Over the past two decades, control efforts have halved malaria cases globally, yet burdens remain high in much of Africa and elimination has not been achieved even where extreme reductions have occurred over many years, such as in South Africa 1,2 . Studies seeking to understand the paradoxical persistence of malaria in areas where surface water is absent for 3-8 months of the year, suggested that certain Anopheles mosquitoes employ long-distance migration 3 . Here, we confirmed this hypothesis by aerial sampling of mosquitoes 40-290 m above ground, providing the first evidence of windborne migration of African malaria vectors, and consequently the pathogens they transmit. Ten species, including the primary malaria vector Anopheles coluzzii, were identified among 235 anophelines captured during 617 nocturnal aerial collections in the Sahel of Mali. Importantly, females accounted for >80% of all mosquitoes collected. Of these, 90% had taken a blood meal before their migration, implying that pathogens will be transported long distances by migrating females. The likelihood of capturing Anopheles species increased with altitude and during the wet seasons, but variation between years and localities was minimal. Simulated trajectories of mosquito flights indicated mean nightly displacements of up to 300 km for 9-hour flight durations. Annually, the estimated numbers of mosquitoes at altitude crossing a 100-km line perpendicular to the winds included 81,000 An. gambiae s.s., 6 million An. coluzzii, and 44 million An. squamosus. These results provide compelling evidence that millions of previously blood-fed, malaria vectors frequently migrate over hundreds of kilometers, and thus almost certainly spread malaria over such distances. Malaria elimination success may, therefore, depend on whether sources of migrant vectors can be identified and

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Long-distance migration of insects impacts food security, public health, and conservation–issues that are especially significant in Africa. Windborne migration is a key strategy enabling exploitation of ephemeral havens such as the Sahel, however, its knowledge remains sparse. In this first cross-season investigation (3 years) of the aerial fauna over Africa, we sampled insects flying 40–290 m above ground in Mali, using nets mounted on tethered helium-filled balloons. Nearly half a million insects were caught, representing at least 100 families from thirteen orders. Control nets confirmed that the insects were captured at altitude. Thirteen ecologically and phylogenetically diverse species were studied in detail. Migration of all species peaked during the wet season every year across localities, suggesting regular migrations. Species differed in flight altitude, seasonality, and associated weather conditions. All taxa exhibited frequent flights on southerly winds, accounting for the recolonization of the Sahel from southern source populations. “Return” southward movement occurred in most taxa. Estimates of the seasonal number of migrants per species crossing Mali at latitude 14°N were in the trillions, and the nightly distances traversed reached hundreds of kilometers. The magnitude and diversity of windborne insect migration highlight its importance and impacts on Sahelian and neighboring ecosystems.

Tracking mosquitoes using current methods of mark–release–recapture are limited to small spatial and temporal scales exposing major gaps in understanding long‐range movements and extended survival. Novel approaches to track mosquitoes may yield fresh insights into their biology which improves intervention activities to reduce disease transmission. Stable isotope enrichment of natural mosquito breeding sites allows large‐scale marking of wild mosquitoes absent human handling. Mosquito larvae that develop in 2H‐enriched water are expected to be detectable for over 4 months using tissue mass fraction 2H measurements, providing opportunities for long‐term mark–capture studies on a large scale. A laboratory study followed by a field experiment of mosquito larval habitat 2H enrichment was conducted in Mali, to evaluate potential labelling of wild mosquitoes. Twelve natural larval sites were enriched using [2H] deuterium oxide (D2O, 99%). Enrichment level was maintained by supplementation following dilution by rains. Availability of 2H to mosquito larvae was enhanced by locally collected and cultured microorganisms (i.e. protozoa, algae and bacteria) reared in deuterated water, and provided as larval diet. Putative natural predators were removed from the larval sites and first instar larvae Anopheles gambiae s.l. larvae were added every other day. Emergence traps enabled collection of eclosing adults. Adult mosquitoes were kept at laboratory conditions for analysis of label attrition with age. Deuterium enrichment of wild mosquitoes above background levels (maximum = 143.1 ppm) became apparent 5–6 days after initial exposure, after which 2H values increased steadily until c. 24 days later (to a mean of c. 220 ppm). Anopheles and Culex mosquitoes showed significantly different 2H values (211 and 194.2 ppm, respectively). Both genera exhibited exponential label attrition (e(‐x)) amounting to 21.6% by day 30 post‐emergence, after which attrition rate continuously decreased. Males of both taxa exhibited a higher mean 2H value compared to females. Deuterium oxide proved useful in marking mosquitoes in their natural larval sites and although costly, may prove valuable for studies of mosquitoes and other aquatic insects. Based on our field study, we provide a protocol for marking mosquito larval sites using deuterium oxide.

Recent results of high-altitude windborne mosquito migration raised questions about the viability of these mosquitoes despite ample evidence that many insect species, including other dipterans, have been known to migrate regularly over tens or hundreds of kilometers on high-altitude winds and retain their viability. To address these concerns, we subjected wild Anopheles gambiae s.l. Giles mosquitoes to a high-altitude survival assay, followed by oviposition (egg laying) and blood feeding assays. Despite carrying out the survival assay under exceptionally harsh conditions that probably provide the lowest survival potential following high altitude flight, a high proportion of the mosquitoes survived for 6- and even 11-h assay durations at 120- to 250-m altitudes. Minimal differences in egg laying success were noted between mosquitoes exposed to high altitude survival assay and those kept near the ground. Similarly, minimal differences were found in the female’s ability to take an additional bloodmeal after oviposition between these groups. We conclude that similar to other high-altitude migrating insects, mosquitoes are able to withstand extended high-altitude flight and subsequently reproduce and transmit pathogens by blood feeding on new hosts.