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1.  The Sequential Aerosol Technique: A Major Component in an Integrated Strategy of Intervention against Riverine Tsetse in Ghana 
Background
An integrated strategy of intervention against tsetse flies was implemented in the Upper West Region of Ghana (9.62°–11.00° N, 1.40°–2.76° W), covering an area of ≈18,000 km2 within the framework of the Pan-African Tsetse and Trypanosomosis Eradication Campaign. Two species were targeted: Glossina tachinoides and Glossina palpalis gambiensis.
Methodology/Principal Findings
The objectives were to test the potentiality of the sequential aerosol technique (SAT) to eliminate riverine tsetse species in a challenging subsection (dense tree canopy and high tsetse densities) of the total sprayed area (6,745 km2) and the subsequent efficacy of an integrated strategy including ground spraying (≈100 km2), insecticide treated targets (20,000) and insecticide treated cattle (45,000) in sustaining the results of tsetse suppression in the whole intervention area. The aerial application of low-dosage deltamethrin aerosols (0.33–0.35 g a.i/ha) was conducted along the three main rivers using five custom designed fixed-wings Turbo thrush aircraft. The impact of SAT on tsetse densities was monitored using 30 biconical traps deployed from two weeks before until two weeks after the operations. Results of the SAT monitoring indicated an overall reduction rate of 98% (from a pre-intervention mean apparent density per trap per day (ADT) of 16.7 to 0.3 at the end of the fourth and last cycle). One year after the SAT operations, a second survey using 200 biconical traps set in 20 sites during 3 weeks was conducted throughout the intervention area to measure the impact of the integrated control strategy. Both target species were still detected, albeit at very low densities (ADT of 0.27 inside sprayed blocks and 0.10 outside sprayed blocks).
Conclusions/Significance
The SAT operations failed to achieve elimination in the monitored section, but the subsequent integrated strategy maintained high levels of suppression throughout the intervention area, which will contribute to improving animal health, increasing animal production and fostering food security.
Author Summary
We document the impact of an integrated strategy of intervention against riverine tsetse flies in the Upper West Region of Ghana within the framework of the Pan-African Tsetse and Trypanosomosis Eradication Campaign, in an area of ≈18,000 km2. The strategy included a sequential aerosol technique (SAT) component, i.e. four applications of low-dosage deltamethrin aerosols, conducted along the three main rivers. The impact of SAT on tsetse densities was monitored in a challenging subsection (dense tree canopy and high tsetse densities) from two weeks before until two weeks after the operations. The SAT operations succeeded in reducing tsetse populations by 98% within one month but fell short of achieving elimination. Insecticide ground spraying, deltamethrin-treated targets and cattle were used as complementary tools to maintain tsetse suppression in the intervention area. An entomological survey conducted one year after SAT operations showed that both target species were still present, albeit at drastically reduced densities as compared to the baseline levels. This integrated strategy of intervention will contribute to improving animal health, increasing animal production and fostering food security in the target area.
doi:10.1371/journal.pntd.0002135
PMCID: PMC3597491
2.  Climate, Cattle Rearing Systems and African Animal Trypanosomosis Risk in Burkina Faso 
PLoS ONE  2012;7(11):e49762.
Background
In sub-Saharan countries infested by tsetse flies, African Animal Trypanosomosis (AAT) is considered as the main pathological constraint to cattle breeding. Africa has known a strong climatic change and its population was multiplied by four during the last half-century. The aim of this study was to characterize the impact of production practices and climate on tsetse occurrence and abundance, and the associated prevalence of AAT in Burkina Faso.
Methodology/Principal Findings
Four sites were selected along a South-north transect of increasing aridity. The study combines parasitological and entomological surveys. For the parasitological aspect, blood samples were collected from 1,041 cattle selected through a stratified sampling procedure including location and livestock management system (long transhumance, short transhumance, sedentary). Parasitological and serological prevalence specific to livestock management systems show a gradual increase from the Sahelian to the Sudano-Guinean area (P<0.05). Livestock management system had also a significant impact on parasitological prevalence (P<0.05). Tsetse diversity, apparent densities and their infection rates overall decreased with aridity, from four species, an apparent density of 53.1 flies/trap/day and an infection rate of 13.7% to an absence at the northern edge of the transect, where the density and diversity of other biting flies were on the contrary highest (p<0.001).
