The WHO Onchocerciasis Control Programme: retrospect and prospects

The history of onchocerciasis control in Africa and the genesis of the WHO Onchocerciasis Control Programme in West Africa (OCP) are briefly reviewed. The importance of experience gained in anti-locust campaigns in helping to plan the OCP is stressed. Members of the Simulium damnosum species complex are the vectors of onchocerciasis, which OCP is controlling with insecticide treatments on the stretches of rivers where the Simulium breed. Migrations of flies have been responsible for reinfestations of controlled areas and the spread of insecticide resistance. The management of these problems and related research are described, but it is emphasized that despite setbacks OCP is achieving its aims. A strategy for the future is outlined: vector control supplemented by chemotherapy is expected to continue until the year 2004.     

Salivary apyrase in New World blackflies (Diptera: Simuliidae) and its relationship to onchocerciasis vector status

Abstract. Salivary gland apyrase is believed to be critical to blood-feeding in arthropod vectors. This enzyme was measured in six New World blackflies representing three taxonomic pairs of non-vectors and vectors of Onchocerca volvulus. In Simulium (Psilopelmia) ochraceum , a highly anthropophilic vector in Mexico and Guatemala, apyrase exhibited maximum activity between pH 8.0 and 9.0, mean 39.8 pM 4.7 milliUnits/pair of gland equivalents (mU), and was enhanced when ATP was used as a substrate. In the zoophilic non-vector Simulium (Psilopelmia) bivittatum maximum activity was significantly less (5.1 pM 0.7 mU) under all conditions examined. Preference for ADP or ATP as substrate was a function of the pH of the reaction for this species. Apyrase activity in Simulium (Simulium) metallicum Bellardi (29.5 pM 11.5 mU), a zoophilic secondary vector in Mexico and Guatemala, resembled that of S.(Ps.)ochraceum (24.8 pM 13.7 mU at pH 8.5) with ADP as substrate, but showed reduced activity with ATP. Both these Central American vectors had higher apyrase activity than found in Simulium (Notolepria) exiguum , a vector of O. volvulus in Ecuador and Colombia. However, maximum apyrase activity, measured at pH 8.0 with ADP as substrate, was greater in S.(N.) exiguum (10.9 pM 0.6mU) than in Simulium (Notolepria) gonzalezi (5.9pM1.9mU), a non-vector species widespread in Central America. Therefore, for the consubgeneric species pairs examined, a positive association was detected between higher concentrations of apyrase activity and their vector status for O. volvulus.     

Phosphoglucomutase and trehalase isoenzymes of Venezuelan simulium vectors of Onchocerca volvulus

Phosphoglucomutase (PGM) and trehalase (Tre) isoenzymes of five species of Simulium blackflies (Diptera: Simuliidae), vectors of onchocerciasis in Venezuela, were investigated by means of a portable electrophoresis field kit. Tre differed between S. incrustatum and S. oyapockense s.l. Electrophoretic variation of Tre in other members of the S. amazonicum and S. incrustatum groups merit further investigation. PGM appears to be more useful for separating populations within species complexes. Multiple populations and/or seasonal changes in population structure of S. guianense s.l., S. exiguum s.l. and S. metallicum s.l. were inferred from elecrophoretic variation of PGM.     

Assessment of rDNA IGS as a molecular marker in the Simulium damnosum complex

For five cytospecies of the Simulium damnosum Theobald complex of blackflies (Diptera: Simuliidae) from West Africa, both ends of the intergenic spacer region (IGS) of the rDNA have been sequenced with the aim of developing specific molecular markers. No specific differences in these two regions were detected between Simulium sanctipauli V. & D., Simulium sirbanum V. & D., Simulium soubrense V. & D., Simulium squamosum Enderlein and Simulium yahense V. & D., except in the number of A subrepeats at the 5' end of the IGS (two in S. squamosum and four or five in the others) and in position 310 of the 3' end (a C in S. squamosum and a G in the others). However, genetic distances within and between species overlapped. These DNA sequences had no strong phylogenetic signal, and the trees obtained were mostly unresolved. Although most sequences from S. squamosum clustered together, a few of them were more similar to those in other cytospecies. These results could be explained either by hybridization with genetic introgression or by ancestral polymorphism and recent speciation.     

