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Knowledge of insect dispersal is relevant to the control of agricultural pests, vector-borne transmission of human and veterinary pathogens, and insect biodiversity. Previous studies in a malaria endemic area of the Sahel region in West Africa revealed high-altitude, long-distance migration of insects and various mosquito species. The objective of the current study was to assess whether similar behavior is exhibited by mosquitoes and other insects around the Lake Victoria basin region of Kenya in East Africa. Insects were sampled monthly from dusk to dawn over 1 year using sticky nets suspended on a tethered helium-filled balloon. A total of 17,883 insects were caught on nets tethered at 90, 120, and 160 m above ground level; 818 insects were caught in control nets. Small insects (<0.5 cm, n = 15,250) were predominant regardless of height compared with large insects (>0.5 cm, n = 2,334) and mosquitoes (n = 299). Seven orders were identified; dipteran was the most common. Barcoding molecular assays of 184 mosquitoes identified 7 genera, with Culex being the most common (65.8%) and Anopheles being the least common (5.4%). The survival rate of mosquitoes, experimentally exposed to high-altitude overnight, was significantly lower than controls maintained in the laboratory (19% vs. 85%). There were no significant differences in mosquito survival and oviposition rate according to capture height. These data suggest that windborne dispersal activity of mosquito vectors of malaria and other diseases occurs on a broad scale in sub-Saharan Africa., (© The Author(s) 2023. Published by Oxford University Press on behalf of Entomological Society of America.)

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., (Published by Oxford University Press on behalf of Entomological Society of America 2020.)

Changes in spatial distribution of mosquitoes over time in a Sahelian village were studied to understand the sources of the mosquitoes during the dry season when no larval sites are found. At that time, the sources of Anopheles gambiae Giles may be local shelters used by aestivating mosquitoes or migrants from distant populations. The mosquito distribution was more aggregated during the dry season, when few houses had densities 7- to 24-fold higher than expected. The high-density houses during the dry season differed from those of the wet season. Most high-density houses during the dry season changed between years, yet their vicinity was rather stable. Scan statistics confirmed the presence of one or two adjacent hotspots in the dry season, usually found on one edge of the village. These hotspots shifted between the early and late dry season. During the wet season, the hotspots were relatively stable near the main larval site. The locations of the hotspots in the wet season and early and late dry season were similar between years. Season-specific, stable, and focal hotspots are inconsistent with the predictions based on the arrival of migrants from distant localities during the dry season, but are consistent with the predictions based on local shelters used by aestivating mosquitoes. Targeting hotspots in Sahelian villages for vector control may not be effective because the degree of aggregation is moderate, the hotspots are not easily predicted, and they are not the sources of the population. However, targeting the dry-season shelters may be highly cost-effective, once they can be identified and predicted.

Reduced survival and future reproduction due to of current reproduction is a trade-off known as the cost of reproduction. Surprisingly, only a few studies have assessed the cost of reproduction in arthropod disease vectors, despite its effect on longevity, and thus on vectorial capacity. We evaluated the cost of reproduction on survival of Anopheles gambiae Giles by comparing mosquitoes that were denied exposure to the other sex, hereafter named virgins, and those that were allowed exposure to the other sex and mating, hereafter named mated. Merely 6 d of exposure to females with mating activity reduced male survival from a median of 17 d in virgins to 15 d in mated, indicating that male mating cost is substantial. The increase in mortality of mated males began several days after the exposure to females ended, indicating that mating is not associated with immediate mortality risk. Notably, body size was negatively correlated with male mortality in mated males, but not in virgins. The rate of insemination declined after 4 d of exposure to females, indicating that male mating capacity is limited and further supporting the hypothesis that mating is costly for males. Consistent with previous studies, female survival on sugar alone (median=16 d) was shorter than on blood and sugar (median=19 d), regardless if she was mated or virgin. Overall, survival of mated females was lower than that of virgins on a diet of blood and sugar, but no difference was found on a diet of sugar only. However, the cost of reproduction in females remains ambiguous because the difference in survival between virgin and mated females was driven by the difference between virgin (median=19 d) and uninseminated females exposed to males (median=17 d), rather than between virgin and inseminated females (median=19 d). Accordingly, sperm and seminal fluid, egg development, and oviposition have negligible cost in terms of female survival. Only exposure to males without insemination decreased female survival. Nonetheless, if exposure to males under natural conditions is also associated with reduced survival, it might explain why females remain monogamous.

