Displaying publications 1 - 20 of 45 in total

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  1. Fulltext Tests of conspecificity for allopatric vectors: Simulium nodosum and Simulium shirakii (Diptera: Simuliidae) in Asia
    Low VL, Adler PH, Sofian-Azirun M, Srisuka W, Saeung A, Huang YT, et al.
    Parasit Vectors, 2015 May 29;8:297.
    PMID: 26022092 DOI: 10.1186/s13071-015-0911-5
    BACKGROUND: Allopatric populations present challenges for biologists working with vectors. We suggest that conspecificity can be concluded in these cases when data from four character sets-chromosomal, ecological, molecular, and morphological-express variation no greater between the allopatric populations than between corresponding sympatric populations. We use this approach to test the conspecificity of Simulium nodosum Puri on the mainland of Southeast Asia and Simulium shirakii Kono & Takahasi in Taiwan. The validity of these two putative species has long been disputed given that they are morphologically indistinguishable.

    FINDINGS: The mitochondria-encoded cytochrome c oxidase subunit I (COI), 12S rRNA, and 16S rRNA genes and the nuclear-encoded 28S rRNA gene support the conspecific status of S. nodosum from Myanmar, Thailand, and Vietnam and S. shirakii from Taiwan; 0 to 0.19 % genetic differences between the two taxa suggest intraspecific polymorphism. The banding patterns of the polytene chromosomes of the insular Taiwanese population of S. shirakii and mainland populations of S. nodosum are congruent. The overlapping ranges of habitat characteristics and hosts of S. nodosum and S. shirakii corroborate the chromosomal, molecular, and morphological data.

    CONCLUSIONS: Four independent sources of evidence (chromosomes, DNA, ecology, and morphology) support the conspecificity of S. nodosum and S. shirakii. We, therefore, synonymize S. shirakii with S. nodosum. This study provides a guide for applying the procedure of testing conspecificity to other sets of allopatric vectors.

  2. Fulltext Seasonal Filarial Infections and Their Black Fly Vectors in Chiang Mai Province, Northern Thailand
    Aupalee K, Saeung A, Srisuka W, Fukuda M, Streit A, Takaoka H
    Pathogens, 2020 Jun 25;9(6).
    PMID: 32630410 DOI: 10.3390/pathogens9060512
    The transmission of zoonotic filarial parasites by black flies has so far been reported in the Chiang Mai and Tak provinces, Thailand, and the bites of these infected black flies can cause a rare disease-human zoonotic onchocerciasis. However, species identification of the filarial parasites and their black fly vectors in the Chiang Mai province were previously only based on a morphotaxonomic analysis. In this study, a combined approach of morphotaxonomic and molecular analyses (mitochondrial cox1, 12S rRNA, and nuclear 18S rRNA (SSU HVR-I) genes) was used to clarify the natural filarial infections in female black flies collected by using human and swine baits from two study areas (Ban Lek and Ban Pang Dang) in the Chiang Mai province from March 2018 to January 2019. A total of 805 and 4597 adult females, belonging to seven and nine black fly taxa, were collected from Ban Lek and Ban Pang Dang, respectively. At Ban Lek, four of the 309 adult females of Simulium nigrogilvum were positive for Onchocerca species type I in the hot and rainy seasons. At Ban Pang Dang, five unknown filarial larvae (belonging to the same new species) were detected in Simulium sp. in the S. varicorne species-group and in three species in the S. asakoae species-group in all seasons, and three non-filarial larvae of three different taxa were also found in three females of the S. asakoae species-group. This study is the first to molecularly identify new filarial species and their vector black fly species in Thailand.
  3. DNA Barcoding of Simulium asakoae (Diptera: Simuliidae) From Northern Thailand
    Low VL, Srisuka W, Saeung A, Tan TK, Ya'cob Z, Yeong YS, et al.
    J Med Entomol, 2020 09 07;57(5):1675-1678.
    PMID: 32333022 DOI: 10.1093/jme/tjaa081
    Previous studies suggested the presence of species complex in the so-called Simulium asakoae Takaoka & Davies (Diptera: Simuliidae) in Thailand due to its high morphological variability and genetic divergence. To investigate whether the true S. asakoae is present in Thailand, we performed a detailed morphological identification of S. asakoae and compared its DNA barcodes with the morphospecies S. asakoae from Myanmar and the typical S. asakoae from Malaysia. Phylogenetic analysis revealed the Thai materials analyzed in this study were indeed genetically similar with those from Myanmar and Malaysia, though genetic distances 0-2.27% were observed. We tentatively regard this divergence as intraspecific variation, and the automatic barcode gap discovery analysis further supports them as a single species.
