon
(Fig.
Male and mature larva. Unknown.
The species name, songense, refers to the district, Song, one of the two localities where this species was collected.
Thailand (Nan and Phrae).
Lacking teeth on the outer margin of the mandible, the female of this new species seems to be most similar to S. myanmarense and S. monglaense from Myanmar (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium klonglanense male, QSBG col. no. 144, Thailand, 27-VI-2013, by W. Srisuka”, collected from a stream of Klong Nam Lai (width 1.4 m, depth 13 cm, bed sandy, moderate flow, pH 6.2, 25.8 °C, exposed to the sun, elevation 196 m, 16°12'28.3"N, 99°15'47.8"E), at Klong Lan District, Kham Phaeng Phet Province, Thailand, 27-VI-2013, by W. Srisuka (Coll. No. 144).
Paratypes : Three females, one male (thorax for DNA analysis) (with their associated pupal exuviae and cocoons), and eight mature larvae (in 80% ethanol), same data as for holotype.
Female: sensory vesicle elongated (Fig.
Female, male, pupa and larva of S. klonglanense sp. nov. A–C female D–I male J–L pupa M larva. A, D sensory vesicles (right side; anterior view A female D male) B mandible (right side) C genital fork E hind basitarsus and second tarsomere (left side; lateral view) F coxites, styles and ventral plate (ventral view) G style (right side; ventrolateral view) H ventral plate and median sclerite (lateral view) I ventral plate (caudal view) J gill filaments (right side; lateral view) K terminal hooks (caudal view) L cocoon (dorsal view) M head capsule (ventral view). Scale bars: 1.0 mm (L); 0.1 mm (E, J, M); 0.02 mm (A, C, D, F–I); 0.01 mm (B, K).
Female (N = 3). Body length 1.8 mm.
Head.
Frontal ratio 1.8:1.0:2.2; frons:head ratio 1.0:4.2. Labrum 0.59 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.0:2.1; sensory vesicle (Fig.
Legs. Foreleg: basitarsus moderately dilated, 5.2–5.4 times as long as its greatest width. Midleg: tarsus light to medium brown though little less than basal half of basitarsus yellow (its border not well defined). Hind leg: coxa light brown; tibia yellowish white on basal two-thirds and light to dark brown on rest; tarsus medium brown except basal two-thirds (though base light brown) and basal half of second tarsomere yellowish white; basitarsus 5.8–6.6 times as long as wide, and 0.7 and 0.6 times as wide as greatest widths of tibia and femur, respectively.
Wing. Length 1.6 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 ochreous (though narrow portion along posterior margin somewhat darkened).
Terminalia.
Sternite 8 with 17 medium-long to long hairs together with four slender short hairs on each side. Genital fork (Fig.
Male (N = 2). Body length 2.0 mm.
Head.
Somewhat wider than thorax. Upper eye medium brown, consisting of large facets in 13 or 14 vertical columns and 14 or 15 horizontal rows on each side. Antenna: first flagellomere elongate, 1.7 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.7; sensory vesicle (Fig.
Thorax. Scutum medium to dark brown except anterolateral calli ochreous, with three dark-brown longitudinal vittae (one medial and two submedial), white pruinose except three longitudinal vittae non-pruinose when illuminated anterodorsally and viewed dorsally. Scutellum medium brown. Postnotum medium to dark brown.
Legs.
Foreleg: basitarsus slightly dilated, 6.8 times as long as its greatest width. Hind leg: coxa light brown; tibia dark brown except little less than basal half whitish yellow; tarsus (Fig.
Wing. Length 1.5 mm. Subcosta with 0–2 hairs.
Genitalia.
Style in ventral view (Fig.
Pupa (N = 5). Body length 2.0–2.4 mm.
Head. Integument light yellow.
Thorax.
Integument light yellow, moderately covered with round tubercles except dorsolateral surface of posterior one-third nearly bare. Gill (Fig.
Abdomen.
Dorsally, all segments unpigmented except segment 9 light yellow; segments 1 and 2 without minute tubercles; segment 9 with pair of small flat terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 8). Body length 3.3–4.0 mm. Body with following color markings: thoracic segment 1 encircled with grey (or greyish ochreous) band (though disconnected ventromedially), thoracic segments 2 and 3 grey or ochreous on ventral surface; abdominal segments 1–4 each encircled with greyish band, abdominal segments 5–8 greyish on dorsal and dorsolateral surfaces; abdominal segment 4 with or without reddish purplish transverse band (though often partially faded medially and ventrally), abdominal segments 5–8 each faintly to moderately overlaid with reddish purplish pigments dorsally and dorsolaterally (though faded medially to various extent), abdominal segments 5 and 6 each with pair of small grey or reddish purplish spots ventrally, and abdominal segment 7 with grey transverse band (overlaid with reddish purplish pigment) ventrally (Fig.
Head.
Head capsule yellow except eye-spot region whitish; head spots indistinct. Antenna: proportional lengths of first, second, and third articles 1.0:0.8:0.8–1.0. Labral fan with 30–34 primary rays. Postgenal cleft (Fig.
Abdomen. Rectal organ compound, each of three lobes with six to eight finger-like secondary lobules. Anal sclerite of usual X-form, with anterior arms nearly as long as posterior ones. Posterior circlet with 71–74 rows of hooklets with up to 14 hooklets per row.
The species name, klonglanense, refers to the district, Klong Lan, where this species was collected.
Thailand (Kham Phaeng Phet).
