Thierry Baldet

Cirad - La recherche agronomique pour le développement, Montpelhièr, Languedoc-Roussillon, France

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Publications (88)158.18 Total impact

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    ABSTRACT: Biting midges of the genus Culicoides (Diptera, Ceratopogonidae) are biological vectors of internationally important arboviruses. To understand the role of Culicoides in the transmission of these viruses, it is essential to correctly identify the species involved. Within the western Palaearctic region, the main suspected vector species, C. obsoletus, C. scoticus, C. dewulfi and C. chiopterus, have similar wing patterns, which makes it difficult to separate and identify them correctly.
    Parasites & Vectors 01/2014; 7:223. · 3.25 Impact Factor
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    ABSTRACT: The swarming behaviour of natural populations of Anopheles gambiae and An. coluzzii (formerly known as An. gambiae S and M forms, respectively) were investigated through longitudinal surveys conducted between July 2006 and October 2009 in two rural areas of south-western Burkina Faso where these forms are sympatric. In both sites, the majority of swarms were recorded above visual markers localised among houses. In Soumousso, a wooded area of savannah, 108 pairs caught in copula from 205 swarms were sampled; in VK7, a rice growing area, 491 couples from 250 swarms were sampled. If segregated swarms were the norm in both sites, many visual markers were shared by the two forms of An. gambiae. Furthermore, mixed swarms were collected annually in frequencies varying from one site to another, though no mixed inseminations were recorded, corroborating the low hybrid rate previously reported in the field. The occurrence of inter-specific mate-recognition mechanisms, which allow individuals to avoid hybridisation, is discussed.
    Acta Tropica. 01/2014; 130:24–34.
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    ABSTRACT: SUMMARY Bluetongue is a disease of major economic concern in Europe. Its causative agent, bluetongue virus (BTV), is transmitted by several Culicoides species (mainly Culicoides imicola and Culicoides obsoletus in Europe). The application of insecticides on animals may reduce transmission of BTV, however, no formulation is currently licensed specifically against Culicoides midges. The present study assesses the susceptibility of C. obsoletus to deltamethrin using an adapted World Health Organization (WHO) susceptibility test. Midges were exposed to different dosages of deltamethrin for 1 h, and mortality after 1 h and 24 h was recorded. Results indicated that deltamethrin is highly toxic to C. obsoletus since a dose of 1·33×10-4% was enough to kill 50% of the population (LD50) in 24 h. The deltamethrin concentration needed to kill 90% of the population (LD90) was 5·55×10-4%. The results obtained in the present work could help to create a system that can be used to assess insecticide resistance and susceptibility of Culicoides biting midges.
    Parasitology 11/2013; · 2.36 Impact Factor
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    ABSTRACT: Rift Valley fever (RVF) is a severe mosquito-borne disease that is caused by a Phlebovirus (Bunyaviridae) and affects domestic ruminants and humans. Recently, its distribution widened, threatening Europe. The probability of the introduction and large-scale spread of Rift Valley fever virus (RVFV) in Europe is low, but localized RVF outbreaks may occur in areas where populations of ruminants and potential vectors are present. In this study, we assumed the introduction of the virus into Italy and focused on the risk of vector-borne transmission of RVFV to three main European potential hosts (cattle, sheep and goats). Five main potential mosquito vectors belonging to the Culex and Aedes genera that are present in Italy were identified in a literature review. We first modelled the geographical distribution of these five species based on expert knowledge and using land cover as a proxy of mosquito presence. The mosquito distribution maps were compared with field mosquito collections from Italy to validate the model. Next, the risk of RVFV transmission was modelled using a multicriteria evaluation (MCE) approach, integrating expert knowledge and the results of a literature review on host sensitivity and vector competence, feeding behaviour and abundance. A sensitivity analysis was performed to assess the robustness of the results with respect to expert choices. The resulting maps include (i) five maps of the vector distribution, (ii) a map of suitable areas for vector-borne transmission of RVFV and (iii) a map of the risk of RVFV vector-borne transmission to sensitive hosts given a viral introduction. Good agreement was found between the modelled presence probability and the observed presence or absence of each vector species. The resulting RVF risk map highlighted strong spatial heterogeneity and could be used to target surveillance. In conclusion, the geographical information system (GIS)-based MCE served as a valuable framework and a flexible tool for mapping the areas at risk of a pathogen that is currently absent from a region.