  4. Three new species of the Simulium (Simulium) striatum species-group (Diptera: Simuliidae) from Thailand, with their phylogenetic relationships
    Aupalee K, Saeung A, Srisuka W, Fukuda M, Junkum A, Pitasawat B, et al.
    Acta Trop, 2020 Nov;211:105625.
    PMID: 32649996 DOI: 10.1016/j.actatropica.2020.105625
    Three new species of black flies from Thailand, Simulium wangkwaiense, S. tadtonense and S. maeklongkeense, are described based on their adults, pupal exuviae and cocoons. All three new species are assigned to the Simulium (Simulium) striatum species-group, bringing its total number in Thailand to seven. Simulium wangkwaiense sp. nov. is the species formerly called S. quinquestriatum in Thailand. Certain male and pupal morphological characteristics are shown to separate all seven Thai species of this species-group. All of the three new species have been analyzed genetically for their phylogenetic relationships, with three known related species (except for S. thailandicum), by using the fast-evolving nuclear big zinc finger (BZF) gene.
  5. A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Northern Thailand, with its genetic relationship in the S. asakoae species-group
    Srisuka W, Aupalee K, Low VL, Yácob Z, Fukuda M, Saeung A, et al.
    Acta Trop, 2021 Jun;218:105889.
    PMID: 33722581 DOI: 10.1016/j.actatropica.2021.105889
    A new black fly species, Simulium (Gomphostilbia) kiewlomense, is described from females, males, pupae and mature larvae in Thailand. This new species is placed in the S. asakoae species-group and is characterized by having a combination of the elongate female sensory vesicle, widened male hind basitarsus, which is much wider than the hind femur, small pupal terminal hooks, and light greenish larval abdominal segments 1-3. Taxonomic notes are given to separate this new species from other related species. A DNA analysis using the COI gene shows that this new species has two genoforms with 1.21% difference. This is the 28th species of the S. asakoae species-group in Thailand, strengthening the evidence for high species diversity of this species-group.
  6. Fulltext Community Structure, Biodiversity and Spatiotemporal Distribution of the Black Flies (Diptera: Simuliidae) Using Malaise Traps on the Highest Mountain in Thailand
    Srisuka W, Sulin C, Aupalee K, Phankaen T, Taai K, Thongsahuan S, et al.
    Insects, 2021 May 31;12(6).
    PMID: 34072677 DOI: 10.3390/insects12060504
    Black flies form a group of small blood-sucking insects of medical and veterinary importance. This study aimed to investigate the community structure, biodiversity and spatial and temporal distribution of adult black flies in tropical rain forests, by using malaise traps in Doi Inthanon National Park, northern Thailand. Malaise traps were placed along six elevational gradients (400 m to 2500 m, above sea level) at Doi Inthanon National Park, Chiang Mai province, from December 2013 to November 2014. A total of 9406 adult female black flies belonging to five subgenera-Daviesellum (2%), Gomphostilbia (23%), Montisimulium (11%), Nevermannia (16%) and Simulium (48%)-were collected. Among 44 taxa found, S. tenebrosum complex had the highest relative abundance (11.1%), followed by the S. asakoae species-group (9.6%), the S. striatum species-group (7.7%), S. inthanonense (6.6%), S. doipuiense complex (6.4%), S. chomthongense complex (5.3%), S. chumpornense (5.1%) and S. nigrogilvum (4.1%). Two human-biting species-S. nigrogilvum and species in the S. asakoae species-group-were found in all of the collection sites with 100% species occurrence. Species richness was highest at mid elevation (1400 m), which is represented by 19 black fly species. The peak and lowest seasonal abundance was observed in the rainy and hot season, respectively. Seasonal species richness was highest in the cold season, except for that from elevation sites at 700 m, 1700 m and 2500 m. This study revealed that the malaise trap is effective in providing important data for further monitoring of the effects of environmental changes and conservation planning on the biodiversity of black flies in Doi Inthanon National Park.