This new species is similar to S. lurauense described from Peninsular Malaysia (
This new species is similar to S. quychauense Takaoka & Chen from Vietnam, which is known only for the male and pupa (
This new species is similar to S. thituyenae Takaoka & Pham from Vietnam, which is known only for the female and pupa (
Holotype : Male (with its associated pupal exuviae and cocoon) (in 80% ethanol) labeled as “Holotype: Simulium namdanense male, QSBG col. no. 59, Thailand, 25-VII-2017, by W. Srisuka”, collected from a medium-sized stream of Nam Khwang (width 4 m, depth 30 cm, bed sandy, fast flow, pH 7.2, 19.5 °C, partially shaded, elevation 1,192 m, 19°11'18.3"N, 101°04'43.7"E), at Nam Dan Village, Pua District, Nan Province, Thailand, 25-VII-2017, by W. Srisuka (Coll. No. 59).
Paratypes : Three females, two males (with their associated pupal exuviae and cocoons), and five mature larvae (in 80% ethanol), same data as for holotype; two females, four males (thorax of one male for DNA analysis) (with their associated pupal exuviae and cocoons), and two mature larvae (one mature larva for DNA analysis) (in 80% ethanol), collected from a medium-sized stream (width 3 m, depth 32 cm, bed sandy and rocky, moderate flow, pH 6.5, 23.2 °C, partially shaded, elevation 503 m, 18°48'44.2"N, 98°56'21.3"E), at Huai Kaew Waterfall, Doi Suthep, Muang District, Chiang Mai Province, Thailand, 30-VII-2017, by W. Srisuka (Coll. No. 94).
Female: elongate sensory vesicle (Fig.
Female, male, pupa and larva of S. namdanense sp. nov. A, C female B, D–I male J, K pupa L larva. A, B sensory vesicles (right side; anterior view A female B male) C genital fork D hind basitarsus and second tarsomere (left side; lateral view) E coxites, styles and ventral plate (ventral view) F, G styles (right side; ventrolateral view) H ventral plate and median sclerite (lateral view) I ventral plate (caudal view) J gill filaments (right side; lateral view) K terminal hooks (caudal view) L head capsule (ventral view). Scale bars: 0.1 mm (D, J, L); 0.02 mm (A–C, E–I); 0.01 mm (K).
Female (N = 5). Body length 1.8–2.0 mm.
Head.
Frons dark brown, densely covered with yellowish white scale-like recumbent short hairs (no dark longer hairs); frontal ratio 1.8–1.9:1.0:2.2–2.4; frons:head ratio 1.0:4.0–4.1. Labrum 0.59–0.63 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:0.9:1.9–2.0; sensory vesicle (Fig.
Legs. Foreleg: trochanter light brown; basitarsus moderately dilated, 5.7–5.8 times as long as its greatest width. Midleg: trochanter dark yellow to light brown. Hind leg: coxa light brown; tibia yellowish white on basal half or little more and light brown to brownish black on rest; basitarsus yellowish white except base and little less than apical one-third dark brown, 6.1–6.4 times as long as wide, and 0.7 and 0.5–0.6 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as width at base, and 0.45–0.56 times as wide as greatest width of basitarsus; claw with large basal tooth 0.47–0.52 times length of claw.
Wing. Length 2.0 mm.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 (except narrow portion along posterior margin darkened) ochreous.
Terminalia.
Sternite 8 with 18–26 medium-long to long hairs together with two or three slender short hairs on each side. Ovipositor valve moderately covered with microsetae interspersed with two to four short hairs. Genital fork (Fig.
Male (N = 7). Body length 2.0 mm.
Head.
Slightly wider than thorax. Upper eye medium brown, consisting of large facets in 15 or 16 vertical columns and 16 or 17 horizontal rows on each side. Antenna: first flagellomere elongate, 1.6–1.8 times length of second. Maxillary palpus light brown, with five palpal segments, proportional lengths of third, fourth, and fifth palpal segments 1.0:1.1–1.3:2.7; third palpomere (Fig.
Legs.
Foreleg: femur light brown except apical cap medium brown (though apical tip yellowish); tibia light brown except median large portion of outer surface whitish and little less than apical one-third dark brown; basitarsus moderately dilated, 6.5–6.9 times as long as its greatest width. Hind leg: coxa light to medium brown; trochanter whitish yellow; femur light to medium brown with base whitish yellow and apical cap dark brown (though apical tip yellow); tibia dark brown to brownish black except little less than basal half yellow; tarsus (Fig.
Wing. Length 1.8–1.9 mm. Subcosta bare or with one to three hairs.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 12). Body length 2.2–2.5 mm.
Head. Integument yellow. Thorax. Integument yellow, moderately covered with round tubercles except posterior half sparsely covered with minute tubercles on each dorsolateral and lateral surface.
Thorax.
Gill (Fig.
Abdomen.
Dorsally, all segments light yellow, or unpigmented except segments 1, 2 and 9 light yellow; segments 1 and 2 bare; segment 9 with pair of conical terminal hooks (Fig.
Cocoon. Whitish yellow, slipper-shaped, thinly woven, moderately extended ventrolaterally; anterior margin not thickly woven medially, with no bulge; 2.5–3.0 mm long by 1.5–2.0 mm wide.
Mature larva (N = 6). Body length 3.9–4.2 mm. Body with following color markings: thoracic segments 1–3 ochreous; abdominal segments 1–4 each encircled with light grey band, abdominal segments 5–8 light grey, overlaid with reddish brown pigment on dorsal and dorsolateral surfaces (though often irregularly faded to varying extent dorsomedially) (Fig.
Head.