Conclusions/Significance
The climatic pressure clearly had a negative impact on tsetse abundance and AAT risk. However, the persistency of tsetse habitats along the Mouhoun river loop maintains a high risk of cyclical transmission of T. vivax. Moreover, an “epidemic mechanical livestock trypanosomosis” cycle is likely to occur in the northern site, where trypanosomes are brought in by cattle transhuming from the tsetse infested area and are locally transmitted by mechanical vectors. In Burkina Faso, the impact of tsetse thus extends to a buffer area around their distribution belt, corresponding to the herd transhumance radius.
doi:10.1371/journal.pone.0049762
PMCID: PMC3498196  PMID: 23166765
3.  A behavioral mechanism underlying ecological divergence in the malaria mosquito Anopheles gambiae 
Behavioral Ecology  2010;21(5):1087-1092.
Disruptive selection mediated by predation on aquatic immature stages has been proposed as a major force driving ecological divergence and fostering speciation between the M and S molecular forms of the African malaria mosquito, Anopheles gambiae. In the dry savannahs of West Africa where both molecular forms co-occur, the S form thrives in temporary pools filled with rainwater, whereas the M form preferentially breeds in permanent freshwater habitats where predator pressure is higher. Here, we explored the proximal mechanisms by which predation may contribute to habitat segregation between molecular forms using progeny of female mosquitoes captured in Burkina Faso. We show that the S form suffers higher predation rates than the M form when simultaneously exposed to the widespread predator, Anisops jaczewskii in an experimental arena. Furthermore, behavioral plasticity induced by exposure to the predator was observed in the M form, but not in the S form, and may partially explain its habitat use and ecological divergence from the S form. We discuss the role of adaptive phenotypic plasticity in allowing successful colonization of a new ecological niche by the M form and highlight further research areas that need to be addressed for a better understanding of the ultimate mechanisms underlying ecological speciation in this pest of major medical importance.
doi:10.1093/beheco/arq114
PMCID: PMC2920295  PMID: 22476108
adaptation; Anopheles gambiae; behavior; habitat divergence; mosquito; notonectidae; phenotypic plasticity, predation; speciation
4.  Restricted Application of Insecticides: A Promising Tsetse Control Technique, but What Do the Farmers Think of It? 
Background
Restricted application of insecticides to cattle is a cheap and safe farmer-based method to control tsetse. In Western Africa, it is applied using a footbath, mainly to control nagana and the tick Amblyomma variegatum. In Eastern and Southern Africa, it might help controlling the human disease, i.e., Rhodesian sleeping sickness as well. The efficiency of this new control method against ticks, tsetse and trypanosomoses has been demonstrated earlier. The invention, co-built by researchers and farmers ten years ago, became an innovation in Burkina Faso through its diffusion by two development projects.
Methodology/Principal Findings
In this research, we studied the process and level of adoption in 72 farmers inhabiting the peri-urban areas of Ouagadougou and Bobo-Dioulasso. Variables describing the livestock farming system, the implementation and perception of the method and the knowledge of the epidemiological system were used to discriminate three clusters of cattle farmers that were then compared using indicators of adoption. The first cluster corresponded to modern farmers who adopted the technique very well. The more traditional farmers were discriminated into two clusters, one of which showed a good adoption rate, whereas the second failed to adopt the method. The economic benefit and the farmers' knowledge of the epidemiological system appeared to have a low impact on the early adoption process whereas some modern practices, as well as social factors appeared critical. The quality of technical support provided to the farmers had also a great influence. Cattle farmers' innovation-risk appraisal was analyzed using Rogers' adoption criteria which highlighted individual variations in risk perceptions and benefits, as well as the prominent role of the socio-technical network of cattle farmers.
Conclusions/Significance
Results are discussed to highlight the factors that should be taken into consideration, to move discoveries from bench to field for an improved control of trypanosomoses vectors.
Author Summary
Restricted application of insecticides to cattle is a cheap and safe farmer-based method to control tsetse and the diseases they transmit, i.e. human and animal African trypanosomoses. The efficiency of this new control method has been demonstrated earlier but no data is available on its perception and adoption intensity by farmers. We studied these two features in Burkina Faso, where the method has diffused thanks to two development projects. The study allowed identifying three groups of farmers with various adoption intensities, of which one was modern and two traditional. The economic benefit and the farmers' knowledge of the epidemiological system appeared to have a low impact on the early adoption process whereas some modern practices, as well as social factors appeared critical. The quality of technical support provided to the farmers had also a great influence on the adoption rate. The study highlighted individual variations in risk perceptions and benefits, as well as the prominent role of the socio-technical network of cattle farmers. The results of the study are discussed to highlight the factors that should be taken into consideration, to move discoveries from bench to field for an improved control of trypanosomoses vectors.
doi:10.1371/journal.pntd.0001276
PMCID: PMC3153426  PMID: 21858241
5.  Contrasting Population Structures of Two Vectors of African Trypanosomoses in Burkina Faso: Consequences for Control 
Background
African animal trypanosomosis is a major obstacle to the development of more efficient and sustainable livestock production systems in West Africa. Riverine tsetse species such as Glossina palpalis gambiensis Vanderplank and Glossina tachinoides Westwood are the major vectors. A wide variety of control tactics is available to manage these vectors, but their removal will in most cases only be sustainable if the control effort is targeting an entire tsetse population within a circumscribed area.