  7. Descriptions of the female and larva of Simulium (Gomphostilbia) udomi (Diptera: Simuliidae) from Thailand, and its transfer to the Simulium asakoae species-group
    Saeung A, Srisuka W, Low VL, Maleewong W, Takaoka H
    Acta Trop, 2017 Aug;172:14-19.
    PMID: 28433572 DOI: 10.1016/j.actatropica.2017.04.014
    The female and larva of Simulium (Gomphostilbia) udomi Takaoka & Choochote from Thailand are described for the first time. The female of this species is similar to those of S. (G.) asakoae Takaoka & Davies from Peninsular Malaysia, Thailand, Hong Kong and Vietnam, and S. (G.) chiangdaoense Takaoka & Srisuka from Thailand. The larva of this species is similar to S. (G.) curtatum Jitklang et al. and S. (G.) nr. asakoae 2 from Thailand in having a medium-long postgenal cleft. Taxonomic notes are given to separate this species from these related species. The COI gene sequence of S. (G.) udomi is compared with those of eight species of the S. asakoae species-group and three species of the S. ceylonicum species-group. This species is transferred from the S. ceylonicum species-group to the S. asakoae species-group based on the adult female and male morphological characters, comparisons of the genetic distances and phylogenetic relationships inferred from the COI gene sequences.
  8. Natural infections with larvae of Onchocerca species type I in the human-biting black fly, Simulium nigrogilvum (Diptera: Simuliidae), in western Thailand
    Saeung A, Srisuka W, Aupalee K, Fukuda M, Otsuka Y, Taai K, et al.
    Acta Trop, 2020 Apr;204:105344.
    PMID: 31954685 DOI: 10.1016/j.actatropica.2020.105344
    Zoonotic onchocerciasis is a human infection caused by Onchocerca species of animal origins and transmitted by black fly vectors. The reported incidence of this disease has increased throughout the world. This study aims to clarify the vectorial roles of black fly species in zoonotic filarial transmission in Tak province, western Thailand. The integrated approach of morphological and DNA sequence-based analyses was used to identify species of both wild-caught female black flies and infective filarial larvae found in the infected black flies. All of 494 female black flies captured were identified as Simulium nigrogilvum, through scanning electron microscopy (SEM) and DNA sequence analyses based on the cytochrome c oxidase subunit I (COI) and subunit II (COII), and the fast-evolving nuclear elongation complex protein 1 (ECP1) genes. Four females of S. nigrogilvum harbored one to three third-stage larvae (infective larvae) in their thoraces, with an infection rate of 0.81% (4/494). All infective larvae were similar in morphology and size to one another, being identified as Onchocerca species type I (= O. sp. type A), a bovine filaria, originally reported from Japan, and also as O. sp. found in S. nodosum in Thailand, based on their body lengths and widths being 1,068-1,346 µm long by 25-28 µm wide, and morphological characters. Comparisons of cytochrome c oxidase subunit I (COI) and 12S rRNA sequences of mitochondrial DNA (mtDNA) and phylogenetic analyses with those of previous reports strongly supported that all larvae were O. sp. type I. This report is the first indicating the presence of O. sp. type I in Thailand and its vector being S. nigrogilvum.
  9. Morphological description and phylogenetic position of a new cryptic species in the Simulium asakoae species-group, Simulium (Gomphostilbia) mokroense, from Western Thailand
    Srisuka W, Aupalee K, Fukuda M, Saeung A, Takaoka H
    Acta Trop, 2022 Jan 14;228:106313.
    PMID: 35038426 DOI: 10.1016/j.actatropica.2022.106313
    Adult females and males reared from pupae of Simulium sp. from Tak Province and those of S. chiangdaoense Takaoka & Srisuka from four localities in Chiang Mai and Lampang Provinces, northern Thailand, were molecularly and morphologically compared. Simulium sp. is morphologically almost indistinguishable from S. chiangdaoense except the hair tuft at the base of the radius of the adult female and male, which is composed of yellow and dark hairs (in place of yellow hairs only). Molecular analysis using COI gene sequences shows that S. sp. formed a distinct clade and was separated from S. chiangdaoense by a genetic distance of 1.56-2.44%. Based on the results of morphological and molecular analyzes, S. sp. is described as a new species, S. mokroense, from females, males, pupae and mature larvae. It is also found that S. chiangdaoense is genetically diverse, with five lineages, and is morphologically variable in the number of male upper-eye (large) facets, which are in 13 to 17 vertical columns and 14 to 17 horizontal rows on each side. This is a first case, in which the difference in the color of the wing tuft hairs is a clue leading to the discovery of a new cryptic species close to S. chiangdaoense.