    Transboundary and Emerging Diseases 11/2013; 60 Suppl 2:14-23. · 2.10 Impact Factor
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    ABSTRACT: The swarming behaviour of natural populations of An. gambiae and An. coluzzii (formerly known as An. gambiae M and S forms, respectively) were investigated through longitudinal surveys conducted between July 2006 and October 2009 in two rural areas of south-western Burkina Faso where these forms are sympatric. In both sites, the majority of swarms were recorded above visual markers localized among houses. In Soumousso, a wooded area of savannah, 108 pairs caught in copula from 205 swarms were sampled; in VK7, a rice growing area, 491 couples from 250 swarms were sampled. If segregated swarms were the norm in both sites, many visual markers were shared by the two forms of An. gambiae. Furthermore, mixed swarms were collected annually in frequencies varying from one site to another, though no mixed inseminations were recorded, corroborating the low hybrid rate previously reported in the field. The occurrence of inter-specific mate-recognition mechanisms, which allow individuals to avoid hybridization, is discussed.
    Acta tropica 10/2013; · 2.79 Impact Factor
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    ABSTRACT: The swarm structure of two sibling species, An. gambiae coluzzii and An. melas, was characterize to explore the ecological and environmental parameters associated with the formation of swarms and their spatial distribution. Swarms and breeding sites were searched and sampled between January and December 2010, and larval and adult samples were identified by PCR. During the dry season, 456 swarms of An. gambiae s.l. were sampled from 38 swarm sites yielding a total of 23,274 males and 76 females. Of these 38 swarming sites, 18 were composed exclusively of An. gambiae coluzzii and 20 exclusively of An. melas, presenting clear evidence of reproductive swarm segregation. The species makeup of couples sampled from swarms also demonstrated assortative mating. The swarm site localization was close to human dwellings in the case of the An. gambiae coluzzii and on salt production sites for An. melas. At the peak of the rainy season, swarms of An. melas were absent. These findings offer evidence that the ecological speciation of these two sibling species of mosquitoes is associated with spatial swarm segregation and assortative mating, providing strong support for the hypothesis that mate recognition is currently maintaining adaptive differentiation and promoting ecological speciation. Further studies on the swarming and mating systems of An. gambiae, with the prospect of producing a predictive model of swarm distribution, are needed to inform any future efforts to implement strategies based on the use of GMM or SIT.
    Acta tropica 10/2013; · 2.79 Impact Factor
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    ABSTRACT: The swarming behavior of natural populations of Anopheles arabiensis was investigated by conducting transect surveys on 10 consecutive days, around dusk, from March to April and from September to October 2012 in Dioulassoba, a district of Bobo-Dioulasso city in Burkina Faso (West Africa). Swarms were observed outside, around identified larval breeding sites on the banks of the Houet River, as well as in the open-air courtyards found at the centre of many homes in the region. Swarms were found to occur in open sunlit spaces, mostly located above physical or visual cues somehow visually distinct from the surrounding area. Overall 67 and 78 swarms were observed, respectively, during the dry season (March-April) and the rainy season (September-October) of 2012, between 1.5 and 4.5 meters above the ground at their centre. 964 mosquitoes were collected and analyzed from dry season swarms, of which most were male, and all were An. arabiensis, as were the few resting mosquitoes collected indoors. Larvae collected from breeding sites found on the banks of the Houet River mostly consisted of An. arabiensis and only a minority of Anopheles coluzzii (formerly identified as An. gambiae M form). Of 1694 mosquitoes analyzed from 78 swarms in the rainy season collections, a few An. gambiae (formerly known as An. gambiae S form) males were identified, and the remainders were An. arabiensis. The majority of larvae collected during the wet season from the same breeding sites were identified as An. arabiensis followed by An. coluzzii and An. gambiae. The same pattern of species composition was observed in resting mosquitoes, though the proportion of An. arabiensis was less overwhelming. These data support the conclusion that An. arabiensis is the most prevalent species in this area, though the difference in species composition when using different population sampling techniques is noteworthy. Further studies are required for more detailed investigations of male dispersal, feeding behaviour and mating patterns in this urban setting.
