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Simulium ledangense, a new species of the Simulium feuerborni species-group of the subgenus Nevermannia (Diptera: Simuliidae) from Mount Ledang, Peninsular Malaysia

Ya'cob Z, Takaoka H, Sofian-Azirun M

Zootaxa, 2014;3881(3):228-36.
PMID: 25543632 DOI: 10.11646/zootaxa.3881.3.2

Simulium (Nevermannia) ledangense sp. nov. is described from females, males, pupae and mature larvae from Peninsular Malaysia. This new species is assigned to the Simulium feuerborni species-group of the subgenus Nevermannia, and is characterized by the pupa having a very long stalk of the ventral paired gill filaments, which is almost five times longer than the interspiracular trunk and female tergites of segments 2 and 5 to 7 shiny. Taxonomic notes are given to distinguish this new species from three known species of the S. feuerborni species-group from Malaysia.
Simulium hiroyukii, a new species of the subgenus Gomphostilbia (Diptera: Simuliidae) from Mount Murud, Sarawak, Malaysia

Ya'cob Z, Takaoka H, Sofian-Azirun M

Zootaxa, 2015;3911(3):424-32.
PMID: 25661622 DOI: 10.11646/zootaxa.3911.3.8

Simulium (Gomphostilbia) hiroyukii is described based on females, males, pupae and larvae collected in Mount Murud, Sarawak, Malaysia. This new species is assigned to the Simulium darjeelingense species-group of the subgenus Gomphostilbia, and is characterized by the darkened fore coxae and the pupal gill with eight long filaments, of which middle and dorsal triplets have elongated primary and secondary stalks, respectively.
Breeding habitat preference of preimaginal black flies (Diptera: Simuliidae) in Peninsular Malaysia

Ya'cob Z, Takaoka H, Pramual P, Low VL, Sofian-Azirun M

Acta Trop, 2016 Jan;153:57-63.
PMID: 26476394 DOI: 10.1016/j.actatropica.2015.10.007

To investigate the breeding habitat preference of black flies, a comprehensive black fly survey was conducted for the first time in Peninsular Malaysia. Preimaginal black flies (pupae and larvae) were collected manually from 180 stream points encompassing northern, southern, central and east coast of the Peninsular Malaysia. A total of 47 black fly species were recorded in this study. The predominant species were Simulium trangense (36.7%) and Simulium angulistylum (33.3%). Relatively common species were Simulium cheongi (29.4%), Simulium tani (25.6%), Simulium nobile (16.2%), Simulium sheilae (14.5%) and Simulium bishopi (10.6%). Principal Component Analysis (PCA) of all stream variables revealed four PCs that accounted for 69.3% of the total intersite variance. Regression analysis revealed that high species richness is associated with larger, deeper, faster and higher discharge streams with larger streambed particles, more riparian vegetation and low pH (F=22.7, d.f.=1, 173; P<0.001). Relationship between species occurrence of seven common species (present in >10% of the sampling sites) was assessed. Forward logistic regression analysis indicated that four species were significantly related to the stream variables. S. nobile and S. tani prefer large, fast flowing streams with higher pH, large streambed particles and riparian trees. S. bishopi was commonly found at high elevation with cooler stream, low conductivity, higher conductivity and more riparian trees. In contrast, S. sheilae was negatively correlated with PC-2, thus, this species commonly found at low elevation, warmer stream with low conductivity and less riparian trees. The results of this study are consistent with previous studies from other geographic regions, which indicated that both physical and chemical stream conditions are the key factors for black fly ecology.
Sexually anomalous individuals of the black fly Simulium trangense (Diptera: Simuliidae) infected with mermithid parasites (Nematoda: Mermithidae)

Ya'cob Z, Low VL, Tan TK, Noor-Izwan A, Lourdes EY, Ramli R, et al.

Parasitol Res, 2021 May;120(5):1555-1561.
PMID: 33655351 DOI: 10.1007/s00436-021-07087-x

Sexually anomalous individuals, typically intersexes or gynandromorphs, bear a mixture of male and female traits. Twelve sexually anomalous individuals of the black fly Simulium (Gomphostilbia) trangense Jitklang, Kuvangkadilok, Baimai, Takaoka & Adler were discovered among 49 adults reared from pupae. All 12 sexually anomalous adults were parasitized by mermithid nematodes, although five additional parasitized adults had no overt external anomalies. Sequence analysis of the 18S rRNA gene revealed that the mermithids, possibly representing a new species, are related to Mesomermis spp., with genetic distances of 5.09-6.87%. All 12 anomalous individuals had female phenotypical traits on the head, thorax, forelegs, midlegs, and claws, but male features on the left and right hind basitarsi. One individual had mixed male and female genitalia. The findings are in accord with the trend that mermithid infections are associated with sexually anomalous adult black flies.
Characterization of Simulium (Simulium) hackeri Edwards (Diptera: Simuliidae) from Malaysia: Morphological description of the pupa and larva, and DNA barcoding

