The Beffa form of Simulium soubrense of the S.damnosum complex in Togo and Benin

The Beffa form of Simulium soubrense Vajime & Dunbar, a member of the S. sanctipauli sub-complex of the S. damnosum complex, was found breeding throughout rivers in the Togo-Benin Gap, as far north as 9 degrees 30'N. Its distribution changed with the season. In southern Togo there were seasonal fluctuations in the relative abundancies of the Beffa form and of S. damnosum/S.sirbanum. There was considerable temporal and regional variation in the frequencies of different colour morphs of adult flies. The flies in Benin tended to be darker. Infections with Onchocerca volvulus (Leuckart) appeared to be independent of the host's colour morph category. Larger flies harboured significantly more first stage Onchocerca larvae but no significant relations with fly size were found for second and third stage larvae.     

Management of insecticide resistance in control of the Simulium damnosum complex by the Onchocerciasis Control Programme, West Africa: potential use of negative correlation between organophosphate resistance and pyrethroid susceptibility

1. Resistance of some populations of the Simulium damnosum complex to temephos (100-fold at the LC50 level), with degrees of cross-resistance to chlorphoxim (14-fold) and other organophosphate insecticides, follows intensive larvicidal control of S. damnosum s.l. in West African river systems since 1975 by the WHO Onchocerciasis Control Programme. 2. Larvae of at least three sibling species of the S. damnosum complex have become organophosphate-resistant: these are the forest species S. sanctipauli Vajime & Dunbar and the savanna species S. sirbanum V. & D. and S. damnosum Theobald sensu stricto. 3. Organophosphate-resistant S. damnosum s.l. larvae show increased susceptibility to some organochlorine and pyrethroid insecticides, especially to permethrin (up to 11-fold) and OMS 3002 (up to 17-fold), as compared with organophosphate-susceptible populations. 4. This differential susceptibility is reflected by increased pyrethroid efficacy in operational use for river treatments against organophosphate-resistant field populations of S. damnosum s.l. larvae. Treatment of 100 km of the lower Bandama River in 1985 showed that permethrin at the highly selective dosage of 10 min exposure to 0.01 mg/l caused reversion towards organophosphate-susceptibility of the target population of S. sanctipauli. This effect was less pronounced when the Comoe River was treated at the lower dosage of 0.005 mg/l for 10 min. 5. To overcome temephos-resistance, it is proposed that the most rational usage of currently available larvicides would involve the following annual sequence of treatments: Bacillus thuringiensis serotype H-14 when river discharge is below 75 m3/s; chlorphoxim for about eight weekly treatment cycles after river discharge rises; permethrin (or alternative pyrethroid) for up to six treatment cycles--this should eliminate any incipient selection for chlorphoxim-resistance; resume chlorphoxim (or perhaps carbosulfan) treatments until river discharge falls below 75 m3/s permitting resumed use of B.t. H-14.     

尼日利亚西南部奥孙河沿线黑蝇Simulium damnosum Theobald complex的形态分类研究(双翅目:蚋科)

黑蝇Simulium damnosum sensu lato是由多个姊妹种组成的复合体,这些种在生态学和盘尾丝虫病的传播方面各不相同.本文对奥孙河沿线的尼日利亚西南部森林区域的黑蝇S.damnosum s.l.复合体的组成以及成虫的形态学特征进行了研究.本研究所用的黑蝇S.damnosum s.l.成虫标本来源于奥孙河沿线的3个区,分别是Osun Eleja,Osun Ogbere和Osun Budepo.标本采集通过人体诱捕的方式,采集时间从2008年2月至2009年6月上午7:00到下午6:00,每两周采集1次.通过观察成虫的翅毛簇(wing tufts)和其他形态特征,对其进行分类研究.结果表明:存在同域分布的森林种和稀树草原种.在奥孙河沿线3个区内,森林种为优势种,占总捕获量的99.18%,而稀树草原种仅占0.82%;森林种和稀树草原种在多度上存在显著差异(P<0.05).所捕获的所有稀树草原种的翅毛簇均为灰白色,而捕获的森林种的翅毛簇颜色存在显著差异(P<0.05).为了更清楚地揭示该地区黑蝇S.damnosum s.l.的物种组成,建议进行更深入的研究.     

