Recent studies on the biology and ecology of adult blackflies in relation to the transmission of human onchocerciasis in Guatemala are reviewed. First, earlier studies on the transmission of the disease since its discovery by Dr. R. Robles in 1915 are outlined. Second, eleven blackfly species caught on humans are evaluated for vector status on the basis of their natural and experimental infections with third-stage larvae of Onchocerca volvulus, and Simulium ochraceum,* S. metallicum* and S. callidum are confirmed as natural vectors of the disease in Guatemala in descending order of importance, whereas S. gonzalezi, S. haematopotum, S. veracruzanum and S. horacioi are potential vectors. Third, the migration and fate of O. volvulus microfilariae ingested by female blackflies are highlighted on the basis of the findings of the cibarial armature of S. ochraceum and physiological incompatibility of S. metallicum as main barriers against microfilariae, both of which greatly decrease the number of ingested O. volvulus microfilariae developing to the third-stage larvae per female. Fourth, among many ecological factors of female blackfly populations, geographical and altitudinal distributions, habits of blood feeding, host preferences for blood feeding, preference for human body parts, parous rates, daily and seasonal fluctuations of biting activities, in particular, of parous females, gonotrophic cycle, longevity, flight range, and annual transmission potential are reviewed, and their influences on the transmission dynamics of the disease agents are considered. Fifth, effects of air temperatures on the O. volvulus-S. ochraceum complex are examined, with a special reference to the characteristic altitudinal distributions of the disease. The importance of reliable identification of both the vector blackfly species and filarial larvae found in female blackflies is emphasized to understand the transmission of the disease. [*It is now known that these two species are actually species complexes.].
Recent studies on the epidemiology and control of Guatemalan onchocerciasis, chiefly made by the Guatemala-Japan Cooperative Project on Onchocerciasis Research and Control, are reviewed. Epidemiological features of Guatemalan onchocerciasis are summarized as to characteristic altitudinal distribution of endemic areas, disease manifestation, vector taxonomy, biology and transmission dynamic of the disease. Extensive insecticide studies in the field and laboratory demonstrate that the characteristic situations of Guatemalan streams where Simulium ochraceum, the main vector of onchocerciasis, breeds require ingenious methods of larviciding. Finally, the feasibility of an area vector control is indicated by the successful control operation in the San Vicente Pacaya Pilot Area, in which a new fixed-dose larviciding method was applied.
Simulium (Gomphostilbia) laosense sp. nov. is described based on adults, pupae, and mature larvae from Laos. This new species is placed in the Simulium batoense species-group of the subgenus Gomphostilbia Enderlein. It is characterized by the pupal gill with eight filaments arranged as 3 + 3 + 2 from dorsal to ventral, of which an inner filament of the ventral pair is slightly longer than its counter filament, and is 1.7-1.8 times as long as filaments of the middle triplet. Taxonomic notes are provided to distinguish this new species from Simulium (G.) johorense Takaoka, Sofian-Azirun & Ya'cob from Peninsular Malaysia and four other related species. The phylogenetic position of this new species in the S. batoense species-group is also presented based on the mitochondrial COI gene. This new species represents the second species known from Laos.
Simulium (Gomphostilbia) fukudae sp. nov. is described based on females, males, pupae, and larvae from Thailand. This new species is placed in the Simulium batoense species-group of Simulium (Gomphostilbia). It is characterized by the female frons almost bare, male paramere covered with minute setae, pupal gill composed of six slender filaments, pupal terminal hooks cone-shaped, and cocoon with an anterodorsal projection. Taxonomic notes are given to separate this new species from the most similar species, Simulium (G.) sextuplum Takaoka & Davies and Simulium (G.) paukatense Takaoka, both from Malaysia.
Simulium (Gomphostilbia) yvonneae sp. nov. is described based on adults, pupae, and mature larvae from Vietnam. This new species belongs to the Simulium duolongum subgroup in the S. batoense species-group of the subgenus Gomphostilbia Enderlein. It is distinguished by having a relatively larger number of male upper-eye facets in 16 vertical columns and 16 horizontal rows and a pupal gill with eight filaments arranged as 3+(1+2)+2 from dorsal to ventral, of which two filaments of the ventral pair are 1.8 times as long as the longest filament of the middle and dorsal triplets. Morphological comparisons are made to distinguish this new species from all 22 related species. The genetic distinctiveness of this new species in the S. duolongum subgroup is also presented based on the DNA barcoding COI gene.
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.