Ya'cob Z, Takaoka H, Low VL, Sofian-Azirun M

Acta Trop, 2018 Sep;185:110-114.
PMID: 29709632 DOI: 10.1016/j.actatropica.2018.04.029

Simulium (Simulium) hackeri Edwards, 1928 of the Simulium variegatum species-group from Malaysia was described initially based on the female specimen from Cameron Highlands, Pahang. In the present study, the pupa and larva of this species are described for the first time. Their morphological characters resemble those of the Simulium variegatum species-group by having six gill filaments per side, abdomen with dorsal spine-combs at least on segments 7 and 8, cocoon with wall-pocket shaped and with or without an anterodorsal projection. Postgenal cleft of the larva medium-sized, rarely small, ventral papillae small or absent. The DNA barcode of this species is also reported herein.
Uncovering the mask of the Simulium feuerborni complex (Diptera: Simuliidae): Description of a new pseudocryptic species Simulium pairoti from Malaysia

Ya'cob Z, Takaoka H, Low VL, Sofian-Azirun M

Acta Trop, 2017 May;169:133-141.
PMID: 28185824 DOI: 10.1016/j.actatropica.2017.02.005

A black fly species of the Simulium feuerborni species-group of Simulium (Nevermannia) from Cameron Highland, Peninsular Malaysia, previously regarded as S. feuerborni Edwards, originally described from East Java, is described as Simulium pairoti sp. nov. based on complete life stages. High intraspecific variations in the arrangement of the six pupal gill filaments, length of the stalk of the ventral paired filaments, and length of the anterodorsal projection of the cocoon, are noted in this species. This new species is readily distinguished from its congeners by having the characters of male genitalia with simple lamellate ventral plate, short inwardly-twisted styles, several parameral hooks, and a simple narrow median sclerite. Morphological data reported herein plus the chromosomal and molecular data presented elsewhere support S. pairoti as a novel pseudocryptic species.
First description of a new cryptic species, Simulium vanluni from Peninsular Malaysia: An integrated morpho-taxonomical and genetic approach for naming cryptic species in the family Simuliidae

Ya'cob Z, Takaoka H, Low VL, Sofian-Azirun M

Acta Trop, 2017 Mar;167:31-39.
PMID: 27986545 DOI: 10.1016/j.actatropica.2016.12.009

In recent decades, the numbers of cryptic taxa have increased significantly with current progress in DNA barcoding, yet, most of these cryptic taxa have not been formally named and recognized as valid species. To address this issue, we provide a guide for applying the procedure of describing new cryptic species in the family Simuliidae. Simulium (Simulium) vanluni from Pahang, Peninsular Malaysia, previously treated as S. nobile De Meijere, is described as a new species by using an integrated morpho-taxonomical and genetic approach. This new species is morphologically identical to S. nobile from Java and S. kiuliense Smart & Clifford from Borneo, but their distinctiveness is supported by an expanded multigene phylogeny analysis.
Description of the female of Simulium (Gomphostilbia) aziruni (Diptera: Simuliidae) and its genetic relationships with members of the Simulium gombakense species-group

Ya'cob Z, Takaoka H, Low VL, Tan TK, Sofian-Azirun M

Acta Trop, 2019 May;193:66-70.
PMID: 30807749 DOI: 10.1016/j.actatropica.2019.02.023

Simulium (Gomphostilbia) aziruni Takaoka, Hashim & Chen was described initially based only on a pupa and a mature larva collected from Peninsular Malaysia. Herein, we describe the morphological characters of the female of S. aziruni for the first time. It resembles those of the other members of the Simulium gombakense species-group by the genital fork with a distinct projection directed medioposteriorly from each arm and claw with a large basal tooth. Cytochrome c oxidase I (COI) barcoding analysis indicates that S. aziruni is the sister species of S. maleewongae, but both are distantly separated by a genetic distance of 4.9%.
A new species of Simulium (Simulium) (Diptera: Simuliidae) from Genting Highlands, Malaysia

Ya'cob Z, Takaoka H, Low VL, Sofian-Azirun M

Acta Trop, 2018 Jun;182:1-3.
PMID: 29453951 DOI: 10.1016/j.actatropica.2018.02.007