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.     

Efficiency of Simulium sanctipauli as a vector of Onchocerca volvulus in the forest zone of Ghana

The role of Simulium sanctipauli Vajime & Dunbar (Diptera: Simuliidae) as a vector of Onchocerca volvulus (Leuckart) (Spirurida: Onchocercidae) in the forest zone of central Ghana was studied in the Upper Denkyira district, where onchocerciasis is hyperendemic. Simulium sanctipauli was found to be a highly efficient vector, with a mean of 377 infective (L3) larvae in the heads of 1000 parous and 122 in the heads of 1000 biting flies. The overall infection rate of 44% of the parous flies with L1, L2 and L3 stages of O. volvulus (identity confirmed by polymerase chain reaction) demonstrates marked anthropophily. Female flies dispersed over a wide area and can transmit onchocerciasis up to at least 10 km away from their breeding sites. Annual community-directed treatments with ivermectin did not have a noticeable effect on the infection rates and parasitic loads of fly populations, which were as high 2 months after as 3 months before the distribution of ivermectin. This failure can be attributed to poor coverage, with treatment taken by only 24.4% of the population of the six study villages.     

Cytotaxonomy,morphology and molecular systematics of the Bioko form of Simulium yahense (Diptera: Simuliidae)

Cytotaxonomic analysis of the polytene chromosomes from larvae of the Simulium damnosum Theobald complex from the island of Bioko in Equatorial Guinea is reported, and a new endemic cytoform is described. Chromosomally this cytoform is close to both S. squamosum (Enderlein) and S. yahense Vajime & Dunbar, but is not identical to either. However, it is morphologically and enzymatically identical to S. yahense. The Bioko form was also found to differ from other cytoforms of the S. damnosum complex in West Africa in the copy number or RFLP pattern of several different repetitive DNA sequences. It is clear that the Bioko form is genetically distinct from other populations of the S. damnosum complex, and whilst it is closest to S. yahense, it shows features that suggest a high degree of geographical and genetic isolation. Such isolation is an important consideration in the assessment of the potential for onchocerciasis vector eradication on Bioko.     

Cytotaxonomic confirmation of two forms of Simulium sirbanum in the eastern part of the Onchocerciasis Control Programme in West Africa

An earlier report (Cheke et al., 1987) of morphological differences between populations of Simulium sirbanum Vajime & Dunbar in West Africa was confirmed cytotaxonomically. Populations with adult males having Type III scutal patterns were associated with male larvae heterozygous for the inversion IS-3, whereas adult males with Type IV scutal patterns were associated with larvae homozygous for IS-3. The taxonomic implications are briefly discussed but the evidence is considered to be inappropriate for resurrecting S. sudanense Vajime & Dunbar. Aspects of the biology of the two forms and their importance in onchocerciasis transmission are discussed.     

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     

尼日利亚西南部奥孙河沿线黑蝇Simulium damnosum Theobald complex的形态分类研究(双翅目: 蚋科)（英文）

黑蝇Simulium damnosum sensu lato是由多个姊妹种组成的复合体,这些种在生态学和盘尾丝虫病的传播方面各不相同。本文对奥孙河沿线的尼日利亚西南部森林区域的黑蝇S. damnosum s.l.复合体的组成以及成虫的形态学特征进行了研究。本研究所用的黑蝇S. damnosum s.l.成虫标本来源于奥孙河沿线的3个区,分别是Osun Eleja, Osun Ogbere和Osun Budepo。标本采集通过人体诱捕的方式,采集时间从2008年2月至2009年6月上午7:00到下午6:00,每两周采集1次。通过观察成虫的翅毛簇（wing tufts）和其他形态特征,对其进行分类研究。结果表明：存在同域分布的森林种和稀树草原种。在奥孙河沿线3个区内,森林种为优势种,占总捕获量的99.18%,而稀树草原种仅占0.82%;森林种和稀树草原种在多度上存在显著差异(P<0.05)。所捕获的所有稀树草原种的翅毛簇均为灰白色,而捕获的森林种的翅毛簇颜色存在显著差异(P<0.05)。为了更清楚地揭示该地区黑蝇S. damnosum s.l.的物种组成,建议进行更深入的研究。     