    Acta tropica 10/2013; · 2.79 Impact Factor
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    ABSTRACT: Before the release of genetically-modified or sterile male mosquitoes in an attempt to control local populations of malaria vectors, it is crucial to determine male traits involved in mating success. The effects of male size and age as determinants of male mating success in Anopheles gambiae s.s. were measured in the field and under laboratory conditions in Burkina Faso. First, the body sizes (estimated by wing length) of mating, swarming, and indoor-resting male mosquitoes were compared over a 3-yr period (2006-2009) from July to October in Soumousso and Vallée du Kou, two villages in western Burkina Faso. Second, the age structure of swarming and resting male mosquitoes were characterized based on the number of spermatocysts and the proportion of sperm in the reservoir of wild-caught male testis. Third, male age effects on the insemination rate of female An. gambiae were investigated in the laboratory. The mean size of males collected in copula was significantly larger than the mean for swarming males and indoor-resting males. The optimum male age for successful insemination of females was 4-8 d. These results suggest that male size is an important trait in determining male mating competitiveness in the field. Although age was not found to be a significant factor in mating competitiveness, it was significantly correlated with swarming behaviors in the field and insemination success in the laboratory. The implications of these results in terms of sexual selection in An. gambiae and vector control programs are further discussed.
    Journal of Medical Entomology 03/2013; 50(2):285-93. · 1.86 Impact Factor
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    ABSTRACT: Since bluetongue outbreaks occurred in november 2007 in northern Spain, an intensive trapping program was carried out to study the diversity and abundance of the Culicoides Latreille (Diptera: Ceratopogonidae) species. Eleven sheep farms and seven natural habitats distributed throughout the Basque country region were sampled using UV light CDC traps between 2008-2011. A total of 348,685 Culicoides specimens belonging to 52 species were collected during 1,480 trappings in 24h periods. An updated checklist of these 52 species (including four new records for the Iberian peninsula) is provided from the Basque country. The most abundant species in sheep farms were the two sibling species of the obsoletus complex (81.8% of the total catches): C. obsoletus (Meigen) and C. scoticus Downes and Kettle. Culicoides lupicaris Downes and Kattle was the next most abundant taxon collected in farms (9.7% of the total). Few specimens of Culicoides imicola Kieffer, the bluetongue vector in Mediterranean Basin and some specimens of Culicoides nubeculosus (Meigen), an important candidate of BTV transmission were collected in northeast and northwest farms, respectively. In natural ecosystems, three species, Culicoides festivipennis Kieffer, Culicoides alazanicus Dzhafarov and Culicoides brunnicans Edwards, comprised 48.5% of the total captures. Culicoides obsoletus/C. scoticus was present throughout the year and even during the winter days in temperate Atlantic areas (Gipuzkoa, Biscay) whereas no catches occurred in winter at the southern farms of Alava, where the climate is much colder in that season. The majority of the species were active from March to November with maximum peaks of catches in summer. Most Culicoides obsoletus/C. scoticus specimens were collected outdoors (65.9%). The study shows greater occurrence and abundance of obsoletus and pulicaris complex in sheep farms and in lesser extent in natural ecosystems. Data about diversity, distribution and seasonal dynamics are also reported improving the knowledge of these Culicoides species in terms of surveillance and prevention for future bluetongue outbreaks.
    Proceedings- Entomological Society of Washington 02/2013; 115:48-69. · 0.39 Impact Factor
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    ABSTRACT: The molecular form composition of Anopheles gambiae Giles s.s. (Diptera: Culicidae) mating swarms and the associated mating pairs (copulae) were investigated during two rainy seasons (July to October, 2005 and July to November, 2006) in the villages of Soumousso and Vallée du Kou (VK7). Although the habitats of these villages differ markedly, sympatric populations of M and S molecular forms of An. gambiae s.s. occur in both places periodically. The main aim was to assess the degree to which these molecular forms mate assortatively. In Soumousso, a wooded savannah habitat, the majority of swarm samples consisted of only S-form males (21/28), although a few M-form males were found in mixed M- and S-form swarms. In VK7, a rice growing area, the majority of swarm samples consisted of only M-form males (38/62), until October and November 2006, when there were nearly as many mixed-form as single-form swarms. Overall, ∼60% of M- and S-form swarms were temporally or spatially segregated; the two forms were effectively prevented from encountering each other. Of the remaining 40% of swarms, however, only about half were single-form and the rest were mixed-form. Of the 33 copulae collected from mixed-form swarms, only four were mixed-form pairs, significantly fewer than expected by random pairing between forms (χ(2) = 10.34, d.f. = 2, P < 0.01). Finally, all specimens of inseminated females were of the same form as the sperm contained within their spermatheca (n = 91), even for the four mixed-form copulae. These findings indicate that assortative mating occurs within mixed-form swarms, mediated most probably by close-range mate recognition cues.