Anthropophilic blackflies (Diptera: Simuliidae) and onchocerciasis transmission in southwest Ethiopia

Abstract. From May 1984 to May 1985, the transmission of onchocerciasis by blackflies (Dipteria: Simuliidae) was studied around Gilgel Ghibe and Gojeb rivers in southwest Ethiopia. The predominant anthropophilic species of Simulium at both localities were S.(Edwardsellum) damnosum sensu law and S.(Lewisellum) ethiopiense. Some S.(Anasolen) dentulosum also landed on human bait at Gilgel Ghibe river.
Dissections of S. damnosum from Gilgel Ghibe and Gojeb revealed mean parous rates of 92% (n = 18,291) and 84% (n = 9530), respectively. S. ethiopiense from Gilgel Ghibe and Gojeb showed mean parous rates of 53% (n = 322) and 93% (n = 14), respectively. Of the parous S. damnosum , 1.3% at Gilgel Ghibe and 0.5% at Gojeb harboured infective third-stage larvae (L3) of Onchocerca volvulus (or morphologically indistinguishable from it). Unknown filariae of animal origin, indicative of zoophily, were found in 0.3% and 0.7% of Simulium damnosum complex females from Gilgel Ghibe and Gojeb, respectively. S. ethiopiense harboured developing (L,) larvae only, with 7.7% infection rate in both localities. In contrast, S. dentulosum did not harbour any filaria larvae. The annual infective biting rate (AIBR) and transmission potential (ATP) of the S. damnosum complex at Gilgel Ghibe river were 858 and 5478, respectively. The AIBR and ATP of S. damnosum s.I. at Gojeb river were 519.5 and 1963, respectively. These results emphasize the predominant role of the S. damnosum complex in the transmission of O. volvulus in southwest Ethiopia     

Blackfly vectors of zoonotic onchocerciasis in Japan

Studies of blackfly vectors of Onchocerca dewittei japonica Uni, Bain & Takaoka (Spirurida: Onchocercidae), a parasite of wild boar implicated in the aetiology of zoonotic onchocerciasis in Japan, and six other zoonotic Onchocerca species of this country are reviewed. Molecular identification of infective larvae found in wild‐caught female blackflies showed that Simulium bidentatum (Shiraki) (Diptera: Simuliidae) is a natural vector of O. dewittei japonica, and also Onchocerca sp. sensu Fukuda et al., another parasite of wild boar. Inoculation experiments demonstrated that Simulium arakawae Matsumura and four other Simulium species are putative vectors. Similarly, S. arakawae, S. bidentatum and Simulium oitanum (Shiraki) are putative vectors of Onchocerca eberhardi Uni & Bain and Onchocerca skrjabini Rukhlyadev, parasites of sika deer. Morphometric studies of infective larvae indicated that Onchocerca lienalis Stiles, a bovine species, is transmitted by S. arakawae, Simulium daisense (Takahasi) and Simulium kyushuense Takaoka, and that Onchocerca sp. sensu Takaoka & Bain, another bovine species, is transmitted by S. arakawae, S. bidentatum, S. daisense and S. oitanum. Prosimulium sp. (Diptera: Simuliidae) and Simulium japonicum Matsumura are suspected vectors of Onchocerca suzukii Yagi, Bain & Shoho and O. skrjabini [Twinnia japonensis Rubtsov (Diptera: Simuliidae) may also transmit the latter], parasites of Japanese serow, following detection of the parasites' DNA genes in wild‐caught blackflies.     

Revision of the Ketaketa subcomplex of blackflies of the Simulium damnosum complex

A revision of the taxonomy of the Ketaketa subcomplex of the Simulium damnosum Theobald complex (Diptera: Simuliidae) is presented including new material from Tanzania, Malawi and South Africa. The cytotaxonomy, morphology and molecular identity of known and new taxa are described. The Ketaketa subcomplex is cytotaxonomically defined by the paracentric inversion 1L-7. We recognize three sibling species, namely Simulium latipollex (Enderlein), Simulium plumbeum Krueger, sp.n. and Simulium kipengere Krueger, sp.n., the latter comprising three cytoforms: 'Typical', 'Linthipe' and 'Mombo'. The cytoforms 'Mwamphanzi', 'Ketaketa' and 'Hammerkopi' are synonymized with S. plumbeum. Identification keys are provided on the basis of chromosomal and morphological characters. In view of their potential role as vectors of human onchocerciasis (river blindness) we also discuss the possible medical importance of the different cytoforms and their geographical distribution.     