Head capsule yellow, moderately covered with unpigmented minute setae (though sparsely on lateral and ventral surfaces); head spots faintly positive (rarely negative or indistinct). Antenna: proportional lengths of first, second, and third articles 1.0:0.7–0.8:0.8–0.9. Labral fan with 32–36 primary rays. Hypostoma: anterior row of nine teeth, of which median tooth somewhat longer than each lateral tooth, with four or five hypostomal bristles per side lying nearly parallel to lateral margin. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle almost bare except abdominal segments 5–8 moderately covered with unpigmented and slightly-darkened unbranched minute setae on dorsal and dorsolateral surface; last abdominal segment moderately covered with unpigmented minute setae on each side of anal sclerite to base of ventral papillae. Rectal organ compound, each of three lobes with 8–10 finger-like secondary lobules. Anal sclerite with anterior arms 1.1 times as long as posterior ones. Posterior circlet with 72–84 rows of hooklets with up to 13 or 14 hooklets per row.
The species name, namdanense, refers to the name of the locality, Nam Dan, where this species was collected.
Thailand (Chiang Mai and Nan).
This new species is similar to S. lurauense described from Peninsular Malaysia (
This new species is almost indistinguishable from S. thituyenae from Vietnam (
This new species is distinguished from S. klonglanense sp. nov. by the female wing length (2.0 mm in this new species versus 1.6 mm in S. klonglanense sp. nov.), and number of male upper-eye facets in 15 or 16 vertical columns and 16 or 17 horizontal rows (13 or 14 vertical columns and 14 or 15 horizontal rows in S. klonglanense sp. nov.).
Simulium (Gomphostilbia) myanmarense
Takaoka, Srisuka & Saeung, in
8 females, 6 males (with their associated pupal exuviae and cocoons), and 20 mature larvae (one mature larva for DNA analysis) (in 80% ethanol), collected from a small stream (width 55 cm, depth 3 cm, bed sandy, moderate flow, pH 7.2, 18.4 °C, partially shaded, elevation 901 m, 19°28'18.3"N, 100°28'00.8"E), at Pha Dang Village, Chiang Kham District, Pha Yao Province, Thailand, 20-III-2018, by W. Srisuka (Coll. No. 75); one female, one male (with their associated pupal exuviae and cocoons), and two mature larvae (in 80% ethanol), collected from a small stream (width 30 cm, depth 5 cm, bed sandy, moderate flow, pH 6.3, 23.6 °C, exposed to the sun, elevation 1,097 m, 18°50'03.7"N, 99°22'32.2"E), at Pa Miang Village, Muang Pan District, Lampang Province, Thailand, 9-VIII-2016, by W. Srisuka (Coll. no. 86).
Female: mandible lacking outer teeth. Male: upper-eye facets in 15 (rarely 14) vertical columns and 15 (rarely 16) horizontal rows on each side. Pupa: cocoon with a short anterodorsal projection. Larva: medium-sized postgenal cleft and abdominal segments 1 and 2 each encircled with a grey transverse band (Fig.
Myanmar and Thailand (Phayao and Lampang).
Simulium (Gomphostilbia) inthanonense Takaoka & Suzuki, 1984: 18–21 (female, pharate male, pupa, and larva).
This species was described from females, pharate males, pupae and larvae collected from Doi Inthanon National park, Chiang Mai Province (
This species is here transferred from the S. ceylonicum species group to the S. asakoae species group, based on the male ventral plate emarginated on both sides when viewed ventrally (Fig.
Female, male, pupa and larva of S. inthanonense. A–C female D–H male I–K pupa L larva. A, D sensory vesicles (right side; anterior view A female D male) B mandible (right side) C spermatheca E hind basitarsus and second tarsomere (left side; lateral view) F coxites, styles and ventral plate (ventral view) G ventral plate and median sclerite (lateral view) H ventral plate (caudal view) I gill filaments (right side; lateral view) J terminal hooks (caudal view) K cocoon (dorsal view) L head capsule (ventral view). Scale bars: 1.0 mm (K); 0.1 mm (E, I, L); 0.02 mm (A, C, D, F–H); 0.01 mm (B, J).
Schematic illustrations of larval body color patterns (dorsal view). A S. teerachanense sp. nov. B S. pitasawatae sp. nov. C S. sutheppuiense sp. nov. D S. asakoae E S. klonglanense sp. nov. F S. namdanense sp. nov. G S. chaowaense sp. nov. H S. loeiense sp. nov. I S. huaimorense sp. nov. J S. nanoiense sp. nov. K S. junkumae sp. nov. L S. myanmarense M S. maewongense sp. nov. N S. phapeungense sp. nov. O S. muangpanense sp. nov. P S. maehongsonense sp. nov. Q S. thungchangense sp. nov. R S. chiangdaoense S S. inthanonense (morph 1) T S. inthanonense (morph 2).
Neighbor-joining tree of the 26 Thai species of the Simulium asakoae species group based on COI gene sequences (586 positions). The numbers at the nodes mean the bootstrap confidence values after 500 replicates. The bootstrap values above 50% are shown. The scale bar indicates the distance in substitutions per nucleotide. The symbols in front of the species names are each the collection number (corresponding to those in the text), developmental stages (F, female; M, male; L, larva), and sample number (if two or more samples were examined) in this order.
Ten females, 10 males (with their associated pupal exuviae and cocoons) (in 80% ethanol), collected from a small stream (width 40 cm, depth 10 cm, bed sandy, moderate flow, pH 7.2, 15.3 °C, partially shaded, elevation 1,314 m, 18°31'01.9"N, 98°28'17.3"E), at Mae Klang Watershed, Doi Inthanon National Park, Mae Cham District, Chiang Mai Province, Thailand, 21-XII-2018, by W. Srisuka (Coll. No. 25); five females and five males and five mature larvae (two mature larvae for DNA analysis) (in 80% ethanol) collected in a stream (width 1.0 m, depth 20 cm, bed sandy, flow moderate, partially shaded, pH 6.8, 14.0 °C, elevation 1,589 m, 18°30'29.8"N, 98°30'37.4"E), at Mae Aum, Doi Inthanon National Park, Chom Thong District, Chiang Mai Province, Thailand, 25-I-2019, by W. Srisuka (Coll. No. 20). One male (thorax for DNA analysis) (with its associated pupal exuviae and cocoon) (in 80% ethanol), collected from a stream (width 70 cm, depth 10 cm, pH 6.5, 13.8 °C, moderate flow, bed sandy, partially shaded, elevation 1,685 m, 18°31'15.4"N, 98°29'59.4"E), before check point 2, Doi Inthanon National parks, Chiang Mai Province, northern Thailand, 20-III-2018, by W. Srisuka (Coll. No. 6).
