Anopheles gambiae complex and disease transmission in Africa

Abstract

Anopheles gambiae complex mosquitoes are present throughout tropical Africa and its offshore islands. Recent work has shown that at least 6 cryptic or "sibling" species are involved. They comprise 2 salt-water species, A. melas and A. merus, 3 freshwater species—provisionally known as species A, B, and C, and a mineralwater species known as species D. Artificial inter-species crosses yield sterile hybrid males. Rarity of hybridization in nature proves the reproductive isolation and valid genetic barriers between these 6 species. Morphological identification of most individuals of both saltwater and the mineralwater species is possible for larvae and adult females, using meristic features and other variable dimensions and ratios. Differential identification of the 3 freshwater species relies almost completely on cytotaxonomic methods. Species A and B occur together in most areas, extending southwards to sub-tropical latitudes and eastward to Mauritius. Proportions of mixed A-B populations may depend directly or indirectly on relative humidity, with A favoured when nocturnal humidities approach saturation. Species B is often absent from areas of highest humidity, but thrives in relatively arid savannas and steppes. Species C and D have relict distributions. Both saltwater species are coastal: melas in West Africa and merus in East Africa and larger islands except Zanzibar; merus also spreads inland. Apart from species C, which is always zoophilic, all members of the complex are proven or probable vectors of human malaria and filariasis, but with some wide contrasts in their levels of vectorial efficiency. Transmission of some arboviruses (Tataguine, O'nyong-nyong) is associated with species B, and perhaps with A also. Species B may transmit setariasis of cattle; melas and merus may also carry enzootic filariae. Much of the confusing ecophenotypic plasticity of A. gambiae sensu lato is attributable to the differential biological characteristics of these 6 species with their dissimilar geographical distributions, behavioural contrasts and asynchronous population dynamics. Shifts in the species balance occur regularly between A and B and between freshwater and saltwater populations. Species C does not interchange so much with B under natural conditions, but may survive at high densities after control of A or B. Additional versatility is caused by genetic polymorphism in some of the species, notably B. This species is the most widespread, and individual females tend to be either endophilic or exophilic, anthropophagic or zoophagic, early biters or late biters, and doubtless other alternatives, according to the arrangement of their floating chromosome inversions. Control measures have to be considered separately for each of the sibling species. House spraying with residual insecticides against endophilic species A is possibly sufficient to break disease transmission (assuming favourable response of ancillary vectors) and even to eradicate A completely. Efficacy of this method against other species in the complex is reduced by their exophily, which can be enhanced by behaviouristic avoidance due to the extremely low threshold of irritability exhibited by gambiae s.l. adults in general. Genetic polymorphism of species B may lead to true behaviouristic resistance. Larvicidal control of species A and B is beneficial, but made operationally difficult by their tendency to utilize temporary breeding-sites. Effectiveness of DDT and cyclodiene insecticides is further limited by the development of physiological insecticide resistance by species A and B. Adequate control of both saltwater species, and probably the mineralwater species also, can be attained by antilarval measures. Prospects of reduction of malaria and filariasis where melas, merus and species A and D are principal vectors may be much better than many people imagine. Widespread prevalence and inherent resilience of species B represents an insurmountable public health obstacle at present. Continued research may provide new control methods for integrated use against these mosquitoes. Concepts such as seeding breeding sites with pathogenic microsporidians or fungi, releasing sterile hybrid males or chemosterilized males, and other even more elaborate genetic control techniques, may be of special relevance to control of gambiae complex mosquitoes and diseases they transmit.