Mosquito swarms are poorly understood mating aggregations. In the malaria vector Anopheles gambiae Giles, they are known to depend on environmental conditions, such as the presence of a marker on the ground, and they may be highly relevant to reproductive isolation. We present quantitative measurements of individual An. gambiae positions within swarms from Donéguébougou, Mali, estimated by stereoscopic video image analysis. Results indicate that swarms in this species are approximately spherical, with an unexpectedly high density of individuals close to the swarm centroid. This high density may be the result of individual males maximizing their probability of encountering a female or a product of mosquito orientation through cues within the swarm. Our analysis also suggests a difference in swarm organization between putative incipient species of An. gambiae with increasing numbers of males. This may be related to a difference in marker use between these groups, supporting the hypothesis that swarming behavior is a mechanism of mate recognition and ultimately reproductive isolation.

Mating in Anopheles gambiae has been observed only in outdoor swarms. Here we evaluate whether mating also occurs indoors. Mark-release-recapture of virgin males and females in natural houses showed that mating occurred over a single day even when mosquitoes can leave the house through exit traps and without adaptation to laboratory conditions. In these experiments, insemination rate in the M molecular form of An. gambiae (and An. arabiensis) was higher than that of the S form (15 versus 6%). Under these conditions, smaller females of the M form mated more frequently than larger females of that form. Sampling mosquitoes throughout the day showed that both sexes enter houses around sunrise and leave around sunset, staying indoors together from dawn to dusk. In an area dominated by the M form, the daily rate of insemination in samples from exit traps was approximately 5% higher than in those from entry traps, implying that mating occurred indoors. Importantly, frequency of cross mating between the molecular forms was as high as that between members of the same form, indicating that, indoors, assortative mating breaks down. Altogether, these results suggest that indoor mating is an alternative mating strategy of the M molecular form of An. gambiae. Because naturally occurring mating couples have not yet been observed indoors, this conclusion awaits validation.

Multiple families representing all possible combinations of crosses between the two molecular forms of Anopheles gambiae sensu stricto Giles and their hybrids were set up using forced mating between offspring of wild-collected females. The results showed that the reproductive output of hybrids and their backcrosses was similar to that of the pure forms as measured by egg batch size, hatching rate, and larval development success. No sex ratio distortion was found among the offspring. We concluded that postmating developmental barriers do not contribute to the isolation between the molecular forms.

Knowledge of ecological differences between the molecular forms of Anopheles gambiae Giles (Diptera: Culicidae) might lead to understanding of their unique contribution to disease transmission, to better vector control, and to identification of the forces that have separated them. We compared female fecundity measured as egg batch size in relation to body size between the molecular forms in Mali and contrasted them with their sibling species, Anopheles arabiensis Patton. To determine whether eggs of different egg batches are of similar "quality," we compared the total protein content of first-stage larvae (L1s), collected < 2 h after hatching in deionized water. Egg batch size significantly varied between An. gambiae and An. arabiensis and between the molecular forms of An. gambiae (mean batch size was 186.3, 182.5, and 162.0 eggs in An. arabiensis and the M and the S molecular form of An. gambiae, respectively). After accommodating female body size, however, the difference in batch size was not significant. In the S molecular form, egg protein content was not correlated with egg batch size (r = -0.08, P > 0.7) nor with female body size (r = -0.18, P > 0.4), suggesting that females with more resources invest in more eggs rather than in higher quality eggs. The mean total protein in eggs of the M form (0.407 microg per L1) was 6% higher than that of the S form (0.384 microg per L1), indicating that the M form invests a greater portion of her resources into current (rather than future) reproduction. A greater investment per offspring coupled with larger egg batch size may reflect an adaptation of the M form to low productivity larval sites as independent evidence suggests.

The M and S molecular forms of Anopheles gambiae sensu stricto Giles are thought to be reproductively isolated through premating barriers. However, the exact mechanisms of recognition of conspecific partners are unknown. Because mating in An. gambiae occurs in swarms, one might expect swarming behavior between the M and S forms to be different and that this probably reduces the risk of contact between males and females of the different forms in areas where they are sympatric. We report the occurrence of four mixed swarms, containing males of M and S forms, out of a total of 26 swarms sampled in Soumousso, a typical savannah village of Burkina Faso, West Africa. However, the frequency of mixed swarms was lower than that expected by chance. This observation suggests partial segregation between the swarms of the molecular forms, which may contribute to their isolation. Because the frequency of mixed swarms seems too high to explain the low frequency of cross-mating and hybrids, we suggest that mate recognition in a swarm is more important than swarm segregation.