Female and male: darkened hair tuft at the base of the radial vein and relatively long sensory vesicle (Fig.
Female (N = 15). Body length 2.0–2.1 mm.
Head.
Frontal ratio 1.6–1.8:1.0:2.3–2.8; frons:head ratio 1.0:4.7–5.1. Labrum 0.55–0.61 times length of clypeus. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:0.9–1.0:2.1–2.2; sensory vesicle (Fig.
Legs. Foreleg: basitarsus moderately dilated, 6.2–6.4 times as long as its greatest width. Hind leg: coxa medium brown; basitarsus 7.1–7.2 times as long as wide, and 0.6–0.7 and 0.5–0.6 times as wide as greatest widths of tibia and femur, respectively; calcipala nearly as long as width at base, and 0.6 times as wide as greatest width of basitarsus; claw with large basal tooth 0.43 times length of claw.
Wing. Length 2.1–2.4 mm. Costa with dark spinules and hairs. Hair tuft on base of radius dark brown.
Abdomen. Dorsal surface of abdomen medium to dark brown except most of segment 2 whitish yellow.
Terminalia.
Sternite 8 with 16–25 medium-long to long hairs together with three or four slender short hairs on each side. Ovipositor valves each covered with microsetae interspersed with three or four short hairs. Paraproct with five to eight sensilla on anteromedial surface; paraproct in lateral view with 26–32 medium-long to long hairs on ventral and lateral surfaces. Cercus in lateral view short, slightly rounded posteriorly, 0.5 times as long as wide. Spermatheca (Fig.
Male (N = 16). Body length 2.3–2.6 mm.
Head.
Slightly wider than thorax. Upper eye medium brown, consisting of large facets in 17 vertical columns and 17 (rarely 18) horizontal rows on each side. Antenna medium to dark brown except scape, pedicel and base of first flagellomere yellow, though apical half or two-thirds of pedicel light brown in some males; first flagellomere 1.7 times length of second. Maxillary palpus: proportional lengths of third, fourth, and fifth palpal segments 1.0:1.2:2.6; third palpomere (Fig.
Legs.
Foreleg: coxa yellow; trochanter light brown; femur medium brown; light brown except apical one-third dark brown and median large portion on outer surface white and shiny; basitarsus moderately dilated, 6.9–7.7 times as long as its greatest width. Midleg: trochanter light brown except base yellow; femur medium brown with base yellowish and apical cap dark brown (though apical tip yellow); tarsus light to dark brown. Hind leg: coxa medium brown; trochanter yellow to dark yellow; tarsus (Fig.
Wing. Length 2.1–2.3 mm. Other characters as in female except subcosta bare. Halter. Light to medium brown except lower portion whitish.
Genitalia.
Coxite in ventral view (Fig.
Pupa (N = 31). Body length 2.5–3.0 mm.
Head. Integument yellow.
Thorax.
Integument yellow, with two somewhat darkened areas in tandem on dorsal surface of posterior half. Gill (Fig.
Abdomen.
Dorsally, segments 1–5 light greyish brown, segment 9 and bases of spine-combs of segments 6–8 light yellow; segments 1 and 2 without minute tubercles; segment 9 with pair of triangular terminal hooks (Fig.
Cocoon
(Fig.
Mature larva (N = 3). Body length 5.6–6.0 mm. Body light ochreous with following color markings: thoracic segment 1 encircled with distinct reddish brown band (though disconnected ventromedially), thoracic segments 2 and 3 dark ochreous on ventral surface; abdominal segments 1 and 3 (or abdominal segments 1–3) greenish grey; abdominal segments 5 and 6 each with three distinct, reddish brown spots (one round dorsomedial spot and two dorsolateral spots) (Fig.
Head.
Head spots faintly to moderately positive. Antenna: proportional lengths of first, second, and third articles 1.0:1.0–1.1:0.8–0.9. Labral fan with 40 primary rays. Hypostoma with four to six hypostomal bristles per side lying nearly parallel to or slightly diverginf from lateral margin. Postgenal cleft (Fig.
Thorax and Abdomen. Thoracic and abdominal cuticle sparsely covered with unpigmented minute setae. Rectal organ compound, each of three lobes with 9–13 finger-like secondary lobules. Anal sclerite with anterior arms nearly as long as posterior ones. Posterior circlet with 82–87 rows of hooklets with up to 15 hooklets per row.
Thailand (Chiang Mai), China and Vietnam.
Simulium (Gomphostilbia) chiangdaoense
One male and two mature larvae, collected from a small stream (width 55 cm, depth 3 cm, bed sandy, moderate flow, pH 7.2, 18.4 °C, partially shaded, elevation 901 m, 19°28'18.3"N, 100°28'00.8"E), at Pha Tea Do mountain, Mae Cham, Chiang Mai Province, northern Thailand, 27-VI-2018, by W. Srisuka (Coll. No. 82).