Simulium (Asiosimulium) saeungae sp. nov. (Diptera: Simuliidae) is described based on females, males, pupae, and mature larvae collected from Nan Province, Northern Thailand. It is characterized by the medium-long cerci in the female, enlarged hind basitarsus, and broad ventral plate with its posterior margin not deeply concave in the male, arborescent pupal gill with 42-56 filaments in the pupa and smaller number of primary rays of the labral fan (30-33) in the larva. This is the fifth species of the subgenus Asiosimulium, the second smallest among 10 subgenera in the Oriental Region. Taxonomic notes are given to distinguish this new species from the three known species from Thailand and one from Nepal.
A new simuliid species, Simulium kalimantanense sp. nov., is described on the basis of females, males, pupae, and mature larvae from East Kalimantan, Indonesia, and is assigned to the Simulium banauense species-group of Simulium (Gomphostilbia). This new species has close similarities to S alienigenum Takaoka from the Philippines, in many characters including the adult antennal color pattern and pupal gill with four long filaments arranged in two pairs each bearing a long stalk, but is distinguished from the latter in the female by the longer sensory vesicle and in the pupa by the gill with an elongate common basal stalk. Simulium kalimantanense sp. nov. is the first member of the S. banauense group in Borneo, and marks the most southerly distribution of the group. Keys to identify 19 Bornean species of the subgenus Gomphostilbia are provided.
A new black fly species, Simulium haiduanense Takaoka, Low & Huang (Diptera: Simuliidae), is described on the basis of females, males, pupae, and mature larvae from Taiwan. This new species is placed in the Simulium argentipes species-group of the subgenus Simulium (Diptera: Simuliidae) and is characterized by the yellowish female legs, ovipositor valves rounded apically and with its inner margin concave, claw with a small subbasal tooth, male style without a basal protuberance, pupal gill with eight filaments, corbicular cocoon, and larval abdomen lacking paired protuberances. It represents the first record of the S. argentipes species-group from Taiwan. Taxonomic notes are given to separate this new species from all eight species in the same species-group. The phylogenetic relationships of this new species with four related species are presented.
Simulium undecimum sp. nov. is described from Thailand. This new species is assigned to the Simulium multistriatum species-group, one of the 20 species-groups of the subgenus Simulium in the Oriental Region. It is characterized by the female cibarium with minute processes, male ventral plate with a narrow body having two vertical rows of distinct teeth on the posterior surface and without setae on the anterior and lateral surfaces, pupal gill with eight short filaments decreasing in length from dorsal to ventral, and divergent at an angle of around 90 degrees when viewed laterally, spine-combs only on abdominal segments 7 and 8, and cocoon wall-pocket shaped with anterolateral windows. Taxonomic notes to separate this new species from related species in Thailand and other countries are given. This new species is the 11th nominal member of this species-group recorded in Thailand. An analysis of the COI gene sequences shows that it is most closely related with S. malayense Takaoka & Davies (cytoform A) from Thailand but they are distantly separated by 3.01-8.87%.
Simulium (Simulium) phraense sp. nov. (Diptera: Simuliidae) is described from females, males, pupae, and larvae from Thailand. This new species is placed in the Simulium striatum species group and is most similar to Simulium (Simulium) nakhonense Takaoka & Suzuki (Diptera: Simuliidae) from Thailand among species of the same species group but is barely distinguished from the latter species by lacking annular ridges on the surface of the pupal gill filaments. The fast-evolving nuclear big zinc finger (BZF) gene has successfully differentiated this new species from its allies, S. (S.) nakhonense and Simulium (Simulium) chiangmaiense Takaoka & Suzuki (Diptera: Simuliidae) of the S. striatum species group. The BZF gene sequences show that this new species is more closely related to S. (S.) nakhonense than to S. (S.) chiangmaiense, further supporting its morphological classification.
A rare non-sex mosaic abnormality represented by genitalia-like appendages on the ventral surface of abdominal segment 8 of a male black fly collected in Hokkaido, Japan, is reported. The appendages consist of a pair of style-like projections each arising from a coxite-like base, inverted-Y shaped ventral plate-like structure, and isolated round structure. This male was morphologically and molecularly identified as an abnormal form of S. (S.) iwatense (Shiraki), the only species in the Simulium (Simulium) ornatum species-group in Japan, although certain morphological characteristics of this male including the reduced number of uppereye (large) facets and elongate cerci are different from those of S. (S.) iwatense.
A multi-locus approach was used to examine the DNA sequences of 10 nominal species of blackfly in the Simulium subgenus Gomphostilbia (Diptera: Simuliidae) in Malaysia. Molecular data were acquired from partial DNA sequences of the mitochondria-encoded cytochrome c oxidase subunit I (COI), 12S rRNA and 16S rRNA genes, and the nuclear-encoded 18S rRNA and 28S rRNA genes. No single gene, nor the concatenated gene set, resolved all species or all relationships. However, all morphologically established species were supported by at least one gene. The multi-locus sequence analysis revealed two distinct evolutionary lineages, conforming to the morphotaxonomically recognized Simulium asakoae and Simulium ceylonicum species groups.