Longitudinal evaluation of repellent activity of Ocimum gratissimum (Labiatae) volatile oil against Simulium damnosum

To determine the repellent activity of Ocimum gratissimum volatile oil against Simulium damnosum (blackflies), a 12 month (January-December 2003) field study was conducted in three onchocerciasis endemic communities (Idomido, Obio camp, and Ikot Adaha) in Ini Local Government Area of Akwa Ibom State, Nigeria. The result revealed that topical application of 20% (v/v) concentration of the oil with liquid paraffin as a base, reduced the biting rate of S. damnosum by 90.2, 81.6, and 79.7%, in Idomido, Obiocamp, and Ikot Adaha respectively. The oil gave protection against the bite of S. damnosum for at least 3 h. A total of 710 adults S. damnosum were caught by individuals treated with Ocimum oil, as against 4296 caught by the control group. When the flies caught by the treated individuals were dissected none of them was infected with microfilariae of Onchocerca volvulus. Human-vector contact and onchocerciasis transmission could be reduced by the topical application of the volatile oil during the peak biting periods of the vector     

Temephos-resistant larvae of Simulium sanctipauli associated with a distinctive new chromosome inversion in untreated rivers of south-western Ghana

Larvae of the Simulium damnosum Theobald complex (Diptera: Simuliidae) were sampled in June 1996 from two sites in south-west Ghana where larviciding has not been applied: Sutri Rapids on the Tano river (05 degrees 23 minutes N 02 degrees 38 minures W) and Sekyere-Heman on the Pra river (05 degrees 11 minutes N 01 degrees 35 minutes W). All specimens were identified as Simulium sanctipauli Vajime & Dunbar sensu stricto (Diptera: Simuliidae). Bioassays with temephos (organophosphorus larvicide employed by the Onchocerciasis Programme for systematic treatment of most rivers across West Africa since the 1970s) showed about five-fold resistance in the Tano population (LC95 2.37-3.14 mg/L) and slight tolerance to temephos in the Pra population (LC95 0.67-0.76 mg/L), vs. the diagnostic concentration of 0.625 mg/L. Larval salivary polytene chromosomes of S. sanctipauli showed fixed inversions 1S-24/24, standard IIL-6 and a new inversion IL/36 polymorphism at Sutri on the Tano. These karyotype characteristics differ from those of temephos-resistant S. sanctipauli in rivers of C te d'Ivoire and other sites on the Tano in Ghana. Thus, temephos resistance in S. sanctipauli at Sutri is associated with distinct chromosomal configurations, showing that immigration was unlikely. This resistance could have been locally selected by exposure of S. sanctipauli larval populations to agrochemicals run-off from cocoa, coffee and oil plantations flanking the rivers.     

Population relationships of Simulium yahense and Simulium sanctipauli in the Firestone Rubber Plantation at Harbel, Liberia

Comparative genetic differences for the phosphoglucomutase and trehalase loci were surveyed in larval and adult blackfly populations of the onchocerciasis vectors Simulium yahense Vajime and Dunbar, and S.sanctipauli Vajime and Dunbar. Genotype frequencies for each stage and all populations were at Hardy-Weinberg equilibrium, indicating that S.yahense and S.sanctipauli remain genetically distinct. S.yahense populations from three different locations were found to be genotypically comparable. The larval population of S.yahense found in closest proximity to a S.sanctipauli population was found to express the phosphoglucomutase allele 1.33, characteristic of S.sanctipauli, with significantly greater frequency than other larval populations of S.yahense. This may constitute evidence of limited genetic introgression of S.sanctipauli with the S.yahense population.     

Poor genetic differentiation but clear cytoform divergence among cryptic species in Simulium damnosum complex (Diptera: Simuliidae)