Simulium (Gomphostilbia) asakoae
One female, reared from a pupa collected from a stream (width 30 cm, depth 5 cm, bed sandy, moderate flow, pH 6.3, 23.6 °C, exposed to the sun, elevation 1,097 m, 18°50'03.7"N, 99°22'32.2"E), at Pa Miang Village, Chae Hom, Lampang Province, Thailand, 9-VIII-2016, by W. Srisuka (Coll. No. 86); one male, reared from a pupa collected from a stream (width 5 m, depth 25 cm, pH 6.9, 22.1 °C, moderate flow, bed sandy, partially shade, elevation 759 m, 16°19'01.2"N, 99°06'19.9"E) near Tao Dam Waterfall, Klong Lan, Kam Phaeng Phet Province, Thailand, 20-XII-2016, by W. Srisuka (Coll. No. 145); one female and one male, reared from pupae collected from a stream (width 2.5 m, depth 30 cm, moderate flow, bed sandy, partially shaded, pH 5.9, 19 °C, elevation 1,293 m, 16°58'49.5"N, 101°03'29.3"E) at Rangkla Village, Nakhon Thai District, Phitsanulok Province, Thailand, 18-VI-2019, by W. Srisuka (Coll. No. 200); one female reared from a pupa and one mature larva collected from a stream (width 50 cm, depth 16 cm, moderate flow, bed sandy, pH 7.3, 21 °C, elevation 1,231 m, 18°32'16.3"N, 98°31'30.5"E) at Khun Klang Village, Doi Inthanon National Park, Chiang Mai Province, northern Thailand, 24-VII-2019, by W. Srisuka (Coll. No. 201).
Female
1 | Hair tuft of base of radial vein brownish | S. inthanonense |
– | Hair tuft of base of radial vein yellow | 2 |
2 | Sensory vesicle elongate, more than 0.5 times length of third palpal segment (Fig. |
3 |
– | Sensory vesicle short to medium-long, less than 0.4 times length of third palpal segment (Fig. |
4 |
3 | Wing length 1.6 mm | S. klonglanense sp. nov. |
– | Wing length 2.0 mm | S. namdanense sp. nov. |
4 | Mandible with distinct teeth on outer margin (Fig. |
5 |
– | Mandible without distinct teeth on outer margin (Fig. |
16 |
5 | Body length 1.9–2.2 mm | 6 |
– | Body length 2.4–2.6 mm | 14 |
6 | Claw tooth 0.50–0.53 times length of claw (Fig. |
7 |
– | Claw tooth 0.43–0.47 times length of claw | 11 |
7 | Claw tooth 0.53 times length of claw | S. udomi |
– | Claw tooth 0.50 times length of claw | 8 |
8 | Sensory vesicle 0.22–0.24 times length of third palpal segment (Fig. |
S. nanoiense sp. nov. |
– | Sensory vesicle 0.26–0.32 times length of third palpal segment | 9 |
9 | Labrum 0.6 times as long as clypeus | S. sutheppuiense sp. nov. |
– | Labrum 0.7 times as long as clypeus 10 | |
10 | Fifth palpal segment 2.6 times as long as third | S. loeiense sp. nov. |
– | Fifth palpal segment 2.1 times as long as third | S. maelanoiense sp. nov. |
11 | Sensory vesicle 0.24 times length of third palpal segment | S. puaense sp. nov. |
– | Sensory vesicle 0.26–0.35 times as long as third palpal segment | 12 |
12 | Sensory vesicle 0.32–0.35 times as long as third palpal segment | S. pitasawatae sp. nov. |
– | Sensory vesicle 0.26–0.29 times length of third palpal segment | 13 |
13 | Spermatheca 1.3 times as long as wide | S. huaimorense sp. nov. |
– | Spermatheca 1.6 times as long as wide | S. nanthaburiense sp. nov. |
14 | Labrum 0.54–0.56 times length of clypeus | S. asakoae |
– | Labrum 0.61–0.68 times length of clypeus | 15 |
15 | Claw tooth 0.50 times length of claw (Fig. |
S. thungchangense sp. nov. |
– | Claw tooth 0.47 times length of claw | S. chiangdaoense |
16 | Sensory vesicle 0.36 times as long as third palpal segment; claw tooth 0.56 times as long as claw | S. maewongense sp. nov. |
– | Sensory vesicle 0.25–0.32 times as long as third palpal segment; claw tooth 0.46–0.50 times as long as claw | 17 |
17 | Fore basitarsus 5.5 times as long as its greatest width | 18 |
– | Fore basitarsus 5.9–6.5 times as long as its greatest width | 19 |
18 | Labrum 0.52 times as long as clypeus | S. maehongsonense sp. nov. |
– | Labrum 0.65 times as long as clypeus | S. songense sp. nov. |
19 | Hind basitarsus 6.5 times as long as its greatest width | S. myanmarense |
– | Hind basitarsus 5.9–6.2 times as long as its greatest width | 20 |
20 | Frons: head ratio 1.0:4.7–5.5 | S. junkumae sp. nov. |
– | Frons: head ratio 1.0:4.0 | S. muangpanense sp. nov. |
Males
1 | Hair tuft of base of radial vein brownish | S. inthanonense |
– | Hair tuft of base of radial vein yellow | 2 |
2 | Upper-eye (large) facets vermilion | 3 |
– | Upper-eye (large) facets medium to dark brown | 4 |
3 | Upper-eye (large) facets in 9 or 10 vertical columns and 12 horizontal rows; fore basitarsus 8.4–8.7 times as long as its greatest width | S. thungchangense sp. nov. |
– | Upper-eye (large) facets in 11 or 12 vertical columns and 13 or 14 horizontal rows; fore basitarsus 7.1–7.4 times as long as its greatest width | S. asakoae |