Simulium (Gomphostilbia) marosense sp. nov. is described based on adults, pupae and mature larvae from Sulawesi, Indonesia. This new species is placed in the Simulium duolongum subgroup of the S. batoense species-group, representing the ninth member for the subgroup in Sulawesi. This new species is characterized by a narrow female frons and pupal gill with eight filaments, of which two filaments of the ventral pair are 1.9-3.0 times as long as the six other filaments. Taxonomic notes are provided to distinguish this new species from related species.
Simulium pumatense sp. nov. is described from Vietnam, and is placed in the Simulium feuerborni species-group of the subgenus Simulium (Nevermannia) Enderlein. Its morphological characteristics include the relatively smaller numbers of the following three numerical features: inner teeth of the female mandible (15-18), minute conical processes (16) on the female cibarium, and male upper-eye facets (in 15 vertical columns and 16 horizontal rows). Keys are constructed to distinguish this species from four species of the same group in Vietnam. Our molecular analysis of the DNA barcoding COI gene shows that this species is most closely related to cytoform A of the S. feuerborni complex from Thailand.
A new black fly species, Simulium taichungense, is described on the basis of one female, one male and two pupal exuviae from Taiwan. This new species is placed in the S. crocinum species-group of Simulium (Simulium) (Diptera: Simuliidae). It is characterized by the pupal gill with six unpigmented short filaments and pupal abdomen with distinct spine-combs on the dorsal surface of segments 7-9, by which it is easily distinguished from most species in the S. crocinum species-group including two species from Taiwan: Simulium serenum Huang Takaoka and S. ufengense Takaoka. It is the third species of the S. crocinum species-group from Taiwan. In addition, the male of S. serenum is described for the first time, and intraspecific variations in a few features of the male of S. chungi Takaoka Huang of the S. chungi species-group are noted.
A new species of black fly, Simulium (Gomphostilbia) isanense, is described based on females, males, pupae and mature larvae from Thailand. This new species is placed in the Simulium epistum species-group of the subgenus Gomphostilbia Enderlein. It is characterized by the pupal gill with eight filaments arranged as 3+3+2 from dorsal to ventral, of which an inner filament of the ventral pair is slightly longer than its counter filament. Taxonomic notes are provided to distinguish this new species from S. (G.) angulistylum Takaoka & Davies from Peninsular Malaysia, and three other related species. The difference between this new species and S. (G.) angulistylum is supported by genetic distances using the mitochondrial COI gene.
A case of zoonotic onchocercosis has been found in a resident who lived in Iizuka City, Fukuoka Prefecture, Japan for some time. A 24-year-old male developed a painful nodule on the middle finger of his right hand. The nodule was surgically removed from the vagina fibrosa tendinis of the finger at Beppu Medical Center, Beppu City, Oita Prefecture in 2012. The causative agent was identified as a female Onchocerca dewittei japonica based on its histopathological characteristics. The identity of the filarioid has been confirmed by sequencing the cox1 gene. The present study indicates that the zoonotic onchocercosis caused by O. dewittei japonica has been concentrated in northeast Kyushu.
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.
Simulium (Asiosimulium) khongchiamense sp. nov. is described based on females, males, pupae, and mature larvae collected from Khong Chiam District, Ubon Ratchathani Province, northeastern Thailand. It is characterized in the female by the medium-long sensory vesicle, scutum with 3 dark longitudinal vittae and elongate cercus; in the male by the number of upper-eye (large) facets in 17 or 18 vertical rows and 18 or 19 horizontal rows, hind basitarsus moderately enlarged and ventral plate with the posterior margin moderately concave medially; in the pupa by the head and thoracic integument sparsely covered with tubercles and gill of arborescent type with 32 or 33 filaments; and in the larva by the postgenal cleft deep, reaching the posterior margin of the hypostoma and sheath of the subesophageal ganglion dark pigmented. DNA analysis based on COI gene of all known species of the subgenus Asiosimulium, except for S. shanense and S. suchitrae, indicated that this new species can be clearly differentiated from all other related species (S. phurueaense, S. oblongum, S. saeungae, S. furvum, and S. wanchaii) with interspecific genetic distances ranging between 4.79% and 19.18%. This is the eighth species of the subgenus Asiosimulium. Taxonomic notes are given to distinguish this new species from the 7 known species members in its same subgenus. Additionally, keys to species of all members in the subgenus Asiosimulium are provided.