    Medical and Veterinary Entomology 01/2013; · 2.21 Impact Factor
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    ABSTRACT: Culicoides species (Diptera: Ceratopogonidae) belonging to the Obsoletus and Pulicaris groups are considered to be the main vectors of bluetongue virus (BTV) in non Mediterranean Europe. Selected terrestrial microhabitats (n = 17) on a farm in northern Spain were sampled repeatedly over a year-long period and characterized for use by Culicoides species for immature development. Concurrent use of CDC light traps showed the presence of 37 species and 66,575 specimens of adult Culicoides. A total of 28 species and 11,396 individuals emerged from laboratory-maintained soil samples. Culicoides obsoletus and Culicoides scoticus (pooled as Obsoletus complex) were particularly abundant (comprising 58.6% and 74.5% of the total collections in light traps and emergence traps respectively). Potential key vectors of animal viruses (such as BTV) were found in two main terrestrial types of microhabitats. In the case of C. obsoletus, different types of manure (old and composted manure, manure mixed with organic matter, and fresh manure) produced most of the specimens. In contrast, larvae of C. scoticus and Culicoides lupicaris were associated with soil substantially comprised of rotting leaf litter that included the parasitic plant Lathraea clandestina. Several species, Culicoides festivipennis, Culicoides punctatus and Culicoides brunnicans, were very common in mud at pond margins. Indeed, pond microhabitats and runoff below barn rooflines supported the greatest species richness. In the pond habitat, 49.4% of Culicoides specimens emerged from mud at the water edge, as opposed to 50 cm above (32.4%) and 1 meter above waterline (18%). Similar species richness, but statistically significant differences in abundance, were observed among the four pond microhabitats. Overall, the majority of the specimens were found in the upper layer (0–3 cm), except in manure, where they preferred deeper layers (>6 cm). Maximum peaks of abundance occurred in both light traps and soil samples in summer months, whereas increased captures in autumn were noticed only in light traps. Both trapping systems failed to collect adult Culicoides midges in the coldest months of December, January and February. The literature on immature habitats of species suspected in BTV transmission in Europe, the Pulicaris group and particularly the Obsoletus group, is briefly reviewed.
    Veterinary Parasitology 01/2013; 191(s 1–2):81–93. · 2.38 Impact Factor
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    ABSTRACT: [This corrects the article on p. e48120 in vol. 7.].
    PLoS ONE 01/2013; 8(4). · 3.53 Impact Factor
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    ABSTRACT: PermaNet 3.0 was evaluated against Culex quinquefasciatus in experimental huts in Lomé. Endpoints were deterrence, exophily, blood feeding inhibition, and mortality. Insecticide susceptibility of Culex quinquefasciatus was assessed with permethrin (1%), DDT (4%), bendiocarb (0.1%), deltamethrin (0.5%, 0.05%), carbosulfan (0.4%), and chlorpyrifos methyl (0.4%). Total of 1,223 Cx. quinquefasciatus females were collected. PermaNet 3.0 unwashed deterred 16.84% Culex mosquitoes. After 20 washes, it deterred 5.79% mosquitoes compared to 6.84% deterrence by unwashed PermaNet 2.0. PermaNet 3.0 induced mosquitoes to exit huts 50.48% and inhibited blood feeding 70.97% in unwashed state. After 20 washes, the net induced 42.91% mosquitoes to exit and inhibited 67.06% mosquitoes from blood feeding. PermaNet 3.0 gave 76% personal protection at zero wash and 69% protection after 20 washes. The net retained 7.1% insecticidal effect in the unwashed state and after 20 washes 6.5%. Cx. quinquefasciatus was resistant to tested insecticides (6%–50% mortality). PermaNet 3.0 is a good control tool against mosquitoes. However, Cx. quinquefasciatus is less affected by PermaNet 3.0. The evaluation depicts the success of vector control innovations using pyrethroids and nonpyrethroids in combination on nets. Additional studies with Culex species are recommended to know whether the difference in blood feeding is interspecific (difference in vector behaviour) and not due to intertrial variability.