4 | Antenna yellowish except few apical flagellomeres slightly to somewhat greyish | S. teerachanense sp. nov. |
– | Antenna brownish except scape, pedicel and base of first flagellomere yellowish | 5 |
5 | Hind basitarsus 0.9 time as wide as hind femur (Fig. |
6 |
– | Hind basitarsus 1.0–1.3 times as wide as hind femur (Fig. |
11 |
6 | Ventral plate with ventral margin rounded when viewed caudally (Fig. |
7 |
– | Ventral plate with ventral margin nearly straight medially when viewed caudally (Fig. |
9 |
7 | Fore basitarsus 6.4 times as long as its greatest width | S. banluangense sp. nov. |
– | Fore basitarsus 7.4–8.8 times as long as its greatest width | 8 |
8 | Wing length 1.6–1.7 mm | S. chaowaense sp. nov. |
– | Wing length 2.0 mm | S. myanmarense |
9 | Upper-eye (large) facets in 13 vertical columns | S. muangpanense sp. nov. |
– | Upper-eye(large) facets in 14–16 vertical columns | 10 |
10 | Upper-eye (large) facets in 14 vertical columns | S. klonglanense sp. nov. |
– | Upper-eye (large) facets in 15 or 16 vertical columns | S. namdangense sp. nov. |
11 | Upper-eye (large) facets in 9 or 10 vertical columns | S. puaense sp. nov. |
– | Upper-eye (large) facets in 11 or more vertical columns | 12 |
12. | Upper-eye (large) facets in 11 vertical columns | 13 |
– | Upper-eye (large) facets in 12–17 vertical columns | 16 |
13 | Head nearly as wide as thorax | 14 |
– | Head slightly wider than thorax | 15 |
14 | Ventral plate rounded ventrally when viewed caudally (Fig. |
S. maewongense sp. nov. |
– | Ventral plate with ventral margin nearly straight when viewed caudally (Fig. |
S. loeiense sp. nov. |
15 | First antennal flagellomere 1.8–1.9 times as long as second | S. sutheppuiense sp. nov. and S. maelanoiense sp. nov. |
– | First antennal flagellomere 1.5–1.6 times as long as second | S. phapeungense sp. nov. |
16 | Upper-eye (large) facets in 16 or 17 vertical columns | 17 |
– | Upper-eye (large) facets in 12–14 vertical columns | 19 |
17 | Subcosta bare or with two hairs; wing length 1.9–2.0 mm | S. huaimorense sp. nov. and S. kiewfinense sp. nov. |
– | Subcosta with 7–12 hairs; wing length 2.2–2.5 mm | 18 |
18 | Ventral plate with ventral margin nearly straight when viewed caudally (Fig. |
S. junkumae sp. nov. |
– | Ventral plate with ventral margin rounded when viewed caudally | S. rampae |
19 | Upper-eye (large) facets in 13 or 14 horizontal rows | 20 |
– | Upper-eye (large) facets in 15 or 16 horizontal rows | 22 |
20 | Ventral plate with ventral margin rounded when viewed caudally (Fig. |
S. nanoiense sp. nov. |
– | Ventral plate with ventral margin nearly straight when viewed caudally (Fig. |
21 |
21 | Upper-eye (large) facets in 12 vertical columns | S. nanthaburiense sp. nov. |
– | Upper-eye (large) facets in 13 vertical columns | S. maehongsonense sp. nov. |
22 | Ventral plate with ventral margin nearly straight when viewed caudally (Fig. |
S. pitasawatae sp. nov. |
– | Ventral plate with ventral margin rounded when viewed caudally | S. chiangdaoense and S. udomi |
Pupae
1 | Cocoon with elongate anterodorsal projection (2.0 mm or little longer) extended far beyond anteroventral tips of cocoon | 2 |
– | Cocoon with or without anterodorsal bulge or short anterodorsal projection (up to ca. 1.0 mm long) extended at most up to anteroventral tips of cocoon | 3 |
2 | Gill with six filaments | S. udomi |
– | Gill with eight filaments | S. chiangdaoense |
3 | Dorsum of abdominal segments 1 and 2 or 1–3 or 1–5 light to medium brown or light greyish brown | 4 |
– | Dorsum of abdominal segments 1 and 2 unpigmented or light yellowish | 5 |
4 | Dorsum of abdominal segments 1 and 2 or 1–3 light to medium brown; cocoon with short bulge | S. asakoae |
– | Dorsum of abdominal segments 1–5 light greyish brown; cocoon with short projection (Fig. |
S. inthanonense |
5 | Dorsum of abdominal segment 1 sparsely covered with minute tubercles | 6 |
– | Dorsum of abdominal segment 1 without minute tubercles | 7 |
6 | Outer margin of terminal hook 2.2 times length of inner margin (Fig. |
S. loeiense sp. nov. |
– | Outer margin of terminal hook 2.7 times length of inner margin (Fig. |
S. thungchangense sp. nov. |
7 | Cocoon with short anterodorsal projection (Figs |
8 |
– | Cocoon without anterodorsal projection | 9 |
8 | Primary stalks of dorsal and middle triplets of gill filaments much shorter than their common stalk (Fig. |
S. huaimorense sp. nov. |
– | Primary stalks of dorsal and middle triplets of gill filaments nearly as long as their common stalk (Fig. |
S. maewongense sp. nov. and S. myanmarense |
9 | Terminal hooks conical, with outer margin as long as or slightly longer than inner margin and not crenulated (Figs |