Simulium (Simulium) rasuli sp. nov. is described from two females collected by a Malaise trap in Genting Highlands, Peninsular Malaysia. This new species is placed in the Simulium christophersi species-group of the subgenus Simulium. The female of this new specie is characterized by the scutum with three longitudinal vittae, dark legs, claw with a small subbasal tooth, and ovipositor valve triangular with its inner margin nearly straight. This new species is distinguished in the female from all the six named species of the species-group by the entirely brownish-black femora and tibiae.
Fulltext Distribution pattern of black fly (Diptera: Simuliidae) assemblages along an altitudinal gradient in Peninsular Malaysia

Ya'cob Z, Takaoka H, Pramual P, Low VL, Sofian-Azirun M

Parasit Vectors, 2016 Apr 19;9:219.
PMID: 27094088 DOI: 10.1186/s13071-016-1492-7

BACKGROUND: Preimaginal black flies (Diptera: Simuliidae) are important components of the stream ecosystem. However, there has been limited research undertaken on the vertical distribution of preimaginal black flies and their associated ecological factors. Stream conditions are generally variable along the altitudinal gradient. Therefore, we conducted an in-depth entomological survey to investigate the simuliid distribution pattern along an altitudinal gradient in Peninsular Malaysia.
METHODS: A total of 432 collections were performed in this study (24 samplings at each of 18 fixed-streams at monthly intervals) from February 2012 to January 2014. Larvae and pupae attached on aquatic substrates such as grasses, leaves and stems, twigs, plant roots and rocks were collected by hand using fine forceps. Stream depth (m), width (m), velocity (m/s), water temperature (°C), acidity (pH), conductivity (mS/cm) and dissolved oxygen (mg/L) were measured at the time of each collection.
RESULTS: A total of 35 black fly species were recorded in the present study. The most frequently collected species were Simulium tani (31.7%) and S. whartoni (21.5%), while the relatively common species were Simulium sp. (nr. feuerborni) (16.2%), S. decuplum (15.5%), S. angulistylum (14.8%), S. bishopi (13.2%) and S. izuae (11.8%). Total estimated species richness ranged between 39.8 and 41.3, which yielded more than 80% of sampling efficiency. Six simuliid species were distributed below 500 m, whereas eight species were distributed above 1400 m. Simulium sp. (nr. feuerborni) and S. asakoae were found from middle to high altitudes (711-1813 m). Simulium whartoni, S. brevipar and S. bishopi were distributed widely from low to high altitudes (159-1813 m). Regression analysis between species richness and PCs revealed that the species richness was significantly associated with wider, deeper and faster streams at low altitude, normal water temperature (23-25 °C), low conductivity, higher discharge, more canopy cover and riparian vegetation and with larger streambed particles (F = 20.8, df = 1, 422, P
New species and records of black flies (Diptera: Simuliidae) from Vinh Phuc Province, Vietnam

Takaoka H, Sofian-Azirun M, Ya'cob Z, Chen CD, Lau KW, Pham HT

Zootaxa, 2014;3838(3):347-66.
PMID: 25081781 DOI: 10.11646/zootaxa.3838.3.6

Four new species of black flies are described, and three others are reported as newly recorded, based on adults reared from pupae, pupae and larvae collected in and near Tam Dao National Park, Vinh Phuc Province, Vietnam. New species include Simulium (Gomphostilbia) hongthaii sp. nov., S. (G.) tamdaoense sp. nov. (both species placed in the asakoae species-group), S. (Simulium) taythienense sp. nov. and S. (S.) xuandai sp. nov. (the two latter species placed in the striatum species-group). Newly recorded species are S. (G.) brinchangense Takaoka, Sofian-Azirun & Hashim, S. (Nevermannia) aureohirtum Brunetti and S. (S.) brevipar Takaoka & Davies. These discoveries increase the number of species of black flies known in Vietnam from 21 to 28.
Three new species of Simulium (Nevermannia) (Diptera: Simuliidae) from Vietnam