DNA probes for the identification of members of the Simulium damnosum complex (Diptera: Simuliidae)

Genomic libraries in plasmid have been constructed from various sibling species of blackflies of the Simulium damnosum complex from West Africa. Three cloned repetitive sequences, which show variation in copy number between sibling species, have been isolated. These clones can be used as probes for the dot-blot identification of larvae, pupae or adults into the three main West African subcomplexes, i.e. damnosum, squamosum and sanctipauli subcomplexes. The sequences also show some intraspecific variation in copy number.     

滋生于海南岛五指山瀑布岩壁上蚋属二新种(双翅目,蚋科)

首次记述我国大陆孳生于海南岛五指山瀑布岩壁上蚋属2新种,即瀑布蚋Simulium ( Simulium ) waterfallum sp. nov.和副瀑布蚋S. ( S. ) parawaterfallum sp. nov.二者均隶属于蚋亚属Simulium的多条蚋组multistriatum-group以其孳生环境命名.文中对其两性成虫、蛹和幼虫进行描述并分别与其近缘种进行分类讨论.模式标本存放于贵阳医学院生物学教研室.     

贵州雷公山蚋属二新种记述(双翅目:蚋科)

记述采自贵州雷公山的蚋属二新种,即细板蚋Ｓ．（Ｓ．）ｔｅｎｕａｔｕｍ ｓｐ．ｎｏｖ．,多枝蚋Ｓ．（Ｓ．）ｒａｍｕｌｏｓｕｍ,ｓｐ。ｎｏｖ。并对其分类地位进行讨论。     

Onchocerciasis transmission levels and Simulium damnosum complex biting activity at riverside and rice field sites in Sierra Leone

Biting densities of the Simulium damnosum Theobald complex of blackflies and their levels of parasitism by Onchocerca volvulus (Leuckart) were compared at two ecologically contrasted sites in the Southern Province of Sierre Leone, West Africa: by the Tabe riverside close to Gbaiima village where onchocerciasis is hyperendemic (63.1% positive skin-snips) and in open rice fields 2 km from the mesoendemic (54.9%) village of Senehun. Using a combination of morphometric and enzyme electrophoresis techniques, the primary vector was found to belong to the S.sanctipauli subcomplex and was most probably S.soubrense 'B' (sensu Post, 1986). In the wet season biting activity took place throughout the day at the village, but had morning and evening peaks in the rice fields. Transmission of O.volvulus was detected at the village in all months except February-April, while in the rice fields it was detected only during September-November and May at one tenth the level of the village. There was a significant difference between the two sites in the proportion of parous flies carrying third stage larvae in the head, but not of other larval stages.     

Elimination of the Djodji form of the blackfly Simulium sanctipauli sensu stricto as a result of larviciding by the WHO Onchocerciasis Control Programme in West Africa

Cytotaxonomic identifications of larvae of members of the Simulium damnosum Theobald (Diptera: Simuliidae) complex collected in forest zones of southeast Ghana and southwest Togo between 1977 and 1996 showed that the Djodji form of Simulium sanctipauli Vajime & Dunbar, a vector of onchocerciasis, was eliminated in 1988 by larvicide operations conducted by the World Health Organization (WHO) Onchocerciasis Control Programme (OCP) in West Africa. No members of the form were identified amongst 997 larvae collected up to 8 years after systematic control operations began in February 1988. The results are discussed in relation to estimates of the numbers of samples required to certify elimination and the possibility that other members of the S. damnosum complex were also eliminated by the OCP.     

Mechanisms of resistance to Onchocerca infection in blackflies

In this article Peter Ham, Hans Hagen, Andrea Baxter and Jorg Grunewald focus on the susceptibility of blackflies to parasitic filarial infection (particularly Onchocera spp, most of the vectors of which belong to the genus Simulium). They outline what is known about, as well as speculating on, the various defence mechanisms of these insects. Investigations have involved the use of natural and surrogate vectors of bovine onchocerciasis as models for the human vector-parasite relationship.     