  10. Fulltext Seasonal biodiversity of black flies (Diptera: Simuliidae) and evaluation of ecological factors influencing species distribution at Doi Pha Hom Pok National Park, Thailand
    Srisuka W, Takaoka H, Otsuka Y, Fukuda M, Thongsahuan S, Taai K, et al.
    Acta Trop, 2015 Sep;149:212-9.
    PMID: 26028177 DOI: 10.1016/j.actatropica.2015.05.024
    This is the first study on the seasonal biodiversity of black flies and evaluation of ecological factors influencing their distribution at Doi Pha Hom Pok National Park, northern Thailand. Larvae were collected from six fixed-stream sites in relation to altitude gradients from May 2011 to April 2013. The water temperature, water pH, conductivity, total dissolved solids (TDS), salt, water velocity, stream width and depth, streambed particle sizes, riparian vegetation, and canopy cover were recorded from each site. Monthly collections from the six sites yielded 5475 last-instar larvae, belonging to 29 black fly species. The most frequently found species from all sites were Simulium asakoae (100%) followed by Simulium yuphae (83.3%), and Simulium chiangdaoense, Simulium gombakense, Simulium phahompokense, Simulium fruticosum, Simulium maeaiense and Simulium fenestratum (66.6%). Of the 5475 last-instar larvae, S. maeaiense (19.3%), S. chiangdaoense (15.8%) and S. asakoae (14.8%), were the three most abundant species. The Shannon diversity index (H) at the six sites with different altitudes of 2100m, 2000m, 1500m, 1400m, 700m, and 500m above mean sea level, were 2.042, 1.832, 2.158, 2.123, 1.821 and 1.822, respectively. The Shannon index and number of taxa in the cold season were higher than those in the rainy and hot seasons. Principal component analysis (PCA) indicated that at least three principal components have eigen values >1.0 and accounted for 93.5% of the total variability of ecological factors among sampling sites. The Canonical correspondence analyses (CCA) showed that most species had a trend towards altitude, canopy cover, riparian vegetation and water velocity.
  11. A new species of the Simulium (Simulium) variegatum species-group (Diptera: Simuliidae) from Thailand
    Takaoka H, Srisuka W, Saeung A
    Acta Trop, 2019 Mar;191:155-161.
    PMID: 30611713 DOI: 10.1016/j.actatropica.2019.01.002
    A new species of black fly, Simulium (Simulium) luculentum, is described from specimens collected in Thailand, and is assigned to the Simulium variegatum species-group of the subgenus Simulium Latreille. This new species is distinguished from most members of this species group in the female by the mid femur and tibia entirely yellowish; and in the pupa by the gill with six filaments, which are somewhat inflated basally. This species represents the fifth member of the S. variegatum species-group recorded from Thailand.
  12. A new species of the Simulium (Gomphostilbia) ceylonicum species-group (Diptera: Simuliidae) from Thailand
    Takaoka H, Srisuka W, Ya'cob Z, Low VL, Saeung A
    Acta Trop, 2021 May;217:105865.
    PMID: 33607063 DOI: 10.1016/j.actatropica.2021.105865
    A new black fly species, Simulium (Gomphostilbia) pangsidaense, is described on the basis of adult female, male, pupal exuviae and mature larvae from Pang Sida National Park, Sa Kaew Province, Eastern Thailand. This new species is placed in the Simulium ceylonicum species-group. It is distinguished from three Thai members of the S. ceylonicum species-group by the following characteristics: from S. (G.) curtatum Jitklang et al. and S. (G.) sheilae Takaoka & Davies by the wide pupal terminal hooks (triangular terminal hooks in the latter two species), and from S. (G.) sheilae and S. (G.) trangense Jitklang et al. by the number of male upper-eye facets in 13 vertical columns and 14 or 15 horizontal rows (10 or 11 vertical columns and 12 or 13 horizontal rows in the latter two species). This is the fourth member of the S. ceylonicum species-group recorded from Thailand.