Takaoka H, Sofian-Azirun M, Ya'cob Z, Chen CD, Lau KW, Pham XD

Zootaxa, 2014;3866(4):555-71.
PMID: 25283675 DOI: 10.11646/zootaxa.3866.4.6

Three new species of black flies, Simulium (Nevermannia) langbiangense, S. (N.) phami, and S. (N.) bachmaense, are described on the basis of females, males, pupae and larvae collected in Vietnam. All three species are assigned to the Simulium feuerborni species-group of Simulium (Nevermannia). Simulium (N.) langbiangense sp. nov. is characterized by the female sensory vesicle with a large opening, and a short common basal stalk of the six pupal gill filaments, S. (N.) phami sp. nov. is most striking in having the pupal gill with five filaments, a character not reported in species of the S. feuerborni species-group, and S. (N.) bachmaense sp. nov. is characterized by the female genital fork with a triangular lobe-like projection pointed posteromedially on each arm, and the small larval postgenal cleft. This represents the first record of the S. feuerborni species-group from Vietnam.
Two new species of the Simulium batoense species-group of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Peninsular Malaysia

Takaoka H, Sofian-Azirun M, Ya'cob Z

Zootaxa, 2014;3774:473-80.
PMID: 24871514 DOI: 10.11646/zootaxa.3774.5.5

Two new black fly species, Simulium (Gomphostilbia) tekamense and Simulium (Gomphostilbia) jerantutense, are described based on adult females emerged from pupae in Peninsular Malaysia, and assigned to the binuanense subgroup of the batoense species-group in the subgenus Gomphostilbia. Simulium (G.) tekamense sp. nov. is characterized in the female by the subcosta with 0-2 hairs, and presence of a deep notch on the apex of the mediolongitudinal ridge of the cibarium, and in the pupa by one of two paired gill filaments of the middle triplet much thicker than the counter filament. Simulium (G.) jerantutense sp. nov. is characterized in the female by the short claw tooth 0.46 times the length of the claw, and in the pupa by the gill filaments arranged as [2+1+(1+2)]+2 filaments from dorsal to ventral. Taxonomic notes are given to distinguish these new species from related species.
A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Malaysia, with keys to 32 species of the Simulium ceylonicum species-group

Takaoka H, Sofian-Azirun M, Ya'cob Z, Hashim R, Otsuka Y

J Med Entomol, 2014 May;51(3):517-28.
PMID: 24897845

Simulium (Gomphostilbia) leparense sp. nov. is described from females, males, and pupae collected from Peninsular Malaysia. This new species is assigned to the ceylonicum species-group of the subgenus Gomphostilbia, and is characterized by the female and male scuta covered with dark-brown short hairs, smaller number of male upper-eye facets, presence of shiny paired spots on the male abdominal segments 2-8, and absence of grapnel-shaped hooklets on the pupal abdominal segment 9. The male and pupa of S. capillatum Takaoka, which was originally described from larvae collected from Sarawak and Sabah, are described for the first time. Keys to identify all 32 species of the Simulium ceylonicum species-group including 27 species from other countries are provided for females, males, pupae, and mature larvae.
Two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Cameron's Highlands, Peninsular Malaysia, with keys to 21 species of the Simulium asakoae species-group

Takaoka H, Sofian-Azirun M, Ya'cob Z, Hashim R

Zootaxa, 2014;3765:54-68.
PMID: 24870884 DOI: 10.11646/zootaxa.3765.1.3

Two new black fly species, Simulium (Gomphostilbia) brinchangense and S. (G.) tanahrataense, are described on the basis of reared adult females, males, pupae and larvae from Cameron's Highlands, Peninsular Malaysia. These new species are assigned to the asakoae species-group within Simulium (Gomphostilbia) and taxonomic notes are given to distinguish each new species from six known species in Malaysia. Revised keys to identify all 21 species including 13 species from other countries are provided for females, males, pupae and mature larvae. The species diversity of the asakoae species-group in Cameron's Highlands is briefly noted.
Two new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from peninsular Malaysia, with keys to 10 peninsular Malaysian species of the Simulium batoense species group

Takaoka H, Sofian-Azirun M, Ya'Cob Z

J Med Entomol, 2014 Jan;51(1):10-26.
PMID: 24605448

Two new blackfly species, Simulium (Gomphostilbia) azhari and Simulium (Gomphostilbia) johorense, are described based on adult females, males, pupae, and larvae collected from Peninsular Malaysia, and assigned to the parahiyangum subgroup and the duolongum subgroup of the batoense species group of the subgenus Comphostilbia, respectively. S. (G.) azhari sp. nov. is characterized in the female by the narrow frons, and in the male by the broad style and the ventral plate moderately produced ventrally. S. (G.) johorense sp. nov. is also remarkable in having the female subcosta lacking hairs or bearing a reduced number of hairs ranging from one to five. The pupae of both new species share a similar arrangement of the eight gill filaments (i.e., stalks of dorsal and middle triplets and ventral pair arising at the same level from the short common basal stalk), although relative lengths of filaments of the ventral pair to those of the dorsal and middle triplets are different between the two new species. Taxonomic notes are given to distinguish these new species from other related species. Keys to identify all 10 species of the batoense species group in Peninsular Malaysia are provided for adult females, males, pupae, and mature larvae.
A new species of Simulium (Gomphostilbia) (Diptera: Simuliidae) from Langkawi Island, Malaysia