Methodology/Principal Findings
In the present study, genetic variation at microsatellite DNA loci was used to examine the population structure of G. p. gambiensis and G. tachinoides inhabiting four adjacent river basins in Burkina Faso, i.e. the Mouhoun, the Comoé, the Niger and the Sissili River Basins. Isolation by distance was significant for both species across river basins, and dispersal of G. tachinoides was ∼3 times higher than that of G. p. gambiensis. Thus, the data presented indicate that no strong barriers to gene flow exists between riverine tsetse populations in adjacent river basins, especially so for G. tachinoides.
Conclusions/Significance
Therefore, potential re-invasion of flies from adjacent river basins will have to be prevented by establishing buffer zones between the Mouhoun and the other river basin(s), in the framework of the PATTEC (Pan African Tsetse and Trypanosomosis Eradication Campaign) eradication project that is presently targeting the northern part of the Mouhoun River Basin. We argue that these genetic analyses should always be part of the baseline data collection before any tsetse control project is initiated.
Author Summary
Tsetse flies are insects that transmit trypanosomes to humans (sleeping sickness) and animals (nagana). Controlling these vectors is a very efficient way to control these diseases. In Burkina Faso, a tsetse eradication campaign is presently targeting the northern part of the Mouhoun River Basin. To attain this objective, the approach has to be area-wide, i.e. the control effort targets an entire pest population within a circumscribed area. To assess the level of this isolation, we studied the genetic structure of Glossina palpalis gambiensis and Glossina tachinoides populations in the target area and in the adjacent river basins of the Comoé, the Niger and the Sissili River Basins. Our results suggest an absence of strong genetic isolation of the target populations. We therefore recommend establishing permanent buffer zones between the Mouhoun and the other river basin(s) to prevent reinvasion. This kind of study may be extended to other areas on other tsetse species.
doi:10.1371/journal.pntd.0001217
PMCID: PMC3125141  PMID: 21738812
6.  Feeding behaviour of potential vectors of West Nile virus in Senegal 
Parasites & Vectors  2011;4:99.
Background
West Nile virus (WNV) is a widespread pathogen maintained in an enzootic cycle between mosquitoes and birds with occasional spill-over into dead-end hosts such as horses and humans. Migratory birds are believed to play an important role in its dissemination from and to the Palaearctic area, as well as its local dispersion between wintering sites. The Djoudj Park, located in Senegal, is a major wintering site for birds migrating from Europe during the study period (Sept. 2008- Jan. 2009). In this work, we studied the seasonal feeding behaviour dynamics of the potential WNV mosquito vectors at the border of the Djoudj Park, using a reference trapping method (CDC light CO2-baited traps) and two host-specific methods (horse- and pigeon-baited traps). Blood meals of engorged females were analysed to determine their origin.
Results
Results indicated that Culex tritaeniorhynchus and Cx. neavei may play a key role in the WNV transmission dynamics, the latter being the best candidate bridging-vector species between mammals and birds. Moreover, the attractiveness of pigeon- and horse-baited traps for Cx. neavei and Cx. tritaeniorhynchus varied with time. Finally, Cx. tritaeniorhynchus was only active when the night temperature was above 20°C, whereas Cx. neavei was active throughout the observation period.
Conclusions
Cx. neavei and Cx. tritaeniorhynchus are the main candidate vectors for the transmission of WNV in the area. The changes in host attractiveness might be related to variable densities of the migratory birds during the trapping period. We discuss the importance of these results on the risk of WNV transmission in horses and humans.
doi:10.1186/1756-3305-4-99
PMCID: PMC3118230  PMID: 21651763
8.  Progress towards the eradication of Tsetse from the Loos islands, Guinea 
Parasites & Vectors  2011;4:18.
Background
The tsetse fly Glossina palpalis gambiensis is the main vector of sleeping sickness (Human African Trypanosomiasis - HAT) in West Africa, in particular in littoral Guinea where this disease is currently very active. The Loos islands constitute a small archipelago some 5 km from mainland Guinea, where G. p. gambiensis is well known as a nuisance and potential disease vector by inhabitants of the three main islands, Fotoba, Room, and Kassa. The National Control Program against HAT of Guinea has decided to eradicate tsetse in Loos islands in order to sustainably protect humans and economic activities. After baseline data collection, tsetse control began on the islands in 2006. On each of the three islands a specific combination of control methods was implemented according to the entomological situation found.