  13. Fulltext Description of male, pupa and larva of Simulium (Asiosimulium) wanchaii (Diptera: Simuliidae) from Thailand, with keys to identify four species of the subgenus Asiosimulium
    Srisuka W, Takaoka H, Saeung A
    Trop Biomed, 2015 Sep;32(3):504-10.
    PMID: 26695212 MyJurnal
    The male, pupa and mature larva of Simulium (Asiosimulium) wanchaii Takaoka & Choochote, one of the four species of the small Oriental black fly subgenus Asiosimulium, are described for the first time based on samples collected from Thailand. The male S. (A.) wanchaii is characterized based on the enlarged hind basitarsus and the ventral plate which is much wider than long. The pupa and larva are characterized by the gill with 19 filaments and the deep postgenal cleft, respectively. Keys are provided to identify all the four species of the subgenus Asiosimulium for females, males, pupae and mature larvae.
  14. Description of the female of Simulium chayamaritae (Diptera: Simuliidae) from Thailand
    Takaoka H, Srisuka W, Saeung A
    Trop Biomed, 2018 Dec 01;35(4):975-980.
    PMID: 33601845
    Simulium chayamaritae Takaoka and Srisuka from Thailand belongs to the Simulium darjeelingense species-group of Simulium (Simulium) (Diptera: Simuliidae). The female of this species is described for the first time based on a female reared from a pupa collected from Chiang Mai, Thailand. It is characterized by the sensory vesicle elongate and the inner margins of the arms of the genital fork divergent, then convergent apically. It is similar to the female of S. eshimai Takaoka and Adler of the same speciesgroup from Vietnam. Taxonomic notes are given to separate it from two other species of the S. darjeelingense species-group from India and Malaysia, and 28 of 31 other species of the subgenus Gomphostilbia recorded from Thailand.
  15. A method for distinguishing the important malaria vectors Anopheles dirus and An. cracens (Diptera: Culicidae) based on antennal sensilla of adult females
    Taai K, Harbach RE, Somboon P, Sriwichai P, Aupalee K, Srisuka W, et al.
    Trop Biomed, 2019 Dec 01;36(4):926-937.
    PMID: 33597464
    Some species of the Anopheles dirus species complex are considered to be highly competent malaria vectors in Southeast Asia. Anopheles dirus is the primary vector of Plasmodium falciparum and P. vivax while An. cracens is the main vector of P. knowlesi. However, these two species are difficult to distinguish and identify based on morphological characters. Hence, the aim of this study was to investigate the potential use of antennal sensilla to distinguish them. Large sensilla coeloconica borne on the antennae of adult females were counted under a compound light microscope and the different types of antennal sensilla were examined in a scanning electron microscope. The antennae of both species bear five types of sensilla: ampullacea, basiconica, chaetica, coeloconica and trichodea. Observations revealed that the mean numbers of large sensilla coeloconica on antennal flagellomeres 2, 3, 7, 10 and 12 on both antennae of both species were significantly different. This study is the first to describe the types of antennal sensilla and to discover the usefulness of the large coeloconic sensilla for distinguishing the two species. The discovery provides a simple, reliable and inexpensive method for distinguishing them.
  16. A New Human-Biting Black Fly Species of Simulium (Simulium) (Diptera: Simuliidae) From Thailand
    Takaoka H, Srisuka W, Saeung A
    J Med Entomol, 2017 Jul 01;54(4):945-948.
    PMID: 28399286 DOI: 10.1093/jme/tjx064
    Simulium (Simulium) umphangense, a new human-biting species of black fly, is described based on females captured while attacking humans in western and central Thailand. The female of this new species is similar to those of S. (S.) indicum Becher, S. (S.) nigrogilvum Summers, and S. (S.) vanellum Huang et al. in the Simulium griseifrons species-group of the subgenus Simulium Latreille in having the frons densely covered with golden-yellow short hairs, a character rarely found in the subgenus Simulium. This new species is readily distinguished from the three related species by having a darkened fore tibia, a bare subcosta (or rarely with a few hairs), and an ovipositor valve not protruded posteriorly. Additional diagnostic characters of this new species are noted. This is the seventh human-biting species of black fly in Thailand.
  17. A new species and a new record of the Simulium (Gomphostilbia) gombakense species-group (Diptera: Simuliidae) from Thailand
    Takaoka H, Srisuka W, Low VL, Saeung A
    Acta Trop, 2018 Sep;185:156-166.