We compared the development of the molecular forms of Anopheles gambiae s.s. in different larval habitats. First stage larvae (L1s) of wild-caught females were placed into cages in natural habitats of the M form (rice fields) or the S form (puddles/ quarries). Each cage was covered with cloth, allowing exchange of water, solutes, and small particles, including microorganisms, and was seeded with 100 L1s of a single form (M or S) or by a mixture of 50:50 of M and S forms. Emergence success of both forms in puddles and quarries was three-fold higher than in the rice fields. The emergence rate of the S form was higher than that of the M form in both habitats, but the form x habitat interaction was not significant. In temporary larval sites such as puddles, emergence success of the M form was lower in mixed cages than in single form cages, whereas the reverse was true for the S form, suggesting competition between the forms. The median developmental time was not significantly different between forms. Although these findings demonstrate differences between forms, they do not suggest that their spatial segregation is determined by differences in their exploitation of the physical and chemical conditions in these environments. These results should be regarded with caution because small numbers of first stage larvae could pass through the cloth of the cages.

Genetic relatedness (kinship) among Anopheles gambiae Giles female mosquitoes was assessed using microsatellite loci in five locations across Africa and in nine samples taken between 1994 and 1999 in western Kenya. We assessed variation among samples in kinship as well as the effect of distance on kinship. Relatedness within populations was low, and differences among samples taken at various times from one locale and from different locales were minimal. Mosquitoes collected from the same compound were slightly more closely related than those collected from different compounds. Our results suggest that newly emerged female siblings move relatively short distances into a few nearby compounds for blood feeding, but that they lay eggs in a more distant location. Kinship decreased nonlinearly with increasing distance. The strongest relationship between kinship and distance was observed for mosquitoes collected 0-3 km apart (-0.014/km, P < 0.001). The effect of distance decreased with increasing distance between mosquitoes; at 7 km or more, the kinship/distance slope approached zero and the intercept became negative, suggesting that beyond this range kinship does not decline with distance. This distance may thus represent the upper limit of the diameter of the basic reproductive unit. Nevertheless, the effect of distance on kinship is weak, reflecting extensive dispersal. Because females mate within days after emergence from larval habitats, where the likelihood of mating with a sibling is presumably highest, we propose a slight inbreeding effect.

The key components of ectoparasite vectorial capacity are poorly known, and no estimates of absolute population size for nest ectoparasites nor average longevity under natural conditions are available. A simple method to estimate these and other population parameters is suggested here, using body infestation data obtained by a specific host sampling design and the reproductive status of female ectoparasites. In contrast to mark-release-recapture techniques, this method requires minimal effort. Data on fleas, Synosternus cleopatrae (Rothschild), infesting gerbils, Gerbilus anderosoni allenbyi Thomas, were used as a preliminary test of this method. Thus, the average absolute density of female S. cleopatrae per host varied from seven in June to 24 in September; average female longevity varied from 17 to 5 d in that period. The quality of the parameter estimates was assessed by computer simulations to evaluate their variance and by comparison with the data available on fleas. The simulation indicated that estimates of absolute population size and proportion of the corporeal subpopulation were robust, but estimates of emergence rate, mortality rate, and average longevity were considerably less. The method's efficiency depends on the infestation rate and was considered suitable for medium to high counts of ectoparasites infesting solitary hosts.

Reproductive activity of Synosternus cleopatrae (Rothschild) infesting Gerbillus andersoni allenbyi Thomas was studied in a natural setting in Israel. Rodents were trapped and measured (weight and length), their sex was identified, and their reproductive status estimated. Their ectoparasites were removed, and fleas were dissected and their oocytes measured. Two indices of flea reproductive activity were analyzed: "reproductive status," which distinguished between gravid and nongravid females, and "reproductive intensity," which was estimated as the sum over the two largest oocytes of the products of oocyte length multiplied by oocyte width. Both indices showed that no reproduction took place between November and January, but reproduction was relatively stable during the rest of the year. Although flea reproductive activity differed significantly among individual hosts, only a small fraction of gerbils (10-15%) carried a significantly different proportion of reproductive fleas than their monthly sample proportion (based on all fleas regardless of hosts). All these hosts carried a lower proportion of reproductive fleas than their monthly sample proportion. The host's sex, but not reproductive status or age, had a significant effect on flea reproduction, expressed as a higher reproductive activity on male gerbils. Infestation burden expressed as ectoparasite counts was included in the statistical analysis. Only lice, Polyplax gerbilli Ferris, but not S. cleopatrae, Stenoponia tripectinata (Tiraboschi), and a total of five mesostigmatid Acari had a significantly negative association with S. cleopatrae reproductive activity. These relationships between S. cleopatrae reproductive activity and the host infestation burden do not support the hypothesis of modulation of S. cleopatrae reproduction by the infestation burden. However, differences in the reproductive activity of ectoparasites between their hosts may play a major role to generate the parasite clumped distribution. Thus, gerbil males probably carry more fleas than gerbil females because of the higher reproductive activity of S. cleopatrae on gerbil males.