S. klonglanense sp. nov. and S. namdanense sp. nov. |
– | Terminal hooks widened, plate-like, with outer margin over 1.5 times length of inner margin, and crenulated | 10 |
10 | Stalk of ventral pair thicker than interspiracular trunk (Fig. |
S. banluangense sp. nov. |
– | Stalk of ventral pair thinner than interspiracular trunk | 11 |
11 | Gill arranged as 3+2+(2+1) or 2+2+(2+1) from dorsal to ventral, with long common basal stalk 1.3–1.5 times length of interspiracular trunk | S. rampae |
– | Gill arranged as (3+3)+2 from dorsal to ventral, with medium-long common basal stalk shorter than interspiracular rrunk | 12 |
12 | Outer margin of terminal hook 2.0–2.9 times as long as inner margin | S. chaowaense , sp. nov., S. junkumae sp. nov., S. maelanoiense sp. nov., S. muangpanense sp. nov., S. nanoiesne sp. nov., S. phapeungense sp. nov., S. puaense sp. nov., S. songense sp. nov., S. sutheppuiense sp. nov. and S. teerachanense sp. nov. |
– | Outer margin of terminal hook 3.3–4.0 times as long as inner margin | 13 |
13 | Filaments of ventral pair 2.1–2.3 mm long (Fig. |
S. maehongsonense sp. nov. |
– | Filaments of ventral pair 2.5–3.0 mm long (Fig. |
S. pitasawatae sp. nov. |
Larvae
1 | Abdominal segments 1–4 unicolorous | 2 |
– | Abdominal segments 1–4 not unicolorous | 12 |
2 | Abdominal segments 1–4 light ochreous (Fig. |
3 |
– | Abdominal segments 1–4 greyish or greenish grey or greenish | 5 |
3 | Postgenal cleft 2.8–3.3 times as long as postgenal bridge (Fig. |
S. teerachanense sp. nov. |
– | Postgenal cleft 1.2–2.4 times as long as postgenal bridge (Fig. |
4 |
4 | Labral fan with 23 or 24 primary rays | S. pitasawatae sp. nov. |
– | Labral fan with 29 or 30 primary rays | S. sutheppuiense sp. nov. |
5 | Pharate pupal gill with elongate common basal stalk | S. rampae |
– | Pharate pupal gill with medium-long common basal stalk | 6 |
6 | Postgenal cleft 1.5–4.0 times length of postgenal bridge | 7 |
– | Postgenal cleft 0.8–1.4 times length of postgenal bridge | 9 |
7 | Head spots moderately positive | S. asakoae |
– | Head spots indistinct or faintly positive | 8 |
8 | Postgenal cleft 2.5 times as long as postgenal bridge (Fig. |
S. namdanense sp. nov. |
– | Postgenal cleft 3.7–4.0 times as long as postgenal bridge (Fig. |
S. klonglanense sp. nov. |
9 | Abdominal segments 5–8 pinkish (Fig. |
S. thungchangense sp. nov. |
– | Abdominal color pattern otherwise | 10 |
10 | Abdominal segments 7 and 8 light grey but abdominal segments 5 and 6 unpigmented (Fig. |
S. huaimorense sp. nov. |
– | Abdominal color pattern otherwise | 11 |
11 | Abdominal segments 5, 7 and 8 grey but abdominal segment 6 unpigmented (Fig. |
S. loeiense sp. nov. |
– | Abdominal segments 5, 6, 7 and 8 greyish (Fig. |
S. chaowaense sp. nov. |
12 | Abdominal segment 1 grey but abdominal segments 2–4 ochreous (Fig. |
S. maehongsonense sp. nov. |
– | Abdominal color pattern otherwise | 13 |
13 | Abdominal segments 1 and 2 grey or green but abdominal segments 3 and 4 unpigmented | 14 |
– | Abdominal color pattern otherwise | 15 |
14 | Abdominal segments 7 and 8 grey with faint reddish brown pigment (Fig. |
S. myanmarense |
– | Abdominal segments 7 and 8 not grey with distinct reddish brown pigment (Fig. |
S. maewongense sp. nov. |
15 | Abdominal segments 1, 2 and 3 grey or greenish grey but abdominal segment 4 unpigmented or light ochreous (Fig. |
16 |
– | Abdominal color pattern otherwise | 18 |
16 | Abdominal segments 7 and 8 grey with reddish brown pigment (Fig. |
S. nanoiense sp. nov. |
– | Abdominal segments 7 and 8 light ochreous (Fig. |
17 |
17 | Abdominal segments 1, 2 and 3 greyish, and segments 7 and 8 with reddish brown pigment (Fig. |
S. junkumae sp. nov. |
– | Abdominal segments 1, 2 and 3 greenish grey but segments 7 and 8 lacking reddish brown pigment (Fig. |
S. inthanonense (morph 1) |
18 | Abdominal segments 1 and 3 greenish grey (Fig. |
S. inthanonense (morph 2) |
– | Abdominal color pattern otherwise | 19 |
19 | Abdominal segments 1, 3 and 4 grey but abdominal segment 2 light ochreous (Fig. |
S. phapeungense sp. nov. |
– | Abdominal color pattern otherwise | 20 |
20 | Abdominal segments 1 and 4 greyish but abdominal segments 2 and 3 light ochreous (Fig. |
S. muangpanense sp. nov. |
– | Abdominal segments 1–3 light green but abdominal segment 4 light ochreous (Fig. |
21 |
21 | Pharate pupal gill with six filament | S. udomi |
– | Pharate pupal gill with eight filaments | S. chiangdaoense |
The genetic relationships of all 26 species (exclusive of S. banluangense sp. nov.) of the S. asakoae species group in Thailand are shown in Fig.
Nucleotide differences (%) among subgroups of the Simulium asakoae species group based on COI sequences.