An epizootic of patent iridescent virus disease in multiple species of blackflies in Chiapas, Mexico

Simulium blackfly larvae (Diptera: Simuliidae) were collected from rivers and streams at 500-1500 m a.s.l. in Chiapas State of southern Mexico. Among 45 sites surveyed over an area of 2300 km2 (around 15 degrees 15'N 92 degrees 20'W), some Simulium larvae from three sites were opalescent violet-blue, interpreted as patent infection with invertebrate iridescent virus (IIV). Dissection confirmed the presence of putative Iridovirus particles, 130nm diameter, but no IIV isolates were obtained from homogenates injected into Galleria mellonella (L) larvae (Lepidoptera: Pyralidae). All Simulium with patent IIV infection died before metamorphosis, whereas approximately 60% of asymptomatic Simulium survived to adulthood in the laboratory. During 1997, standard monthly samples from two parallel rivers 42-50 km north-west of Tapachula comprised the following species proportions (and rates of patent IIV infection): 41.8% (47%) Simulium mexicanum Bellardi complex, 31.3% (31.4%) S. rubicundum Knab, 10.1% (13.1%) S. paynei, 6.5% (2.9%) S. callidum (Dyar & Shannon), 6.3% (5.1%) S. ochraceum Walker complex, 3.1% (0.7%) S. downsi Vargas et al., 0.7% S. samboni Jennings and 0.2% S. metallicum Bellardi complex, showing a strong correlation between blackfly abundance and the prevalence of patent infection. An epizootic of IIV in January and February (infection rates 41-100%) was followed by absence of larvae (March-August) until the end of the rainy season, when numbers collected on nylon strings rose to approximately 1/cm with patent IIV infection rates of 0-12.5% during September-December. Further investigations are underway to isolate this IIV and assess its potential usefulness for biological control of Simulium pests and vectors of onchocerciasis.     

Detection of Onchocerca volvulus (Nematoda: Onchocercidae) infection in vectors from Amazonian Brazil following mass Mectizan distribution

Detection of Onchocerca volvulus in Simulium populations is of primary importance in the assessment of the effectiveness of onchocerciasis control programs. In Brazil, the main focus of onchocerciasis is in the Amazon region, in a Yanomami reserve. The main onchocerciasis control strategy in Brazil is the semi-annually mass distribution of the microfilaricide ivermectin. In accordance with the control strategy for the disease, polymerase chain reaction (PCR) was applied in pools of simuliids from the area to detect the helminth infection in the vectors, as recommended by the Onchocerciasis Elimination Program for the Americas and the World Health Organization. Systematic sampling was performed monthly from September 1998 to October 1999, and a total of 4942 blackflies were collected from two sites (2576 from Balawaú and 2366 from Toototobi). The molecular methodology was found to be highly sensitive and specific for the detection of infected and/or infective blackflies in pools of 50 blackflies. The results from the material collected under field conditions showed that after the sixth cycle of distribution of ivermectin, the prevalence of infected blackflies with O. volvulus had decreased from 8.6 to 0.3% in Balawaú and from 4 to 0.1% in Toototobi.     

Cryptic biodiversity in the cytogenome of bird‐biting blackflies in North Africa

Bird‐biting blackflies in the Simulium (Eusimulium) aureum group (Diptera: Simuliidae) are widespread vectors of Leucocytozoon and Trypanosoma parasites. The polytene chromosomes of 619 larvae of the three nominal members of the S. aureum group in North Africa were evaluated cytogenetically for cryptic biodiversity. Seven chromosomal segregates were discovered among 29 populations in Algeria and Morocco. This diversity was based primarily on two chromosomal inversions, which have assumed unique roles in different lineages, including sex linkage, fixation, loss and autosomal polymorphism. Reproductive isolation was demonstrated for six of the seven segregates, doubling the number of species known in the area. Four species were linked with existing names: (a) Simulium mellah Giudicelli & Bouzidi, which is known only from North African high‐salinity habitats; (b) Simulium petricolum (Rivosecchi), which is tentatively conspecific with continental European populations; (c) Simulium rubzovianum (Sherban) and its synonym Simulium latinum (Rubtsov), which is widely distributed from North Africa across Europe into Western Asia, and (d) Simulium velutinum (Santos Abreu) and its new synonym Simulium tenerificum Crosskey, which is restricted to North Africa and the Canary Islands. Of the remaining entities, two are new species precinctive to North Africa and one, known only from Morocco, is of undetermined taxonomic status.     