A complex cryptic species taxonomy has been developed for the flies that vector Onchocerca volvulus L euckart in A frica. This taxonomy names cytoforms of Simulium damnosum T heobald s.l. based on chromosomal inversion polymorphisms. Researchers have linked variation in cytoforms to variation in geographical distributions and biological traits, but have failed to demonstrate that many of these cytoforms are differentiated genetically or morphologically. This may be explained by recent divergence and incomplete lineage sorting of ancestral polymorphisms. Alternatively, it may be the outcome of ongoing gene flow, with inversions maintaining sets of co‐adapted alleles and selection driving inversion frequency differences across habitats. Here, our goals were to test species delimitations in the S. damnosum complex, and determine if the poor differentiation between these species is better explained by the ancestral polymorphisms or gene‐flow hypotheses. Geometric morphometric analysis and more intensive genetic sampling of populations in N igeria only reinforced the lack of clear divergence among S. damnosum complex cytoforms. By contrast, the first statistical estimates of evolutionary relationships based on chromosomal inversion polymorphism data revealed clear signs of divergence, which lends support to the ancestral polymorphisms hypothesis. But the latter analyses may be biased by non‐neutral evolution of inversion polymorphisms; more rigorous tests of cytoform‐based species hypothesis will require deeper sampling across populations and neutral genetic markers along with explicit modelling of genetic isolation and migration.     

Variability according to the individual of the efficacy of repellents with a base of vegetable oils in the individual protection against Simulium damnosum s. l. bites

In Ivory Coast, treatments against black flies larva are interrupted since December 1998 and these flies are back everywhere in abundance. They are now a severe source of discomfort, so it is therefore very important to propose to rural populations a personal protection method against their bites. This article is about the individual attraction of fly boys on the efficiency of lotions and creams of the different repellents (coconut, palm nut, gobi). Our works were carried out in savanna zone (Niakaramandougou) and in forest zone (Soubré) in Ivory Coast. The methodology consisted in capturing black flies on volunteers whose legs were rubbed with repellent. Coconut, palm nut and gobi formulas (lotions and creams) were tested on Simulium damnosum s.l. During each session, which lasted from 07:00 am to 12:00 and from 03:00 pm to 06:00 pm, one person without repellent was used as a control person. The results show that the least efficient repellent (gobi) ensures nonetheless a reduction of 90% of the number of bites that an individual would have received if he was not protected by the repellent. Experimentations allows to say that for a given repellent and formulas (lotion and cream), there is no significant difference between the number of Simulium damnosum s. l. bites received by fly boys.     

Immunity to Onchocerca lienalis microfilariae in mice. II. Effects of sensitization with a range of heterologous species

The model of Onchocerca lienalis microfilariae (mff) injected into CBA and T.O. strains of mice has been used to examine immunity to skin-dwelling microfilariae following exposure to a range of species of helminths. Mice which had received a primary infection with O. cervicalis mff were significantly resistant to challenge with O. lienalis mff (58% reduction relative to challenge controls). Immunization with the uterine contents (eggs and mff) of O. lienalis, O. gutturosa or O. volvulus conferred equivalent levels of protection against challenge with O. lienalis mff (66 to 75%). Similar results were obtained with immunizations in mice that employed either fresh or freeze-killed eggs of O. gutturosa. Significant reductions in the recoveries of O. lienalis mff were also demonstrated following the intraperitoneal implantation of adult male worms of O. gutturosa (30 to 52%), the adults of either sex of Dipetalonema viteae (60%), or after infection with Trichinella spiralis (27 to 81%). Infections with the trematode, Schistosoma mansoni, had a negligible effect on mff recoveries. It is concluded that partial resistance in mice to Onchocerca mff may be stimulated by factors, yet to be determined, that are neither stage nor species-specific.     

Ecology of Stomoxys flies (Diptera: Muscidae) in Gabon. I. First survey in different ecological areas

The stomoxyine flies are hematophagous diptera and potential vectors of various pathogenic agents. Like those of the Afrotropical Region, the stomoxyine flies of Gabon remain nearly unknown. For these reasons, an entomological survey was conducted in a transverse way in eight localities representative of the various ecological zones of Gabon. The survey was based on the use of Vavoua traps. Various environmental factors able to influence the captures were noticed and included into a canonical correspondence analysis. In total, 15,966 Stomoxys spp., belonging to seven species or subspecies, were captured. The apparent densities (DAP) expressed as the number of flies per trap and per day, were highest in Franceville (41), Bakoumba (40), Makokou (25) and Mouila (21). The most abundant species were S. n niger (33.4%), S. transvittatus (33%), then S. calcitrans (17%). The principal factors that could explain the variability of the captures were the degree of anthropisation, the botanical facies (savanna or forest), the presence of wild and domestic fauna and the nature of the vegetal cover of the ground. S. calcitrans, S. niger niger were abundant in the areas where human presence was manifest. S. xanthomelas was present in forest belts. S. transvittatus, S. omega, S. inornatus were ubiquitous species. S. niger bilineatus was found in savannas areas.     