    ISRN Infectious Diseases. 12/2012; 2013.
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    ABSTRACT: We investigated the occurrence of the L1014F and L1014S kdr mutations in malaria vector populations in Burkina Faso (West Africa). A cross sectional survey was conducted at 10 sites all located in cotton cultivation areas which are assumed to be the major insecticide resistance selection foci in Burkina Faso. The hot ligation method was used to detect the two kdr mutations in field collected Anopheles gambiae s.l. samples. For the first time in Burkina Faso the L1014S mutation was identified in both M and S forms of An. gambiae s.s. populations collected from the site of Koupela in the central-eastern region at low frequency. Furthermore, the L1014S mutation was also found in one specimen of An. arabiensis collected from the Dano site. The data generated in this study provides additional evidence of the spread of the L1014S mutation into An. gambiae s.l. populations in West Africa. It is now important to evaluate the role of the L1014S mutation in the pyrethroid resistance phenotype and assess its potential impact on the efficacy of pyrethroid-based control measures in West Africa where several resistance mutations now coexist.
    Acta tropica 11/2012; · 2.79 Impact Factor
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    ABSTRACT: Much effort is being devoted for developing new indicators to evaluate the human exposure to Aedes mosquito bites and the risk of arbovirus transmission. Human antibody (Ab) responses to mosquito salivary components could represent a promising tool for evaluating the human-vector contact. To develop a specific biomarker of human exposure to Aedes aegypti bites, we measured IgG Ab response to Ae. aegypti Nterm-34 kDa salivary peptide in exposed children in 7 villages of Southern Benin (West Africa). Results showed that specific IgG response presented high inter-individual heterogeneity between villages. IgG response was associated with rainfall and IgG level increased from dry (low exposure) to rainy (high exposure) seasons. These findings indicate that IgG Ab to Nterm-34 kDa salivary peptide may represent a reliable biomarker to detect variation in human exposure to Ae. aegypti bites. This preliminary study highlights the potential use of Ab response to this salivary peptide for evaluating human exposure to Ae. aegypti. This biomarker could represent a new promising tool for assessing the risk of arbovirus transmission and for evaluating the efficacy of vector control interventions.
    PLoS Neglected Tropical Diseases 11/2012; 6(11):e1905. · 4.57 Impact Factor
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    ABSTRACT: Feeding success depends on host availability, host defensive reactions and host preferences. Host choice is a critical determinant of the intensity at which pathogens are transmitted. The aim of the current study was to describe host preferences of Palaearctic Culicoides species (Diptera: Ceratopogonidae) Latreille using traps baited with the five different host species of poultry, horse, cattle, sheep and goat. Collections were carried out nightly in July and August 2009 in western France with three replicates of a 5 × 5 randomized Latin square (five sites, five hosts). Moreover, an ultraviolet (UV) light/suction trap was operated during host-baited collections to correlate Culicoides biting rates and UV light/suction trap catches. A total of 660 Culicoides belonging to 12 species, but comprised mainly of Culicoides scoticus Downes and Kettle, Culicoides dewulfi Goetghebuer and Culicoides obsoletus Meigen, were collected on animal baits. Abundance was highest for the horse, which accounted for 95% of all Culicoides caught, representing 10 species. The horse, the largest bait, was the most attractive host, even when abundance data were corrected by weight, body surface or Kleiber's scaling factor. Culicoides obsoletus was the only dominant species attracted by birds. Both C. scoticus and C. dewulfi were collected mainly from the upper body of the horse. Finally, the quantification of host preferences allows for discussion of implications for the transmission of Culicoides-borne pathogens such as bluetongue virus.