Progress in controlling the reinvasion of windborne vectors into the western area of the Onchocerciasis Control Programme in West Africa

Since vector control began in 1975, waves of Simulium sirbanum and S. damnosum s.str., the principal vectors of severe blinding onchocerciasis in the West African savannas, have reinvaded treated rivers inside the original boundaries of the Onchocerciasis Control Programme in West Africa. Larviciding of potential source breeding sites has shown that these 'savanna' species are capable of travelling and carrying Onchocerca infection for at least 500 km northeastwards with the monsoon winds in the early rainy season. Vector control has, therefore, been extended progressively westwards. In 1984 the Programme embarked on a major western extension into Guinea, Sierra Leone, western Mali, Senegal and Guinea-Bissau. The transmission resulting from the reinvasion of northern C?te d'Ivoire and Burkina Faso has been reduced by over 95%, but eastern Mali has proved more difficult to protect because of sources in both Guinea and Sierra Leone. Rivers in Sierra Leone were treated for the first time in 1989 and biting and transmission rates in Sierra Leone and Guinea fell by over 90%. Because of treatment problems in some complex rapids and mountainous areas, flies still reinvaded Mali, though biting rates were approximately 70% lower than those recorded before anti-reinvasion treatments started. It was concluded that transmission in eastern Mali has now been reduced to the levels required to control onchocerciasis.     

Blackflies (Diptera: Simuliidae) of Southern Guyana with keys for the identification of adults and pupae--a review

A revision is made of the previously poorly studied blackfly fauna from the south-western border of Guyana with Brazil. Notes on the biosystematics of the species found are provided, together with keys and illustrations based on their morphology. Of the 14 species recorded, eight are anthropophilic and two of these (Simulium oyapockense s.l. and S. guianense s.l.) are proven vectors of human onchocerciasis in the nearby Amazonia focus of the disease in neighbouring Brazil.     

Biting behaviour and potential vector status of anthropophilic blackflies in a new focus of human onchocerciasis at Minaçu, central Brazil

Monthly collections were made of man‐biting female blackflies: Simulium auripellitum Enderlein, S. guianense Wise, S. minusculum Lutz and S. nigrimanum Macquart (Diptera: Simuliidae) from four catching stations in the newly discovered focus of human onchocerciasis at Minaçu (13°35′ S 48°18′ W), 300 km north of Brasília in Goiás State. These provided baseline data on biting habits, population density and seasonal prevalence during the year before completion of the Serra da Mesa hydroelectric dam on the Rio Tocantins near Minaçu, in a project investigating the effect of dam construction on onchocerciasis transmission in the area. All four simuliid species were most abundant during the dry season, and only bit in low numbers (S. auripellitum S.minusculum, S. nigrimanum) or were absent (S. guianense) in the wet season. Simulium minusculum was the predominant species at all catching stations, being particularly abundant by the large River Tocantins. The other three species were mainly associated with smaller rivers. In the dry season, biting rhythms of S. minusculum varied with catching site, while S. nigrimanum showed peaks of activity in early morning and during the afternoon. Experimental infection with Onchocerca volvulus (Leuckart) (Nematoda: Onchocercidae), from a human volunteer, showed that this parasite could develop fully in the four simuliid species, which are all considered to be potential vectors in the area.     

The thermal constant of the onchocerciasis vector Simulium damnosum s.l. in West Africa

The minimum water temperature for development (t(0)) and the thermal constant (K) for the development of immature stages of Simulium damnosum s.l. (Diptera: Simuliidae) in West Africa were estimated as 20.1 °C and 93 day-degrees, respectively, based on analyses of published data on development rates of eggs, larvae and pupae at different water temperatures (24.0 °C and 31.5 °C). Thus, at a constant water temperature of 30.0 °C (approximately 10 °C above t(0)), adult flies would emerge about 9 days after oviposition. Analysis of a dataset probably restricted to S. damnosum s.s., but for which the temperature for the egg stage varied, revealed a much lower t(0) (16.3 °C) and a much higher K (181 day-degrees), suggesting that the insects' thermal relations may be cytoform-specific. The results will aid control decisions and predictions of possible effects of climate change on sizes and geographic distributions of populations of onchocerciasis vectors in West Africa.