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.
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 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.
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 (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.
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.
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.
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.
Two new species of black flies, Simulium (Simulium) murudense and Simulium (Simulium) cheedhangi, are described on the basis of females, males, pupae, and larvae collected in Mount Murud, Sarawak, Malaysia. Both species belong to the Simulium melanopus Edwards species group. S. (S.) murudense sp. nov. is distinguished from most known species by a combination of the haired basal portion of the radial vein and the darkened fore coxae, and S. (S.) cheedhangi sp. nov. is characterized in the female by having a medium-sized claw tooth and in the pupa by six somewhat inflated gill filaments. Notes are given on the S. melanopus species-group in Sarawak and Sabah.
The classification, annotated list and keys for the black fly species from Peninsular Malaysia are updated. The number of black fly species increased from 38 in 1995 to 62 (including a new species herein described) in 2018. The 62 species are classified in four subgenera of the genus Simulium: one species in Daviesellum, 35 species in Gomphostilbia, five species in Nevermannia and 21 species in Simulium. Species in the latter three subgenera are further placed in species-groups or subgroups. Keys are provided for females, males, pupae and mature larvae. Simulium (S.) perakense sp. nov. is described and placed in the S. striatum species-group. The males of S. (G.) decuplum, S. (G.) tahanense, S. (S.) malayense, the female of S. (G.) adleri, and the female and larva of S. (G.) varicorne are described for the first time. The female, male, pupa and larva of S. (G.) trangense, the male and pupa of S. (G.) varicorne and the pupa of S. (G.) adleri are redescribed. Simulium (G.) sp. A is identified as S. (G.) pegalanense. Brief notes for each species are given on morphological characteristics, aquatic habitats and geographical distributions.
Simulium (Gomphostilbia) okinawense Takaoka and S. (G.) tokarense Takaoka, both from the Nansei Islands, Japan, were morphologically reexamined and genetically analysed by using the COI gene sequences. The female, male, pupa and mature larva of the two species are redescribed. Morphological reexamination shows that both species are more similar to species in the S. asakoae species-group than to those in the S. ceylonicum species-group, by having a medium-long female sensory vesicle, yellow tuft hairs (S. (G.) okinawense) or yellow tuft hairs mixed with a few to several dark hairs (S. (G.) tokarense) at the base of the radial vein in the female and male, and medium-long larval postgenal cleft. However, the body of the male ventral plate (viewed ventrally) is parallel-sided (S. (G.) okinawense) or parallelsided or slightly narrowed (S. (G.) tokarense) and not emarginated basally, differing from those of most species in the S. asakoae species-group. Our genetic analysis shows that S. (G.) tokarense is in the S. asakoae species-group, and S. (G.) okinawense formed a separate sister clade with other members of the S. asakoae species-group with high bootstrap support. From the results of morphological and genetic analysis combined, S. (G.) okinawense and S. (G.) tokarense are transferred from the S. ceylonicum species-group to the S. asakoae species-group.
Simulium takahasii (Rubtsov), which was originally described from Japan, and recorded from Korea and China, is the first among the 19 species of the subgenus Wilhelmia Enderlein recorded from East Asia. It is striking in mating, blood-feeding and ovipositing in captivity and in experimentally transmitting Dirofilaria immitis (Leidy) and Brugia pahangi (Buckley & Edeson), and it is a severe biter of cattle and horses, rarely of humans. Nevertheless, updated information about its morphological characteristics was lacking, making comparisons with related species described from China difficult, since species of the subgenus Wilhelmia are almost indistinguishable from one another, in particular, in their female terminalia, male genitalia and most of larval features. In this study, as many morphological characteristics as possible of S. takahasii based on specimens from Japan are redescribed. New information about many features of this species including the length of the female sensory vesicle against the third palpal segment, number of male upper-eye (large) facets, arrangement of the eight pupal gill filaments, presence or absence of tiny dark setae on the dorsum of the larval abdomen and the number of rows and hooklets of the larval posterior circlet will be useful in evaluating the species status of several Wilhelmia species in China including the species regarded as S. takahasii.
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.
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.
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.
Simulium ( Simulium) rufibasis Brunetti originally described from India was once considered a geographic generalist widely distributed in the Oriental Region. In this study, the species previously regarded as S. (S.) rufibasis in Taiwan was morphologically re-evaluated and found to be distinguished in the male and pupa from true S. (S.) rufibasis from India. This new species is described as S. ( S.) hehuanense sp. nov. based on a female, a male and their pupal exuviae in Taiwan. This new species is placed in the S. rufibasis subgroup of the S. tuberosum species-group, and is similar to S. (S.) yamatoense Takaoka, Adler & Fukuda from Japan and Korea, but it is barely distinguished by the slenderer forebasitarsi of the female and male. Another new, related species, S. (S.) xiulinense sp. nov., is described based on a male and its pupal exuviae. This new species is distinguished from S. (S.) hehuanense sp. nov. by the number of male upper-eye (large) facets in 20 or 21 vertical columns and 22 horizontal rows (19 vertical columns and 19 horizontal rows in the latter species), and presence of a pair of dorsolateral shiny spots on male abdominal segment 5 (absence in the latter species). Both new species inhabit streams at high elevations (ca, 2,600 m). They differ from S. (S.) sp. (probably S. (S.) arisanum Shiraki) from Taiwan, of the same subgroup, which breeds in streams at low elevations (433-685 m), by the pupal head and thorax covered with tubercles (bare in the latter species).
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.
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.
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.
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.