Takaoka H, Sofian-Azirun M, Ya'cob Z

J Med Entomol, 2013 Jul;50(4):701-8.
PMID: 23926767

ABSTRACT A new black fly species, Simulium (Comphostilbia) langkawiense, is described based on adult female, adult male, pupal, and larval specimens collected from Langkawi Island, Malaysia. This new species is similar in the configuration of the pupal gill to Simulium (Comphostilbia) gombakense Takaoka & Davies, 1995, originally described from Peninsular Malaysia, but differs from the latter species by the female genital fork with an anterolaterally angulated plate on each arm, the female tarsal claw tooth shorter than one half of the claw, the small number of male upper-eye large facets, the ventral plate with its ventral margin nearly flat in the middle when viewed posteriorly, and the inflated structure of the pupal gill with a less produced middle portion (width of middle inflated portion: length of inflated structure = 0.24). Taxonomic notes are also given to separate this new species from two other related species from Nepal and India. This represents another example of a unique species of black fly on one of the continental islands of Peninsular Malaysia.
Fulltext Two new species of Simulium (Gomphostilbia) (Diptera, Simuliidae) from Peninsular Malaysia, with keys to Peninsular Malaysian members of the Simulium ceylonicum species-group

Takaoka H, Sofian-Azirun M, Hashim R, Ya'cob Z

Zookeys, 2011.
PMID: 21998510 DOI: 10.3897/zookeys.118.1552

Two new species of black flies, Simulium (Gomphostilbia) roslihashimisp. n. and Simulium (Gomphostilbia) lurauensesp. n., are described on the basis of reared adult, pupal and larval specimens collected from Peninsular Malaysia. These two new species are placed in the ceylonicum species-group within the subgenus Gomphostilbia. Simulium (Gomphostilbia) roslihashimisp. n. is most distinctive with the male having almost entirely yellow antennae, and Simulium (Gomphostilbia) lurauensesp. n. is characterized in the female by having the elongate sensory vesicle and the yellowish-white hairs on the base of the costal vein and on the stem vein, in the male by the greater number of large upper-eye facets and the spindle-shaped hind basitarsi which are much narrower than the hind tibiae and femora and in the pupa by the small terminal hooks. Keys to species of the ceylonicum species-group reported from Peninsular Malaysia are provided for females, males, pupae and mature larvae.
A new species of Simulium (Simulium) (Diptera: Simuliidae) from Langkawi Island, Peninsular Malaysia

Takaoka H, Sofian-Azirun M, Ya'cob Z

J Med Entomol, 2012 Nov;49(6):1198-205.
PMID: 23270147

Simulium (Simulium) kisapense sp. nov. is described on the basis of reared adult, pupal, and larval specimens collected from Langkawi Island, Peninsular Malaysia, and placed in the multi-striatum species-group in the subgenus Simulium. This new species is characterized by the bare basal section of the female radial vein, the male ventral plate with setae, the eight pupal gill filaments divergent at an angle of >90 degrees, and the shoe-shaped cocoon with small lateral window(s). Taxonomic notes are given to separate this new species from S. (S.) hirtinervis Edwards and S. (S.) malayense Takaoka and Davies, both from Peninsular Malaysia, and several other known species from Bhutan, India, Nepal, and Thailand.
Simulium (Gomphostilbia) merapiense sp. nov. (Diptera: Simuliidae) from Java, Indonesia

Takaoka H, Sofian-Azirun M, Ya'cob Z, Chen CD, Low VL, Zaid A

J Med Entomol, 2016 Jan;53(1):76-82.
PMID: 26516192 DOI: 10.1093/jme/tjv154

Simulium (Gomphostilbia) merapiense sp. nov. is described based on females, males, pupae, and mature larvae from Yagyakarta, Java, Indonesia. This new species is placed in the Simulium epistum species-group, and is characterized by the pupal gill with eight short filaments all arising at the same level from a short stalk, somewhat enlarged basal fenestra, entirely bare pupal head and thoracic integument, and small and short larval postgenal cleft. These characters rarely are found in the subgenus. Taxonomic notes are given to separate this new species from related species of the S. epistum species-group.

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