    PMID: 29772215 DOI: 10.1016/j.actatropica.2018.05.011
    Larvae and adults reared from pupae of Simulium (Gomphostilbia) gombakense Takaoka & Davies from eight streams in five provinces of Thailand were genetically analyzed by using cytochrome c oxidase subunit 1 sequences. The material was composed of four lineages, of which lineages 2, 3 and 4 are genetically closely related to one another, but more distantly related to lineage 1, which is equivalent to typical S. (G.) gombakense from Peninsular Malaysia. Lineages 2, 3 and 4, which are morphologically indistinguishable, are designated as genoforms A, B and C within a species complex of S. (G.) paiense sp. nov., which is described based on specimens of lineage 2 (=genoform A). Simulium (G.) paiense sp. nov. is similar to S. (G.) gombakense from Peninsular Malaysia but appears to be barely distinguished by the relative length of the female fore basitarsus. Simulium (G.) thuathienense Takaoka & Sofian-Azirun is newly recorded from Thailand and its male is described for the first time. The female of S. (G.) gombakense is briefly described based on females reared from pupae collected from Peninsular Malaysia.
  18. Two New Species of the Simulium (Simulium) variegatum Species-Group of Black Flies (Diptera: Simuliidae) From Thailand
    Takaoka H, Srisuka W, Saeung A
    J Med Entomol, 2017 09 01;54(5):1213-1223.
    PMID: 28505301 DOI: 10.1093/jme/tjx080
    Two new species of black flies, Simulium (Simulium) srisukai and S. (S.) kiewmaepanense, are described from specimens collected in Thailand. Both species are assigned to the Simulium variegatum species-group of the subgenus Simulium (Simulium) Latreille. They are characterized by the darkened female femora and tibiae and six inflated pupal gill filaments, and the darkened female tibiae and six ordinary thread-like pupal gill filaments, respectively. Taxonomic notes are given to separate these new species from 10 related species among the group. These new species represent the third and fourth species of the S. variegatum species-group from Thailand.
  19. Two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Myanmar, and their phylogenetic relationships with related species in the S. asakoae species-group
    Takaoka H, Srisuka W, Low VL, Maleewong W, Saeung A
    Acta Trop, 2017 Dec;176:39-50.
    PMID: 28746824 DOI: 10.1016/j.actatropica.2017.07.024
    Two new species of Simulium (Gomphostilbia), S. (G.) myanmarense and S. (G.) monglaense, are described from females, males, pupae and larvae from Myanmar. The two new species are placed in the S. asakoae species-group, and are similar to each other in the female and male but distinguished in the pupa by the presence or absence of an anterodorsal projection of the cocoon, and in the larva by a unique pattern of colored markings on the abdomen. Taxonomic notes are given to separate these species from related species. The COI gene sequences of both species are compared with those of eight species of the S. asakoae species-group and three species of the S. ceylonicum species-group. Both new species are most closely related to each other, further supporting their morphological classification in the S. asakoae species-group.
  20. Morphological and Molecular Evidence for Multiple Taxa in Simulium parahiyangum (Diptera: Simuliidae), a Geographic Generalist Black Fly in Southeast Asia
    Takaoka H, Low VL, Srisuka W, Ya'cob Z, Saeung A
    J Med Entomol, 2018 Oct 25;55(6):1453-1463.
    PMID: 30060220 DOI: 10.1093/jme/tjy109
    Seven populations of Simulium parahiyangum Takaoka & Sigit (Diptera: Simuliidae), a geographically widespread nominal species of black fly in Southeast Asia, were morphologically and molecularly studied. Three morphoforms based on male and pupal morphological features, and two primary lineages based on the COI gene sequence analysis were recognized. Morphoform 1 (lineage 1) from Sarawak, Malaysia, is identified as S. parahiyangum sensu stricto and morphoform 2 (lineage 2) from Thailand and Vietnam, and morphoform 3 (lineage 1) from Peninsular Malaysia are each regarded as distinct species, although morphoform 3 is partially homosequential for the COI gene with morphoform 1. Morphoforms 2 and 3 are described as Simulium ngaoense sp. nov. and Simulium sazalyi sp. nov., respectively. Overall, S. parahiyangum is not a single geographic generalist but a composite of multiple species.
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