Completion of the sequence of the nuclear ribosomal DNA subunit of Simulium sanctipauli,with descriptions of the 18S, 28S genes and the IGS

We describe the IGS-ETS, 18S and 28S ribosomal gene sequences of Simulium sanctipauli Vajime & Dunbar, a member of the S. damnosum Theobald (Diptera: Simuliidae) complex of blackflies (Diptera: Simuliidae). These regions, together with the ITS-1, ITS-2 and 5.8S rDNA presented elsewhere (accession number U36206), constitute the composite sequence of the entire rDNA unit, making S. sanctipauli the second dipteran species of medical importance for which the entire rDNA has been sequenced. Despite the lack of sequence identity, the IGS of S. sanctipauli showed some structural similarities to other Diptera, i.e. the mosquito Aedes albopictus Skuse (Culicidae), the fruitfly Drosophila melanogaster Meigen (Drosophilidae) and the tsetse Glossina (Glossinidae). Two blocks of tandemly repeated subunits were present in the IGS of S. sanctipauli and, unlike other species of Diptera, they contained no duplications of promoter-like sequences. However, two promoter-like sequences were identified in the unique DNA stretches of the IGS by their sequence similarity to the promoter of Aedes aegypti L. (Diptera: Culicidae). The observed sequence variation can be explained, as in the case of Drosophila spp., by the occurrence of slippage-like and point mutation processes, with unequal crossing-over homogenizing (to a certain extent) the region throughout the gene family and blackfly population. The 18S and 28S rDNA genes show more intraspecific variability within the expansion segments than in the core regions. This is also the case in the interspecific comparison of these genes from S. sanctipauli with those of Simulium vittatum, Ae. albopictus and D. melanogaster. This pattern is typical of many eukaryotes and likely to be the result of a more relaxed functional selection in the expansion segments than on the core regions. The A + T content of the S. sanctipauli genes is high and similar to those of other Diptera. This could be the result of a change in the mutation pressure towards AT in the Diptera lineage.     

Sibling species distributions of the Simulium damnosum complex in the West African Onchocerciasis Control Programme area during the decade 1984–93, following intensive larviciding since 1974

Abstract .During the decade from 1984 to 1993, nine species of the Simulium damnosum complex of blackflies (Diptera: Simuliidae) were identified from the area covered by the Onchocerciasis Control Programme. These were S. damnosum s.s., S. dieguerense, S. konkourense, S. leonense, S. sanctipauli, S. sirbanum, S. soubrense, S. squamosum, and S. yahense. Some of these species were found to consist of two chromosomal variant populations. These were S. konkourense‘Konkouré' and ‘Menankaya' forms, S. sanctipauli sensu stricto and‘Djodji' form, S. soubrense‘Chute Milo' and ‘Beffa' forms. The distribution of these twelve cytological taxa was assessed in relation to the two main vegetation zones of West Africa (forest and savanna), topography, river size and other factors. The range of each species was influenced by seasonal climatic changes in wind movement and river water level. The most widely distributed species were S. sirbanum and S. damnosum s.s., associated with savanna areas, recorded from all river basins. Simulium dieguerense was restricted mainly to Western Mali on the Rivers Bafing and Bakoye in the Senegal River basin. Simulium squamosum was identified from rivers draining mountainous areas in both the forest and savanna zones. Simulium yahense was found in small permanent rivers along a wide forested band parallel to the coast and was absent from the plains of Togo and Benin. Members of the S. sanctipauli subcomplex had restricted distributions except for S. sanctipauli s.s., which was widespread in large rivers of the forest zone from Sierra Leone to the Volta Lake in Ghana. Simulium soubrense‘Beffa' form occurred in Togo and Benin, S. soubrense‘Chutes Milo' form in Guinea, both ‘Konkouré' and ‘Menankaya' forms of S. konkourense occurred predominantly in Guinea and S. leonense in Sierra Leone. The relevance of the distribution maps and the importance of the data bank to vector control larvicidal operations are discussed.     

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.