    Medical and Veterinary Entomology 09/2012; · 2.21 Impact Factor
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    ABSTRACT: Historical studies have indicated that An. gambiae s.s. is the predominant malaria vector species in Bobo-Dioulasso the second biggest city of Burkina Faso (West Africa). However, over the last decade, An. arabiensis appears to be replacing An. gambiae s.s. as the most prevalent malaria vector in this urban setting. To investigate this species transition in more detail the present study aims to provide an update on the malaria vector composition in Bobo-Dioulasso, and also the Plasmodium infection rates and susceptibility to insecticides of the local An. gambiae s.l. population. An entomological survey was carried out from May to December 2008 in Dioulassoba and Kodeni (central and peripheral districts respectively), which are representative of the main ecological features of the city. Sampling consisted of the collection of larval stages from water bodies, and adults by monthly indoor residual spraying (IRS) using aerosol insecticides. Insecticide susceptibility tests were performed using the WHO filter paper protocol on adults emerged from larvae. PCR was used to determine vector species and to identify resistance mechanisms (kdr and ace-1(R)). The Plasmodium infection rate was estimated by ELISA performed on female mosquitoes collected indoors by IRS. An. arabiensis was found to be the major malaria vector in Bobo-Dioulasso, comprising 50 to 100% of the vector population. The sporozoite infection rate for An. arabiensis was higher than An. gambiae s.s. at both Dioulassoba and Kodeni. An. gambiae s.l. was resistant to DDT and cross-resistant to pyrethroids at the two sites with higher levels of resistance observed in An. gambiae s.s. than An. arabiensis. Resistance to 0.1% bendiocarb was observed in the An. gambiae s.s. S form but not the M form or in An. arabiensis. The L1014F kdr mutation was detected in the two molecular forms of An. gambiae s.s. at varying frequencies (0.45 to 0.92), but was not detected in An. arabiensis, suggesting that other mechanisms are involved in DDT resistance in this species. The ace-1(R) mutation was only detected in the S molecular form and was observed at the two sites at similar frequency (0.3). Over the last ten years, An. arabiensis has become the major malaria vector in Bobo-Dioulasso city where it was formerly present only at low frequency. However, the ecological determinant that enhances the settlement of this species into urban and peri-urban areas of Bobo-Dioulasso remains to be clarified. The impact of the changing An. gambiae s.l. population in this region for vector control including resistance management strategies is discussed.
    Parasites & Vectors 06/2012; 5:127. · 3.25 Impact Factor
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    ABSTRACT: Culicoides (Diptera: Ceratopogonidae) associated with livestock were captured using CDC blacklight traps at three BTV-infected farms in Basque country between November 2007 and December 2008. Twenty seven and twenty Culicoides species were respectively caught at outdoors and indoors habitats. Indoors insect community represented 86.11% of the whole captured individual biting midges. Culicoides obsoletus/C. scoticus (two sibling species of the Obsoletus complex) was dominant throughout all months and sexes with maximum phenological peaks in November 2007 and June-July 2008. Culicoides lupicaris was the second dominant species followed by C. pulicaris (both species of the Pulicaris complex). Few specimens of C. imicola, the principal Afro-Mediterranean vector of BTV, as well as four new species records for the Iberian Peninsula, were also caught. BTV was detected by RT-PCR from pools of C. obsoletus/C. scoticus, C. lupicaris and C. pulicaris parous females. DL-Lactic acid significantly attracted more C. obsoletus/C. scoticus females and males, C. lupicaris females, C. pulicaris females and C. punctatus females and males; whereas acetone increased only the captures of C. achrayi.
    Veterinary Parasitology 05/2012; 186:415-424. · 2.38 Impact Factor
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    02/2012; , ISBN: 978-953-307-895-3
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    01/2012: pages 77-102;

Publication Stats

1k Citations
158.18 Total Impact Points

Institutions

  • 2003–2014
    • Cirad - La recherche agronomique pour le développement
      • Centre de Coopération Internationale en Recherche Agronomique pour le Développement (CIRAD)
      Montpelhièr, Languedoc-Roussillon, France
  • 2013
    • University of Abomey-Calavi
      Kotonu, Littoral, Benin
  • 2012–2013
    • Neiker-Tecnalia Basque Institute for Agricultural Research and Development
      Vitoria, Basque Country, Spain
  • 2008–2013
    • Centre de Recherche Entomologique de Cotonou
      Kotonu, Littoral, Benin
  • 2008–2012
    • Research Institute of Health Sciences
      Wagadugu, Centre, Burkina Faso
  • 2009–2011
    • Intelligent Insect Control
      Castelnau, Languedoc-Roussillon, France
    • London School of Hygiene and Tropical Medicine
      Londinium, England, United Kingdom
  • 1997
    • Pasteur Center of Cameroon
      Jaúnde, Centre Region, Cameroon