Results
Starting densities before control operations were 10, 3 and 1 tsetse/trap/day in Kassa, Room and Fotoba respectively, but by July 2010, tsetse were no longer caught in any of the sentinel traps used for monitoring. The reduction rate was faster where several control methods were implemented as a combination (impregnated traps and targets ITT, selective groundspraying, epicutaneous insecticide treatment of pigs, and impregnated fences around pig pens), whereas it was slower when ITT were used as the only control method.
Conclusions
This 100% suppression is a promising step in the eradication process, but G. p. gambiensis may still occur at very low, undetectable, densities on the archipelago. Next step will consist in assessing a 0.05 probability of tsetse absence to ascertain a provisional eradication status. Throughout these operations, a key factor has been the involvement of local teams and local communities without whom such results would be impossible to obtain. Work will continue thanks to the partners involved until total eradication of the tsetse on Loos islands can be declared.
doi:10.1186/1756-3305-4-18
PMCID: PMC3048576  PMID: 21310074
9.  Population Genetics as a Tool to Select Tsetse Control Strategies: Suppression or Eradication of Glossina palpalis gambiensis in the Niayes of Senegal 
Background
The Government of Senegal has initiated the “Projet de lutte contre les glossines dans les Niayes” to remove the trypanosomosis problem from this area in a sustainable way. Due to past failures to sustainably eradicate Glossina palpalis gambiensis from the Niayes area, controversies remain as to the best strategy implement, i.e. “eradication” versus “suppression.” To inform this debate, we used population genetics to measure genetic differentiation between G. palpalis gambiensis from the Niayes and those from the southern tsetse belt (Missira).
Methodology/Principal Findings
Three different markers (microsatellite DNA, mitochondrial CO1 DNA, and geometric morphometrics of the wings) were used on 153 individuals and revealed that the G. p. gambiensis populations of the Niayes were genetically isolated from the nearest proximate known population of Missira. The genetic differentiation measured between these two areas (θ = 0.12 using microsatellites) was equivalent to a between-taxa differentiation. We also demonstrated that within the Niayes, the population from Dakar – Hann was isolated from the others and had probably experienced a bottleneck.
Conclusion/Significance
The information presented in this paper leads to the recommendation that an eradication strategy for the Niayes populations is advisable. This kind of study may be repeated in other habitats and for other tsetse species to (i) help decision on appropriate tsetse control strategies and (ii) find other possible discontinuities in tsetse distribution.
Author Summary
Tsetse flies transmit trypanosomes to humans (sleeping sickness) and animals (nagana). Controlling these vectors is a very efficient way to contain these diseases. There are several strategies and methods that can be used for control, each being more or less efficient depending on several factors. The Government of Senegal wants to sustainably eliminate trypanosomosis from the Niayes region by controlling the tsetse vector, Glossina palpalis gambiensis. To reach this objective, two different strategies may be used: suppression (decrease in tsetse densities) or eradication (remove all the tsetse in the region until last one). For eradication, the approach has to be area-wide, i.e. the control effort targets an entire pest population within a circumscribed area, to avoid any possible reinvasion. Three different tools (microsatellite DNA, mitochondrial DNA and morphometrics) were used, and all showed an absence of gene flow between G. p. gambiensis from the Niayes and from the nearest known population in the south east of the country (Missira). This genetic isolation of the target population leads to the recommendation that an eradication strategy for the Niayes populations is advisable. This kind of study may be extended to other areas on other tsetse species.
doi:10.1371/journal.pntd.0000692
PMCID: PMC2876113  PMID: 20520795
10.  Learning influences host choice in tsetse 
Biology Letters  2006;3(2):113-117.
A learning capacity for feeding is described in many insect species including vectors of diseases, but has never been reported in tsetse flies (Diptera, Glossinidae), the cyclic vectors of human (sleeping sickness) and animal trypanosomoses in Africa. Repeated feeding on the same host species by a disease vector is likely to increase the within-species disease-transmission risk, but to decrease it between species.
An experiment with cattle and reptiles in a stable provides evidence that the species of host selected for the second blood meal in tsetse flies depends on the host encountered for the first blood meal when the between-meal interval is 2 days. This preference disappears when the between-meal interval is extended to 3 days. The energetic advantages of this acquired preference and its importance in trypanosomoses epidemiology are discussed.
doi:10.1098/rsbl.2006.0578
PMCID: PMC2375919  PMID: 17251119
tsetse flies; trypanosomosis; feeding preference; learning

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