I | II | III | IV | V | VI | VII | VIII | IX | |
---|---|---|---|---|---|---|---|---|---|
I | 0–3.41 | ||||||||
II | 7.68–9.04 | 0–2.22 | |||||||
III | 7.34–9.39 | 5.97–7.68 | 0–2.39 | ||||||
IV | 6.66–9.22 | 6.31–7.85 | 5.46–7.17 | 0–5.29 | |||||
V | 6.66–7.85 | 5.63–6.31 | 5.12–5.97 | 5.12–6.14 | 0 | ||||
VI | 6.31–8.87 | 6.48–7.34 | 5.80–7.85 | 4.95–6.66 | 3.92–4.78 | 0–4.61 | |||
VII | 7.51–9.73 | 7.34–8.36 | 5.46–7.00 | 5.46–7.51 | 4.27–4.78 | 3.41–5.12 | 0–2.90 | ||
VIII | 7.51–9.73 | 6.48–8.87 | 5.80–6.83 | 5.29–6.83 | 4.27–5.12 | 4.10–5.12 | 2.90–4.27 | 0–3.41 | |
IX | 7.17–9.22 | 6.83–8.02 | 5.46–7.00 | 5.12–6.66 | 4.27–5.12 | 3.24–4.61 | 2.05–3.92 | 1.88–3.58 | 0–2.56 |
In subgroup I, two new species, S. klonglanense and S. namdanense, and two known species, S. lurauense and S. sofiani (
Subgroup I is an assemblage of species that is morphologically characterized by the combination of an elongate female sensory vesicle and small pupal terminal hooks.
Subgroup II is represented only by S. asakoae. A similar distinct genetic status of this species within the species group was also shown by
None of six samples from four localities of Thailand was identical to those of S. asakoae from Peninsular Malaysia. The differences between six samples from Thailand and S. asakoae from Peninsular Malaysia were 1.22–1.98 % and 1.37–2.22 %, based on the sequences of 685 bp and 586 bp, respectively. These differences are regarded as intraspecific variation because four Thai populations examined in this study are morphologically indistinguishable from S. asakoae from Peninsular Malaysia. Relatively high intraspecific variation is likely to reflect a wide range of its aquatic habitats from low to high elevations (196 m to 1,610 m) in Thailand.
In subgroup III, one new species, S. thungchangense, and three known species, S. chiangdaoense (of which the COI gene sequences were newly added in this study), S. rampae, and S. udomi (
In subgroup IV, nine new species and one known species, S. brinchangense from Peninsular Malaysia, are included. A high similarity of COI gene sequences of four new species, S. huaimorense, S. junkumae, S. kiewfinense, and S. maewongense, is observed. Notably, the first three of these new species are morphologically similar in sharing a higher number of male upper-eye large facets, suggesting that they have recently been derived from a common ancestor.
Subgroup V is represented only by S. inthanonense, which was formerly treated as a member of the S. ceylonicum species group (
In subgroup VI, S. sutheppuiense sp. nov. forms a close relationship with S. tanahrataense from Peninsular Malaysia, although both species differ morphologically from each other.
Subgroup VII is formed by two taxa, one being S. maelanoiense sp. nov. and the other being identified morphologically and genetically as S. myanmarense, originally described from Myanmar. Simulium myanmarense was suggested to exist in Thailand based on COI gene sequences of adult females (
In subgroup VIII, one new species, S. teerachanense, is included together with two known species from Peninsular Malaysia, S. roslihashimi and S. izuae. Simulium teerachanense sp. nov. is morphologically similar to S. roslihashimi in sharing the yellowish male antennae. It is demonstrated that both species are genetically close to each other, with a difference of 2.05–2.56%.
In subgroup IX, three new species, S. nanthaburiense, S. chaowaense, and S. muangpanense, and one known species, S. monglaense from Myanmar, are included. Simulium nanthaburiense sp. nov. is almost identical to one of two haplotypes (MF101845) of S. monglaense, although both species are morphologically distinguished from each other by the number of male upper-eye (large) facets that are in 12 vertical columns and 13 or 14 horizontal rows in S. nanthaburiense sp. nov. but in 15 vertical columns and 15 horizontal rows in S. monglaense (
Our morphological study reveals a high rate of species radiation of the S. asakoae species group in Thailand, as evidenced by the increased species number from four to 27. Most of these species are morphologically distinguished from one another as females, males, and larvae but not pupae, as shown in the keys. Ten of 27 species were indistinguishable in the pupal stage due to similarities of key characters such as the gill, terminal hooks and cocoon, the phenomenon being contrary to other species groups of the subgenus Gomphostilbia, in which species are usually identified in the pupal stage by their distinct expression of the number and arrangement of gill filaments and/or shape of the inflated structure of the gill (
A COI gene sequence-based analysis shows that all (but one species) in the S. asakoae species group in Thailand are divided into nine subgroups, of which subgroups II, V and VIII are represented by S. asakoae, S. inthanonense, and S. teerachanense sp. nov., respectively, and the other six subgroups are represented by two to nine species, suggesting a relatively recent radiation, accompanied by some morphological differentiation but few COI gene changes.
In the latter six subgroups, genetic differences between certain new species and known species are too small or even unrecognized, as in the cases of S. thungchangense sp. nov. versus S. udomi, S. nanthaburiense sp. nov. versus S. monglaense, and S. sutheppuiense sp. nov. versus S. tanahrataense. Certain members of the S. asakoae species group were previously reported to share the same or closely similar COI gene sequences, e.g., S. udomi versus S. rampae (
DNA barcoding is generally useful for resolving the phylogenetic relationships among species of most species groups of the subgenus Gomphostilbia and for uncovering cryptic species. However, caution is required when members of the S. asakoae species group are identified based only on similarities of the COI gene sequences.
We are grateful to Peter H. Adler (Professor Emeritus, Clemson University, Clemson, SC, U.S.A.) for reading the current manuscript and providing valuable comments. We would also like to thank all staff of the Section of Entomology of the Botanical Garden Organization for their kind help in working on black flies in the field and laboratory.
This research was funded by Anandamahidol Foundation (Year 2016–2019) and Chiang Mai University through the Center of Insect Vector Study to AS. Thanks are due to the Botanical Garden Organization, Chiang Mai, Thailand, for providing a research fund to WS. This work was partly supported by JSPS KAKENHI (Grant Number 19K06421) to MF.